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VOLUME SEVENTY SIX
THEPSYCHOLOGY OF
LEARNING AND
MOTIVATION

Series Editor
KARA D. FEDERMEIER
Department of Psychology, Program in Neuroscience,
and the Beckman Institute for Advanced Science and Technology,
University of Illinois Champaign, IL, United States

VOLUME SEVENTY SIX
THEPSYCHOLOGY OF
LEARNING AND
MOTIVATION
Edited by
KARA D. FEDERMEIER
Department of Psychology, Program in Neuroscience,
and the Beckman Institute for Advanced Science and
Technology, University of Illinois Champaign, IL,
United States

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Contents
Contributors ix
1. Understanding cerebellar function through network
perspectives: A review of resting-state connectivity of the
cerebellum 1
Jessica A. Bernard
1.Introduction 2
2.The human cerebellum 3
3.Information super highways: Methods for studying cerebellar networks 12
4.The human cerebellum at rest 26
5.Implications for understanding cerebellar function 38
6.Conclusions 40
Acknowledgments 41
References 42
2. What if they're just not that into you (or your experiment)? On
motivation and psycholinguistics 51
Kiel Christianson, Jack Dempsey, Anna Tsiola, and Maria Goldshtein
1.Introduction 52
2.The“ideal speaker-hearer” 56
3.Online vs. offline data 59
4.Comprehension probes 64
5.Feedback during experiments 68
6.Open ended questions and qualitative analyses 70
7.Modulating motivation: Taking cues from second language acquisition (SLA) 73
8.Conclusion 79
References 81
Further reading 88
3. Spectral characteristics of visual working memory in the monkey
frontoparietal network 89
Bryan D. Conklin
1.Introduction 90
2.Memory 90
3.Working memory models 91
v

4.Large-scale networks supporting cognition 95
5.Data acquisition 95
6.Time-frequency analysis 96
7.Working memory nonhuman primate tasks 97
8.Task-dependent oscillations 99
9.Frequency bands 100
10.Choice of tasks 102
11.Limitations 103
12.Frequency band activity in spatial vs non-spatial working memory 104
13.Frequency band activity in visuospatial working memory 109
14.Conclusion 112
References 116
4. The N400in silico: A review of computational models 123
Samer Nour Eddine, Trevor Brothers, and Gina R. Kuperberg
1.General introduction 124
2.Word-level models 127
3.Sentence-level models 143
4.Summary of models in relation to cognitive and biological constraints 167
5.A predictive coding account of the N400 173
6.Future directions 189
Acknowledgments 195
References 195
5. Dynamic decision making: Empirical and theoretical directions 207
Jared M. Hotaling and David Kellen
1.Introduction 208
2.Empirical review 211
3.Candidate explanations and future directions 226
4.Conclusion 233
References 233
6. Connecting movement and cognition through different modes of
learning 239
Elizabeth B. Torres
1.Introduction 240
2.Disparate delays in sensory transduction, transmission and sensory motor
transformations for action could play a role in screening developmental
disorders earlier than current social-cognitive criteria 242
vi Contents

3.Coordinate transformation and frames of refence for purposeful and
exploratory actions 248
4.Kinesthetic reafference bridging movement and cognition 256
5.The degrees of freedom problem: Kinematics of natural motions 262
6.Fundamentally different modes of learning and motivation 266
7.Considering contextual variations for learning, transfer and generalization 270
8.Spontaneous autonomy, exploratory learning and human agency: The role
of reafference in learning to own one's actions and their consequences 275
9.Concluding thoughts 278
References 278
viiContents

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Contributors
Jessica A. Bernard
Department of Psychological and Brain Sciences; Texas A&M Institute for Neuroscience,
Texas A&M University, College Station, TX, United States
Trevor Brothers
Department of Psychology and Center for Cognitive Science, Tufts University, Boston, MA,
United States
Kiel Christianson
University of Illinois, Urbana-Champaign, Champaign; Beckman Institute for Advanced
Science and Technology, Urbana, IL, United States
Bryan D. Conklin
School of Psychology, Georgia Institute of Technology, Atlanta, GA, United States
Jack Dempsey
University of Illinois, Urbana-Champaign, Champaign, IL, United States
Maria Goldshtein
Arizona State University, Tempe, AZ, United States
Jared M. Hotaling
Department of Psychology, University of Illinois, Urbana-Champaign, Champaign, IL,
United States
David Kellen
Department of Psychology, Syracuse University, Syracuse, NY, United States
Gina R. Kuperberg
Department of Psychology and Center for Cognitive Science, Tufts University; Department
of Psychiatry and the Athinoula A. Martinos Center for Biomedical Imaging, Massachusetts
General Hospital, Harvard Medical School, Boston, MA, United States
Samer Nour Eddine
Department of Psychology and Center for Cognitive Science, Tufts University, Boston, MA,
United States
Elizabeth B. Torres
Psychology Department, Rutgers, The State University of New Jersey; Computer Science
Department, Rutgers University Center for Cognitive Science, Computational Biomedicine
Imaging and Modeling Center, Piscataway, NJ, United States
Anna Tsiola
University of Illinois, Chicago, IL, United States
ix

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CHAPTER ONE
Understandingcerebellarfunction
through network perspectives: A
review of resting-state
connectivity of the cerebellum
Jessica A. Bernard
a,b,
∗
a
Department of Psychological and Brain Sciences, Texas A&M University, College Station, TX, United States
b
Texas A&M Institute for Neuroscience, Texas A&M University, College Station, TX, United States
∗
Corresponding author: e-mail address: [email protected]
Contents
1.Introduction 2
2.The human cerebellum 3
2.1Historical context and perspectives: Motor function 5
2.2A brief history of the cerebellum in cognition 7
2.3Cerebellar evolution: Comparative perspectives from non-human primates 9
2.4Cerebellar internal models 10
3.Information super highways: Methods for studying cerebellar networks 12
3.1Comparative insights via tract tracing 13
3.2Quantifying in vivo white matter in the human brain 17
3.3Resting-state functional connectivity 20
4.The human cerebellum at rest 26
4.1Seed-based measures 27
4.2ICA and parcellations 33
4.3Graph theory 35
4.4Cerebellar connectivity in the era of big data 37
5.Implications for understanding cerebellar function 38
6.Conclusions 40
Acknowledgments 41
References 42
Abstract
The human cerebellum, though relatively small in total volume, makes up for it in its
neuronal density and immense computational power. As we seek to understand com-
plex higher order human behavior, it is critical to consider how this structure may con-
tribute to these domains. While historically conceptualized as a motor structure, likely
in large part due to the overt motor deficits often experienced by those with cerebellar
damage, it is now known to play a critical role in cognition. In the last decade in
Psychology of Learning and Motivation, Volume 76 Copyright #2022 Elsevier Inc.
ISSN 0079-7421 All rights reserved.
https://doi.org/10.1016/bs.plm.2022.03.001
1

particular there has been a great deal of growth in the literature in this regard (though
these ideas have been percolating since the 1980s). The development of resting-state
functional connectivity magnetic resonance imaging (fcMRI) also resulted in a boom of
literature seeking to clarify cerebellar interactions with the cortex. In this chapter,
cerebellar anatomy and function are reviewed, with a particular focus on how fcMRI
has impacted our understanding of the human cerebellum and what this has meant
for our accounts of cerebellar processing (that is, the underlying computations). This
work has broadened our appreciation of the cerebellum’s networks linked to higher
order processing, and resulted in thought provoking findings with respect to the
functional organization of the cerebellum that may in the future impact our under-
standing of how the“little brain”helps the cortex produce nuanced and complicated
human behavior.
1. Introduction
When was the last time you rode a bike? If it has been a while, imagine
what it might be like to hop back on and go for a ride. Perhaps at first you
may feel a little shaky and unsteady. However, after a short spin around the
block or down the road you notice that you fairly quickly begin to feel com-
fortable and the ability to ride has “come back to you” without very little
thought at all. This ability to ride a bike, a procedural memory, is engrained
due to practice and has become automatic in a way that you often cannot
describe how to do it. Such is the case with many other motor behaviors
that are automatic and efficient like smoothly reaching for a cup of coffee
across the table, or a bottle of water. In typically developing or healthy indi-
viduals, these smooth movements and procedures occur, thanks in large
part, to a well-functioning cerebellum. The “little brain,” which is not actu-
ally all that little, engages in precise computations that allow for fluid action
often without much thought.
However, it is not just movement that is processed by the cerebellum. As
our understanding of this structure has grown, it has been proposed that in
the same way the cerebellum allows for smooth and precise movement, it
also contributes to our thought processes. Some of the most compelling
evidence comes from instances of disease or lesions that impact the cerebel-
lum, wherein individuals are less capable of producing coherent thought or
have deficits in cognitive processes. Perhaps most famously, the cerebellum
has been implicated in cognitive dysmetria and the associated symptoms
seen in individuals with schizophrenia (
Andreasen et al., 1996;Andreasen,
Paradiso, & O’Leary, 1998;Bernard & Mittal, 2015;Picard, Amado,
Mouchet-Mages, Olie, & Krebs, 2008). This understanding of non-motor
2 Jessica A. Bernard

processing in the cerebellum has also been bolstered by neuroimaging, partic-
ularly functional magnetic resonance imaging (fMRI), though functional
lesion mapping has also been very informative (Ilgetal.,2013;Richter
et al., 2007;Timmann et al., 2008). Methods of measuring cerebellar net-
works (both functional and structural) in particular have also gained popularity
in the past two decades, and have been used extensively to investigate the
cerebellum and its interactions with the rest of the brain.
As will be reviewed in the first part of this chapter, converging evidence
indicates that the cerebellum is a crucial contributor to behavior across func-
tional domains. In addition, classic frameworks for understanding the neural
computations in the cerebellum, in the form of internal model processing
will be discussed. In parallel to our growing understanding of cerebellar
function came the rise of modern neuroimaging methodologies. Indeed,
these approaches made many of these new insights possible. In the last
decade and a half, cerebellar networks have been of particular interest.
Resting-state functional connectivity (fcMRI) has become a widely used
technique that has grown exponentially in its usage since the year 2000. A
recent search on PubMed of the term “resting-state connectivity” gave
15,720 results (as of December 1, 2021), with 2416 in 2021 alone. When
appending “cerebellum” to the above search, 1198 results were returned,
with 199 in 2021. This explosion of research with this technique, parti-
cularly looking at the cerebellum, has provided vast amounts of new
knowledge. A complete and detailed overview of these nearly 1200 papers
is beyond the scope of this chapter (and certainly, some of those studies may
be irrelevant or false positives). However, the latter part of this review will
focus on different approaches for understanding cerebellar networks with
the cortex, primarily networks quantified with fcMRI. In this context,
the discussion will be framed to probe whether and how this explosion in
research has changed and advanced our understanding of cerebellar function
and computations, as we have grown in our understanding of how the
cerebellum and cortex interact.
2. The human cerebellum
The cerebellum is a processing powerhouse. With a volume that is
approximately 10% of the total brain, the cerebellum is home to more than
half of the brain’s neurons (
Andersen, Korbo, & Pakkenberg, 1992). Put
another way, there are more neurons in the cerebellum than there are in
the rest of the brain. This astounding number of neurons is contained in
3A review of resting-state connectivity of the cerebellum

a relatively small volume due to the tight folding of the cerebellar folia that
make up its lobules (subdivisions that are analogous to lobes in the cerebral
cortex). Indeed, thanks to high resolution structural brain imaging, recent
evidence has demonstrated that the human cerebellum, with all of its fold-
ing and convolutions, has a surface area that is 78% of the size of the surface
area of the cerebral cortex (Sereno, Diedrichsen, Tachrount, & Testa-silva,
2020). Because of the large number of neurons that are together in a
relatively small space, the computational power of the cerebellum is vast.
Within the structure there is a well categorized cytoarchitecture. The
cerebellum is home to perhaps the most recognizable of neurons, the
Purkinje cell (Fig. 1). The dense and vast dendritic branching of these cells
are innervated by parallel fibers from granule cells (originating in the granule
cell layer) as well as by climbing fibers originating in the inferior olive.
The Purkinje cells in turn send inhibitory inputs to the deep cerebellar
nuclei that serve as the output units of the cerebellum. Granule cells are also
influenced by mossy fibers (and can in turn influence Purkinje cells) with
Fig. 1Drawn schematic of the Purkinje cell showing innervation by the parallel fibers.
The Purkinje cell structure is one of the most notable in the human brain. Artist credit:
Akshay Markanday via SciDraw.doi.org/10.5281/zenodo.3926003
4 Jessica A. Bernard

excitatory projections (Kandel, Schwartz, Jessell, Siegelbaum, & Hudspeth,
2013). What is especially notable, however, is that cerebellar cytoarchi-
tecture remains consistent across regions whereas patterns of cell
types and their distributions vary across regions in the cerebral cortex.
Thus, the cerebellum differs from the cerebral cortex in cytoarchitectonics
and cortical layering patterns. This suggests that the computations of the
cerebellum are consistent across subunits and regions (
Diedrichsen, King,
Hernandez-castillo, Sereno, & Ivry, 2019;Ito, 2008;Ramnani, 2006;
Schmahmann, 2018). This then raises the question: what is the role of this
computational powerhouse? How does the cerebellum contribute to
behavior?
2.1 Historical context and perspectives: Motor function
The initial framework and functional conceptualization of the cerebellum
was as a motor structure. Some of the earliest work in this area was based
on patients with lesions to the cerebellum. In his foundational paper
Holmes (
Holmes, 1939) described the motor signs and symptoms experi-
enced by individuals with cerebellar damage. These deficits included
decreased muscle tone (hypotonia) as well as a variety of symptoms related
to motor control and coordination. This bias toward motor function and
deficits may have been due, in part, to the immediately observable and quan-
tifiable nature of such deficits. That is, depending on the location of the
lesion, these individuals may have also experienced cognitive deficits that
went relatively unnoticed, particularly in comparison to the observed motor
deficits.
This early characterization in motor function resulted in decades of work
investigating the motor contributions of the cerebellum. While the deficits
first described by
Holmes (1939)were in terms of gross motor systems,
animal models have further demonstrated the contributions of the cerebel-
lum, perhaps most famously in the field of eye-blink conditioning. The
eye-blink reflex, a blink response when the nictitating membrane is stimu-
lated (often with a puff of air, or small piece of cotton), is an automatic
response. This has been well-studied in rabbits, and cerebellar circuits
have been consistently implicated (
Christian & Thompson, 2003;Kim &
Thompson, 1997). Damage to the cerebellum, including to the cerebellar
hemispheres, negatively impacts eye-blink conditioning, such that condi-
tioned responses are not developed (Yeo & Hardiman, 1992). In humans,
the eye-blink conditioning paradigm also demonstrates this pattern of
5A review of resting-state connectivity of the cerebellum

associative learning (Woodruff-Pak, Papka, & Ivry, 1996;Woodruff-Pak &
Thompson, 1988), though the degree to which the associations are learned is
lessened in advanced age (Woodruff-Pak et al., 2010;Woodruff-Pak, Papka,
Romano, & Li, 1996;Woodruff-Pak & Thompson, 1988). Initial specula-
tion was that this was due to differences in the cerebellum (though perhaps
the hippocampus as well, given that it also plays a role in this paradigm)
that occur with the process of aging (e.g.,
Bernard & Seidler, 2014;
Woodruff-Pak et al., 2010). Subsequent work in lesion patients has further
supported cerebellar contributions to this associative learning process in
the human brain (
Gerwig et al., 2006;Gerwig, Kolb, & Timmann,
2007), and populations with known differences in cerebellar structure and
function, such as those with schizophrenia (Picard et al., 2008) and their
first-degree relatives, also show differences in performance on eye-blink
conditioning relative to healthy matched controls (
Bolbecker et al., 2014;
Kent et al., 2020). With the advent of neuroimaging, these links have been
strengthened via data showing relationships between cerebellar volume and
associative learning (
Woodruff-pak et al., 2001), and functional activation
in the cerebellum during the conditioning process (Cheng, Disterhoft,
Power, Ellis, & Desmond, 2008;Cheng et al., 2014;Lundin et al., 2021).
Together, this has further cemented a role for the cerebellum in motor
processes, but also more generally in processes related to learning.
The stimulus-response associations seen in eye-blink conditioning
paradigms rely upon well-defined cerebellar circuitry, but are in many ways
much simpler than the more complex processes associated with learning a
new sequence of movements or a procedure. In addition to the role in
muscle tone and control of movement as described by
Holmes (1939)the
cerebellum has been implicated in the learning of new tasks (Doyon,
Gabitov, Vahdat, Lungu, & Boutin, 2018;Imamizu et al., 2000). While
there has been some suggestion that the cerebellum is not involved in the
process of learning, and rather is critical for the expression of learned motor
behaviors (
Seidler et al., 2002), activation in the cerebellum has been seen
across numerous studies during various motor learning paradigms (reviewed
inBernard & Seidler, 2013), and it is considered to be an important part
of the network involved in motor learning (reviewed inDoyon et al.,
2018). Indeed, much like there are differences observed in eye-blink con-
ditioning in populations with known differences in cerebellar structure and
function, there are also differences in motor learning (e.g.,Dean et al.,
2014). Together, this provides further evidence and support for the role
of the cerebellum in coordinated motor function, and in processes associated
6 Jessica A. Bernard

