The role of the cerebellum in adaptation: ALE meta‐analyses on sensory feedback error

Abstract It is widely accepted that unexpected sensory consequences of self‐action engage the cerebellum. However, we currently lack consensus on where in the cerebellum, we find fine‐grained differentiation to unexpected sensory feedback. This may result from methodological diversity in task‐based human neuroimaging studies that experimentally alter the quality of self‐generated sensory feedback. We gathered existing studies that manipulated sensory feedback using a variety of methodological approaches and performed activation likelihood estimation (ALE) meta‐analyses. Only half of these studies reported cerebellar activation with considerable variation in spatial location. Consequently, ALE analyses did not reveal significantly increased likelihood of activation in the cerebellum despite the broad scientific consensus of the cerebellum's involvement. In light of the high degree of methodological variability in published studies, we tested for statistical dependence between methodological factors that varied across the published studies. Experiments that elicited an adaptive response to continuously altered sensory feedback more frequently reported activation in the cerebellum than those experiments that did not induce adaptation. These findings may explain the surprisingly low rate of significant cerebellar activation across brain imaging studies investigating unexpected sensory feedback. Furthermore, limitations of functional magnetic resonance imaging to probe the cerebellum could play a role as climbing fiber activity associated with feedback error processing may not be captured by it. We provide methodological recommendations that may guide future studies.

studies reported cerebellar activation with considerable variation in spatial location.
Consequently, ALE analyses did not reveal significantly increased likelihood of activation in the cerebellum despite the broad scientific consensus of the cerebellum's involvement. In light of the high degree of methodological variability in published studies, we tested for statistical dependence between methodological factors that varied across the published studies. Experiments that elicited an adaptive response to continuously altered sensory feedback more frequently reported activation in the cerebellum than those experiments that did not induce adaptation. These findings may explain the surprisingly low rate of significant cerebellar activation across brain imaging studies investigating unexpected sensory feedback. Furthermore, limitations of functional magnetic resonance imaging to probe the cerebellum could play a role as climbing fiber activity associated with feedback error processing may not be captured by it. We provide methodological recommendations that may guide future studies.

K E Y W O R D S
cerebellum, fMRI, forward model, meta-analysis, prediction, sensory feedback

