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Evolution of the Cerebellar
Sense of Self
Evolution of the
Cerebellar Sense
of Self
John Montgomery
University of Auckland, New Zealand
David Bodznick
Wesleyan University, Connecticut, United States of America
1
1
Great Clarendon Street, Oxford, OX2 6DP,
United Kingdom
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Preface
that includes the familiar sharks, skates, and rays, and lesser known groups such
as chimaeras and ghost sharks. There is a long tradition in biology about being
very particular with names, but, for a book covering the collective disciplines
that make up comparative neuroscience, each with its own language and vocab-
ulary, we will relax that tradition to the extent possible. New words (for non-
biologists) will be introduced where necessary to avoid too much repetition of
alternate definitions. But new words and TLAs (three-letter acronyms) will be
kept to a minimum in the interest of preserving accessibility for non-biologists.
Thirdly, when we ask the question: what does the cerebellum do?—the answer
will inevitably get us into the territory of how brains are wired up, and the ter-
minology necessary to describe the brain structure and function. Fortunately,
we are more interested in ‘how nerves work animals’ than how ‘nerves work
per se’, so we can substantially pare down the level of cellular and molecular
detail we require. Put simply, it is important to understand that neurons are
cells that convey information from one part of the body, or brain, to another,
and that information is conveyed as electrical impulses along cell structures
called axons. Also fundamental is knowing that nerve axons largely interface
with other neurons (or with muscles) through chemical connections called syn-
apses, which come in two flavours: excitatory synapses which tend to nudge
the following cell towards firing an impulse; and inhibitory synapses that tend
to reduce the likelihood of an impulse. And finally, that learning in the brain
occurs through changes in the strength of those synaptic connections. So, in
three sentences, we now have the bulk of neuroscience background to under-
stand the storyline. However, it is worth noting some additional terminology
here. Neurons have distinctive shapes, with a cell body that contains the nucleus
of the cell, and two types of cell structures: dendrites (from the Greek for ‘tree-
like’ branching structure) that receive the excitatory and inhibitory input from
other cells; and axons that transmit the impulses to axon terminals that connect
(synapse) with another neuron or a muscle cell. Additional background will be
introduced, where necessary, in the main text of the book, but our focus and
challenge are to address the question of what the cerebellum does and how it
does it at the level of local circuits, without delving into the underlying cellular
or molecular mechanisms of neuron function.
It is hoped that restricting cellular detail to the minimum necessary will open
up the accessibility of this subject area to non-neuroscience readers with a range
of backgrounds and interests. Although it would be desirable to have interest-
specific navigation guides for the text, the range of reader backgrounds we aim
to attract makes this a difficult task. Specialist cerebellar researchers will inevi-
tably zoom in on their area of interest and find it under-represented. Our hope
is that they would find more value in the overall scaffold portrayed in the book
Preface vii
and the opportunity to test the utility and limitations of the scaffold against
their specialist knowledge. Engineers and IT readers will no doubt strug-
gle with the terminology and anatomical descriptions, just as biologists will
feel unfamiliar with the engineering analogies. Behavioural psychologists will
struggle with the detail of the model systems, as will all mammalian researchers
with the intricacies of electric fishes. Mindful of these reader-specific fairways
and hazards, we have attempted to provide bullet-point introductions to each
chapter in a way that will signal essential concepts. Hopefully, these will allow
different readers to access the text in a way that works best for them. In addi-
tion, our objective has been to provide figures for each of the critical structures,
or concepts, of the story. The figures and legends alone should provide a useful
guide to the flow of ideas in the main text. Finally, in covering such a swathe of
material, it is not surprising that some areas, especially those in which we have
a particular interest or experience, will be covered in more depth and detail
than others. To attempt to balance this effect and not disrupt the flow of the
narrative, we have introduced some additional material in figure legends and
indicated sections that can be skimmed or skipped. These are areas that are
not essential to the main theme but are covered nevertheless for their intrinsic
interest and the strength we hope they add to the main story.
Acknowledgements
Index 207
List of Figures
◆ In the human brain, the cerebellum occupies only 10% of the volume
but accounts for a surprising 80% of the nerve cells; it is a neural net-
work of 69 billion neurons.
◆ Understanding the cerebellum depends on both its intriguing struc-
ture, made up of a repeated neuronal motif, and the functional opera-
tion of that motif as an adaptive filter.