with learning (and execution) of motor behaviors. With that said, in recent
decades, our understanding of the cerebellum has grown to indicate a role
in non-motor behavior as well.
2.2 A brief history of the cerebellum in cognition
Prior to the advent of human neuroimaging and its widespread use, our
knowledge of brain structure–function relationships resulted from work
with lesion patients. As previously noted, patients with cerebellar lesions
often show motor impairments. However, as early as the 1980s, it was
hypothesized that in humans, there are also non-motor contributions of
the cerebellum, particularly in the domain of language (
Leiner, Leiner, &
Dow, 1986, 1989, 1991). By the late 1990s, this idea gained substantial
traction due to two findings that emerged in this time period. First, foun-
dational work from Nancy Andreasen introduced the concept of cognitive
dysmetria in individuals with schizophrenia (Andreasen et al., 1996, 1998).
Using positron emission tomography (PET), Andreasen and colleagues
(
Andreasen et al., 1996) demonstrated different patterns of brain activation
in the frontal cortex, thalamus, and cerebellum when comparing patients
to neurotypical controls, and suggested that this may result in “dysmetria
of thought” or uncoordinated thought which is seen in the cognitive deficits
and disorganized thought that are present in schizophrenia. This highlighted
the non-motor contributions of the cerebellum.
In near parallel to this first work on cognitive dysmetria,
Schmahmann
and Sherman (1998)carefully characterized cognitive and affective deficits in
a sample of 20 patients with cerebellar lesions. These lesions were primarily
localized to the posterior lobe and the vermis, and deficits were seen across
domains, but were particularly robust for executive function, reasoning, and
affect. These findings resulted in the coining of the cerebellar cognitive
affective syndrome (
Schmahmann & Sherman, 1998), highlighting and
emphasizing that the neurological deficits associated with cerebellar damage
extend beyond the motor domain. Subsequent work from Schamahmann
and colleagues has further expanded our understanding of this pattern of
deficits and reinforced our understanding of cerebellar contributions to
behavior outside the motor domain (e.g.,
Schmahmann, 2004, 2018;
Schmahmann, Guell, Stoodley, & Halko, 2019).
As such, by the late 1990s, there were multiple parallel lines of work
clearly implicating the cerebellum in non-motor behavior. Increased use
of neuroimaging at this time further advanced our understanding in this
7A review of resting-state connectivity of the cerebellum

regard. Lesion mapping across participants using brain imaging has allowed
for the quantification of overlap across patients (Timmann et al., 2008). In
these instances, the degree of overlap in lesions and the locations can be
quantified (
Richter et al., 2007;Timmann et al., 2009), and investigated
in the context of deficits. This approach has provided further insights into
non-motor contributions of the cerebellum and insights into the regions
where damage relates to cognitive deficits. Generally speaking, this work
has also suggested that the lateral and posterior aspects of the cerebellum
are important for non-motor processing.
Not only was neuroimaging instrumental in methods that allow for
more detailed quantification of lesions, but functional magnetic resonance
imaging (fMRI) has further opened the door to our understanding of the
cerebellum in non-motor behaviors. Certainly, since the advent of fMRI,
studies were reporting cerebellar activations as part of their results, but it
was also not uncommon to see these findings largely ignored in discussions.
But, about 10 years ago the tenor shifted to some degree, thanks in part to
foundational work using meta-analysis.
Stoodley and Schmahmann (2009)
took advantage of the fact that many studies across functional domains
had shown activation in the cerebellum. They concatenated the activation
foci in the cerebellum across different task domains and investigated patterns
of overlap in activation across studies (
Stoodley & Schmahmann, 2009).
Most notably, they reported that in anterior regions and in the posterior
lobules VIIIa and VIIIb, activation was consistently seen across motor tasks
(
Stoodley & Schmahmann, 2009). This is consistent with what was known
about the primary and secondary motor representations in the human cer-
ebellum (
Grodd, H€ulsmann, Lotze, Wildgruber, & Erb, 2001;Mottolese
et al., 2013;Rijntjes, Buechel, Kiebel, & Weiller, 1999;Wiestler,
McGonigle, & Diedrichsen, 2011). Further, activation overlap was also
found across tasks investigating working memory, language, and executive
function, localized, perhaps not surprisingly, to more lateral and posterior
aspects of the cerebellum (
Stoodley & Schmahmann, 2009). Notably, they
subsequently replicated this work using fMRI both in an individual subject
(Stoodley & Schmahmann, 2010), and in a larger group analysis (Stoodley,
Valera, & Schmahmann, 2012). This pattern has also replicated in subse-
quent meta-analyses in both healthy young adults, in older adults, and
in patient populations (
Bernard & Mittal, 2015;Bernard et al., 2020;
Keren-Happuch, Chen, Ho, & Desmond, 2014). Finally, there have also
been additional targeted investigations of the cerebellum in higher-order
cognitive processing using both imaging and non-invasive brain stimulation
8 Jessica A. Bernard

(Balsters, Whelan, Robertson, & Ramnani, 2013;Desmond, Chen, &
Shieh, 2005;Ferrucci et al., 2008;Jackson, Maldonado, Eakin, Orr, &
Bernard, 2020;Pope & Miall, 2012). Most recently, King took a detailed
look at the functional organization of the human cerebellum (King,
Hernandez-Castillo, Poldrack, Ivry, & Diedrichsen, 2019) using 28 different
task conditions including memory, spatial processing, motor function, atten-
tion, and emotion. This work showed the same general patterns wherein
motor activation was associated with the anterior cerebellum and the
secondary motor representation (lobules VIIIa/VIIIb) while non-motor
tasks predominantly showed activation in lateral and posterior lobular
regions. Most importantly, King demonstrated that functional territories
are not limited to lobular boundaries, and within a lobule there may exist
different regions associated with performance of distinct tasks/cognitive
domains (
King et al., 2019). This has provided deeper insights into cerebellar
functional organization, and importantly, further cemented the role of the
cerebellum in non-motor processing.
2.3 Cerebellar evolution: Comparative perspectives from
non-human primates
The cerebellum itself is phylogenetically ancient, seen in a variety of species,
including birds, reptiles, amphibians, and primates (
Larsell, 1934). However,
in primates in particular, through evolution, the cerebellum seems to have
expanded (
Balsters et al., 2010;Barton, 2012;Barton & Venditti, 2014).
There are two particularly salient factors to consider in thinking about
evolution of the cerebellum. First, is that in primates this occurred rather
quickly (
Barton & Venditti, 2014). Second, is the idea that this may be
related to cognitive abilities (Balsters et al., 2010;Barton, 2012) and
co-evolution with the expansion of regions of the cortex also associated with
cognitive processing. In a comparative analysis of primate species including
humans, Balsters and colleagues investigated cerebellar volume at a lobular
level to determine whether or not there was selective expansion in regions
associated with non-motor processing (
Balsters et al., 2010). Crus I and II,
which are lateral areas associated with the prefrontal cortex and cognitive
processing (e.g.,King et al., 2019;Stoodley & Schmahmann, 2009), are
selectively larger in the human brain than other areas of the cerebellum
(
Balsters et al., 2010). This further implicates these regions in non-motor
processing, but also suggests that there was selective evolution favoring these
regions that contribute to cognitive processing, further cementing the role
of the cerebellum in this domain.
9A review of resting-state connectivity of the cerebellum

2.4 Cerebellar internal models
It is established that the cerebellum is active during and presumably involved
in cognitive processing, in addition to motor behaviors. However, what
does the cerebellum do to facilitate behavior? Evidence from fMRI has
demonstrated cerebellar activation during performance (
Balsters et al.,
2013;Imamizu et al., 2000;Jackson et al., 2020;King et al., 2019;
Stoodley et al., 2012), and work with lesion patients and in those with
disease that impacts the cerebellum has suggested that an intact cerebellum
is key for optimal performance, given the deficits that have been observed
(e.g.,
Andreasen et al., 1998;Dean et al., 2014;Gerwig et al., 2006;Picard
et al., 2008;Richter et al., 2007;Schmahmann & Sherman, 1998). As
previously discussed, there is also a consistent cytoarchitecture across the
cerebellum, which has led to the idea of the cerebellar universal transform
(
Diedrichsen et al., 2019;Ito, 2008;Ramnani, 2006;Schmahmann et al.,
2019). That is, because the cellular architecture is identical throughout
the cerebellum, the cellular computations are suspected to be the same
across the structure.
Cerebellar function has been conceptualized in terms of internal models
(Wolpert, Miall, & Kawato, 1998), both forward and inverse that are
processed due to an efference copy of a cortical command. This can be most
easily conceptualized in the motor domain. When a motor command is
sent from the brain to the body, a copy of the command is then sent to
the cerebellum. The cerebellum then acts as a comparator, wherein the
predicted motor command is compared to feedback from the periphery
about where a particular effector (limb, finger, etc.) is in space relative to
the goal movement. Put more concretely, if you would like to reach for
a cup of water in front of you, you are generally able to do so with a great
deal of ease. Your motor cortex sends the final command to the effectors
(in this case your arm and hand) to reach for the water. In parallel a copy
of this command is sent to the cerebellum, which monitors where your
arm is in space during the movement process, and provides feedback to
the cortex so your movement can be updated in subtle ways in near real
time. As a result, if you have a healthy and intact cerebellum, reaching
for that glass of water is a relatively straightforward undertaking that you
can complete with relatively little thought. Evidence in support of this ini-
tially came from neurological patients with cerebellar damage (e.g.,
Holmes,
1939) but it is also supported by more recent work on the tickle sensation.
To experience a tickle, the somatosensory cortex is active. Blakemore and
colleagues (
Blakemore, Wolpert, & Frith, 1998) investigated differences in
10 Jessica A. Bernard

brain activation when a person is tickled by an external source, and when
they tried to tickle themselves. Indeed, it is the case that one typically cannot
produce a tickle sensation in themselves. In this investigation, they demon-
strated that when a participant tickled the palm of their hand themselves,
there was no activation in the somatosensory cortex, and patterns of cere-
bellar activation differed as compared to an external source of a tickle
(
Blakemore et al., 1998). This suggests that when one produces an efference
copy of a motor behavior, in this instance, tickling oneself, the cerebellum is
processing the efference copy of that motor command, and as such the
brain knows what to expect via feedback to the cortex and there is no
somatosensory activation as a result (
Blakemore et al., 1998). The monitor-
ing of ongoing movement means the brain expects what is coming and there
is no tickle sensation.
Motor contributions of the cerebellum were the first target for this
functional theorization. However, as our understanding of cerebellar con-
tributions to non-motor behaviors grew, extensions of this framework
were made to apply to non-motor behavior as well. Ramnani (
Ramnani,
2006) suggested that similar computations may be performed for informa-
tion that comes from the prefrontal cortex, instead of from the motor cortex.
Effectively, cerebellar control theory was applied to information from
non-motor regions of the brain, given this idea of a “universal transform”
(
Ramnani, 2006). That is, the same computations are performed across
regions of the cerebellum, but the inputs differ. Shortly thereafter, Ito
(Ito, 2008) further conceptualized this in terms of forward and inverse
models and suggested that the cerebellum may act on efference copies of
thought, along with those of motor commands. In this case, we can use
verbal working memory as an example. It is not uncommon for many indi-
viduals to automatically start mentally repeating a string of digits, such as a
phone number, that needs to be briefly remembered. During this process,
the prefrontal cortex can send a copy of the command of sorts, that allows
the brain to do this. The cerebellum can then support this process, allowing
individuals to maintain the numbers and detect instances where they may be
losing track. However, instead of relying upon external feedback such as
sensory information about limb position, instead feedback related to the list
of numbers and its accuracy is compared.
Support for this came from co-activation of the cerebellum and prefron-
tal cortex during the performance of a variety of cognitive tasks, including
working memory, language and attention, but also from instances of cere-
bellar dysfunction wherein thought is impacted (
Ito, 2008). More recently,
11A review of resting-state connectivity of the cerebellum

Ramnani (Ramnani, 2014) further expanded upon these ideas, to also incor-
porate circuitry related to feedback sources for the processing of cognitive
information. Although the cortical areas and systems for cognition and
motor function are interacting in the cortex, Ramnani argues that autom-
atization of cognitive processing and behavior can also occur independently
from that in the motor domain (though interactive aspects may occur as
well) (
Ramnani, 2014).
While these ideas are now fairly noncontroversial, particularly the idea
that the cerebellum plays a role in cognition and may do so via automatiza-
tion of these behaviors via internal models, the notion of the “universal
transform” has been called into question (
Diedrichsen et al., 2019). Our
knowledge of the cerebellum’s cellular architecture remains the same, and
that itself remains unchanged; however, thanks to careful detailed fMRI
work, it is recognized that different functional activation patterns are seen
topographically in the cerebellum for different tasks (
King et al., 2019).
Diedrichsen and colleagues in turn suggest that there could perhaps be
differences in the processing in those regions (
Diedrichsen et al., 2019).
Instead of a universal transform there may be “multifunctionality” in the
cerebellum. Critically however, this is highly speculative and theoretical,
and there is no evidence from cellular work or animal models to suggest that
this may be the case.
As noted in the introduction, connectivity, particularly resting-state con-
nectivity has rapidly grown as a method for understanding the organization
of the human brain, and the cerebellum specifically is no different. As our
understanding of cerebellar function has expanded, particularly to involve
an understanding of its contributions to non-motor behavior, with an
increase in work on connectivity that occurred in parallel, it is worth
probing whether and how this advanced our understanding of the cerebel-
lum and our understanding of cerebellar function. The next section will pro-
vide a broader overview of connectivity methods and analysis, including
structural methods, and a particular focus on resting-state connectivity.
3. Information super highways: Methods for studying
cerebellar networks
Communication between brain regions occurs through structural
connections and the transmission of information from neuron to neuron
via action potentials. There are structural connections, myelinated axons,
from region to region that underlie neural communication, and in turn,
12 Jessica A. Bernard

complex thought and behavior. Studying these networks and communica-
tion between brain regions provides insights into how regions of the brain,
sometimes somewhat disparate in terms of location, may be engaged
together to produce behavior. These neural highways are the routes that
signals follow when transmitting messages from one brain area to another.
As we try to understand the function and organization of the human brain,
and in cognitive neuroscience how this results in behavior, network
perspectives stand to be especially informative. It has been understood for
quite some time now that the complexity seen in human behavior is due
to interactions between regions and that it is rare for an individual brain
region to act alone. While the primary motor cortex may be the “final com-
mon pathway” to the initiation of a motor command from the cortex to the
periphery, many other regions work together as a network to get to that
point. Thus, studying cortical and cerebellar networks can help us under-
stand the wiring of the brain, and in turn may help us to understand their
function as well as contributions to behavior.
Networks can be studied in multiple ways. Structure can be looked at via
mapping of white matter pathways. But, functional interactions can be
investigated as well, predicated on the idea that areas of the brain that have
a similar pattern of signal at rest are likely to be working together as part of
a brain network. This section will provide an overview of these methods,
and in the cases of comparative approaches and in vivo quantification of
white matter in the human brain, this will be discussed in the context of
our understanding of the human cerebellum. Resting-state approaches will
be described, but a more detailed discussion of resting-state investigations
of the cerebellum will be covered in detail later.
3.1 Comparative insights via tract tracing
Prior to the widespread usage of neuroimaging in the human brain, substan-
tial advancements in our understanding of cerebellar networks came from
work in animal models. Non-human primates in particular have been a
key model for advancing our understanding of these networks, given the
relative similarity of cortical and cerebellar organization. Of course, as
already noted, aspects of the lateral posterior cerebellum and prefrontal
cortex have undergone additional evolution in the human brain and as such
are more expanded in terms of volume (
Balsters et al., 2010), but there are
regional homologs that allow for reasonable relative comparisons and for
the generation of general organizational principles.
13A review of resting-state connectivity of the cerebellum

Motivated by a desire to find structural underpinnings to support
cerebellar contributions to non-motor behaviors, consistent with findings
in human lesion patients, Schmahmann and Pandya (Schmahmann &
Pandya, 1997) investigated cortico-pontine projections in non-human pri-
mates. They investigated different regions within the prefrontal cortex in
reference to the pons to determine potential substrates for heterogeneous
deficits in patients. Ultimately this work provided initial evidence in
support of inputs to the pons from the prefrontal cortex, and inputs were
particularly prominent for the lateral prefrontal cortex (in this case area
9/46d in the primate brain) (
Schmahmann & Pandya, 1997).
Following this work, additional studies took on the challenge of under-
standing cerebello-thalamo-cortical connections using viral tract tracing
methods. The use of viral tract tracing allows for the mapping of connections
between brain regions by injecting a particular region, for example, Crus I
in the lateral cerebellum with a virus, often a rabies virus tracer, and follow-
ing the neurons that result in inputs to that area (
Kelly & Strick, 2000,
reviewed broadly inStrick, Dum, and Fiez (2009)). This can be completed
bidirectionally to determine inputs and outputs, ultimately mapping circuits
in the brain. For our knowledge of cerebellar circuitry, this has been an
invaluable approach and has been particularly informative for the formation
of frameworks for cerebellar function.
These viral tracing studies looked to map the locations of cerebellar
inputs to the cortex (
Hoover & Strick, 1999;Kelly & Strick, 2003;
Middleton & Strick, 2002). Some of the first studies with this approach used
the herpes simplex virus to investigate connections between both the cere-
bellum and basal ganglia to the primary motor cortex (
Hoover & Strick,
1999), though subsequent work used the rabies virus for tracing (Kelly &
Strick, 2000). In this work with the herpes virus,Hoover and Strick
(1999)found outputs to the primary motor cortex in the cerebellar dentate
nucleus, one of the key output regions of the structure, and the primary such
output region for cortical connections. Generally speaking, these outputs
were also localized to the dorsal regions of the cerebellar dentate nucleus
(
Hoover & Strick, 1999). Looking separately at the prefrontal cortex, both
conventional and viral tracers were used together to map connections with
the cerebellum, providing further support for these anatomical connections
(
Middleton & Strick, 2002). The tracing methods used here extended our
understanding of cerebello-frontal connections beyond the pontine map-
pings fromSchmahmann and Pandya (1997), and provided further support
for neural substrates and connections allowing for cerebellar contributions
to cognition.
14 Jessica A. Bernard

Subsequently, the primary motor cortex and lateral prefrontal cortex
were used as viral injection sites, and it was found that neurons in the pri-
mary motor cortex receive inputs from lobules IV and V in the cerebellum
(more anterior), while those in the lateral prefrontal cortex (in the model
used here, area 46) received inputs from Crus II (Kelly & Strick, 2003).
This mapping suggested that there are distinct areas of the cerebellar cortex
that are associated with the motor and prefrontal cortices, respectively, in
the primate brain. Further work also demonstrated that this circuit dissoci-
ation is present in the cerebellar dentate nucleus as well (Dum & Strick,
2003). WhileHoover and Strick (1999)found evidence to suggest that
the dorsal dentate is a source of cerebellar output to the primary motor
cortex, this subsequent investigation further refined our understanding of
these cerebellar circuits and found evidence for outputs to the prefrontal
cortex as well. Using a similar mapping approach, the dorsal dentate was
shown to have projections (via the thalamus) that connect with the primary
motor cortex, while the more ventral region is connected to the lateral
prefrontal cortex (again, area 46 in the monkey model) (Dum & Strick,
2003).Fig. 2provides a schematic overview of these dissociated loops on
a human anatomical image.
Fig. 2Cerebello-thalamo-cortical connections sketched on a template brain MRI. The
circuit from the lateral posterior cerebellum to the frontal cortex is presented in purple,
while the circuit from the anterior cerebellum to the motor cortex is presented in gold.
Here, only the projections from cerebellum to cortex are shown, and the way points in
the thalamus do not reflect detailed subregional placement. They are solely to reflect
the findings that there are distinct synapses within the thalamus for these circuits.
Inputs from the respective cortical regions reach the cerebellum via the pontine nuclei,
closing these parallel loops.
15A review of resting-state connectivity of the cerebellum