| INTRODUCTION
To successfully act within a dynamic environment, we continuously monitor sensory feedback associated with our own movements to ensure our actions have the desired outcomes. Even the simplest movements require complex coordination between multiple effectors.
The continuous monitoring of sensory feedback helps to refine motor plans and adjust them to contextual and environmental changes. The forward model is a computational process that compares expected to actual sensory consequences of an action (see Figure 1; Jordan & Rumelhart, 1992;Miall & Wolpert, 1996;Wolpert, 1997). This comparison is essential to motor control and relies partly on the cerebellum (Blakemore, Frith, & Wolpert, 2001;Ishikawa, Tomatsu, Izawa, & Kakei, 2016;Ito, 1984a;Kawato, Furukawa, & Suzuki, 1987;Miall, Weir, Wolpert, & Stein, 1993;Wolpert, Miall, & Kawato, 1998). However, there is still no strong consensus on where in the cerebellum unexpected sensory feedback is processed. To this end, the current meta-analysis systematically explores patterns of cerebellar activation in neuroimaging studies of sensory feedback manipulations. Such manipulations create an artificial mismatch between intended and perceived sensory consequences of an action, constituting methods commonly used to probe forward models in brain imaging studies.
A fundamental component of the forward model is the efference copy: when the motor cortex sends a command to the peripheral nervous system for the execution of motor behavior, a copy of the command is fed forward to provide an estimate of the sensorimotor feedback predicted from the movement (von Holst & Mittelstaedt, 1950). The predicted feedback is compared to the actual feedback from the proprioceptive (Miall & Wolpert, 1996;Wolpert et al., 1998), visual (Leube et al., 2003), auditory (Hashimoto & Sakai, 2003), and tactile (Blakemore, Wolpert, & Frith, 1998) sensory periphery. Any discrepancy between predicted and actual feedback constitutes an output known as a corollary discharge (Feinberg, 1978). The resultant error signal is processed by the cerebellum, ultimately resulting in an output to cortical areas to adaptively fine-tune behavior. As both the state of the organism and the environment are dynamic, forward models are employed continuously to reduce any discrepancy between predicted and actual feedback through constant monitoring and adjustment of behavior (Desmurget & Grafton, 2000;Jordan & Rumelhart, 1992).
The cerebellar cortex comprises overlapping functional zones that process input from specific sensory modalities (Witter & De Zeeuw, 2015). Specific areas respond to auditory or visual stimuli (O'Reilly, Beckmann, Tomassini, Ramnani, & Johansen-Berg, 2009;Petacchi, Laird, Fox, & Bower, 2005;Sang et al., 2012). Likewise, portions of the cerebellum segment into somatomotor topographies associated with the control of different body parts (Buckner, Krienen, Castellanos, Diaz, & Yeo, 2011;Mottolese et al., 2012). These divisions are coupled to associated subdivisions of motor cortex responsible for controlling the same body parts. The motor areas contribute cortical input to communication loops between the cortex and cerebellum employed in ongoing motor control. Therefore, the cerebellar cortex receives two principle types of afferents: mossy fibers primarily via the pons which relay information such as the efference copy from corresponding cortical regions (Raymond, Lisberger, & Mauk, 1996), while medullary nuclei relay bottom-up sensory feedback-related signals and induce changes in the influence of cortical top-down signals to the cerebellar cortex (Ito, Sakurai, & Tongroach, 1982). The inferior olive monitors the discrepancy between predicted and actual sensory input and relays an error signal to the cerebellum via climbing fibers (Ito, 1984b;Kawato & Gomi, 1992). The cerebellum also receives signals of unpredicted auditory feedback from the dorsal cochlear nuclei of the medulla . In turn, the cerebellum continuously signals the discrepancy back to the cerebral cortex to induce adaptation of motor behavior until the expected feedback matches the actual sensory feedback.
A common experimental approach to study neural substrates that implement processes related to the forward model is to manipulate the sensorimotor feedback of self-generated movements. In these F I G U R E 1 Components of a forward model. A diagram which outlines five major components of a Forward Model. The "Motor Plan" incorporates the components (1) "Efference Copy" and (2) "Motor Command." The implantation of the Motor Command leads to (3) "Observed Sensory Feedback." The efference copy is an expectation of sensory consequences of the enactment of the motor plan, providing (4) "Predicted Sensory Feedback." The Observed Sensory Feedback and the Predicted Sensory Feedback are compared. If they do not match, an (5) "Error Signal" indicating violation of the expected consequences is returned for updating of the motor plan. The "Observed Sensory Feedback" can also notify the "Predicted Sensory Feedback" with contextual information of body or environment in order to make temporary changes to the prediction rather than updating the motor plan. This is denoted as "Dynamic assessment of state" experiments, participants typically produce articulatory or manual movements while feedback is altered. For example, studies have investigated how the brain responds to unpredicted feedback by manipulating the acoustic properties of one's own voice (Tourville, Reilly, & Guenther, 2008;Zheng et al., 2013), by introducing illusory visual displacement of the hand or a mechanically controlled avatar (David et al., 2007;Diedrichsen, Hashambhoy, Rane, & Shadmehr, 2005;Schnell et al., 2007), or by applying an unpredicted external physical force (Diedrichsen et al., 2005;Golfinopoulos et al., 2011). Therefore, there can be much variability in how this mechanism is studied in neuroimaging research in terms of form of motor production, feedback manipulation, and sensory modality of feedback. It was our intention to evaluate over the body of literature eliciting activity in response to various manipulations of self-generated sensory feedback any common areas reported in the brain, with specific interest in finding consensus on the regions of the cerebellum involved. We conducted three activation likelihood estimation (ALE) meta-analyses of functional neuroimaging studies to identify patterns of neural activation that are reliably affected by these manipulations. A primary analysis was expected to yield a modality independent but anatomically precise global impression of cerebellar contributions to processes related to the forward model.
Two secondary ALE analyses were conducted to differentiate this impression in terms of potential modality-specific components, as well as a contrast analysis between auditory and visual feedback results.
This distinction is made as although higher processing cortical areas may be responsive irrespective of the sensory modality, there is cerebellar and cortical distinction between areas responsive to auditory and visuomotor feedback manipulations. In doing so, we may shed light on a consensus of where the cerebellum is involved in processing feedback error, and specifically if the cerebellum is more reliably probed in areas segregated for auditory or visual sensory input. Additionally, due to the high diversity in methods across this body of literature, we aimed to further investigate the dependency of cerebellar activity reported on the factors which most commonly varied across the experiments selected for our meta-analyses.