◆ The cerebellum evolved in early sharks, providing further insights that
illuminate both the nature of the adaptive filter and an emerging view
of what the cerebellum does.
◆ The function of shark cerebellum-like structures is to discriminate
‘self ’ from ‘other’ in sensory inputs, and is the fingerpost for an evolu-
tionary account of our cerebellar sense of self.
One of the most extraordinary things about the cerebellum is that, although it only
occupies 10% of the brain volume, it contains more nerve cells than all the rest of
the brain put together! By recent accounts (Azevedo et al., 2009), the human brain
contains something like 86 billion nerve cells, with 69 billion (80%) of them in the
cerebellum. The disproportionate number of neurons clearly indicates this densely
packed part of the brain is important. But what does the cerebellum do, and why
does it need so many neurons to do it? These are simple questions to ask, but how do
we even begin to understand such an intricate and complex structure? Single nerve
cells are complex entities, but that complexity compounds with the sheer numbers
of cells involved and the way in which neurons connect and interact in networks.
This book seeks to address the question of cerebellar function, and argues
that this is made possible by three recent developments:
1. The insight that the functional contribution of the cerebellum to behaviour
is achieved through a core computational element called an adaptive filter.
2. Our perspective on the evolution of the cerebellum, particularly that its origin
and function in early sharks provide insights that illuminate both the nature
of the adaptive filter and the emerging view of what the cerebellum does.
2 Evolution of the Cerebellar Sense of Self
3. The view that the cerebellum provides a sophisticated, but subliminal, sense
of self, which we have termed the ‘cerebellar self ’.
Our goal is to use an evolutionary perspective to open up cerebellar research
to a wider audience than neuroscience specialists. Neuroethology is the endeav-
our to understand the neural basis of animal behaviour (ethology is the study of
behaviour). The broad perspective of neuroethology leads us to look across the
full spectrum of vertebrate animals with a cerebellum, and ask questions about
the origin of the cerebellum and the function of the cerebellum across the wide
range of vertebrate behaviour. We don’t underestimate the complexity of the
cerebellum or the complexity of the body of cerebellar research work. But from
this broad perspective, we identify an emergent concept of the cerebellar sense
of self as a scaffold on which to build a picture of what the cerebellum is, what
it does, and how it does it.
Historically, the function of the cerebellum has been an enigma. Despite the
fact that it contains so many nerve cells, it tends to be under-represented in
modern neuroscience texts and in our thinking about the brain. As we gain a
better understanding of the cerebellum, this disparity between neuron num-
bers and attention starts to resolve. Once we lay out the architectural structure
of the cerebellum and the common links that exist across evolution and across
the model systems used to understand cerebellar function, we can address the
question: why are so many neurons tied up in cerebellar circuits, and what does
this mean for cerebellar function?
As part of the explanation of our tendency to ignore the cerebellum, we will
also find a common thread of sophisticated, yet quite subtle, contributions that
the cerebellum makes. Both the evolutionary perspective and our understand-
ing of experimental models illuminate the idea that we can, and do, ignore
much of what the cerebellum does, and that we do this precisely because the
cerebellum operates largely at a subconscious level. But, even though it oper-
ates subliminally, as we begin to understand the cerebellar self, we also start to
appreciate how important it is to our perception of our surroundings, how we
move, and even the implicit sense of agency we have in our interactions with
the world.
One of the most significant recent insights into cerebellar function is the idea
that the cerebellum is made up of a massively repeated series of neuronal circuits
functionally equivalent to adaptive filters. Adaptive filters are electronic devices
that will be familiar to some, particularly engineers, but not to most other read-
ers. Gaining an intuitive understanding of what an adaptive filter is and how it
works is central to the understanding of the cerebellum that we portray in this
book. This is a challenge we will address multiple times through the book. As
we progress through the chapters, we will illustrate the structure and versatility
Introduction to the cerebellar sense of self 3
of adaptive filters: how the neuronal circuits of the cerebellum provide for adap-
tive filter function; how they can be used to provide useful predictions; and how
adaptive filters can be put to multiple use. But by way of introduction, it is worth
sketching the generic idea of an adaptive filter (e.g. Haykin, 1984). Firstly, filters
process signals; the output signal of a filter is computed from input signals. The
‘adaptive’ in adaptive filters does not refer to their selective advantage in evolu-
tion, but rather their most important property—the computation of adaptive
filters can be changed, refined, or updated through experience to alter the filter’s
input–output relationship. Finally there is some adaptive algorithm, or rule,
that describes how the computation is adjusted through time to improve the fil-
ter output in some defined way. That, in essence, is an adaptive filter, and, from
an engineering perspective, they have proved to be highly useful and versatile
across a very wide range—from control systems to adaptive noise cancelling.