While these dissociable loops with the prefrontal cortex and primary
motor cortex have been especially informative and have provided the
primary anatomical basis for the expansion of cerebellar internal model
frameworks to the cognitive domains (
Ito, 2008;Ramnani, 2006), addi-
tional connections have also been discovered. The inferior parietal lobule
also receives projections from the cerebellar dentate nucleus which pro-
vides additional support for non-motor contributions of the cerebellum
(
Clower, West, Lynch, & Strick, 2001). Finally, in the last decade, this
methodology has furthered our understanding of the cerebellar vermis,
which had been conceptualized as being separate from the cerebral cortex
in terms of connectivity, by demonstrating that there are inputs from pri-
mary and pre-motor cortical regions (
Coffman, Dum, & Strick, 2011).
From a functional perspective this suggests that the cerebellar vermis may
also be processing information from the cortex, though historically it was
referred to as the “spinocerebellum.”
Together, this work in primate models was foundational for our
understanding of cerebellar networks with the cortex, and mapped out
closed-loop (named because the loops from distinct cerebellar regions to
distinct cortical regions are separate from one another and in parallel)
cerebello-thalamo-cortical circuits that provide structural circuit-based
support for cerebellar processing in non-motor behavior (
Strick et al.,
2009). Indeed, this understanding of these parallel circuits is a key element
to the internal model frameworks for both motor and non-motor processing
(Ito, 2008;Ramnani, 2006, 2014). These circuits allow for the segregated
and distinct inputs from the cortex, a mechanism by which the efference
copy may be transmitted to the cerebellum. It is also not at all surprising
that the functional topography in the cerebellum, and deficits seen after
damage in lateral posterior regions (such as Crus II) (
Keren-Happuch
et al., 2014;King et al., 2019;Schmahmann & Sherman, 1998;Stoodley
& Schmahmann, 2009;Stoodley et al., 2012) map on to the relative inputs
from the motor and lateral prefrontal cortex, respectively.
In addition to the mapping of these cerebello-thalamo-cortical circuits,
these viral tract tracing approaches have also expanded our understanding
of broader cerebellar connectivity. In addition to its networks with the
cortex, there is a set of direct connections linking the cerebellum to another
key subcortical system, the basal ganglia (
Bostan, Dum, & Strick, 2010;
Bostan & Strick, 2018;Hoshi, Tremblay, Feger, Carras, & Strick, 2005).
Again, work using viral tract tracing showed that there are bidirectional
connections linking these regions directly to one another, potentially
16 Jessica A. Bernard

allowing for complex interactions that mediate behaviors (across the motor
and non-motor domains) outside of the cerebellar contributions via cortical
interactions and processing of internal models. Both the cerebellum and
basal ganglia have been impacted in learning processes (
Ashby, Turner, &
Horvitz, 2010;Caligiore, Arbib, Miall, & Baldassarre, 2019;Doyon et al.,
2018;Imamizu et al., 2000), though the mechanisms by which this occurs
differs. Bostan and Strick (Bostan & Strick, 2018) argue that together the
cerebellum and basal ganglia contribute to integrated networks from the
subcortex to the cortex with these connections, though the specific interac-
tions between these two systems (the cerebellum and basal ganglia) are
unclear. However, this additional understanding of cerebellar networks
in non-human primates has further contributed to our knowledge of the
cerebellum and its function more broadly, and adds to the foundation of
work that can serve as a starting point for follow-up investigations in the
human brain. While at this point the implications of these cerebellar
connections with the basal ganglia are unknown and have not necessarily
changed our theories and frameworks for understanding cerebellar function,
as we gain additional understanding of these connections there may be
additional theoretical impacts.
3.2 Quantifying in vivo white matter in the human brain
Because of the toxic nature of viral tract tracing methods (Kelly & Strick,
2000), this approach to investigate white matter connections in the human
brain is not feasible. This methodology is only an option in animal models.
However, with modern neuroimaging we are able to measure and trace
these white matter connections in vivo using specialized parameters during
data acquisition. Diffusion tensor imaging (DTI) is a methodology that
investigates the myelin that wraps the axons of the neurons in the brain.
With this approach, we can quantify the health of white matter, and we
can also trace the connections between different brain regions. DTI
measures the diffusion of white matter along an axon (
Jones & Leemans,
2010;Le Bihan et al., 2001) based on the idea that intact myelin will result
in the streamlined diffusion of water along that track. Myelin is an insulator
of the axon, much like a plastic coating on housing wires, and when it is
intact, water diffuses along the axon. When myelin is damaged as can occur
in some diseases and as it degrades in advanced age, water diffusion is less
constrained (
Sullivan & Pfefferbaum, 2006). With tractography measures
(e.g.,Gong et al., 2009), in addition to extracting measures related to the
17A review of resting-state connectivity of the cerebellum

diffusion of water, we can also trace the connections between brain regions
in safe ways that parallel the viral tract tracing methods used in non-human
primates. Here, a summary of key findings will be provided in the context
of what this has meant for our understanding of cerebellar function.
However, Habas and Manto (
Habas & Manto, 2018) recently wrote a com-
prehensive overview of this work for those that are interested in further
detail.
The use of DTI approaches has further advanced our understanding
of cerebello–cortical interactions in the human brain, and has allowed for
researchers to conduct investigations that parallel those completed in
non-human primates. Work looking at white matter from the primary
motor cortex, using a seed based on functional activation during tapping
of the hands, showed that there are tracts connecting to the cerebellum,
in addition to cortical regions (
Guye et al., 2003). This cortically focused
work confirmed the ability to map connections between the motor cortex
and cerebellum.
Habas and Cabanis (Habas & Cabanis, 2007) conducted a detailed map-
ping of the dense white matter of the cerebellum and brainstem. They were
successfully able to resolve pontine inputs, as well as white matter connec-
tions from the deep cerebellar nuclei to the cerebellar peduncles (inferior,
middle and superior). Notably, thanks to the methodological approach
they used, they were also able to differentially map cortico-pontine tracts
from the prefrontal cortex (
Habas & Cabanis, 2007), providing in vivo
evidence of the cortico-ponto-cerebellar tracts delineated in non-human
primates (Kelly & Strick, 2003;Middleton & Strick, 2002). Additional
white matter mapping within the cerebellum itself has also been completed,
resulting in further confirmation of the white matter connections within
the structure, including connections from the cerebellar cortex to the
deep cerebellar nuclei and peduncles (
Granziera et al., 2009;Salamon
et al., 2007), and there is now a probabilistic atlas of cerebellar white matter
available (van Baarsen et al., 2016).
While motor connections from the cortex to the cerebellum had
been mapped, investigating those from non-motor regions was also of
interest. Jissendi and colleagues (
Jissendi, Baudry, & Baleriaux, 2008) used
tractography approaches to investigate both the prefrontal and parietal
cortices with respect to their inputs to the cerebellum. Given the existing
work in non-human primates (
Clower et al., 2001;Kelly & Strick, 2003;
Middleton & Strick, 2002), it is perhaps not surprising to see these tracts
resolved in the human brain. Shortly thereafter, Kamali and colleagues
18 Jessica A. Bernard

(Kamali, Kramer, Frye, Butler, & Hasan, 2010) also published work
delineating the cortico-ponto-cerebellar pathways, though in this instance
they did so from the frontal, parietal, occipital, and temporal cortices, expan-
ding our understanding of cerebellar inputs beyond the existing motor,
prefrontal, and parietal circuits. The inclusion of temporal and occipital
inputs provides further structural underpinnings of cerebellar processing
across domains (
King et al., 2019), and is also consistent with recent novel
work implicating the cerebellum in visual attention, including a cerebellar
map of the visual field (
Brissenden et al., 2018).
Taking an approach looking at whole-brain circuits, work conducted
by Salmi and colleagues (Salmi, Pallesen, & Neuvonen, 2010) combined
task-based fMRI with diffusion imaging. While work to this point had
largely looked at these circuits separately in different samples, here the full
closed-loop circuits were investigated, and multiple loops were mapped
in the same sample in conjunction with behavior. They demonstrated a
topographical dissociation in activation tasks for a motor task and a working
memory task within the cerebellum that is consistent with other work in
this regard (e.g.,
King et al., 2019;Stoodley et al., 2012), and they also
found dissociable white matter tracts connecting lateral posterior areas of
the cerebellum to the prefrontal cortex and more anterior regions to the
motor cortex, respectively (
Salmi, Pallesen, Neuvonen, Brattico, et al.,
2010). In this carefully executed study, the authors were able to link func-
tional activation patterns that include topographically distinct cerebellar
and cortical regions with the purported structural circuits that underlie
the networks for this processing. Notably, they traced the complete circuit,
including the cortical inputs to the cerebellum through the pons as well
as cerebellar outputs to the cortex through the cerebellar peduncles and
via the thalamus (
Salmi, Pallesen, Neuvonen, Brattico, et al., 2010), com-
pleting the closed-loop circuits mapped in non-human primates (e.g.,
Kelly & Strick, 2003). Further, they demonstrated that the connections to
the prefrontal and motor cortices are parallel to one another both via the
thalamus to the cortex, and in the pons when inputs are coming to the
cortex, supporting the non-overlapping and closed-loop nature of this com-
munication (
Salmi, Pallesen, Neuvonen, Brattico, et al., 2010). This work
demonstrated the structural connections that purportedly allowed for the
functional activation patterns that were described by using the activation
in the cerebellum as a starting point for tractography (
Salmi, Pallesen,
Neuvonen, Brattico, et al., 2010). Subsequently, this dissociation has also
been replicated in the human brain in an adolescent and young adult
19A review of resting-state connectivity of the cerebellum

population, further supporting the presence of these networks, and our abil-
ity to resolve them in humans (Bernard, Orr, & Mittal, 2016).
As techniques for mapping white matter in the human brain have
advanced, along with improved imaging acquisition techniques, and higher
field imaging, we have been able to better understand cerebello-cortical
white matter connections in a more fine-grained manner. Taking advantage
of high-resolution imaging and a 7 T scanner Steele and colleagues were able
to map the motor and cognitive output circuits from the dorsal and ventral
cerebellar dentate nucleus, respectively in the human brain (
Steele et al.,
2017). This provided confirmatory evidence for circuits initially mapped
in non-human primates (Dum & Strick, 2003) and provided additional
support for parallel loops for processing information from disparate regions
of the cortex. Finally, though cerebellar-basal ganglia circuits have a less
clearly delineated role in our understanding of cerebellar function, these
have also been mapped in the human brain (
Pelzer et al., 2013), and may
provide a foundation for future work seeking to better understand this
circuit and its contributions to behavior.
Together, work in the human brain provides evidence suggesting
strong parallels in networks as those mapped in non-human primates using
invasive methods. That is, the neuronal connections are present that link
the cerebellum to multiple disparate areas of the cortex, but in parallel
ways. These loops are seemingly non-intersecting, though cortico-cortical
communication could also impact the information being shared with the
cerebellum via these circuits. Furthermore, the cerebellar-basal ganglia con-
nections in both human and non-human primates (
Bostan et al., 2010;
Hoshi et al., 2005;Pelzer et al., 2013) may serve to further modify and mod-
ulate cerebellar (and basal ganglia) processing. Our knowledge of cerebellar
white matter and networks has provided key insights into the pathways
that allow for communication between the cerebellum and the rest of the
brain. And while this has been critical knowledge, perhaps the largest growth
in literature investigating the cerebellum has been with fcMRI.
3.3 Resting-state functional connectivity
During functional brain imaging paradigms, the blood oxygen level depen-
dent (BOLD) signal is measured and used to identify brain regions that are
purportedly involved in the processing of a particular task (relative to a rel-
evant control condition). This measure is based upon the idea that blood
flow can serve as a proxy for activation, given that the metabolic needs of
20 Jessica A. Bernard

the brain are likely to increase in regions that are involved in task process-
ing (again, relative to an appropriate control condition). The process of
thinking demands energy, and blood flow increases to accommodate this.
In addition to looking at task activation patterns, the BOLD signal has also
been harnessed as a way of investigating brain networks. Resting-state func-
tional connectivity (fcMRI) looks at similarities in the BOLD signal in the
absence of a task (e.g.,
Biswal et al., 2010;Shehzad et al., 2009), resulting in
functional networks. Ultimately, this is based on the idea that areas that
are part of networks are likely to have similar patterns in the BOLD signal
at rest, due to their functional coupling during tasks (
Van Dijk et al., 2010).
The first investigations in this regard were conducted looking at the motor
network and revealed that areas in premotor cortex, the basal ganglia, and
cerebellum had BOLD signals at rest that were highly correlated with that
in the primary motor cortex (
Biswal, 2012;Biswal, Van Kylen, & Hyde,
1997). Put another way, when looking at areas that are not thought to work
together in known networks, like for example, the motor cortex and ipsi-
lateral visual cortex, it has also been demonstrated that the correlation
between the BOLD signal in these regions is low (
Van Dijk et al., 2010),
further suggesting that this is indicative of brain networks for regions that
work together. Here, an overview of resting state will be provided, and
in the next section, we will delve deeper into the data that resulted from
investigating the cerebellum. In
Fig. 3, example time courses from the cer-
ebellum are shown, overlaid on one another for areas that are significantly
correlated with one another at rest, and those that have weaker connections.
As noted earlier, this methodology has become increasingly popular in
the past two decades. There are several reasons that have contributed this.
First, fcMRI provided a novel way to investigate brain networks function-
ally. While structural connections are critical for our understanding of
neuroanatomy, further insights into function are of particular interest in
cognitive neuroscience as we seek to understand how the brain underlies
complex behavior. Functional networks can provide insights into the brain’s
functional organization and in turn inform our understanding of behavior.
Second is the relative ease with which data can be collected, given the
absence of a task. Paradigms vary in terms of eyes opened vs closed, but
in both cases, participants are asked to lay still. This eliminates the potential
need for additional training on tasks outside of the scanner, and also elim-
inates any possible transforms and confounds that occur from doing training
in an upright manner at a computer and then moving to a scanner environ-
ment where one is laying supine. Finally, particularly early on, the fcMRI
21A review of resting-state connectivity of the cerebellum

protocols were relatively short, sometimes as little as 5–6min. While there is
data indicating that the stability of resting-state measures stabilizes to a
relative asymptote after about 8min (Van Dijk et al., 2010), more recent
approaches have suggested longer scan times for optimal data collection
and analysis. Indeed, there are also some groups doing “deep” imaging
wherein hours of resting-state data are collected from smaller samples of
participants (Gordon et al., 2017). Particularly early on, the shorter scans
allowed for data collection to happen relatively easily, often in conjunction
with already existing task-based scans. Though longer scans are increasingly
becoming the norm, these paradigms, and the ability to concatenate across
scans within a person still provide advantages, particularly in aging and
clinical populations where task performance or participant comfort are big-
ger concerns and challenges. Multiple shorter runs of resting-state data can
be concatenated relatively easily in cases where breaks are needed.
Fig. 3Time series data from Right Crus I (black) and left dorsolateral prefrontal cortex
(dlPFC; dark gray) overlaid together (top panel), and Right lobule V (black) overlaid with
left dlPFC (dark gray; bottom panel). Data come from an individual participant from the
HCP data set and show the greater relative similarity in the timeseries from Crus I and
dlPFC which are significantly correlated at rest. They are more dissimilar when looking at
lobule V and dlPFC, which are not part of known networks and circuits together. The
vertical axis represents the BOLD signal in arbitrary units, while the horizontal axis
represents time.The idea of this figure is modeled after an example from Van Dijk KRA,
Hedden T, Venkataraman A, Evans KC, Lazar SW, Buckner RL. 2010. Intrinsic functional
connectivity as a tool for human connectomics: Theory, properties, and optimization.
Journal of Neurophysiology 103:297–321.
22 Jessica A. Bernard