| MATERIALS AND METHODS
We generally followed recent best practice guidelines for the conducting of neuroimaging meta-analyses (Müller et al., 2017). These guidelines have been put forward to improve the transparency and replicability of meta-analyses. We accordingly report information advised such as research question, inclusion and exclusion criteria, detailed information for all experiments, and a step-by-step flowchart (see Figure 2).

| Study selection
The PubMed (www.ncbi.nlm.nih.gov/pubmed) database was searched for human neuroimaging studies using combinations of relevant keywords (e.g., functional magnetic resonance imaging [fMRI], positron emission tomography [PET], sensorimotor learning, adaptation, and shifted-, delayed-, altered-, masked-, incongruent-, and distorted feedback). We further cross-referenced the articles produced from the search term to corroborate that no relevant articles were overlooked.
Studies were selected if they reported activation contrasts of unpredicted (manipulated) compared to predicted (non-manipulated) self-generated feedback, data acquisition covered the whole brain encompassing the cerebellum, and included tables listing peak activations in standard stereotaxic space. Only data from healthy adult participants were selected. Results reported in Talairach space (Talairarch & Tournoux, 1988) were converted to Montreal Neurological Institute (Holmes et al., 1998) using the GingerALE (2.3.6) icbm2tal conversion F I G U R E 2 Meta-analysis flowchart. A flowchart diagram recommended in the best practice guidelines for the conducting of neuroimaging meta-analyses (Müller et al., 2017)  Visual studies were subject to temporal and spatial shifts, and random mismatch between action and feedback, auditory studies were subject to temporal and acoustic shifts, noise masking, and random mismatch between action and feedback. Imaging method: acquisitions from MRI or PET imaging equipment. Number of foci: amount of contributing foci of significant activity from each study. Abbreviation: MRI, magnetic resonance imaging.  Significance thresholds were set at p < .05.

| Tests of independence
3 | RESULTS

| Manipulated feedback: ALE analyses
The primary ALE identified five clusters (Table 2a and Table S1).

| Relationship between experimental design and cerebellar activation
To determine whether certain methodological differences were linked with being more likely to activate the cerebellum we conducted chi-

| DISCUSSION
We conducted three meta-analyses of neuroimaging studies that altered sensory feedback during ongoing movements with the aim of localizing the cerebellar contributions to processes related to the forward model. In doing so, we attempted to reconcile an apparent lack of consensus of the role of the cerebellum in experiments of unexpected changes to sensory consequences of our own action. Contrary to our expectations and to broad scientific consensus suggesting that the cerebellum is involved in this process, we did not observe a convergence of activation foci in the cerebellum, although the analyses successfully identified the expected network of cortical areas.
We therefore systematically assessed methodological factors that could potentially increase the likelihood of specific experimental designs to activate the cerebellum. Our findings confirm that studies that require adaptation of behavior in response to sensory feedback manipulations most reliably evoke cerebellar activation.

