Adaptive behavior is facilitated by an attentional system that can proactively modify the state of perceptual systems. While the majority of research on the underlying mechanisms has focused on spatial orienting, the brain is also anticipatory in time (Nobre and van Ede, 2018). In temporal anticipation, the brain exploits various temporal regularities in the environment to predict the timing of upcoming sensory events. These temporal expectations can guide temporal orienting, the adjustment of attention in time to modulate perceptual sensitivity, for example, increasing it at expected compared to unexpected times (Correa et al., 2005; Rohenkohl et al., 2012; Davranche et al., 2011; Denison et al., 2017; Samaha et al., 2015; Fernández et al., 2019) (also referred to in the literature as ‘temporal attention’). This form of anticipation has been often associated with left inferior parietal and ventral premotor cortices (Davranche et al., 2011; Coull and Nobre, 1998; Bolger et al., 2014), a network that overlaps with frontal-parietal components of the cortico-thalamic network traditionally implicated in attentional orienting in space (Posner and Petersen, 1990; Corbetta and Shulman, 2002; Fiebelkorn and Kastner, 2020).

Recently, we reported that individuals with cerebellar degeneration (CD) fail to exhibit reaction time benefits from temporal cues on a simple detection task (Breska and Ivry, 2018; Breska and Ivry, 2020). These findings implicate the cerebellum in temporal preparation, but do not enable to determine whether it has a role in attentional modulation of perceptual systems, in motor preparation, or in both. This is because in speeded detection tasks, reaction time benefits from temporal cues could result from adjustment of perceptual sensitivity and/or of motor preparation (Rohenkohl et al., 2012; Sanabria et al., 2011; Morillon et al., 2016; van Ede et al., 2020). Given the well-documented role of the cerebellum in precisely timed movement (Ivry and Keele, 1989; Perrett et al., 1993), the impairments we observed in the CD group may merely reflect a novel manifestation of the cerebellar role in the temporal control of motor preparation (Shalev et al., 2019). However, in addition to its role in motor timing, the cerebellum could also be essential to attentional orienting in time to proactively modulate perception. Determining the functional domain of the cerebellum in temporal anticipation is critical to both models of attention and of cerebellar function.

To address this question, we compared the ability of individuals with CD and healthy controls to use temporal cues to benefit performance in a challenging non-speeded perceptual discrimination task, in which the benefits of attention are assumed to reflect modulation of perceptual sensitivity (Correa et al., 2005; Davranche et al., 2011; Raymond et al., 1992). Participants judged the orientation of a briefly presented visual target whose contrast was set to be near-threshold, making the task perceptually demanding. In each trial, a temporal cue indicated that the interval between the target and a preceding warning signal (WS) would either be 600 or 1000 ms (Figure 1A). Targets mostly appeared at the cued time (valid trials) and rarely at the uncued time (invalid trials). Critically, participants were only queried for a response after a substantial random delay following target offset. In tandem with our instructions that only emphasized accuracy, the inclusion of the delay eliminated the need for motor preparation. With these manipulations, we assume that the expected performance advantage on valid compared to invalid trials (validity effect; Correa et al., 2005; Coull and Nobre, 1998; Miniussi et al., 1999; Rohenkohl et al., 2014; Breska and Deouell, 2017) will reflect temporally focused enhancement of perceptual sensitivity. If the cerebellum has a causal role in attention orienting in time at a non-motor level, the validity effect should be reduced in CD patients relative to controls.

Figure 1

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In addition to manipulating the validity of the temporal cue, we also varied the manner in which this information was presented. In our previous work (Breska and Ivry, 2018; Breska and Ivry, 2020), we found that the cerebellar involvement was limited to when temporal anticipation required encoding and recalling an isolated interval, but not when temporal anticipation could be based on synchronization with a stream of rhythmic sensory events. We employ a similar manipulation in the present study, comparing perceptual benefits from interval and rhythmic temporal cues (Figure 1B).

Perceptual sensitivity was quantified using d’, a measure derived from signal detection theory (Green and Swets, 1966). Temporal orienting should manifest as higher d’ for valid compared to invalid trials. A four-way omnibus analysis of variance (ANOVA) revealed a significant validity effect across groups, tasks, and target intervals (main effect of cue validity: F(1,23)=13.42, p=0.001, ${\displaystyle {\eta }_{\mathrm{p}}^2}$=0.37). Prior work has shown that in two-interval designs as implemented here, the validity effect is less robust or even absent for late targets (van Ede et al., 2020; Miniussi et al., 1999; Correa et al., 2006). Thus, we employed an a priori analysis plan that focused on trials in which the target appeared at the short interval (600 ms, ‘early targets’) to increase sensitivity for detecting attenuation of the validity effect (see Materials and methods). As expected, the validity effect was larger when the target appeared at the short interval (cue validity × interval interaction: F(1,23)=3.71, one-tailed p=0.033, ${\eta }_p^2$=0.14). Indeed, neither group showed a significant validity effect in either task for the late onset targets (all uncorrected p’s > 0.09, Figure 2—figure supplement 1).