Further adding to the benefits and popularity of fcMRI is its high degree
of replicability across scanners (Biswal et al., 2010). In a large proof of
principle study, Biswal and colleagues concatenated resting-state data
from across 35 different sites to investigate the replicability of these net-
works. They demonstrated what they describe as a “universal functional
architecture” in the resting state, along with the feasibility of data sharing
across multiple sites. They did, however, note that age and sex are both
factors that can influence the signal (
Biswal et al., 2010), and perhaps not
surprisingly, subsequent work has been dedicated to investigating these
factors further (e.g.,
Bernard, Ballard, & Jackson, 2021;Bernard et al.,
2013;Ferreira & Busatto, 2013;Ferreira et al., 2016;Fitzgerald, Pritschet,
Santander, Grafton, & Jacobs, 2020;Pritschet et al., 2020). However, this
ability to concatenate across data collection sites has also led to data sharing
through public websites (examples include 1000 Functional Connectomes,
NITRIC, SchizConnect, among others), and arguably most well-known
with the Human Connectome Project (HCP) (
Smith et al., 2013;Van
Essen et al., 2013, 2012). This effort has resulted in the acquisition of
resting-state data from over 1000 people, including detailed phenotypic
data on behavior, health history, and background. The data are also publicly
available, allowing for researchers to explore these functional networks,
even if they do not have a scanner themselves. Subsequently, off-shoots
focusing on development and aging have also begun (
Bookheimer et al.,
2019;Harms et al., 2018) allowing for an increased understanding of
network dynamics in development and senescence.
As fcMRI has become a popular method for providing insights into brain
networks and functional organization, the question as to what it is in fact
measuring is important. The general assumption is that resting-state corre-
lations in the BOLD signal reflect, at least to some degree, the underlying
functional connections and networks linking brain regions. For example,
in development, resting-state connectivity in what is known as the default
mode network is associated with measures of white matter in the cingulum
bundle (
Gordon et al., 2011). Furthermore, when looking at development
over 12 months in adolescents, changes in white matter connecting cere-
bellar Crus I to the thalamus was associated with differences in resting-state
connectivity from the cerebellum to frontal cortical regions (
Bernard et al.,
2016). At baseline, associations between resting state and white matter were
also present (Bernard et al., 2016). In a review of this field from the last
decade, Damoiseaux and Grecius noted that in most cases these associations
23A review of resting-state connectivity of the cerebellum

are present. That is, white matter connections do seem to be underlying the
associations seen in fcMRI (Damoiseaux & Greicius, 2009). However, they
also discuss case studies of both a callosotomy and commissurotomy patient,
and in both instances found the presence of some bilateral resting-state
connectivity. In the callosotomy patient, where scans were available before
and after the surgery, there were notably fewer bilateral connections (while
ipsilateral within hemisphere connections remained), but some were still
present. Damoiseaux and Greicius therefore suggest that resting-state net-
works, though largely related to underlying white matter connections, are
also perhaps indicative of more than just the structural networks of the brain
(
Damoiseaux & Greicius, 2009).
Further highlighting this idea is more recent work investigating
resting-state connectivity in non-human primates (macaque monkeys),
when awake, and at different levels of anesthesia (
Barttfeld, Uhrig, Sitt,
Sigman, & Jarraya, 2015). First, and perhaps not surprisingly given other
work in this area, they demonstrated that connectivity differed across awake,
medium, and heavy sedation. They note, however, that under sedation,
the patterns of connectivity seem to be those more locked to and related
to the underlying anatomy connecting disparate brain regions. In the awake
state, there were changes in the functional organization that often differed
from what would be expected due to the underlying anatomy. The authors
suggest that this may be due to consciousness and the associated cognitive
processing (
Barttfeld et al., 2015). That the resting-state signal when awake
may in part reflect consciousness and thought would also help explain why
resting-state correlations may still be present in cases where the underlying
anatomical connections have been severed (
Damoiseaux & Greicius, 2009).
Structural connectivity is certainly related to the functional architecture
of the brain, but at a more basic level, there has also been work interrogating
the origin of these correlations in the BOLD signal at rest (Leopold & Maier,
2012). That is, are they related to underlying neuronal activity? The first
work in this area looked at direct recordings from the primate visual cortex
in conjunction with MRI (Shmuel & Leopold, 2008). This investigation
revealed correlations between the BOLD signal and fluctuations in neuronal
firing from the direct neural recordings. The authors argued then that the
correlations seen in human fcMRI may be linked to underlying neuronal
activity. With that said, this work was not without controversy. In a
follow-up, Logothetis and colleagues (
Logothetis et al., 2009) contended
that the findings reported byShmuel and Leopold (2008)were inherently
flawed and confounded due to a flickering screen that provided visual
24 Jessica A. Bernard

stimulation. While this stimulation was not perceptible, there was a flicker-
ing present, and in a reanalysis of the data, they determined the results
reported byShmuel and Leopold (2008)were inappropriate, and they could
not conclude that rates of neuronal firing were associated with the BOLD
signal (
Logothetis et al., 2009). Shortly thereafter, Sch€olvinck further took
on this important question (Sch€olvinck, Maier, Ye, Duyn, & Leopold,
2010). In a separate study, they recorded local field potentials from different
lobes of the monkey brain (macaque monkeys) again in conjunction with
the collection of MRI data. Here, they definitively demonstrated correla-
tions between the BOLD signal and that which was recorded from local
field potentials. There are two key additional findings of note. First, they
saw similar correlations when looking at LFPs from different lobes in the
brain. Second, they found that these correlations were state dependent (eyes
open vs closed though awake) (
Sch€olvinck et al., 2010). Thus, there also
appear to be neuronal underpinnings to the resting-state signal.
While resting-state connectivity methods are based on the BOLD signal
in the absence of a task, and data collection approaches are reasonably
consistent and straightforward, the ways in which the data can be analyzed
are varied (for a detailed overview see
Behrens & Sporns, 2012). Arguably
the most common approach is referred to as seed-based, or seed-to-voxel
connectivity. A particular region of interest acts as a seed region, and from
that region, the BOLD signal from across all voxels within that seed is
extracted. Correlations between the signal in the seed region and all other
voxels in the brain are then computed. These correlation coefficients are
converted to z-scores, which are then thresholded and corrected for multiple
comparisons at the group level, to account for the many comparisons that
are conducted in a whole-brain analysis. In most cases, this is a somewhat
hypothesis-driven approach, in that researchers typically have an area in
mind and are hoping to better understand its functional networks. In
contrast independent components analysis (ICA) takes a more data-driven
approach and determines consistent components in the resting-state signal.
Groups of regions that have similar patterns over time in the BOLD signal
together form a distinct component, revealing resting-state brain networks
(
Calhoun & Pekar, 2000). At this point there are several canonical net-
works seen in the resting state (motor network, default mode network,
fronto-parietal control network), and these are revealed using ICA, but
can also be mapped using seed-based approaches. Finally, graph theory
approaches have also been applied to understand the functional architecture
of the human brain (
Bullmore & Sporns, 2009;Rubinov & Sporns, 2010).
25A review of resting-state connectivity of the cerebellum

Network nodes are delineated, and the architecture of these connections is
defined in terms of edges and interactions. Areas may cluster together, and
may reveal key network hubs, and more generally, the properties of these
networks can be quantified (
Bullmore & Sporns, 2009). This approach
has been very successful in describing the functional architecture of the
human brain and has been done on a relatively large scale (
Power et al.,
2011). Graph theory has also been useful in development to quantify the
emergence of brain network architecture as maturation processes occur
(
Fair et al., 2009).
Finally, in parallel to these approaches parcellation has been of great
interest (e.g.,Buckner, Krienen, Castellanos, Diaz, & Thomas Yeo, 2011;
Power et al., 2011;Yeo et al., 2011). Ultimately, the goal is to define the
functional architecture of the brain at rest. Parcellation approaches also
take a data driven approach (though distinct from that used in ICA) to define
networks in the cortex (or subcortex or cerebellum) with similar signals at
rest. The approaches themselves have been a distinct area of methodological
inquiry, but broadly speaking, they remain agnostic to anatomy and white
matter connections, and focus solely on the BOLD signal at rest. Together,
with graph theory, ICA, and seed approaches, all of these methodologies are
different means to one goal—that is, understanding the human brain, its
functional architecture, and, certainly at least in some cases, how this results
in complex behavior. These methods have also been applied to the cerebel-
lum, which will be explored next, along with questions as to whether
(or not) these approaches have served to advance our understanding of
cerebellar function, and theories.
4. The human cerebellum at rest
Resting-state connectivity quickly opened up new and uncharted
territory in terms of understanding the brain, and in turn behavior.
Application of the methodologies to map functional brain networks have
been applied to the whole brain, but also to the cerebellum. As noted earlier,
at this point, when searching PubMed for “resting-state connectivity
cerebellum,” there are approximately 1200 hits. Of course, these are not
all original investigations and include reviews, or only cursory mentions
of the cerebellum; however, there have also been many studies focused
on cerebellar networks in an attempt to advance our knowledge and under-
standing of the function of this region and its role in behavior. The different
methodologies for investigating the BOLD signal at rest have all been
26 Jessica A. Bernard

applied to the cerebellum, providing a variety of insights into functional
organization of the cerebellum in the human brain. In some cases, as dis-
cussed below, the functional networks parallel what is known about cere-
bellar anatomy, particularly with respect to white matter circuits with the
cortex. In others, however, the specific underlying white matter projections
have not yet been mapped. However, it is possible that this is more due to
the challenges of resolving these connections due to the dense white matter
and the limitations of neuroimaging. In this next section, an overview of
the key findings across methodologies is provided. Notably, this is primarily
focused on studies of healthy young adults, though some work on develop-
ment and aging is referenced. There is also a large literature on cerebellar
connectivity in clinical populations (both psychiatric and neurological) that
is beyond the scope of this chapter. In this section, the different analysis
approaches are discussed separately, though comparisons are drawn across
sections to some extent, and in multiple instances, investigations included
multiple resting-state analysis techniques. Further, it is notable that around
2010 (and shortly before) there was a quick increase in research in this
area, likely tied, at least in part to work from Stoodley and Schmahmann
(
Stoodley & Schmahmann, 2009) describing the cerebellar functional
topography, and a more general re-emergence in interest in the cerebellum
in non-motor behavior. As such, many of these results were published
in parallel across research groups working on similar questions. In the last
section of the chapter, whether and how these investigations have impacted
our understanding of cerebellar function is addressed.
4.1 Seed-based measures
Perhaps not surprisingly, seed-based connectivity measures are very com-
monly used in investigations of cerebellar functional networks. Relatively
speaking, this started fairly early in 2005, with work from Allen and col-
leagues (
Allen et al., 2005). This is seemingly the first study using fcMRI
in a manner focused on the cerebellum. This study placed a seed in the
cerebellar dentate nucleus and demonstrated that both frontal and motor
networks are connected to the cerebellum, demonstrating that areas of
the cortex known to have structural connections with the cerebellum
(
Dum & Strick, 2003;Kelly & Strick, 2003) are connected to the structure
functionally at rest. Importantly, they also included control analyses of the
auditory and visual cortices, areas that are not known to have white matter
connections with the cerebellum. In these instances, there were no areas of
27A review of resting-state connectivity of the cerebellum

connectivity, demonstrating the specificity of this work, and supporting
links between structural and functional connectivity (Allen et al., 2005).
This was foundational work demonstrating the feasibility of using fcMRI
to investigate cerebello-cortical networks in the human brain, and the
dissociability of motor and non-motor circuits at rest.
Perhaps not surprising, there was, and still is, a broad interest in mapping
and understanding these dissociable motor and non-motor networks, as they
purportedly parallel the closed-loop circuits seen structurally in non-human
primates (
Clower et al., 2001;Dum & Strick, 2003;Kelly & Strick, 2003;
Strick et al., 2009), and subsequently delineated in the human brain (Salmi,
Pallesen, Neuvonen, Brattico, et al., 2010). As such, several papers were
published within quick succession of one another tackling these networks
in more detail, building off the work from Allen and colleagues (
Allen
et al., 2005). Krienen and Buckner focused on fronto-cerebellar circuits
specifically, as they had not yet been carefully mapped out (Krienen &
Buckner, 2009). They placed seeds in the dorsolateral prefrontal cortex,
medial prefrontal cortex, and anterior prefrontal cortex, in addition to seeds
in the primary motor cortex and in their analysis were able to dissociate
those cerebello-cortical networks (
Krienen & Buckner, 2009). Notably,
they found connectivity in topographically distinct regions of the cerebel-
lum that was largely consistent with the dissociations shown in non-human
primate tracing work (
Kelly & Strick, 2003;Krienen & Buckner, 2009).
Shortly thereafter, work from O’Reilly and colleagues was published
also investigating dissociable motor and prefrontal connectivity patterns
with the cerebellum (O’Reilly, Beckmann, Tomassini, Ramnani, &
Johansen-Berg, 2010). In this investigation, the authors looked at how each
voxel in the cerebellum mapped on to pre-defined regions of interest in
the cortex. Ultimately, the dissociations demonstrated in this investigation
(O’Reilly et al., 2010) largely parallel those found byKrienen and Buckner
(2009)adding further support to the dissociability of cerebello-thalamo-
motor and prefrontal networks and parallel functional circuits in the human
brain. These parallel, but still distinct analysis approaches yielded compar-
able findings, also underscoring how relatively robust this pattern is in the
human brain.
Fig. 4demonstrates the relative dissociation of frontal/
association networks with the cerebellum and those associated with motor
circuits using Crus I and Lobule V, respectively, as seeds.
Following these initial investigations looking somewhat generally at
delineating the resting-state analogs of the closed-loop motor and frontal
cerebellar circuits came a series of investigations looking in more detail at
28 Jessica A. Bernard

the whole cerebellum. Somewhat simultaneously, Bernard and colleagues
(Bernard et al., 2012) and Sang and colleagues (Sang et al., 2012) published
more detailed investigations of lobular cerebellar connectivity. Bernard and
colleagues (Bernard et al., 2012) took advantage of, what was at that time, a
fairly new cerebellar normalization and structural parcellation available via
the SUIT toolbox (Diedrichsen, 2006;Diedrichsen, Balsters, Flavell,
Cussans, & Ramnani, 2009). This allowed them to look at individual cer-
ebellar lobules as seeds. All cerebellar resting-state data was processed sepa-
rately from the cortex to normalize specifically to a cerebellar template
and, as best as possible, minimize the impact of signal from ventral visual
areas on the signal in the cerebellum. They investigated both within cere-
bellar connectivity, as well as cerebello-cortical connectivity on a lobule by
lobule basis. While the general principles of the cerebellar functional topog-
raphy and close-loop patterns were revealed (Stoodley & Schmahmann,
2009;Stoodley et al., 2012), this investigation also expanded our general
understanding of cerebellar connectivity on a more granular level, produc-
ing detailed maps for each hemispheric and vermal lobule. Similarly, Sang
and colleagues also produced detailed connectivity maps at the lobular level
(Sang et al., 2012). However, they also computed comparisons between the
connectivity patterns of vermal lobules relative to their hemispheric coun-
terparts. Interestingly, and consistent with the general idea that the vermis is
arguably less linked with cortical processing (though not entirely dissociated
from the cortex as evidenced by animal tracing work, seeCoffman et al.,
2011), they found greater connectivity with the cortex in hemispheric
Fig. 4Resting-state networks of Crus I (orange/yellow) and Lobule V (purple/blue) in a
large sample of participants (n¼590) from Cam-CAN (Shafto et al., 2014;Taylor et al.,
2017). These unpublished analyses followed methods and thresholding used in our
recent work on the dentate nucleus (Bernard et al., 2021). The dissociable networks
for Crus I and Lobule V are notable, and this general pattern has been consistent across
the studies reviewed here, despite different methods or approaches to defining the
resting-state seeds.
29A review of resting-state connectivity of the cerebellum

lobules relative to those in the vermis. However, this is not to say that vermal
lobules had no connectivity with the cortex at rest; but, the patterns and
extent differed (Sang et al., 2012). Finally, to further link these lobular
resting-state signals with cortical connectivity, the authors defined cortical
resting-state networks using ICA and then looked at the lobular associations
with these networks. In brief, they demonstrated that different cerebellar
lobules were differentially associated with cortical functional networks in
topographically organized ways, consistent with prior resting-state cerebel-
lar connectivity investigations (
Habas et al., 2009;Krienen & Buckner,
2009;O’Reilly et al., 2010). In both of these investigations, cerebellar
connectivity with the basal ganglia and medial temporal lobe regions was
reported (Bernard et al., 2012;Sang et al., 2012), and Sang and colleagues
noted that the functional coupling they observed with the amygdala was
particularly novel (
Sang et al., 2012).
Finally, taking this same general lobular approach, Kipping and col-
leagues looked more specifically at motoric networks of the cerebellum
(
Kipping et al., 2013) with a focus on five lobules in the more anterior
aspects of the cerebellum. They too took careful steps to minimize the influ-
ence of the BOLD signal from adjacent ventral visual areas. This analysis
furthered our understanding of the primary and premotor connectivity
patterns within the cerebellum, demonstrating that the more anterior
medial lobules I–IV are connected to the primary motor cortex, which is
largely expected given the somatotopic representations seen in this region
(
Grodd et al., 2001). Lobule V was connected with supplementary motor
regions, while lobule VI was associated with more prefrontal cortical
regions. However, across all networks both the medial prefrontal cortex
and cingulate were present, indicative of some degree of similarity across
networks as well (
Kipping et al., 2013). While these findings are broadly
consistent with other lobular work (Bernard et al., 2012;Sang et al.,
2012), the added detail related to cerebellar connectivity with cortical motor
regions represented a novel addition to the literature.
Subsequent seed-based investigations of cerebello-cortical resting-state
connectivity have been more targeted to better understand specific net-
works and cortical interactions. While Allen and colleagues first mapped
the resting-state connectivity networks from the dentate (
Allen et al.,
2005), they did so using a central seed encompassing much of the nucleus.
However, distinct patterns of connectivity have been dissociated between
the dorsal and ventral dentate in non-human primates (Dum & Strick,
2003). With this in mind, we attempted to determine whether these networks
can also be dissociated at rest in the human brain (Bernard et al., 2014).
30 Jessica A. Bernard

This approach again took advantage of the cerebellar specific methods we
used to look at the cerebellar lobules (Bernard et al., 2012), but also used
smaller seeds, and the resting-state signal in the cerebellum was unsmoothed
to prevent signal from outside the dentate from mixing (
Bernard et al.,
2014). In this healthy young adult sample, we were indeed able to dissociate
connectivity from the two seeds. The dorsal dentate was more strongly
associated with motor and pre-motor regions in the cortex, while the
ventral dentate showed correlations atrest with prefrontal and association
cortices (Bernardetal.,2014). This initial investigation was conducted in a
relatively small sample of healthy young adults (n¼39) (Bernard et al.,
2014). However, we recently replicated this dissociation in a much larger
sample (n¼590) (Bernardetal.,2021), using data from the Cambridge
Center for Aging and Neuroscience (Cam-CAN) (Shafto et al., 2014;
Taylor et al., 2017). The same robust patterns for the dorsal and ventral
dentatewererevealed,butthistimeinasamplespanningtheadultlifespan
(ages 18–88) and with traditional analysisapproaches, further showing
that this is a robust and replicable dissociation at rest in the human brain
(Bernard et al., 2021). Independently of this work, Guell and colleagues
also looked at functional territoriesin the dentate, and in their analysis
identified a third subregion (Guell et al., 2020). These territories are
associated with the default network, salience and motor related regions,
and finally visual networks. While these networks do not map on to the
existing dissociations as clearly,this may be due to methodological
differences.
In a similar, more targeted fashion Brissenden and colleagues were
especially interested in understanding cerebellar contributions to the dorsal
attention network (
Brissenden, Levin, Osher, Halko, & Somers, 2016). In
this investigation, participants completed task-based fMRI using visual
working memory and visual attention. Seeds were then determined based
on the functional activation patterns and were placed in lobules VIIb and
VIIIa. The seed-based connectivity results demonstrated connectivity
between these regions that was consistent with the dorsal attention network,
as delineated in a parcellation by Yeo and colleagues (
Yeo et al., 2011). The
resting-state connectivity patterns also showed a great deal of similarity to
the task-based activation. Together, this work provided novel insights into
cerebellar contributions to visual attention and the underlying networks
at rest, which to this point had not been given much focal attention.
Further, this provides additional confirmation for the idea that many of
the networks revealed at rest are related to those involved in task processing
as measured by the BOLD signal during task performance.
31A review of resting-state connectivity of the cerebellum