| Cerebral cortex
The primary ALE analysis of the manipulated sensory feedback dataset indicated clusters in the SMA, preCG, IFG, STG, and TPJ. Two of the clusters, the SMA and preCG, were centered in the right secondary motor cortex, regions associated with the production of an efference copy for assessing sensory consequences of movement (Christensen et al., 2006;Ellaway, Prochazka, Chan, & Gauthier, 2004;Haggard & Whitford, 2004). The cluster centered at the IFG and extending into the prefrontal frontal regions incorporates areas which lend themselves to a broader self-awareness network and are thought to reflect agency over action (David, Newen, & Vogeley, 2008;Fink et al., 1999;Jardri et al., 2007;Leube et al., 2003;Nahab et al., 2010).
The right IFG plays a role in detecting cues which are relevant to inhibiting motor activity (Corbetta & Shulman, 2002;Hampshire, Chamberlain, Monti, Duncan, & Owen, 2010) and in subsequent reorienting and updating of action plans (Levy & Wagner, 2011). The cluster centered in the TPJ extends into much of the IPL, which is associated with monitoring motor outflow (Desmurget & Sirigu, 2009;Sirigu et al., 1996), as well as with awareness of consistency of intended and actual motor consequences (Farrer et al., 2008), where activity is higher when another agent is active (Decety, Chaminade, Grezes, & Meltzoff, 2002;Farrer & Frith, 2002;Ruby & Decety, 2001 studies, suggesting a multisensory network. This asserts that some cortical areas process error in both auditory and visual self-generated feedback. The cluster centered in the left STG however was an exception, with only foci contributed from auditory studies. Analyses of the auditory subset ALE revealed bilateral STG activation. These auditory cortical processing areas have shown a stronger response to auditory stimuli initiated by others than by oneself (Christoffels et al., 2007;Curio et al., 2000; Heinks-Maldonado, F I G U R E 3 Legend on next page.  Curio, 1999). Increased activation of auditory cortex has also been correlated with the degree of delay in auditory feedback in speech production (Hashimoto & Sakai, 2003) and in response to sound masking compared to expected auditory feedback (Christoffels et al., 2007). Likewise, there was a large medial cluster in the auditory subset in the right SMA. The SMA has been suggested to play a role in auditory sensorimotor associations, for example, in using auditory information to elicit automatic motor responses (Lima, Krishnan, & Scott, 2016) and auditory conditioning in a motor task (Kurata, Tsuji, Naraki, Seino, & Abe, 2000).
Studies that manipulated visual feedback were more likely to activate the premotor cortex of the right preCG, and a cluster in the EBA extending into the IPL. The EBA is involved in the visual processing of perceiving the human body (Downing, Jiang, Shuman, & Kanwisher, 2001;Peelen & Downing, 2007), of goal-directed action of body parts (Astafiev, Stanley, Shulman, & Corbetta, 2004), and in processing incoherent human biological motion sequences (Downing, Peelen, Wiggett, & Tew, 2006). David et al. (2007) suggest that the EBA may be part of a larger network including posterior parietal cortex, premotor cortex, and the cerebellum, that is involved in correcting sensorimotor discrepancy. The IPL engages in the monitoring and comparison of one's own action and the visual feedback that it generates (Schnell et al., 2007), and in visuomotor incongruencies (Balslev et al., 2006), reported as well more broadly in the parietal cortex (Fink et al., 1999;Shimada, Hiraki, & Oda, 2005).

| Cerebellum
Despite a strong consensus suggesting that forward models generally rely on the cerebellum (Bastian, 2006;Ishikawa et al., 2016;Ito, 2005;Kawato & Gomi, 1992;Wolpert et al., 1998), cerebellar activations were reported in only half of the 38 experiments that formed the dataset for the current analyses. The majority of these activations were localized in lobules VI and VIII. Cerebellar lobule VI contains overlapping functional zones that are sensitive to auditory and visual stimulation while a functional zone of lobule VIII is associated with sensorimotor processing (O'Reilly et al., 2009;Sang et al., 2012). The cerebellum receives climbing fiber inputs from the sensory periphery and corticopontocerebellar mossy fiber inputs from cortical motor areas.
The majority of activations in the cerebellum reflecting fine-tuning motor control will be predominately in either of these two somatotopically organized lobules. However, in the ALE analyses, we do not see clustering at each of those regions. The reason for this may be due to the histological organization of the cerebellum. Its structure is completely uniform in its cortex, made up of repeating modules intermixed and overlapping with modules of separate function, having no integral borders (Ito, 1984b). This can lead to a lack of clear separation of focal activity for one specific function. For instance, electrical F I G U R E 3 (a) Sensory feedback error ALE. Illustrates results from a meta-analysis on studies which report data reporting areas of the brain that increase in activity when self-initiated sensory feedback is experimentally manipulated compared to regular conditions of expected feedback. Images: six slices at MNI space x axis 49, 3, −60, y axis 55, −24, and z axis 8. Units of measurement: ALE scores with a minimum value of 0.008 and maximum of 0.029. A threshold of likelihood calculated from a cluster-forming threshold of p < .001, with a cluster-level correction of 0 Illustrates results from a meta-analysis on studies which report data reporting areas of the brain that increase in activity when self-initiated visual feedback is experimentally manipulated compared to regular conditions of expected feedback. Images: four slices at MNI space where there may be multiple representations for movement of one body part, and if so where these representations may be sparsely distributed.
For these reasons although many studies will report cerebellar activation in response to the same contrast, attempting to find meaningful clusters of functional localization common across studies may be limited.