Figure 2 presents d’ values for short interval targets. A three-way ANOVA revealed a significant validity effect across tasks and groups (F(1,23)=14.75, p=0.001, ${\eta }_p^2$=0.39). Across tasks, the validity effect was larger in the control compared to the CD group (cue validity × group interaction, F(1,23)=4.83, p=0.038, ${\eta }_p^2$=0.17), but this effect was qualified by a significant cue validity × group × task interaction (F(1,23)=4.76, p=0.04, ${\eta }_p^2$=0.17). We used a series of planned contrasts to evaluate the validity effect within each task. For the interval task (Figure 2A), the validity effect was significantly smaller in the CD group relative to the control group (two-way ANOVA, cue validity × group interaction: F(1,23)=7.11, p=0.014, ${\eta }_p^2$=0.24). The control group showed a higher mean d’ for valid compared to invalid trials (t(11)=3.08, p=0.011, Cohen’s d=0.89), while d’ values in the CD group were not significantly different, and even numerically higher on invalid trials (t(12) = −0.43, p=0.67, BF₁₀=0.21, moderate evidence against a directional hypothesis of a validity effect). Thus, the CD group failed to show an enhancement in perceptual sensitivity from an interval-based temporal cue.

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A different pattern was observed on the rhythm task (Figure 2B). Here, the magnitude of validity effect did not differ between the two groups (F(1,23)=0.053, p=0.82, ${\eta }_p^2$<0.01, BF₁₀=0.32). The control group again showed a validity effect (t(11)=2.67, p=0.022, Cohen’s d=0.77). However, unlike in the interval task, the CD group also showed a validity effect (t(12)=3.45, p=0.005, Cohen’s d=0.95). Thus, the ability to increase perceptual sensitivity at a specific time point based on a rhythm cue was preserved in the CD group. Furthermore, the presence of the three-way interaction implies that the attenuation of the validity effect in the CD group relative to controls was significantly larger in the interval task than in the rhythm task.

Our results provide compelling evidence that the integrity of the human cerebellum is necessary for proactive modulation of perceptual processing based on temporal expectations. The ability to use a temporal cue to enhance perceptual sensitivity at specific times was abolished in individuals with CD when prediction was based on an interval cue. In contrast, the CD group showed a comparable benefit to controls when temporal orienting was based on a rhythmic cue. Critically, the use of a non-speeded, challenging perceptual discrimination task argues strongly against the idea that the impairment is related to motor preparation processes. Rather, the results point to an essential context-specific role of the cerebellum in the temporal orienting of visual attention, adding to the substantial literature highlighting cerebellar involvement in a broad range of cognitive functions beyond the motor domain (Sokolov et al., 2017).

These findings point to the need for an expanded picture of the neuroanatomical network involved in attentional control of perceptual sensitivity. Attention research has traditionally emphasized cortico-thalamic networks, and in particular, dorsal and ventral fronto-parietal networks associated with top-down and stimulus-driven attention control, respectively (Posner and Petersen, 1990; Corbetta and Shulman, 2002; Fiebelkorn and Kastner, 2020). For attention orienting in the time domain, human imaging studies consistently reveal activations of the left inferior parietal cortex and the ventral premotor cortex (Davranche et al., 2011; Coull and Nobre, 1998; Bolger et al., 2014). Similarly, neural signatures of temporal anticipation identified in human electrophysiology studies have been localized to cortico-thalamic circuits (Gómez et al., 2001; Praamstra, 2006). In general, the cerebellum has not featured in this work, perhaps due to the difficulty in neuroimaging studies to separate the effects of temporal prediction on neural activations from the effects of other task parameters, or the difficulty of electrophysiology studies to measure cerebellar sources. One exception here is a study by O'Reilly et al., 2008 who observed greater cerebellar activation for detecting violations of spatio-temporal predictions compared to spatial predictions, pointing to cerebellar involvement in temporal prediction (O'Reilly et al., 2008). The current findings demonstrate that the cerebellum has a causal role in temporal anticipation and establish its role in using temporal information to modulate attention for the perceptual analysis of visual features.