Seed-based approaches have also provided novel insights into networks
with subcortical regions. As described earlier, non-human primate tract
tracing has provided evidence for reciprocal connections between the
cerebellum and basal ganglia (
Bostan et al., 2010;Bostan & Strick, 2018;
Hoshi et al., 2005), and diffusion imaging has provided evidence for these
connections in the human brain as well (Pelzer et al., 2013). Initial lobular
resting-state work indicated some degree of subcortical connectivity,
including with the basal ganglia (Bernard et al., 2012;Sang et al., 2012),
and work in aging has suggested that these connections at rest differ in older
adults relative to young adults (
Bernard et al., 2013). Using a targeted
seed-based approach, we sought to map the connections between these
regions in young and older adults. Most notably, with seeds placed in the
cerebellum, as well as those in the basal ganglia (based off of prior work
on basal ganglia connectivity;
Di Martino et al., 2008), we demonstrated
robust connectivity between the cerebellum and basal ganglia (Hausman,
Jackson, Goen, & Bernard, 2020). We also found that these were topograph-
ically specific, wherein areas of the cerebellum associated with motor
function and cortical motor areas were associated with parallel regions
in the basal ganglia. There was a similar topographical segregation for
cognitive/associative regions as well (
Hausman et al., 2020). Notably, these
patterns changed in advanced age, such that there were anti-correlations
between the cerebellum and basal ganglia, and the topographical distinction
of the circuits was less robust (
Hausman et al., 2020). Thus, we were able to
use resting-state connectivity to identify new connections at rest, and due
to the approach, we further showed some degree of functional specificity,
perhaps allowing for domain specific coordinated control of behavior in
these two subcortical regions.
Finally, using a large data set from the Human Connectome Project
(HCP), Guell and colleague used task-based activity to define seeds for use
in resting-state connectivity analysis (
Guell, Gabrieli, & Schmahmann,
2018). Working memory, language, social, and emotional processing tasks
were used, and in their resting-state analysis they found distinct resting-state
network representations for each. In many ways this was expected, given
prior work demonstrating a topographical organization in the cerebellum
(
Stoodley & Schmahmann, 2009;Stoodley et al., 2012). What was unex-
pected, however, and especially novel was that these representations were
in triplicate, such that they were repeated within the cerebellum three
times. This triple representation has potential implications for our thinking
in terms of cerebellar function, and the authors suggest that this may be a
32 Jessica A. Bernard

result of the cellular organization of the cerebellum (Guell, Gabrieli, &
Schmahmann, 2018). However, it is thanks to this resting-state analysis
that there are exciting new directions for thinking about cerebellar function,
and this redundancy in representations may lead to new work, and in turn a
better understanding of cerebellar function and processing in both motor
and non-motor behaviors.
4.2 ICA and parcellations
While seed-based approaches are often driven by questions rooted in
anatomy or existing functional networks, data-driven approaches are also
highly informative. In parallel to the earlier seed-based analyses of cerebel-
lar connectivity, both ICA and parcellation approaches were used to better
understand cerebellar functional organization and networks. As another
methodological approach to investigating parallel cerebello-thalamo-
cortical circuits in the human brain, Habas and colleagues used ICA to deter-
mine how different cerebellar regions are associated with known cortical
networks (
Habas et al., 2009). In this investigation, they found differential
associations with cortical networks. Crus I and II were associated with
the executive control network, lobule VI with the salience network, and
Lobule IX with the default mode network. They also demonstrated a dis-
sociation from the motor regions (lobules I–IV and V), and found overlap
in both the dentate and pontine nuclei, which are key output and input
regions for the cerebellum (
Habas et al., 2009). Thus, they found converging
evidence for dissociable motor and frontal/association networks in the
human cerebellum, and further support for the lateral posterior cerebellum
being involved with regions of the cortex associated with higher order
thought and cognitive processing.
More recently, Habas again took advantage of the ICA approach, this
time to further investigate associations between the cerebellum and amyg-
dala (
Habas, 2018). While some prior work has demonstrated connectivity
between these regions (Sang et al., 2012), cerebellar-amygdaloid connec-
tions are largely lesser-known, despite being of great interest particularly
in the context of psychiatric disorders. In this small exploratory study, they
reported the emergence of a cerebellar-amygdaloid network that included
the dentate nucleus, lobule VI, and lobules VIIIa and VIIIb (
Habas,
2018).Habas (2018)suggests that this may relate to sensory processing
and the processing of noxious stimuli. While the sample used was relatively
small (n¼15), this is a particularly novel finding that stands to further open
33A review of resting-state connectivity of the cerebellum

up research investigating cerebellar-amygdaloid interactions. Resting-state
work was a catalyst that allowed for this investigation, and suggests that
we should further expand our frameworks of cerebellar function and
processing to include these interactions with the amygdala.
ICA-based approaches have generally been used to look at existing
cortical networks, though they certainly have provided insights into cere-
bellar network connectivity in large part by linking the cerebellum to these
cortical networks. In the early 2010s, functional network parcellation
became increasingly popular as a new methodological approach to under-
standing network organization (
Power et al., 2011;Yeo et al., 2011).
Perhaps not surprisingly, this has also been applied to the cerebellum.
One of the first cerebellar functional parcellations was conducted in a
large sample (n¼1000) and sought to determine how a cortical network
parcellation (
Yeo et al., 2011) mapped on to the cerebellum (Buckner
et al., 2011). Yeo and colleagues had previously created network
parcellations of the cortex resulting in two network solutions with 7 and
17 networks, respectively (
Yeo et al., 2011). Using a split-half approach,
Buckner then mapped these parcellations to the cerebellum, and completed
a within sample replication. Both the 7 and 17 networks solution were
effectively mapped to the human cerebellum suggesting that the structure
contributes to these known cortical networks (
Buckner et al., 2011).
This work also revealed several key new findings about the functional orga-
nization of the cerebellum. First, the cerebellar contributions to cortical
networks are proportional to their size in the cortex. Second, they also
demonstrated that there are at least two mirrored maps of the cortical
networks in the cerebellum, with the potential for a third map of the
cortical network representations (
Buckner et al., 2011). This latter point
in particular is notable in the context of recent seed-based work described
above from Guell and colleagues (Guell, Gabrieli, & Schmahmann, 2018)
describing a triple representation. They provided converging evidence for
the idea of multiple cortical representations initially described by
Buckner
et al. (2011), which has possible implications for our understanding of
cerebellar function and processing. It is also notable that this work mapped
a precise motor somatotopy in the cerebellum, and demonstrated that
network boundaries do not follow lobular boundaries (
Buckner et al.,
2011). That the functional boundaries extend beyond cerebellar lobules is
an interesting consideration, particularly when thinking about the under-
lying structural circuits with the cortex and how those are mapped and
resolved.
34 Jessica A. Bernard

In our own work investigating lobular cerebellar connectivity, we also
incorporated a self-organizing map (SOM) approach to parcellate the
resting-state signal in the cerebellum (Bernard et al., 2012). Unlike the
approach used by Buckner and colleagues where the networks defined in
the cortex were mapped on to the cerebellum, here we focused solely on
the cerebellar signal and sought to see how different voxels organize and
group based on the resting BOLD (
Bernard et al., 2012). The SOM resulted
in 20 clusters, which is relatively on par with the 17 network parcellation
used by Buckner (
Buckner et al., 2011). In visually comparing the
parcellations there were certainly some areas of overlap, particularly with
respect to general motor areas grouping together in the anterior cerebellum
and lateral posterior groupings that were distinct. Again, like in the work
from
Buckner et al. (2011), these regions did not necessarily follow the ana-
tomical boundaries of the cerebellar lobules (Bernard et al., 2012). Finally,
the regions defined by the SOM were then used to conduct a seed-to-voxel
analysis with the whole-brain revealing their cortical network connections.
While not all of the cerebellar regions produced significant associations
with cortical voxels, many of these regions were correlated with signals in
frontal and association cortices (
Bernard et al., 2012), consistent with the
idea that the cerebellum has a large representation of networks and regions
with frontal and association cortices (
Buckner et al., 2011).
Subsequent parcellations have also been conducted looking at the cere-
bellum. Most notably, Wang and colleagues used sparse dictionary learning
clustering and they found what they deemed “biologically reasonable” func-
tional breakdowns within the human cerebellum (Wang, Kipping, Bao,
Ji, & Qiu, 2016). Finally, Seitzman and colleagues used a winner-take-all
approach to partition resting-state data to create regions of interest in the
cerebellum and subcortex that map on to existing cortical parcellations
(Seitzman et al., 2020). This work effectively extended cortical network
organization to the cerebellum in a way that can be effectively applied in
the future through careful validation across data sets. Together, ICA and
parcellation have further added to our understanding of the functional orga-
nization of the cerebellum, in a way that complements and extends what was
learned from seed-based approaches.
4.3 Graph theory
Generally, by their nature, graph theory approaches look more at whole
brain interactions. As such, while they have not necessarily been employed
35A review of resting-state connectivity of the cerebellum

to look specifically at the cerebellum, work using these methods has pro-
vided important novel insights with respect to the cerebellum and how it
interacts with the rest of the brain. Developmental work from Fair and col-
leagues (
Fair et al., 2009, 2007) has been particularly informative. First,
looking at the process of network development, with a particular focus
on networks for cognitive control (fronto-parietal and cingulo-opercular
networks) they demonstrated that connectivity of the cerebellum in these
networks emerged later in age (
Fair et al., 2007). They argue that the
cerebellar involvement in these networks is important for the processing
of feedback and errors, things that do not emerge until later in children.
These are “long-range connections” and emerge later over the course of
the development (
Fair et al., 2007). Second, they demonstrated that net-
works in children move from more local connections to more long-range
communication over the process of development (a shift from local to dis-
tributed processing) (
Fair et al., 2009). In this context, the cerebellum is
again later integrated in cortical networks, and generally speaking we see this
when comparing adults to children. Seemingly then, cerebellar network
development and integration with the networks of the whole brain takes
some time (
Fair et al., 2009, 2007), and this maturation pattern may suggest
that cerebellar processing itself has not yet matured, though alternatively, it
may just be that it has not yet been integrated with the cortex. With
this latter idea in mind, it may be the case that children are less effectively
using cerebellar processing to modify their behavior, because the networks
allowing for this have not developed.
Finally, recent work from Fitzgerald and colleagues revealed that
hormone variations across the menstrual cycle also impact cerebellar con-
nectivity (
Fitzgerald et al., 2020). Sensitivity of resting-state networks to
hormone levels across menstruation had been previously demonstrated in
the cortex (Pritschet et al., 2020), but this follow-up focused specifically
on the cerebellum. In brief, over a 28-day cycle graph metrics of cerebellar
connectivity were shown to be related to hormone levels wherein there
were associations with both estradiol and progesterone and global efficiency
in multiple networks (
Fitzgerald et al., 2020). This insight into hormonal
impacts on cerebellar connectivity metrics does not necessarily change
our understanding of cerebellar function or functional organization in that
the networks themselves do not change. However, it provides key new
insights into factors that may influence these networks and their behavior,
and suggest the importance of considering broader factors at play that
influence brain function and in turn behavior.
36 Jessica A. Bernard

4.4 Cerebellar connectivity in the era of big data
In the last decade with the creation of the HCP (Van Essen et al., 2013) and
coordinated data sharing efforts, a greater emphasis has been placed on “big
data” approaches. These approaches use large samples to better capture indi-
vidual variability and phenotypic variability that may relate to connectivity
measures. Notably, the work from Buckner and colleagues (2011) investigat-
ing cortical network parcellations in the cerebellum was very much in
this vein. Generally speaking, these large data sets allow for improved statis-
tical power and can reveal important factors influencing connectivity that
would not necessarily be measurable in smaller samples.
For instance, in our recent replication of dorsal and ventral dentate sub-
regions, we took advantage of the large Cam-CAN data set (
Bernard et al.,
2021). This allowed us to investigate this dissociation in a well-powered
sample providing a more robust replication, and we were able to investigate
sex differences in dentate subregion connectivity. This latter analysis
indeed revealed both sex differences, and age by sex interactions such that
the relationships between connectivity and age differ (
Bernard et al.,
2021). The large sample and statistical power allowed us to effectively inves-
tigate these questions, leading to a more nuanced understanding of factors
that impact cerebellar connectivity, in this case sex, consistent with prior
work (Biswal et al., 2010).
The large samples used by Guell and colleagues (2018) also may have
been instrumental in their ability to resolve the triple representation of
functional networks in the cerebellum. In this instance, they used HCP data,
and as such had a large sample and a high degree of statistical power. This
may have allowed for them to discover this novel triple representation
(
Guell, Gabrieli, & Schmahmann, 2018). Subsequently, they also used this
data set to investigate functional gradients within the cerebellum (Guell,
Schmahmann, Gabrieli, & Ghosh, 2018) and discovered two gradients
within the cerebellum. There is a primary gradient that moves from motor
to more association processing, and a secondary axis that relates to task focus
during processing (
Guell, Schmahmann, et al., 2018). Across these investi-
gations, and in conjunction with the work from Buckner and colleagues
(Buckner et al., 2011) there are clearly multiple representations and
functional hierarchies within the cerebellum that are key for processing.
In this era of big data, it is not just objectively large data sets with
hundreds or thousands of participants that are of interest. On the flip side,
there has also been a great deal of interest in “deep imaging” wherein a large
quantity of data (on the order of 45min to an hour or more) is collected from
37A review of resting-state connectivity of the cerebellum

a smaller number of participants. Though not the first to take this approach,
the Midnight Scan Club has done extensive work with deep imaging data
(Gordon et al., 2017) to complete “precision imaging” analyses on an indi-
vidual level. Though initially implemented to look at the cortex, this meth-
odology has also been applied to look at the cerebellum and subcortical
structures (reviewed in
Marek & Greene, 2021). Taking this approach to
cerebellar data at rest was highly informative (Marek et al., 2018). Using
a sample of 10 people with extensive data collection they investigated
cerebellar networks with the cortex and the resting-state signal. Much
like in the cortex, the data are specific to each individual though there
are fluctuations around what they refer to as a “central tendency.”
Further, though there is individual variability, similar patterns of organiza-
tion can be seen across the cerebellar networks of these participants (
Marek
et al., 2018). Ultimately, they demonstrated that there is a degree of
common organization across the group, but as compared to the cortex, there
is more individual variability. This later point is of particular importance
when investigating the cerebellum in clinical populations (
Marek et al.,
2018). Generally, they suggested that reliable individual analysis in the
cerebellum requires a great deal of data (more than 30min). Finally, one
of the most notable findings from this investigation was with respect to
the degree to which the cerebellum contributes to cortical networks. In
many cases, the cerebellar regions were fairly proportional to those in the
cortex, but the fronto-parietal control network was a notable exception.
The cerebellar representation for this network was larger than the motor
and pre-motor networks combined, and much larger than what would
be expected based on the size of its cortical representation. That is, it is
largely over-represented in the cerebellum (
Marek et al., 2018). This
over-representation certainly has implications for how we think about
and understand the role of the cerebellum more broadly.
5. Implications for understanding cerebellar function
It has been 16 years since the first resting-state paper focused on the
cerebellum was published in the literature (Allen et al., 2005), and this first
paper was followed by many as researchers attempted to improve their
understanding of cerebellar function and how this subcortical structure
interacts with the brain to contribute to motor and complex higher-order
processing. With this burst of research activity, and progress forward in
the last decade and a half, the question then becomes, what has this meant
for our understanding of cerebellar function and processing?
38 Jessica A. Bernard

The studies reviewed above have resulted in several key insights, some
of which likely would not have been revealed with task-based fMRI or
structural imaging alone. First, is the idea of multiple functional representa-
tions or gradients of function within the cerebellum (
Buckner et al., 2011;
Guell, Gabrieli, & Schmahmann, 2018;Guell, Schmahmann, et al., 2018).
The mirrored and repeated representation of networks of the cortex in
the cerebellum was possible due to resting-state approaches. It is in fact plau-
sible that this would not have been noticed with functional imaging alone, as
these representations were revealed at rest. Similarly, this methodology
was critical for revealing functional gradients (
Guell, Schmahmann, et al.,
2018). At this point, the task now is to parse and understand why this
organization exists, and how it can be integrated with the predominant
functional models based upon efference copies and internal models. Are
multiple efference copies being shared from different cortical regions that
work in concert during task performance, resulting in processing of those
copies in different, parallel cerebellar regions? That is, human behavior is
inherently complex. Any given task could include multiple subcomponents
of processing that occurs in parallel, and instead of just one efference copy
associated with a cognition for the task in its entirety, multiple smaller copies
may be shared that relate to diffuse processing across multiple cortical
regions. This could result in multiple functional representations revealed
in the resting state. This is of course speculative, but as we seek to understand
how the cerebellum functions during task performance, particularly for
complex cognition, an interesting area of research in the future.
Second, resting-state connectivity has suggested that a substantial portion
of the cerebellum is linked with networks that relate to cognitive and asso-
ciative cortices (
Bernard et al., 2012;Buckner et al., 2011;Marek et al.,
2018;Sang et al., 2012). While it had long been recognized that there are
closed-loop circuits linking the cerebellum to the cortex, these were rela-
tively proscribed and limited in their extent (
Clower et al., 2001;Kelly &
Strick, 2003), likely due to methodological challenges. Similarly, human
diffusion imaging work followed this relatively proscribed approach
mapping dissociable tracts linking the cerebellum to the motor and prefron-
tal cortices (
Salmi, Pallesen, & Neuvonen, 2010), but more detailed tracts
have yet to be resolved. In this case, the methodological challenges associ-
ated with many crossing fibers and dense white matter certainly may be a
hindrance. But, resting state has been especially useful. Thanks to the growth
of work in this regard, the extent of cerebellar interactions with non-motor
regions in the cortex can be more fully appreciated. It is clear based on fMRI
that there are areas of activation in the cerebellum spanning cognitive
39A review of resting-state connectivity of the cerebellum

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But, in the meantime, affairs had been becoming every day more
and more interesting and critical. On the one hand the sympathy for
the Greeks had been increased by the unexpected resolution they
had displayed; they had a loan, a government, and able and
enterprising foreigners had entered into their service. So much was
encouraging for their cause. But on the other hand the Egyptian
army of Ibrahim Pasha had achieved cruel triumphs, and a great
part of the Morea, devastated and depopulated, had submitted to his
arms.
During these events the Czar Alexander died; and for some little
time there was hesitation in the Imperial counsels. Alexander’s
successor, however, soon pursued the policy which his accession to
the empire had interrupted, and propositions (not unlike those
formerly contemplated) were now submitted to our Minister,
propositions in the carrying out of which Great Britain and Russia
were alone to be combined. The circumstances of the moment
showed that the period of action had arrived, and Mr. Canning no
longer shrank from accepting a part which there appeared some
hope of undertaking with success.
An alliance between two powers, indeed, afforded a fairer chance
of fixing upon a definite course, and maintaining a common
understanding, than the various counsels amongst which union had
previously been sought. The Greeks also, who had formerly rejected
all schemes of compromise (May, 1826), now requested the good
offices of England for obtaining a peace upon conditions which
would have recognised the supremacy of the Sultan, and entailed a
tribute upon his former subjects. Finally (and this affords an
interpretation to the whole of that policy which prevailed in the
British counsels, from the first to the last moment of negotiation),
the treaty of alliance into which Mr. Canning felt disposed to enter,
contained this condition:
“That neither Russia nor Great Britain should obtain any
advantage for themselves in the arrangement of those affairs which
they undertook to settle.”