| Considerations for reliable probing of the forward model
Although all studies included in this meta-analysis contrasted experimental conditions of manipulated feedback with non-manipulated feedback, there was a considerable degree of diversity in methodology. Experiments varied in terms of feedback modality, the quality and quantity of the feedback manipulation, whether participants were able to adapt behavior, and whether similar experimental trials were blocked together or intermixed.

| Choice of feedback manipulation
There was considerable variability in the way that different studies implemented manipulations of sensory feedback from self-produced action. The most common feedback manipulations employed delays, a mismatch leading to an abrupt loss of control over feedback, noise masking, spectral shifting of auditory feedback or spatial shifting of visual feedback, and the physical application of an external force. The relative amount of cerebellar foci contributing to the study pool from different forms of manipulations differed accordingly. For example, shifted and feedback mismatch studies were twice as likely to elicit cerebellar activity than not (12/18 and 4/6), while temporal manipulations only elicited cerebellar activity in a quarter of the respective experiments (2/8).
High magnitude manipulations may cause feedback to be perceived as entirely outside a range of control of the actor, and thus no longer triggering an automatic compensation in motor production.
This stems from the theory that our sense of agency over sensory feedback from the environment is dependent on the magnitude of discrepancy between the predictable consequences of our own action and the unpredictable external influences of sensory input (David et al., 2008). For example, singers can successfully suppress the automatic compensation response when the pitch of their voice is shifted by a large amount, but not when a smaller shift is applied (Zarate & Zatorre, 2008). The duration of manipulated feedback also influences compensation responses. Pitch shifts with short durations prompted automatic adjustments while pitch shifts with longer durations were more easily ignored (Burnett, Freedland, Larson, & Hain, 1998;Hain et al., 2000;Zarate et al., 2010;Zarate & Zatorre, 2008). Some forms of feedback can disrupt movement altogether. By applying delayed auditory feedback (DAF), speech and musical performance are interrupted (Black, 1951;Fukawa, Yoshioka, Ozawa, & Yoshida, 1988;Havlicek, 1968;Howell & Powell, 1987;Lee, 1950;Mackay, 1968;Siegel, Schork, Pick, & Garber, 1982). The magnitude of delay is also an important consideration, with DAF of approximately 200 ms being the most disruptive (Fairbanks & Guttman, 1958;Hashimoto & Sakai, 2003;Stuart, Kalinowski, Rastatter, & Lynch, 2002). Further increase of delay may lead to similar disregard of feedback as irrelevant to the agency of the actor.

| Adaptation to changes in feedback
Unpredicted feedback informs not only adjustments to ongoing movements, but also updates predictions for future movements by means of adaptation. This response to changes in environmental feedback is a form of motor learning and differs qualitatively from motor sequence learning (Doyon, Penhune, & Ungerleider, 2003). This type of motor learning therefor must not be seen as planning new coordinated motor plans, and instead viewed specifically as reoptimization that seeks to minimize future costs to the motor system by forming more accurate predictions of existing motor plans (Izawa, Rane, Donchin, & Shadmehr, 2008). Across the studies in our analyses, cerebellar foci were most common in studies driving adaptation in response to physical perturbation of the mouth (Golfinopoulos et al., 2011) and arm (Diedrichsen et al., 2005), learning new associations between the spatial consequences of movement when visual feedback is shifted (Anguera et al., 2010;Brand et al., 2017;Diedrichsen et al., 2005;Grafton et al., 2008;Graydon et al., 2005;Inoue et al., 2000;Krakauer et al., 2004;Seidler et al., 2006;Zheng et al., 2013), and when vocal pitch was shifted during continuous speech (Tourville et al., 2008;Zheng et al., 2013). All of these manipulations evoke the fine-tuning of accurately predicting movement outcomes in response to changes in the environment (Ishikawa et al., 2016). Indeed, adaptation was found to be the only factor in our analyses that showed a significant dependency with the elicitation of cerebellar activation.
The cerebellum plays an important role in adapting future predictions in light of error. Cerebellar patients are able to react to changes to feedback (Morton & Bastian, 2006;Smith, Brandt, & Shadmehr, 2000), but are unable adapt by calibrating their predictions for subsequent behavior (Maschke, Gomez, Ebner, & Konczak, 2004;Morton & Bastian, 2006;Smith & Shadmehr, 2005). This suggests that adjustments to feedback, which inform subsequent fine-tuning, require cerebellar engagement. Monkeys with experimental lesions to areas of the cerebellum which receive mossy fibers from cortex such as the posterior lobe parafloculus and uvula are unable to adapt to changes in feedback (Baizer, Kralj-Hans, & Glickstein, 1999). Inactivation of deep cerebellar nuclei impairs adaptation to physical and visuomotor perturbation (Kerr, Miall & Stein, 1993). Cerebellar activity may change over time as the system moves from a state of adapting predictions that have failed, to executing predictions that have been adapted (Gilbert & Thach, 1977).
This has strong implications for the choice of design in experiments seeking to probe the cerebellum's involvement in the forward model.