Given that animal athleticism is dependent on sophisticated neural control and
that effective sensing of the world is critically dependent on the detection of
biologically important signals in a sea of sensory noise, it would be surprising if
there wasn’t a biological equivalent to an adaptive filter.
In electronics, an adaptive filter is typically implemented as an electronic
chip. The adaptive filter analogy for the cerebellum equates the neuronal com-
putational circuit to a ‘cerebellar chip’. One of the key questions for the adaptive
filter proposal is: how would it be implemented in the neuronal circuitry of
the cerebellum? In nature, structure and function are inextricably intertwined,
so this question leads us directly to cerebellar structure. Neuroscientists have
for decades been encouraged in the quest to understand the cerebellum by its
intriguing microscopic structure. Early anatomical work showed that the cer-
ebellum is made up of relatively few nerve cell types, or neurons, and moreo-
ver that these are put together in a highly ordered and intriguing latticework.
Essentially the same repeated motif of interconnected neurons occurs across
the entire structure of the cerebellum in a way not seen in other brain areas. The
obvious idea is that, if we could understand the function of this cerebellar motif,
this would unlock a more general understanding of cerebellar function. What
if the cerebellar motif were indeed functionally equivalent to an adaptive filter?
The cerebellar neural circuit motif and details of the nerve cell types and inter-
connections have been well characterized for more than half a century. They are
beautifully portrayed in a classic book by John Eccles and his co-authors (Eccles
et al., 1967). On the basis of the repeated cerebellar motif, Eccles characterized
the cerebellum as a ‘Neuronal Machine’. However, although the cerebellum was
demonstratively machine-like in its structure, the Eccles et al., 1967 book was
almost silent on what the ‘Neuronal Machine’ might do. Indeed, the function
of the cerebellum has for years been frustratingly elusive. The final chapter of
4 Evolution of the Cerebellar Sense of Self
(B)
spiny tega
lamprey ratfish dogfish trout frog lizard cormorant dog
Brains
Mammals
200 Neocortex
Innovations
Time (M years)
300
Tetrapods Paired limbs
400
is the question: what is the relationship between mind and body? The sim-
ple answer is: we don’t know. The search for an answer does, however, have a
long history—a history that has been framed somewhat differently at different
times and by different cultures. Recently, modern neuroscience has delicately
shifted the language from the ‘seat of the soul’ to the ‘seat of the self ’ and used
new brain scanning methods to provide evidence to ‘pinpoint the “seat of self-
consciousness” ’ in the cerebral cortex (e.g. Ionta et al., 2011). Perhaps more
interesting than identifying the location, these same studies relied on manipu-
lating sensory input to trick the subject into an out-of-body experience. 3D
goggles projected a recorded image of a person’s back a short distance in front
of them. The patient was then stroked by a robot at precisely the same time that
the body in the virtual world was stroked, tricking the mind into believing that
the image of the body in front of the patient was the patient’s own body! The
brain scans showed that, during such manipulations, the brain area most acti-
vated was a small portion of the cerebral cortex. These findings allow us to say
with some confidence that particular parts of the brain are directly involved
in self-consciousness, and show it is even possible to use illusions to literally
dislocate the self, but they still don’t give us the essential nature of the relation-
ship between body and mind. The ‘mind body problem’ remains a profound
and tantalizing paradox between our consciousness, as central to how we per-
ceive and understand the world, and our inability to understand the nature of
consciousness itself.