France became subsequently a party to this scheme of
intervention, and it was hoped that a confederacy so powerful would
induce the Turks to submit quietly to the measures which it had
been determined, at all events (by a secret article), if necessary, to
enforce.
But whilst these projects were being carried out, these hopes
entertained, that dread King, more potent than all others, held his
hand uplifted over the head of the triumphant and still ardent
statesman.
XI.
On the 2nd of July Parliament had been prorogued; on the 6th the
triple alliance was signed. This celebrated treaty was the last act of
Mr. Canning’s official life. The fatigues of the session, short as it had
been, had brought him near the goal to which the enterprising mind
and assiduous labours of our most eminent men have too often
prematurely conducted them. Of a susceptibility which the slightest
word of good or evil keenly affected, and of that sanguine and
untiring temperament which would never suffer him to repose during
circumstances in which he thought his personal honour, his public
opinions, and the welfare of his political friends required his
exertions: tortured by every sneer, irritated by every affront, ready
for every toil; in the last few months in which he had risen to the
heights of power and ambition—such are human objects—was
concentrated an age of anxiety, suffering, and endurance. His
countenance became more haggard, his step more feeble, and his
eye more languid. Yet at this moment, jaded, restless, and worn, he
held in the opinion of the world as high and enviable a position as
any public man ever enjoyed. All his plans had succeeded; all his
enemies had been overthrown. By the people of England he was
cherished as a favourite child; on the Continent he was beloved as
the tutelary guardian of Liberal principles, and respected as the
peaceful and fortunate arbiter between conflicting interests. Abroad,
one of the most formidable alliances ever united against England

had been silently defeated by his efforts. At home, the most
powerful coalition that a haughty aristocracy could form against
himself had been successfully defied by his eloquence and good
fortune. The foes of Don Miguel, in Portugal; the enemies of the
Inquisition in Spain; the fervent watchers after that dawn of
civilization, which now opened on the vast empires of the New
World, and which promised again to shine upon the region it most
favoured in ancient times; the American patriot, the Greek
freedman, and last of all, though not the least interested (whether
we consider the wrongs he had endured, the rights to which he was
justly born, the links which should have joined him to, and the
injustice which had severed him from, the national prosperity of
Great Britain), last of all, the Irish Catholic, dwelt fondly and
anxiously on the breath of the aspiring statesman at the head of
affairs. His health was too precious, indeed, for any one to believe it
to be in danger.
The wound, notwithstanding, was given, which no medicine had
the power to cure. On the 1st of August the Prime Minister gave a
diplomatic dinner; on the 3rd he was seized with those symptoms
which betokened a fatal crisis to be at hand. At this time he was at
the Duke of Devonshire’s villa at Chiswick, where he had resided
since the 20th of July, for the sake of greater quiet and purer air.
The room in which he lay, and in which another as proud and
generous a spirit, that of Mr. Fox, had passed away, and towards
which the eyes of the whole Liberal world were now turned with
agonizing suspense for five days, has since become a place of
pilgrimage. It is a small low chamber, once a kind of nursery, dark,
and opening into a wing of the building, which gives it the
appearance of looking into a courtyard. Nothing can be more simple
than its furniture or decorations, for it was chosen by Mr. Canning,
who had always the greatest horror of cold, on account of its
warmth. On one side of the fireplace are a few bookshelves;
opposite the foot of the bed is the low chimneypiece, and on it a
small bronze clock, to which we may fancy the weary and impatient
sufferer often turning his eyes during those bitter moments in which

he was passing from the world which he had filled with his name,
and was governing with his projects. What a place for repeating
those simple and touching lines of Dyer:
“A little rule, a little sway,
A sunbeam on a winter’s day,
Is all the proud and mighty have
Between the cradle and the grave.”
After passing some time in a state of insensibility, during which
the words “Spain and Portugal” were frequently on his lips, on the
8th of August Mr. Canning succumbed. His remains sleep in
Westminster Abbey; a peerage and a pension were granted to his
family; and a statue is erected to his memory on the site of his
parliamentary triumphs.
The generation amidst which Mr. Canning died, attended his
hearse, and crowned his funeral with honours. What is the place he
ought to hold in the minds of future generations of his countrymen?
Part V.
One must judge men by a real and not ideal standard of mankind.—Criticisms
on Mr. Canning’s conduct.—His faults when in a subordinate position.—His better
qualities developed in a superior one.—Nature of faculties.—Influence on his own
time and the succeeding one.—Foreign policy considered.—Person; manners;
specimens of his various abilities; eloquence; art; and turn for drollery and satire.
—Style of speaking of despatches.—Always young, and inspiring admiration and
affection, even when provoking censure.
I.

In estimating the character of public men, the biographer or critic,
if he descend from the sublimity of unbounded panegyric, is often
apt to elevate himself at the expense of the person of whom he
speaks; and to treat with artificial severity any dereliction from that
perfection of conduct which he sees nowhere attained. Thanks to
this affected severity or paltry envy, we have hardly a great man left
to us. Bolingbroke is nothing but a quack; the elder Pitt only a
charlatan; Burke himself a declaimer and a renegade; Fox an
ambitious politician out of place; all of which things these great men
to a certain degree were, being still great men; and deserving the
admiration of a posterity which can hardly hope to furnish their
equals.
“No one should write history,” said Montaigne, “who has not
himself served the State in some civil or military capacity.” By which
this shrewd and impartial observer meant, that no man is fit to judge
the conduct of men of action who is not himself a man of action,
and can judge it practically, according to what men really are in the
world, and not according to any imaginary theory which he may
adopt in the obscure nook of his own chimney corner, as to what
they might and ought to be.
“We are not,” says Cicero, “in the Republic of Plato, but in the mud
of Romulus;” and they who have observed and meditated upon the
vicissitudes of empires, will have seen that such have risen or fallen
according to the number of eminent men, endowed with lofty
intelligences and daring spirits, whom they have produced. And
where have such eminent men existed without defects? Human
nature is too imperfect for us to expect to find extraordinary abilities
and energies under the constant control of moderate virtues.
To those, then, who have read the preceding pages, the whole of
Mr. Canning’s career may be shortly summed up in the words of Lord
Orford (Horace Walpole), who, speaking of Lord Chatham, says:
“His ambition was to be the most illustrious man in the first
country in the world, and he thought that the eminence of glory

could not be sullied by the steps to it being passed irregularly” (vol.
iv. p. 243).
In the same manner Canning was less scrupulous than he should
have been to obtain power and fame. But, in the most memorable
part of his life, he made a noble use of the one and well deserved
the other. Desirous of office and distinction, he attached himself, on
entering life, to that minister by whom office and distinction were
most likely to be conferred. The circumstances of the time afforded
him not merely an apology, but a fair reason for doing this; still,
there seems no injustice in adding that, in ranging himself under the
banner of the great commoner’s great son, he thought of his own
personal prospects as well as of the public interests.
Mr. Pitt died; Mr. Canning was, as he declared himself, henceforth
without a leader. Some of his opinions inclined him to unite with his
early friends and recent opponents (the Whigs), who then came into
office; and this, it seems, he was on the point of doing, when, by a
sudden whirl of Fortune’s wheel, the persons he was seceding from
were jerked into power, and those he was about to join jerked out of
it. A young man, conscious of his own abilities, and satisfied in his
own mind that, however he might obtain influence, he would use it
for the public advantage, he did not refuse a high situation from the
party to which he still publicly belonged, in order to follow a party
just driven from the Administration, and with which he had but
begun to treat.
There are things to say in excuse of this conduct, and I have said
them; but no one who wishes that Mr. Canning’s life had been
without a flaw, can do otherwise than regret that the statesman who
made so many subsequent sacrifices for the Catholics, should have
joined, at this juncture, a Ministry which rallied its partisans under
the cry of “No Popery!”
It is likewise to be regretted that having so frequently expressed
his sense of the incapacity of Lord Castlereagh, he should
nevertheless have consented first to serve as a subordinate under
him when he was mismanaging foreign affairs; and, secondly, to

serve as a colleague with him when he was alike lowering us abroad
and misgoverning us at home.
During four years he did not shrink from the promulgation of any
arbitrary edict—from the suppression of any popular right; and
though I admit that many liberal and prudent persons (influenced, I
cannot but think, by most exaggerated apprehensions) considered
that the strongest measures were necessary at that time to control a
spirit of insurrection, which the mingled harshness and incapacity of
the ruling Administration had provoked; still, there is a great
difference between men who sanction bad laws which a bad
government, in which they have had no share, may render
momentarily necessary, and men who bring forward bad laws as the
result of a bad government which has been carried on by
themselves.
It is hardly an excuse to say his errors were committed in an
inferior situation, with the idea of rising to a commanding one; but,
at all events, when he reached the eminence towards which he had
so long been toiling, he made, as I have shown, the best use of that
power which had not always been sought for by the best means.
Thus, from first to last, we see a man anxious to have power and to
use it well; but as anxious to have it as to use it well. That he was
blamed and praised with exaggeration was natural; for amidst
confronting arrays he was seen for ever in the first rank with the
most glittering arms, exciting the admiration of friends and the
hatred of foes by his scornful air and ostentatious attitude of
defiance.
His talents, by nature showy, were given their peculiar turn by his
early education, and his career was shaped to the paths which
offered to lead him most easily to distinction. Trained to the juvenile
task of writing a foreign language in polished periods, he was at
times less anxious to find solid arguments than striking expressions.
Not brought up in communication with the uneducated classes, he
was more keenly alive to the opinion of the cultivated and refined.
Too accommodating as to the temporary suspension of national

freedom at home, he was constantly anxious and determined to
maintain the power and prestige of the country abroad—throughout
his whole life he exhibited the effects of the public school and the
close borough.
Like most men who have become illustrious, Mr. Canning owed
much to fortune. Lucky in the time of his decease, lucky in the times
at which many of those with whom he had hitherto acted deserted
him. If he had lived longer, it would have been difficult for him to
have kept the station to which he had risen: if he had not been left
when he was by a great portion of his party, he would never have
obtained the popularity by which his death was hallowed. To few has
it happened to be supported by a set of men just as long as their
support was useful,—to be quitted by them just when their alliance
would have been injurious. The persons who as friends gave Mr.
Canning power, as enemies conferred on him reputation. That
reputation was above all others, at the time of his demise, amongst
his countrymen and contemporaries; and it still retains its
predominance, though the influence which he exercised over our
domestic policy, and over the events which succeeded his death, is
not yet, perhaps, sufficiently recognised. I have already observed
that if he had not been Prime Minister in 1827, it is not likely that
Lord Grey would have been Premier in 1830. I may add that had not
his appointment at the former period brought together all the
elements of a great Liberal party, who were allied under the cry of
Catholic Emancipation, thus giving a hope and a spirit to the
Catholics which they had not previously possessed, the Duke of
Wellington would not within a year or two afterwards have been
forced to acknowledge that further resistance to them was
impossible. Furthermore, if such men as Lord Melbourne, Lord
Palmerston, the Grants, and a large party in the country looking up
to these statesmen as safe as well as liberal guides—had not been
already connected with the Whigs, and alienated from the Tories,
under the influence of Mr. Canning in 1827, the Reform Bill would
hardly have been proposed in 1830, and would certainly not have
been carried in 1832. The more minutely, in short, that we examine

the events of the last thirty-six years, the more we shall perceive
how much their quiet development has been owing to Mr. Canning,
and to the class of men whom Mr. Canning formed, and in his later
days represented.
In determining his merits as director of the foreign policy of Great
Britain, I have stood, I confess, by the old doctrines, and argued
upon the assumption that England is a great state, disposed to
maintain that greatness; that the English people is a proud,
generous, and brave people, prepared to assert its principles and its
position, and to assume its part in the affairs of the world—a nation
that takes its share in the general policy of nations—that feels it has
a common interest in the maintenance of justice, in the limitation of
unscrupulous ambition, in the progress of civilization. I have
supposed that the collective wisdom and experience of past ages,
have taught us that human nature is ever, though under different
forms, guided by the same rules; that the strong, unless they are
adequately restrained, insult and oppress, and finally vanquish the
weak; that those who under all circumstances are determined to be
at peace, become eventually the certain victims of aggression and
war; that the spirit of a people cannot with impunity be allowed to
droop and languish without dimming the brightness of its genius and
losing the force of its character. That a mere money-making
population, which, lapped in the luxury of commercial prosperity,
begins to disregard its nice sense of honour, its admiration for valour
and daring, becomes daily weaker against the spoiler, and a greater
temptation to spoliation. I have ventured to believe that a noble
people has a heart open to noble emotions—that such a heart is not
dead to pity for the unfortunate, to sympathy with the brave—to the
love of glory inspiring to great deeds, and to the love of power, with
the intention to use it for the public good. I do not think it wise to
exchange the principles of action derived from these sentiments for
a colder, less generous, and, as I feel convinced, a less sound code
of political philosophy. The same sentiments which make one man
considered and beloved above others, must distinguish the State
aspiring to be great and beloved; but it does not follow that if you

feel compassion for a drowning man, you are to plunge into the sea
to save him if you cannot swim; that if you see two men valiantly
struggling against two regiments, you are to rush into the middle of
the combat with the certainty of not vanquishing the assailants, and
with that of losing your own life. I condemn nations that interfere
needlessly with the international affairs of others, as I should the
lady who pretended to dictate to her neighbour how she should have
her drawing-room swept, or her chimneys cleaned. I condemn
governments which threaten heedlessly, and then fail to strike in
spite of their threats; but I esteem governments which look carefully
after their honour and interests, and do interfere when it is
necessary or expedient to do so, in order either to defend that
honour, or to maintain those interests; governments cautious to
speak, but bold in acting up to their words.
It is with these views that I look upon the foreign policy of Mr.
Canning,—a policy for giving England a great and proud position,—
for giving to Englishmen a glorious and respected name; for
safeguarding our shores by the universal prestige of our bravery and
our power; for limiting the ambition of rival states, without
needlessly provoking their animosity; for showing a wish to conciliate
wherever moderation is displayed, and for displaying a resolution to
resist when conciliation is repulsed—as a great English policy, with
which the people of England will ever sympathize, and by which the
permanent interests of England will best be preserved.
There are men who are anxious for civil commotion, which they
think may be more easily brought about by concentrating the public
mind on domestic grievances; there are men who are indifferent to
the pride of country—who would as soon be Portuguese, Mexicans,
or Moldo-Wallachians, as Englishmen. There are men who, though
fame and consideration are the great objects of their countrymen,
hold they ought not to be objects for their country. These will
repudiate my opinion. But every Briton who is justly proud of his
race, who will inquire from a small and despised state the value of
being a great and renowned one, will, I believe, recognise the
foreign policy I have been describing to be the true policy for

maintaining the dignity and authority, without rashly risking the
peaceful prosperity, of the British empire.
In person Mr. Canning was favoured by nature, being of a good
height, of a strong frame, and of a regular and remarkably intelligent
countenance. The glance of his eye when excited, and the smile of
his lip when pleased, were often noted by his contemporaries.
“And on that turtle I saw a rider,
A goodly man, with an eye so merry,
I knew ’twas our foreign secretary,
Who there at his ease did sit and smile
Like Waterton on his crocodile;
Cracking such jokes, at every motion,
As made the turtle squeak with glee,
And own that they gave him a lively notion
Of what his own forced-meat balls would be.”
A Dream of a Turtle.—T. Moore.
Charming in manner, as I have said, constant in attachments, it
was observed of him at one period, that he was as dear to his
friends as odious to the public.
[125]
Ever ready to praise his subordinates, and to consult the tastes of
his associates, he was honoured as a chief as much as he was
relished as a companion. His accomplishments were various, and of
a kind which may leave disputes open as to the degree of their
excellence, but they were all of that brilliant and genial description
which was sure to attract sympathy and procure reputation. How
many must have chuckled over the following light and lazy piece of
satire:

“I am like Archimedes for science and skill,
I am like the young prince who went straight up the hill;
And to interest the hearts of the fair be it said,
I am like a young lady just bringing to bed.
If you ask why the eleventh of June I remember
So much better than April, or March, or December,
’Tis because on that day, as with pride I assure ye,
My sainted progenitor took to his brewery.
On that day in the month he began making beer;
On that night he commenced his connubial career.
On that day he died when he had finished his summing,
And the angels all cried ‘here’s old Whitbread a coming.’
So that day I still hail with a smile and a sigh,
For his beer with an e and his bier with an i;
And that day every year, in the hottest of weather,
The whole Whitbread family dine altogether.
My Lords, while the beams of the hall shall support
The roof which o’ershades this respectable court
(Where Hastings was tried for oppressing the Hindoos),
While the rays of the sun shall shine in these windows
My name shall shine bright as my ancestor’s shines,
Emblazoned on journals as his upon signs.”
How many must have felt their minds respond and their hearts
bound at the following argumentative and spirited declamation:
“When the elective franchise was conceded to the Catholics of
Ireland, that acknowledgment and anticipation, which I now call
upon the House formally to ratify and realize, was, in point of fact,
irrevocably pronounced. To give the latter the elective franchise was
to admit him to political power; for, to make him an elector and at
the same time to render him incapable of being elected, is to attract
to our sides the lowest orders of the community, at the same time
that we repel from us the highest orders of the gentry. This is not
the surest or safest way to bind Ireland to the rest of the Empire in
ties of affection. And what is there to prevent our union from being

wrought more closely? Is there any moral—is there any physical
obstacle? Opposuit natura? No such thing. We have already bridged
the channel! Ireland now sits with us in the Representative Assembly
of the Empire; and when she was allowed to come there, why was
she not also allowed to bring with her some of her Catholic children?
For many years, alas! we have been erecting a mound, not to assist
or improve the inclinations of Providence, but to thwart them. We
have raised it high above the waters, and it has stood there frowning
hostility and effecting a separation. In the course of time, however,
chance and design—the necessities of man and the sure workings of
nature—have conspired to break down this mighty structure, till
there remains of it only a narrow isthmus standing
‘between two kindred seas,
Which mounting view each other from afar,
And long to meet.’
What, then, shall be our conduct? Shall we attempt to repair the
breaches, and fortify the ruins? A hopeless and ungracious
undertaking! or shall we leave them to moulder away by time and
accident? a sure but distant and thankless consummation! Or shall
we not rather cut away at once the isthmus that remains, allow free
course to the current which our artificial impediments have
constructed, and float upon the mighty waters the ark of our
common constitution?”
And we are now to be told that this same man, so playful and
jocose, so ornamented and brilliant, was a close arguer, and
indefatigable in attendance at his office. But though always ready for
business, he would not scruple to introduce a piece of drollery into
the most serious affairs. For instance:
The embassy at the Hague is in earnest dispute with the King of
Holland; a despatch addressed to Sir Charles Bagot arrives—it is in
cypher. The most acute of the attachés set to work to discover the
meaning of this particular document; they produce a rhyme! they
are startled, thrown into confusion; set to work again, and produce

another rhyme. The important paper (and it was important) contains
something like the following doggrel:
“Dear Bagot, in commerce the fault of the Dutch
Is giving too little, and asking too much,
So since on this policy Mynheer seems bent,
We’ll clap on his vessels just 20 per cent.”
As a specimen of his more private and trivial pleasantries may be
mentioned his observation to, I believe, Lord Londonderry, who had
been telling a story of some Dutch picture he had seen, in which all
the animals of antediluvian times were issuing from Noah’s Ark,
“and,” said Lord Londonderry, “the elephant was last.” “That of
course,” said Mr. Canning; “he had been packing up his trunk.”
In his celebrated contest with Lord Lyndhurst (then Sir John
Copley), that noble lord having appeared in it with a speech
borrowed for the most part from a popular pamphlet, written by the
late Bishop of Exeter (then Doctor Philpotts), he was overthrown
amidst shouts of laughter, by the appropriate recollection of the old
song:
“‘Dear Tom, this brown jug that now foams with mild ale,
Out of which I now drink to sweet Nan of the Yale,’
Was once Toby Philpot.”
Again, who does not remember the celebrated sketch of Lord
Nugent
[126]
—who went out to join the Spanish patriots when their
cause was pretty well lost—a sketch which furnished Mr. Canning’s
most effective defence of the neutral policy he had adopted towards
Spain, during the French expedition.
“It was about the middle of last July that the heavy Falmouth
coach”—(here Mr. Canning was interrupted with loud and continued
laughter)—“that the heavy Falmouth coach was observed travelling
to its destination through the roads of Cornwall with more than its
wonted gravity (very loud laughter). The coach contained two inside

passengers—the one a fair lady of no inconsiderable dimensions, the
other a gentleman who was conveying the succour of his person to
the struggling patriots of Spain. I am further informed—and this
interesting fact, sir, can also be authenticated—that the heavy
Falmouth van (which honourable gentlemen, doubtless, are aware is
constructed for the conveyance of cumbrous articles) was laden,
upon the same memorable occasion, with a box of most portentous
magnitude. Now, sir, whether this box, like the flying chest of the
conjuror, possessed any supernatural properties of locomotion, is a
point which I confess I am quite unable to determine; but of this I
am most credibly informed—and I should hesitate long before I
stated it to the House, if the statement did not rest upon the most
unquestionable authority—that this extraordinary box contained a
full uniform of a Spanish general of cavalry, together with a helmet
of the most curious workmanship; a helmet, allow me to add,
scarcely inferior in size to the celebrated helmet in the castle of
Otranto (loud laughter). Though the idea of going to the relief of a
fortress, blockaded by sea and besieged by land, in a full suit of light
horseman’s equipments was, perhaps, not strongly consonant to
modern military operations, yet when the gentleman and his box
made their appearance, the Cortes, no doubt, were overwhelmed
with joy, and rubbed their hands with delight at the approach of the
long-promised aid. How the noble lord was received, or what effects
he operated on the councils of the Cortes by his arrival, I (Mr.
Canning) do not know. Things were at that juncture moving rapidly
to their final issue; and how far the noble lord conduced to the
termination by throwing his weight into the sinking scale of the
Cortes, is too nice a question for me just now to settle.”
[127]
Mr. Canning’s wit, it is true, was not unfrequently too long and too
laboured, and a happy combination of words would almost always
seduce him into an indiscretion. The alliteration of “revered and
ruptured,” as applied to the unfortunate Mr. Ogden, cost him more
abuse, and procured him for a time more unpopularity, than the
worst of his acts ever deserved. His description of the American navy
(in 1812) as “half a dozen fir-frigates, with bits of bunting flying at

their heads,” excited the American nation more than any actual
grievance, and caused in a great measure the bitterness of that
contest in which we were so insolent and so unsuccessful. His
propensity to jokes made him also many enemies in private life. The
late Duke of Bedford told a friend of mine that Mr. Canning, when
staying with a party at Lord Carrington’s (a few weeks after Lord C.
had been made a peer by Mr. Pitt), wrote in chalk, on the outside of
the hall-door, the following lines:—
“One Bobby Smith lives here,
Billy Pitt made him a peer,
And took the pen from behind his ear.”
This unnecessary impertinence, I have heard, Lord Carrington
never forgave.
In the art of speaking, our orator’s progress, like that of Pulteney,
Fox, and all our great parliamentary debaters, with the exception of
the two Pitts, Bolingbroke, and Lord Derby, was slow and gradual;
and though I have heard Lord Lansdowne (once known as Henry
Petty) observe that he considered Canning in his best days even
more effective than Fox or Pitt, he had at an earlier period been
often accused, by no mean judges, now of being wordy and tedious,
now of being rather elegant than argumentative. To time, practice, a
proud spirit, and a continually developing understanding, he owed
his triumph over these defects. Then it was that his eloquence
approached almost to perfection, as we consider the audience, half
lounging and sleepy, half serious and awake, to which it was
addressed. Quick, easy, and fluent, frequently passionate and
sarcastic, now brilliant and ornamented, then again light and playful;
or, if he wished it, clear, simple, and incisive; no speaker ever
combined a greater variety of qualities, though many have been
superior in each of the excellences which he possessed. Remarkable
as a general rule for the polish of his language (we have proof, even
to the last, of the pains he bestowed upon it), those who knew him
well assert that he would sometimes purposely frame his sentences

loosely and incorrectly, in order to avoid the appearance of
preparation. “Erat memoriæ nulla tamen meditationis suspicio.” His
action exhibiting when calm an union of grace and dignity, became,
as he warmed, unaffectedly fervent; and made natural by its vigour
and animation the florid language and figurative decorations in
which he rather too fondly indulged. His arguments were not placed
in that clear, logical form, which sometimes enchains, but more often
wearies, attention; neither did he use those solemn perorations by
which it is attempted to instil awe or terror into the mind. His was
rather the endeavour to charm the ear, to amuse the fancy, to excite
the feelings, to lead and fascinate the judgment; and in these
different attributes of his great art he succeeded in the highest
degree, insomuch that though he might be said to want depth and
sublimity, the faculties he possessed were elevated to such a pitch,
that at times he appeared both profound and sublime.
A great merit, which he finally possessed, was that of seizing and
speaking the general sense of the popular assembly he addressed.
Sir Robert Peel, his distinguished rival, told me one day, in speaking
of Mr. Canning as to this particular, that he would often before rising
in his place, make a sort of lounging tour of the House, listening to
the tone of the observations which the previous debates had excited,
so that at last, when he himself spoke, he seemed to a large part of
his audience to be merely giving a striking form to their own
thoughts.
Neither were his despatches, though not so elaborately perfect as
those of his successor (Lord Dudley), inferior to his orations;
possessing precision, spirit, and dignity, they remain what they were
justly called by no incompetent authority, “models and masterpieces
of diplomatic composition.”
[128]
There are critics who have said that there was something in his
character which tended to diminish our respect for his talents,
though it softened our censure for his defects. And it is true that the
same unstately love for wit—the same light facility for satire—the
same imprudent levity of conduct, that involuntarily lowered our

estimate of his graver abilities—involuntarily led us to excuse his
graver errors. We at one time blame the statesman for being too
much the child—at another we pardon the veteran politician in the
same humour in which we would forgive the spoiled and high-
spirited schoolboy.
Mr. Canning, indeed, was always young. The head of the sixth
form at Eton—squibbing “the doctor,” as Mr. Addington was called;
fighting with Lord Castlereagh; cutting jokes on Lord Nugent; flatly
contradicting Lord Brougham; swaggering over the Holy Alliance; he
was in perpetual personal quarrels—one of the reasons which
created for him so much personal interest during the whole of his
parliamentary career. Yet out of those quarrels he nearly always
came glorious and victorious—defying his enemies, cheered by his
friends—never sinking into an ordinary man,—though not a perfect
one.
No imaginative artist, fresh from studying his career, would sit
down to paint this minister with the broad and deep forehead—the
stern compressed lip—the deep, thoughtful, concentrated air of
Napoleon Bonaparte. As little would the idea of his eloquence or
ambition call to our recollection the swart and iron features—the
bold and haughty dignity of Strafford. We cannot fancy in his eye the
volume depth of Richelieu’s—the volcanic flash of Mirabeau’s—the
offended majesty of Chatham’s. Sketching him from our fancy, it
would be as a few still living remember him, with a visage rather
marked by humour and intelligence than by meditation or sternness;
with something of the petulant mingling in its expression with the
proud; with much of the playful overruling the profound. His nature,
in short, exhibited more of the genial fancy and the quick irritability
of the poet who captivates and inflames an audience, than of the
inflexible will of the dictator who puts his foot on a nation’s neck, or
of the fiery passions of the tribune who rouses a people against its
oppressors.
Still, Mr. Canning, such as he was, will remain one of the most
brilliant and striking personages in our historical annals. As a

statesman, the latter passages of his life cannot be too deeply
studied; as an orator, his speeches will always be models of their
kind; and as a man, there was something so graceful, so fascinating,
so spirited in his bearing, that even when we condemn his faults, we
cannot avoid feeling affection for his memory, and a sympathetic
admiration for his genius.

SIR ROBERT PEEL.
Part I.
Family.—Birth.—Formation of character.—Education at Harrow and Oxford.—
Entry into Parliament.—Line adopted there.—Style of speaking.—Becomes
Secretary of Colonies.—Secretary for Ireland.—Language on the Catholic question.
—Returned as member for the University of Oxford.—Resigned his post in Ireland.
I.
The family of the Peels belonged to the class of yeomanry, which
in England, from the earliest times, was well known and reputed,
forming a sort of intermediate link between the gentry and the
commonalty, as the gentry formed an intermediate link between the
great barons and the burghers or wealthy traders. The yeoman was
proud of belonging to the yeomanry, and if you traced back the
descent of a yeoman’s family, you found it frequently the issue of the
younger branch of some noble or gentle house. For some
generations this family of Peel had at its head men of industry and
energy, who were respected by their own class, and appeared to be
gradually rising into another. The grandfather of the great Sir Robert
inherited a small estate of about one hundred pounds a year, called
Peel’s Fold, which is still in the family. He received a fair education at
a grammar-school, and married (1747) into a gentleman’s family
(Haworth, of Lower Darwen).
Beginning life as a farmer of his little property, he undertook, at
the time that the cotton manufacture began to develop itself in
Lancashire, the business of trader and printer.
The original practice had been to send up the fabricated article to
Paris, where it was printed and sent back into this country for sale.

Mr. Peel started a calico printing manufactory, first in Lancashire and
afterwards in Staffordshire, and his success was the result of the
conviction—that “a man could always succeed if he only put his will
into the endeavour,” a maxim which he often repeated in his later
days, when as a stately old gentleman he walked with a long gold-
headed cane, and wore the clothes fashionable for moderate people
in the days of Dr. Johnson.
The first Sir Robert Peel was a third son. Enterprising and
ambitious, he left his father’s establishment, and became a junior
partner in a manufactory carried on at Bury by a relation, Mr.
Haworth, and his future father-in-law, Mr. Yates. His industry, his
genius, soon gave him the lead in the management of this business,
and made it prosperous. By perseverance, talent, economy, and
marrying a wealthy heiress—Miss Yates, the daughter of his senior
partner—he had amassed a considerable fortune at the age of forty.
He then began to turn his mind to politics, published a pamphlet
on the National Debt, made the acquaintance of Mr. Pitt, and got
returned to Parliament (1790) for Tamworth, where he had acquired
a landed property, which the rest of his life was passed in increasing.
He was a Church and King politician in that excitable time, and his
firm contributed no less than ten thousand pounds in 1797 to the
voluntary subscriptions for the support of the war. So wealthy and
loyal a personage was readily created a baronet in 1800.
His celebrated son was born in 1788, two years before he himself
entered public life, and on this son he at once fixed his hopes of
giving an historical lustre to the name which he had already invested
with credit and respectability.
II.
It was the age of great political passions, and of violent personal
political antipathies and partialities. The early elevation of Mr. Pitt
from the position of a briefless barrister to that of prime minister had
given a general idea to the fathers of young men of promise and

ability that their sons might become prime ministers too. The
wealthy and ambitious manufacturer soon determined, then, that his
boy, who was thought to give precocious proofs of talent, should
become First Lord of the Treasury. He did not merely bring him up to
take a distinguished part in politics, which might happen to be a high
position in opposition or office, he brought him up especially for a
high official position. It was to office, it was to power, that the boy
who was to be the politician was taught to aspire; and as the
impressions we acquire in early life settle so deeply and
imperceptibly into our minds as to become akin to instincts, so
politics became instinctively connected from childhood in the mind of
the future statesman with office; and he got into the habit of looking
at all questions in the point of view in which they are seen from an
official position; a circumstance which it is necessary to remember.
To say nothing of the anecdotes which are told in his family of the
early manifestations which Mr. Peel gave of more than ordinary
ability, he was not less distinguished at Harrow as a student for his
classical studies, than he was as a boy for the regularity of his
conduct. I remember that my tutor, Mark Drury, who, some years
previous to my becoming his pupil, had Peel in the same position,
preserved many of his exercises; and on one occasion brought some
of them down from a shelf, in order to show me with what terseness
and clearness my predecessor expressed himself, both in Latin and
English.
Lord Byron says: “Peel, the orator and statesman that was, or is,
or is to be, was my form-fellow, and we were both at the top of our
remove, in public school phrase. We were on good terms, but his
brother was my intimate friend. There were always great hopes of
Peel amongst us all, masters and scholars, and he has not
disappointed them. As a scholar, he was greatly my superior; as a
declaimer and actor, I was reckoned at least his equal; as a
schoolboy out of school, I was always in scrapes—he never.” This
character as a lad developed itself, without altering in after life.

At the University of Oxford the young man was the simple growth
of the Harrow boy. He read hard, and took a double first-class,
indicating the highest university proficiency both in classics and
mathematics. But it is remarkable that he studiously avoided
appearing the mere scholar: he shot, he boated, he dressed
carefully, and, without affecting the man of fashion, wished evidently
to be considered the man of the world.
As soon as he became of age, his father resolved to bring him into
Parliament, and did so, in 1809, by purchasing a seat for him at
Cashel.
III.
The great men of the Pittite day were passing away. The leading
men at the moment were Grey, Liverpool, Petty, Perceval, Tierney,
Whitbread, Romilly, Horner, Castlereagh, Canning: the genius of
Sheridan had still its momentary flashes; and Grattan, though rarely
heard, at times charmed and startled the House of Commons by his
peculiar manner and original eloquence.
Brougham, Palmerston, Robinson, were Peel’s contemporaries.
The Duke of Portland was prime minister; Perceval, the leader of the
House of Commons; Canning, minister of foreign affairs; and Lord
Castlereagh, secretary of war. But this ministry almost immediately
disappeared: the Duke of Portland resigning, Lord Castlereagh and
Canning quarrelling, and Mr. Perceval, as prime minister, having to
meet Parliament in 1810 with the disastrous expedition of Walcheren
on his shoulders. Young Peel, not quite twenty-two, was chosen for
seconding the address, and did so in a manner that at once drew
attention towards him. He was then acting as private secretary to
Lord Liverpool, who had become minister of war and the colonies.
The condition of the Government was but rickety: Lord Carnarvon
carried against it a motion for inquiry into the conduct and policy of
the expedition to the Scheldt; and, subsequently, it could only obtain
a vote of confidence by a majority of twenty-three, which, in the

days of close boroughs, was thought equivalent to a defeat. Peel
spoke in two or three debates, not ill, but not marvellously well;
there was, in fact, nothing remarkable in his style; and its fluency
and correctness were more calculated to strike at first than on
repetition. He never failed, however, being always in some degree
beyond mediocrity.
In the meantime his business qualities became more and more
appreciated; and it was not long before he was appointed to the
under-secretaryship of the colonies.
It was no doubt a great advantage to him that the government he
had joined wanted ability.
Mr. Perceval’s mediocrity, indeed, was repulsive to men of
comprehensive views; but, on the other hand, it was peculiarly
attractive to men of narrow-minded prejudices. The dominant
prejudice of this last class—always a considerable one—was at this
time an anti-Catholic one; some denouncing Romanists as the pupils
of the devil, others considering it sufficient to say they were the
subjects of the Pope. Mr. Peel joined this party, which had amongst it
some statesmen who, sharing neither the bigotry nor the folly of the
subalterns in their ranks, thought, nevertheless, that it would be
impossible to satisfy the Catholics in Ireland without dissatisfying the
Protestants in England, and were therefore against adding to the
strength of a body which they did not expect to content.
IV.
Mr. Perceval’s unexpected death was a great blow to the anti-
Catholics, and appeared likely to lead to the construction of a new
and more liberal Cabinet. The general feeling, indeed, was in favour
of a Cabinet in which the eminent men of all parties might be
combined; and a vote in favour of an address to the Regent, praying
him to take such measures as were most likely to lead to the
formation of a strong administration, passed the House of Commons.