| Neurovascular coupling in the cerebellum
The blood-oxygen-level-dependent (BOLD) response that is measured by fMRI may not be sensitive to some of the neural processes of sensory feedback error in the cerebellum. The BOLD response is correlated with local field potentials (LFP) rather than the spiking rate of neurons (Ekstrom, 2010) (Diedrichsen, Verstynen, Schlerf, & Wiestler, 2010). This is consistent with the view that the mossy fiber input system is more strongly associated with processes of motor learning in adapting to sensorimotor prediction errors (Giovannucci et al., 2017;Ito, 2000;Thach, 1998).

| Choice of experimental design
Our findings suggest that some experimental fMRI designs are more appropriately suited to elicit BOLD responses in the cerebellum.
Designs that prevent participants from habituating to altered feedback, and continually cause them to adapt their motor responses, may be most effective in eliciting a detectable BOLD response. McGuire et al. (1996) illustrate habituation in block designs as they observed increased activation of the cerebellum during the first half of their study, but not in the latter half. This is consistent with the broader finding that the cerebellum may be more strongly engaged in adjusting to altered feedback than applying adjustments that have already been computed (Andersson & Armstrong, 1987;Flament, Ellermann, Kim, U gurbil, & Ebner, 1996;Horn, Pong, & Gibson, 2004;Imamizu et al., 2000;Moberget, Gullesen, Andersson, Ivry, & Endestad, 2014). However, the arrangement instead of fast cycling between trials of different conditions in event-related designs may hinder adaptation responses if feedback is not consistent from trial to trial. Long events of consistent perturbation (e.g., Christoffels et al., 2007;Grafton et al., 2008;Limanowski et al., 2017) or similarly with short blocks (e.g., Inoue et al., 2000;McGuire et al., 1996;Seidler et al., 2006) may be best suited for probing processes related to the forward model associated BOLD response in the cerebellum.

| CONCLUSIONS
We performed three ALE meta-analyses and one contrast analysis of functional neuroimaging studies that manipulated predicted selfinitiated auditory, visual, and sensory feedback with the primary aim to identify cerebellar areas responsive to prediction error. No cerebellar clusters were produced as a result of these analyses. Contrary to common presumptions, we found that not all studies that used such approach show significant activation of the cerebellum, as well as variability in where in the cerebellum activations were reported. Our study suggests that this discrepancy stems from differential sensitivity and specific limitations of the experimental paradigms employed across MR neuroimaging altered sensory feedback experiments. These method-specific characteristics can restrict compatibility with other frameworks, which overwhelmingly support the involvement of the cerebellum in responding to errors in predicted feedback as part of the forward model. We therefore assessed methodological variations that may determine the success of brain imaging experiments in evoking cerebellar activation. The results indicate that experimental designs which most reliably evoked cerebellar activation employed continuous feedback manipulations relevant for adapting motor plans for future action.
Due to constraints of neurovascular coupling in cerebellar activity, it is possible that only mossy fiber inputs in response to adaptation elicit demonstrable BOLD signals, while error signals conveyed via climbing fiber spike firing increase may not suitable for fMRI testing. The results further suggest that short-blocked designs may offer the most effective approach, engaging a period of adaptation to changes in feedback without reaching a state of habituation, leading to reliable activation of the cerebellum.

CONFLICT OF INTEREST
The authors state no conflict of interest.

DATA AVAILABILITY
The data that support the findings of this study are available from the corresponding author upon reasonable request.