Our focus in this book is not on the conscious self, but mostly on a deeper,
unconscious self that distinguishes ‘self ’ from ‘other’ in our interactions
with the physical world. Our subject of interest is not our conscious pilot,
but rather the autopilot—our subliminal self. Clearly the subliminal-self
and consciousness interact. But perhaps like the autopilot on an overnight
flight, the subliminal self frees our conscious pilot to contemplate the stars
or perhaps the comforts (or otherwise) that awaits the journey’s end. So
although our topic interfaces with the question of consciousness, its core
focus is the subliminal self and an emerging view that the cerebellum is
instrumental in constructing this sophisticated, yet subliminal, sense of
self-and-other. But it is not simply the sense of self that is important, but
also the fundamental, but largely hidden, contribution it makes to our sen-
sory and physical interactions with the world. The autopilot of our sublimi-
nal self monitors the sensors and runs the control surfaces through which
we interact with our environment. This is the meaning implicit in our use
of the ‘cerebellar sense of self ’. What is particularly compelling about this
view of the cerebellum is that we can go well beyond simply identifying the
cerebellum as the responsible part of the brain. Such is the nature of the
8 Evolution of the Cerebellar Sense of Self
cerebellar neuronal machine that we can outline in some detail what it does
and how it does it. Unlike consciousness, our understanding of the cerebel-
lar self can be directly related back to the operation of single neurons and
their associated circuits.
So our goal is to portray the current understanding of the cerebellum
across vertebrates from sharks to humans, but in particular to provide an
accessible insight into what it is that our cerebellum does for us. Outlining
the conservation of cerebellar structure throughout vertebrate animals pro-
vides a strong argument that neuroethological study of the cerebellum across
the full range of vertebrate evolution will inform our understanding of the
role it plays in our interactions with the world. But before progressing to a
more mechanistic consideration of cerebellar function, we will profile one
high-level role of the cerebellum in humans. To gain a view of the poten-
tial higher-level function of the cerebellum, we will start with an illustra-
tion of the critical importance of the contribution of the cerebellar self to
our cognitive experience. Although our core focus is the subliminal self, we
have already indicated that this does interface with consciousness, and what
is particularly fascinating is how this interface becomes apparent when the
cerebellum misfunctions. The example we will profile is one demonstrated
role of the cerebellum in neuropsychiatric disorders. This case provides quite
remarkable insights into underlying neural mechanisms of psychoses, and a
window into a role of the cerebellum which crosses the line into our cogni-
tive sense of self.
Between sharks and psychoses, there is a massive body of work on specific
models of cerebellar function; these too add to the coherent picture of the cer-
ebellum as instrumental to our sense of self. In a recent synopsis of the cer-
ebellum, Masao Ito, one of the pre-eminent cerebellar researchers, calls it the
‘brain for the implicit self ’ (Ito, 2012). This view portrays an unconscious self
that distinguishes ‘self ’ from ‘other’ in our interactions with the physical world.
We have developed and extended the concept of the implicit self to include the
early origins of the cerebellum and aspects of how the cerebellum interacts with
consciousness. It is for this overarching view of cerebellar function that we have
coined the term ‘cerebellar self ’.
To reiterate, this developing view of the cerebellum helps to explain the rela-
tively low profile of the cerebellum in our collective consciousness—‘it works so
well it can largely be ignored’. It is, in some respects, the brain’s equivalent to our
immune system. Distinguishing ‘self ’ and ‘other’ is as important to our brain in
our daily interactions with the physical world as it is to our immune system in
recognizing our own tissues as different from an invading microbe. And, like
the immune system, we can largely ignore it until things go wrong. This potent
Introduction to the cerebellar sense of self 9
idea of the cerebellar self links across the full spectrum of known cerebellar
function—at one end of the spectrum, it provides a fascinating insight into
some human psychotic states that fail to distinguish ‘self ’ from ‘other’; at the
other end of the spectrum, it captures the known function of cerebellum-like
structures in the shark brain.
We have chosen to begin with a consideration of the role of the cerebellum
in our sense of agency (Chapter 2). This high-level operation of the cerebellum
and its interaction with conscious experience provide a clear example of the
potency of cerebellar function to contribute to prediction and expectation. It
also provides a salutary illustration of the consequences of misfunction. For
those readers with a non-neuroscience background, it also seems a kinder
place to start than a headlong dive into cerebellar anatomy. Nevertheless,
understanding the anatomical structure of the cerebellum is essential to under-
standing function. The process of evolution in biological systems dictates that
structure and function (= physiology) in nature are necessarily intertwined. To
rephrase a quote from the philosopher of science Lakatos: ‘physiology with-
out anatomy is blind; anatomy without physiology is empty’. Chapter 3 pro-
vides a succinct overview of cerebellar anatomy and how the cerebellar motif
relates back to the structure of the cerebellum as seen in a scan of the human
brain. This chapter also shows how similar the basal shark cerebellum is to our
own. Consideration of the shark cerebellum leads directly to the considera-
tion of the evolutionary precursor to the cerebellum (Chapter 4). These are the
cerebellum-like structures that have a function equivalent to noise-cancelling
headphones. They provide a means to separate biologically important signals
from self-generated sensory noise and a direct way of illustrating the structure
and function of a biological adaptive filter. The shark brain also provides us
with an intriguing hypothesis as to the evolutionary origins of the cerebellum
(Chapter 5).