Our neuropsychological approach provides a more direct method to evaluate the contribution of the cerebellum to attention. Previous work, focusing on the spatial domain, has proven inconclusive. Some studies have found that individuals with focal cerebellar lesions show reduced benefits from cues indicating the spatial location of a forthcoming stimulus (Townsend et al., 1999; Allen et al., 1997). However, it has been proposed that these impairments may be motoric in nature, with the tasks confounding attentional demands with demands on eye movements and/or response preparation (Ravizza and Ivry, 2001; Haarmeier and Thier, 2007). These concerns do not apply to the current study given that the spatial aspects of the task were fixed and the motor requirements were minimal, delayed until well after stimulus offset. We note that the current results do not address the question of whether the cerebellum, in addition to its role in temporal orienting of attention, is also involved in other spheres of attention.

The dissociation between the impairment in the interval task and the preserved performance in the rhythm task in the CD group has two important implications. First, a longstanding debate in the timing literature concerns whether temporal anticipation in a rhythmic context is mediated by rhythm-specific mechanisms (e.g., entrainment) or by the repeated operation of an interval-based mechanism (Breska and Deouell, 2017; Haegens and Zion Golumbic, 2018; Drake and Botte, 1993). Our results are at odds with the latter hypothesis given that the CD group failed to benefit from the interval cues yet showed normal benefits from the rhythm cues. Beat-based timing may rely on cortico-striatal circuits even in the absence of movement (Cannon and Patel, 2021), consistent with previous findings of impaired rhythm-based temporal prediction in Parkinson’s disease (Breska and Ivry, 2018).

Second, the selective contribution of the cerebellum in interval-based but not rhythm-based contexts has been observed in multiple timing domains, including duration judgments, timed movement, and timed motor preparation (Breska and Ivry, 2018; Breska and Ivry, 2020; Grube et al., 2010; Spencer et al., 2003). Our findings extend this functional specificity to the attentional domain, pointing to a generalized role for the cerebellum in interval timing, at in the sub-second range. Notably, while inferences from single dissociations such as that observed in the present study can be limited by concerns about differences in task difficulty, this concern is alleviated by the comparable benefits observed in healthy controls from interval- and rhythm-based cues (Breska and Ivry, 2018; Breska and Ivry, 2020; Breska and Deouell, 2017) (also observed in current dataset: cue validity × task interaction within the control group, F(1,11)=0.86, p=0.37, ${\eta }_p^2$=0.07).

Computationally, how might the cerebellum contribute to the attentional control of perceptual sensitivity in time? Given the cerebellar involvement in interval-based timing across timing domains, an intuitive hypothesis is that the cerebellum is necessary for the temporal processing of isolated intervals (Ivry and Keele, 1989; Ivry and Schlerf, 2008). This could reflect a central role in one or both of two putative functional subcomponents in interval timing models (Addyman et al., 2016; Gibbon et al., 1984), the formation of a temporal representation or retrieval processes related to interval memory. By this view, predictive processing in non-cerebellar circuits (e.g., prefrontal cortex) relies on cerebellar interval timing capacities to parameterize the temporal dimension of the prediction. In rhythm-based orienting, an interval-based mechanism would not be required as the temporal parameters are contained within ongoing neural dynamics. However, a broader hypothesis is that the interval-based prediction itself is formed within the cerebellum, part of the cerebellar role in prediction in the motor domain and beyond (Sokolov et al., 2017; Wolpert et al., 1998; Miall et al., 1993). By this view, these cerebellar temporal predictions guide proactive modulation in non-cerebellar circuits according to task goals (e.g., to prepare perceptual or motor systems). In rhythm-based orienting, dedicated prediction mechanisms are not necessary due to the self-sustaining limit cycle properties of the putatively entrained oscillatory dynamics. Future work should aim to explore the separability of timing and prediction, identifying the cerebellar computations that provide the essential information for temporal orienting.

De-identified source data files for all figures and analyses in the article have been provided. Additional demographic information was not uploaded as it was not used in any analysis reported in the text, and can be provided upon request in personal communication with the corresponding author, without additional restrictions.

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Author details

1. Assaf Breska

   Department of Psychology and Helen Wills Neuroscience Institute, University of California, Berkeley, Berkeley, United States

   Contribution

   Conceptualization, Data curation, Software, Formal analysis, Validation, Investigation, Visualization, Methodology, Writing - original draft, Project administration, Writing - review and editing

   For correspondence

   assaf.breska@berkeley.edu

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0002-6233-073X

2. Richard B Ivry

   Department of Psychology and Helen Wills Neuroscience Institute, University of California, Berkeley, Berkeley, United States

   Contribution

   Conceptualization, Resources, Supervision, Funding acquisition, Project administration, Writing - review and editing

   Competing interests

   Senior editor, eLife

   "This ORCID iD identifies the author of this article:" 0000-0003-4728-5130

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