But it may almost be said that eminent men are natural enemies,
who can rarely be united in the same Cabinet, and are pretty sure to
destroy or nullify each other when they are. The attempt at such an
union was, at all events, on this occasion a signal failure.
Thus, luckily for the early advancement of Mr. Peel, Lord Liverpool
had to construct a government as best he could out of his own
adherents, and the under-secretary of the colonies rose at once to
the important position of Secretary for Ireland, to which the Duke of
Richmond, a man more remarkable for his joviality than his ability,
and a strenuous anti-Catholic, was sent as Lord Lieutenant.
V.
The Catholic question was to be considered an open one in the
new Cabinet, but the Irish Government, as I have shown, was
altogether anti-Catholic. This was in fact the strong bias of the
administration, and also of the Prince Regent, who, regardless of
former promises and pledges, had now become an avowed
opponent of the Catholic claims. These claims, moreover, were
strongly opposed by the feelings, at that time greatly excited, of the
English clergy, and, speaking generally, of the English people.
Under such circumstances, a Catholic policy was at the moment
impracticable; that is, it could not be carried out: for to carry out a
policy opposed by the sovereign, opposed by the premier (who had
been selected because his most able opponents could not form a
Cabinet), opposed by the English clergy, opposed by the general
sentiment of the English people, was impracticable, whatever might
be said theoretically in its favour.
Mr. Peel then, in taking up the anti-Catholic policy, took up the
practical one.
The Catholics themselves, indeed, destroyed for a while all hope in
their cause, for when the most considerable of their supporters, in
order to dissipate the alarm of their co-religionists, proposed certain
guarantees for maintaining the authority of the King and the State

over the Catholic priesthood, although the English Catholics and the
highest orders of Catholics in Ireland willingly agreed to these
guarantees, the more violent of the Irish Catholics, with Mr.
O’Connell at their head, joining the most violent anti-Catholics,
vehemently opposed them. Moderate people were, therefore,
crushed by the extremes. Even Grattan was for a moment put on
one side.
This was unfortunate for Mr. Peel, who would willingly have been
as moderate as his situation would permit him, but could only at
such a crisis live with violent people, and thus obtained the
nickname of “Orange Peel,” so that after different altercations with
Mr. O’Connell—altercations which nearly ended in a duel—he found
himself, almost in his own despite, regarded by both Protestants and
Catholics as the great Protestant champion.
It was in this position that he made, in 1817 (on an unsuccessful
motion of Mr. Grattan’s), a very remarkable speech, the success of
which Sir James Mackintosh attributes to its delivery.
“Peel,” he says, “made a speech of little merit, but elegantly and
clearly expressed, and so well delivered as to be applauded to
excess. He now fills the important place of spokesman to the
intolerant faction.”
The speech, however, had other merits than those Sir James
acknowledged, and I quote a passage which subsequently formed
the groundwork of all Mr. Peel’s anti-Catholic speeches.
“If you give them” (the Catholics) “that fair proportion of national
power to which their numbers, wealth, talents, and education will
entitle them, can you believe that they will or can remain contented
with the limits which you assign to them? Do you think that when
they constitute, as they must do, not this year or next, but in the
natural, and therefore certain order of things, by far the most
powerful body in Ireland—the body most controlling and directing
the government of it; do you think, I say, that they will view with
satisfaction the state of your church or their own? Do you think that

if they are constituted like other men, if they have organs, senses,
affections, passions, like ourselves; if they are, as no doubt they are,
sincere and zealous professors of that religious faith to which they
belong; if they believe your intrusive church to have usurped the
temporalities which it possesses; do you think that they will not
aspire to the re-establishment of their own church in all its ancient
splendour? Is it not natural that they should? If I argue from my
own feelings, if I place myself in their situation, I answer that it is.
May I not then, without throwing any calumnious imputations upon
any Roman Catholics, without proclaiming (and grossly should I
injure them if I did) such men as Lord Fingal or Lord Gormanston to
be disaffected and disloyal, may I not, arguing from the motives by
which men are actuated, from the feeling which nature inspires, may
I not question the policy of admitting those who must have views
hostile to the religious establishments of the State to the capacity of
legislating for the interests of those establishments, and the power
of directing the Government, of which those establishments form so
essential a part?”
VI.
Have we not seen that every word I have been quoting is
practically true? Are we not beginning to acknowledge the difficulty
of maintaining a Protestant Church establishment in Ireland in the
face of a large majority of Irish Catholic representatives? Are we not
beginning to question the possibility of upholding an exclusive
church belonging to a minority, without a government in which that
minority dominates? Do we not now acknowledge the glaring
sophistry of those who contended that the Catholics having once
obtained their civil equality would submit with gratitude to religious
inferiority? Mr. Peel saw and stated the case pretty clearly as it
stood; the whole condition of Ireland, as between Catholic and
Protestant, was involved in the question of Catholic emancipation,
and as the avowed champion of Protestant ascendancy, he said, “do
not resign your outworks as long as you can maintain them, if you

have any serious design to keep your citadel.” But the very nature of
his argument showed in the clearest manner that we were ruling
against the wishes and interests of the large majority of the Irish
people; that we were endeavouring to maintain an artificial state of
things in Ireland which was not the natural growth of Irish society;—
a state of things only to be maintained by force, and which, the day
that we were unable or unwilling to use that force, tumbled naturally
to pieces. It is well to bear this in mind.
The anti-Catholic party, however, accepted Mr. Peel’s argument;
they did not pretend to say that they governed by justice; and they
applauded their orator for showing that, whenever there was an
attempt to govern justly, as between man and man, and not
unjustly, as between Protestant and Catholic, their cause would be
lost.
His reward was the one he most valued. Mr. Abbott, then Speaker,
represented the University of Oxford. Mr. Abbott was made a peer,
and Mr. Peel, through the interest of Lord Eldon and of the party that
Sir James Mackintosh calls the intolerant one, was elected in his
place, in spite of the well-known and favourite ambition of Mr.
Canning.
With this result of his Irish administration Mr. Peel was satisfied.
All the duties attached to his place he had regularly and punctually
fulfilled. His life had been steady and decorous in a country where
steadiness and decorum were peculiarly meritorious because they
were not especially demanded. In all matters where administrative
talents were requisite he had displayed them: the police, still called
“Peelers,” were his invention. He protected all plans for education,
except those which, by removing religious inequalities and
animosities, and infusing peace into a discordant society, would have
furnished the best; and with a reputation increasing yearly in weight
and consideration, resigned his post, and escaped from a scene, the
irrational and outrageous contentions of which were out of harmony
with his character.

Part II.
Currency.—Views thereupon.—Chairman in 1859 of Finance Committee.—
Conduct as to the Queen’s trial.—Becomes Home Secretary.—Improvement of
police, criminal law, prisons, &c.—Defends Lord Eldon, but guards himself against
being thought to share his political tendencies, and declares himself in favour in
Ireland of a general system of education for all religions, and denounces any
attempt to mix up conversion with it.—Begins to doubt about the possibility of
resisting the Catholic claims.—The Duke of York dies, and Lord Liverpool soon after
follows.—Question of Premiership between the Duke of Wellington and Mr.
Canning.—Peel sides with the Duke of Wellington.
I.
The great practical question at issue, on Mr. Peel’s return from
Ireland, was the currency.
The Bank, in 1797, declared, with the consent of the Government,
that its notes would not be converted, on presentation, into gold.
At the time this was, perhaps, a necessary measure. It enabled
the Bank to make large advances to the State, which it could not
have made otherwise, and without which the Government would
have found it difficult to maintain the struggle of life and death it
was engaged in. We did, in fact, in our foreign war, what the United
States lately did in their domestic war; but the commercial
consequences of such a measure were inevitable.
If the Bank gave a note convertible into gold on presentation it
gave gold: if it gave paper, which simply specified the obligation to
pay gold for it some day or other, the value of the note depended on
the credit attached to the promise. The promise to do a thing is
never entirely equivalent to doing it; consequently, it was utterly
impossible that a bank-note, not immediately convertible into gold,
could have precisely the same value as gold. Gold, therefore, would
have a value of its own, and a bank-note a value of its own.

Moreover, as the value of the bank-note depended on the faith
placed in it, if it had been merely required for home trade, the
decrease in value would have been small; because the English
people had confidence in the Bank of England and in the
Government which sustained it; but in all foreign transactions the
case was different. If an English merchant had to purchase goods on
the Continent and he sent out bank-notes, the merchant at St.
Petersburg would have less confidence in the English bank-note than
the Manchester merchant, and he would therefore say, “No, pay me
in gold; or if you want to pay me in bank-notes, I will only take them
at the value I place on them.” In proportion, therefore, to the extent
of purchases abroad was the natural abasement of paper money at
home, and the increase in the value of gold as compared with paper.
Besides, paper money, resting on credit, partook of the nature of the
public funds, depending also on credit. As the one fell naturally, in a
long and critical war, so the other fell from the same cause, though
not in the same degree; all our dealings were thus carried on in a
money which had one real value and one nominal one; and the real
value depending, in a great measure, on matters beyond our control.
Efforts on the part of our legislature to sustain it were useless. We
forbade persons giving more for a guinea than twenty-one shillings
in paper money, and we forbade persons exchanging a twenty-
shilling bank-note for less than twenty shillings. We tried, in short, to
prevent gold and silver getting the same price in England that they
could get out of it.
The inevitable consequence was, that the precious metals, in spite
of stupid prohibitions against their exportation, went to those
countries in which it could obtain its real value. In this manner there
was, first, the transmission of coin for the maintenance of our
armies; secondly, its exportation for the purposes of our commerce;
and, lastly, its escape from the laws which deteriorated its value, all
operating to drain England of its gold and silver; and in proportion as
they became scarcer, their comparative value with paper increased,
insomuch that fifteen shillings in coin became at last equivalent to
twenty shillings in paper bank-notes.

Much was said as to the over-issue of bank-notes. It may always
be taken for granted that where there is an inconvertible paper,
there is an over-issue of bank-notes; because the over facility of
having or making money will naturally tend to the over-advance of it.
But we must remember, that a currency must be in proportion to the
transactions which require it; that our trade increased almost, if not
quite, in proportion to the increased issue from the Bank; that the
absence of coin necessitated a large employ of paper, and that there
did not appear to be that multitude of bubble schemes which are the
usual concomitants of a superabundant circulation. There were, in
fact, quite sufficient reasons, without attributing indiscretion to the
Bank, to account for the difference between its paper and the coin it
was said to represent; nor is there any possibility of keeping paper
money on an equality with metallic money, except by making the
one immediately exchangeable for the other.
The inequality, then, between paper money and metallic money
could only be remedied by re-establishing that immediate exchange.
But this was not an easy matter.
II.
For many years in England every transaction had been carried on
in paper. Individuals had borrowed money in it, and had received
this money in bank-notes. If they were called upon to repay it in
gold, they paid twenty-five per cent. beyond the capital they had
received. On the other hand, if individuals had purchased annuities,
the seller, whether the Government or an individual, had to pay
them twenty-five per cent. more than they had purchased.
The resumption of cash payments, therefore, could not take place
without great individual hardship and great public loss. There can be
no doubt, also, that paper money afforded great facilities for trade;
and that the sudden withdrawal of these facilities might be felt
throughout every class of the population.

Thus, although Mr. Horner brought the subject before the House
of Commons with great ability in 1811, it was not till 1819, when the
war had ceased, and the public mind in general had been gradually
prepared for terminating a situation which could not be indefinitely
prolonged, that the ministers intimated their intention to deal with it
by the appointment of a select committee, of which Mr. Peel was
named the chairman.
Up to this period, it is to be observed, the resumption of cash
payments could not have been carried; and up to this period Mr. Peel
and his father, who both voted against Mr. Horner, had opposed the
resumption. But the question was probably now ripe, so to speak,
for being dealt with. It was a matter, therefore, of practical
consideration, and Mr. Peel reconsidered it; and on the 20th of May
it was curious to see the venerable Sir Robert representing the ideas
of his time, and coming forward with a petition in favour of paper
money; and his son, the offspring of another epoch, rising, after the
father had sat down, to propose a measure by which paper money (I
speak of paper money not immediately convertible into gold) was to
be abolished; and avowing, as he said, “without shame and
remorse,” a thorough change of opinion.
His proposals compelled the Government to repay the sums which
it owed to the Bank, and compelled the Bank to resume cash
payments at a date which the Bank anticipated by resuming them in
1821.
Of the necessity of these measures there can be no doubt; at the
same time they were calculated, as I have said, to produce
momentary discontent and distress, and already much discontent
and distress existed.
There was, indeed, a dark period in our history to which I have
already alluded in these biographical sketches, but Peel (luckily for
him) was out of office during the greater portion of that gloomy
time, and never made himself prominent in it except once, when
called upon as a neighbour to defend the character of the
magistrates on that day still memorable, in spite of all excuses and

palliations, as the day of the “Manchester massacre.” He undertook
and performed his very delicate task on this occasion with tact and
discretion. No one, indeed, ever spoke in a less unpopular manner
on an unpopular subject. Far superior to Mr. Canning, in this respect,
from that calm, steady, and considerate tone which never gives
offence, and which, laying aside the orator, marks the statesman, he
neither attempted to excite anger, nor ridicule, nor admiration; but
left his audience under the impression that he had been performing
a painful duty, in the fulfilment of which he neither expected nor
sought a personal triumph.
III.
From the proceedings against the Queen, which shortly followed
(the old King dying in 1820), he kept as much as possible aloof. On
one occasion, it is true, he defended the legal course which the
Ministry had adopted for settling the question of the Queen’s guilt or
innocence; but he blamed the exclusion of her Majesty’s name from
the litany; the refusal of a ship of war to bring her to England, and
of a royal residence on British soil; in short, he separated himself
distinctly from any scheme of persecution, manifesting that he would
not sacrifice justice to Royal favour.
The Government at this time was so weak, having suffered, even
previous to the Queen’s unfortunate business, which had not
strengthened it, several defeats, that Lord Liverpool saw the
necessity of a reinforcement, and, faithful to the system of a double-
mouthed Cabinet, took in Mr. Wynn (the representative of the
Grenvilles), to speak in favour of the Catholics, and Mr. Peel (as
successor to Lord Sidmouth, who gave up the Home Office, but
remained in the ministry), to speak against them.
The change, nevertheless, considerably affected the
administration, both as to its spirit and its capacity. The Grenvillites
were liberal, intelligent men generally, as well as with respect to the

Catholics, and Peel was generally liberal, though hostile to the claims
of the Catholic body.
Lord Sidmouth, at the Home Office, had moreover been a barrier
against all improvement. His career, one much superior to his merits,
had been owing to his having all George III.’s prejudices without
George III.’s acuteness. He was, therefore, George III.’s ideal of a
minister, and on this account had been stuck into every ministry,
during George III.’s lifetime, as a kind of “King’s send,” representing
the Royal mind. Uniting with Lord Eldon against every popular
concession, and supporting in a dry, disagreeable manner every
unpopular measure, he was as much hated as a man can be who is
despised. Peel, at all events, wished to gain the public esteem. His
abilities were unquestioned. He was much looked up to by his own
party, much respected by the opposing one; and, as it was known
that Mr. Canning had at this time engaged himself to accept the
Governor-Generalship of India, every one deemed that, if the Tories
should remain in power, Peel would be Lord Liverpool’s inevitable
successor.
The moderate and elevated tone of his language, his indefatigable
attention to business, a certain singleness and individuality which
belonged to him, foreshadowed the premiership. Even the fact that
his father had, undisguisedly, intended him for this position, though
the idea was quizzed at Peel’s entry into public life, tended
eventually to predispose persons to accept it; for people become
accustomed to a notion that has been put boldly and steadily before
them, and it is rare that a man of energy and ability does not
eventually obtain a distinction for which it is known, during a certain
number of years, that he is an aspirant.
But one of those accidents which often cross the ordinary course
of human life—the sudden death of Lord Castlereagh and the
appointment of Mr. Canning as his successor—retained the Home
Secretary in a second-rate position, over which the great and
marvellous success of the new foreign secretary threw a certain
comparative obscurity. He was obliged, therefore, to be satisfied with

continuing to pursue a subordinate, but useful career, which might
place him eventually in men’s minds, side by side with his more
brilliant competitor.
IV.
The subject to which he now particularly devoted himself was the
most useful that he could have chosen. We had at the time he
entered office a police that was notoriously inefficient; prisons,
which by their discipline and condition were calculated rather to
increase crime than to act as a corrective to it; and laws which
rendered society more criminal than the criminals it punished. One
can scarcely, in fact, believe that such men as Lord Eldon and Lord
Ellenborough did not think it safe to abolish the punishment of death
in the case of privately stealing six shillings in a shop; and it is with a
shudder that one reads of fourteen persons being hanged in London
in one week in 1820, and of thirty-three executions in the year 1822.
No one reflected whether the punishment was proportionate to
the offence; no one considered that the alleged criminal himself was
a member of the community, and had as much right to be justly
dealt with and protected against wrong as the community itself.
Satisfied with the last resort of hanging, the State neglected to take
suitable precautions against the committal of those acts which led to
hanging; nor did it seem a matter of moment to make places of
confinement places of reformation, as well as places of atonement.
To Bentham, Romilly, Mackintosh, Basil Montagu, and others, we
owe that improvement in the public mind which led finally to an
improvement in our laws. Mr. Peel had marked and felt this gradual
change of opinion; and almost immediately after he became invested
with the functions of the Home Department, he promised to give his
most earnest attention to the state of the police, the prisons, and
the penal laws; a promise that, in the four or five succeeding years,
he honourably fulfilled; thus giving to philanthropic ideas that
practical sanction with men of the world, which theories acquire by
being taken up by men in power.

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The Psychology Of Learning And Motivation Volume 76 Kara D Federmeier

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The Psychology Of Learning And Motivation Volume 76 Kara D Federmeier

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