Chapter 6 addresses many of the key model systems that have been so influ-
ential in generating our understanding of the cerebellum. These model sys-
tems include: the reflex systems that stabilize the eyes during head movement;
the eye blink reflex as an example of protective and predictive motor learn-
ing; rapid target-directed eye movements; and the role of the cerebellum in the
actions and reactions that underlie animal athleticism. In Chapter 7, we will use
these model systems to directly address the question of what the cerebellum
does and challenge the generality of the ideas of adaptive filter operation and
cerebellar self.
Science is an engine of understanding. The progress of science in the last
100 years has been impressive, and the process of science is, in many ways,
as interesting as the outcomes it produces. In Chapter 8, we will profile a
10 Evolution of the Cerebellar Sense of Self
condensed history of cerebellar research set in the wider context of key neuro-
science discoveries. This history is important to gain an understanding of how
progress has been made, to reinforce the understanding of the cerebellum, and
to speculate on where we are on the trajectory of addressing the question of
cerebellar function.
Chapter 9 addresses potential applications that might arise from an under-
standing of cerebellar function. As we approach a better understanding of the
human cerebellum, it should prove possible to utilize that understanding in
enhancing cerebellum-related performance. Hopefully, this progress can be
used to moderate the adverse effects of cerebellar injury and disease, but per-
haps also to enhance performance in sport and other complex motor actions.
There are also the rapidly growing areas of bioinspiration and biomimetics.
Biomimetics is literally the idea of building new technology that mimics the
best in nature. As an idea, biomimetics is not new. Alessandro Volta invented
the battery after working with electric rays (their biological name is Torpedo),
in the attempt to find out how they stored electricity underwater. Although he
didn’t use the word biomimetics, he is quoted as saying ‘nature has found the
way to succeed with this in the electric organs of Torpedo. And perhaps we are
not far from the possibility that our art could imitate them’. Lord Cavendish,
who gives his name to the Cavendish Laboratory in Cambridge, also drew
inspiration for his work on electricity from electric rays (Cavendish, 1776). If
one cartilaginous fish, the Torpedo ray, could inspire the invention of the bat-
tery over 200 years ago, what other biomimetic inventions from sharks and
other animals await discovery?
Finally, as we review the evolution and function of the cerebellum, it
becomes possible to make several rather dramatic claims. Firstly, the evolu-
tion of the cerebellum, along with paired fins, in early jawed vertebrates paved
the way for the subsequent command of the three dimensional environments
of sea, land, and air. In other words, the cerebellum is the source of the athleti-
cism and movement finesse that we see in swimming, running, climbing, and
flying vertebrates. Secondly, as Ito notes in his recent book, work on the cer-
ebellum has the potential to be at the forefront of endeavours to understand
the mechanistic details of how the brain can ‘accomplish its most complex
and sophisticated actions’. Finally, the conserved structure of the cerebel-
lar ‘chip’ over such a long evolutionary period and the dramatic diversity of
functional utility enabled by the ‘chip’ provide inspiration for technological
mimics. The idea of copying the best of nature to inspire new technologies is
gaining currency, and the cerebellar ‘chip’ provides real potential for a diverse
array of applications.
Introduction to the cerebellar sense of self 11
References
Azevedo, F. A., Carvalho, L. R., Grinberg, L. T., et al. (2009). Equal numbers of neuronal
and nonneuronal cells make the human brain an isometrically scaled–up primate brain.
Journal of Comparative Neurology, 513(5), 532–41.
Buckner, R. L. (2013). The cerebellum and cognitive function: 25 years of insight from
anatomy and neuroimaging. Neuron, 80(3), 807–15.
Cavendish, H. (1776). An account of some attempts to imitate the effects of the Torpedo by
electricity. Philosophical Transactions of the Royal Society of London, 66, 196–225.
Eccles, J.C., Ito, M., and Szentágothai, J. (1967). The cerebellum as a neuronal machine.
Berlin-Heidelberg-New York: Springer.
Haykin, S. S. (1984). Introduction to adaptive filters. New York: Macmillan.
Ionta, S., Heydrich, L., Lenggenhager, B., et al. (2011). Multisensory mechanisms in
temporo-parietal cortex support self-location and first-person perspective. Neuron,
70(2), 363–74.
Ito, M. (2012). The cerebellum: brain for an implicit self. Upper Saddle River, New
Jersey: Pearson Education Ltd.
Chapter 2
The plan is to first take a high-level look at our own cerebellum and one of its
contributions to consciousness to get a better idea of why the ‘subliminal self ’
view of the cerebellum is so compelling. This offers a glimpse of the potential
subtlety and sophistication of what the cerebellum can do, before we delve into
the intriguing anatomical structure of the cerebellum and some of the more
basic tasks of the cerebellum in sharks and other model systems.
To interact with cognition, the cerebellum must be connected with, and
inform, our conscious self, even if this connection is to the lowest form of con-
scious self that philosophers sometimes refer to as the ‘minimal self ’ (Gallagher,
2000). The example chosen to illustrate this interactive functionality is our
sense of agency, or the level of command and control we experience generating
our own body movements. That sense of agency is the compelling, but fragile,
experience ‘of generating and controlling actions in order to influence events in
the outside world’. This quote comes from a recent review of how this sense of
agency may be constructed and how it can fail in certain disease states (Moore
and Fletcher, 2012).
Cerebellar sense of self and sense of agency 13
Not only does our sense of agency operate at the interface of our subliminal
and minimally conscious self, but it also seems that this is one example where
we may be able to draw something of a straight line between a cognitive human
facility and the detailed workings of the cerebellum neuronal machine. Let’s see
how that might work.
Self-tickle
The starting point might seem like a trivial question: why can’t you tickle your-
self? Or perhaps the more precise question: why is self-tickle less effective? The
answer is really interesting. Self-tickle is less effective because the part of your
brain that is ‘tickle-aware’ doesn’t get such a strong message when the tickler is
yourself. You can try this out. Tickle the palm of your hand, and pay attention
to how ticklish that feels. Ask a friend to repeat the same tickle movement in the
same place. For most people, the heightened effect of the external tickle is very
evident. The same phenomenon applies not just to tickle, but to any touch on our
skin. Under controlled conditions in the laboratory, brain-wave signals can be
compared in subjects either when they were touched by another person with a
brush or when they touched themselves with the same brush. The signals in the
primary somatosensory cortex, the area of the brain that contributes to touch,
or tickle ‘awareness’, were about one-fifth weaker in the case of self-produced
stimuli, in comparison with externally applied stimuli (Hesse et al., 2010).
At the level of the receptors, a sensory stimulus to the skin is the same, irrespec-
tive of whether the stimulus is self-generated or caused by an external agent. The
difference is that, in the self-generated case, the brain has knowledge in advance
of the precise details of the pending movement and hence knows the source of
that movement to be ‘self ’ (Figure 2.1). The biological perspective is that, in many
circumstances, the predicted consequences of our own actions are not useful
and so should not tie up the ‘limited’ capacities of the brain systems that mediate
our sensory awareness. The signals that are biologically important are often the
unpredicted, or unexpected, events that are part of our interaction with the exter-
nal world. Folding our hands behind our head may stimulate the touch receptors
on our neck but doesn’t require the sort of response we might give to a cockroach
scuttling across the same receptors. So it is perhaps not surprising there is con-
siderable evidence for weaker cortical brain responses to similar self-produced
versus externally generated stimuli across a range of different sensory modalities.
Self-generated stimuli are inherently predictable, so it makes sense that
they should be handled differently from the same sensory stimulus not self-
generated. Attenuation of the cortical response to your own actions makes it
possible to ignore expected, hence less relevant, consequences of your own
14 Evolution of the Cerebellar Sense of Self
Cerebellum + –
Predicted Sensory feedback
sensory (reafference)
feedback
(forward model)
actions and to enhance unanticipated sensory inputs. As we will see later, this
same differentiation between self and other may also help with the active explo-
ration of one’s environment and with the ability to correct errors in execution
of an intended movement.
So what are the neural mechanisms underlying the reduced cortical response
to self-tickle? As we have already noted, the brain has knowledge in advance
of an impending movement. For self-generated movement, the parts of the
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reluctantly, but still with a secret alacrity. She was left alone with the
mentor, who had so often brought her advice or semi-reproof.
‘You have something to tell me? Oh, Mr. Sommerville, what is it?’ she
cried.
‘It is nothing very bad. You must not be alarmed—there is no ill news,’
he said.
The anxious mother looked at him with a wistful entreaty in her eyes. Ill
news was not what she feared. When a woman has had neither
companionship nor help from her husband for a dozen years or so, naturally
her sensitiveness of anxiety about him gets modified, and it is to be feared
that she would have taken information of Mr. Meredith’s serious illness, for
instance, more easily than the summons which she feared for one of her
boys. She watched every movement of her visitor’s face with anxious
interest.
‘Edward cannot go till the settled time. You know that,’ she said,
instinctively following the leading of her own thoughts.
‘It is not Edward that I have come to speak of; it is neither of the boys.’
‘Ah!’ said Mrs. Meredith, with a sigh of involuntary relief; and she
turned to him with cheerful ease and interest, delivered from her chief fear.
This evident ignorance of any other cause for animadversion moved the old
Spy in spite of himself.
‘What I am going to say to you, my dear lady, is not exactly from
Meredith—though he has heard of the subject, and wishes me to say
something. I hope you will believe there is no harm meant, and that what I
do, I do from the best feeling.’
‘I have never doubted your kind feeling, Mr. Sommerville; but you half
frighten me,’ she said, with a smile. ‘If it is not the boys, what can there be
to be so grave about? Tell me quickly, please.’
Mr. Sommerville cleared his throat. He put his hat upon the head of his
cane, and twirled it about. It did not often happen to the old Scotch nabob to
be embarrassed; but he was so now.
‘You’ll understand, my dear lady, that in what I say I’m solely actuated
by the thought of your good.’
‘How you alarm me!’ said Mrs. Meredith. ‘It is something, then, very
disagreeable?’
‘Oh, yes. I’ve no doubt it will be disagreeable. Medicines are seldom
sweet to the palate. Mrs. Meredith, I will out with it at once, not to keep you
in suspense.’
Here, however, he paused to take out his handkerchief, and blew his nose
with a very resounding utterance. After he had finished this operation he
resumed:
‘I don’t presume to teach a lady of your sense what is her duty; and I
don’t need to tell you that the world exercises a great supervision over
women who, from whatever cause, are left alone.’
‘What have I done?’ cried Mrs. Meredith, half frightened, half laughing.
‘I must have made some mistake, or you would not speak so.’
‘I doubt if it could be called a mistake; perhaps it would be better to say
a misapprehension. Mrs. Meredith, there is one of your friends who pays
you a visit every day.’
‘Several,’ she said, relieved. ‘You know how kind people are to me.
Instead of supervision, as you say, I get a great deal of sympathy——’
Mr. Sommerville waved his hand, as if to ward off her explanation. ‘I am
speaking of one person,’ he said: ‘a man—who is here every evening of his
life, or I’m mistaken—your neighbour, Mr. Beresford, next door.’
‘Mr. Beresford!’ she said, with a thrill of disagreeable surprise; and there
came to her instantaneously one of those sudden realisations of things that
might be thought or said, such as sometimes overwhelm the unsuspecting
soul at the most inappropriate moment; her colour rose in spite of herself.
‘Just Mr. Beresford. He means no harm and you mean no harm; but he
should be put a stop to, my dear lady. You gave me your word you would
not be angry. But, madam, you’re a married lady, and your husband is at a
distance. It’s not for your credit or his good that he should visit you every
night.’
‘Mr. Sommerville! stop, please! I cannot let you talk so—or anyone.’
‘But you must, my dear lady, unless you want everybody to talk, and in a
very different spirit. The world is a wicked world, and takes many things
into its head. You’re a very attractive woman still, though you’re no longer
in your first youth——’
‘Mr. Sommerville, what you say is very disagreeable to me,’ said Mrs.
Meredith, offended. ‘Poor Mr. Beresford! since he lost his wife he has been
miserable. Nobody ever mourned more truly; and now, when he is trying to
learn a little resignation, a little patience——’
‘He should not learn those virtues, madam, at your expense.’
‘At my expense!’ she said, with sparkling eyes; ‘at what expense to me?
I allow him to come and sit with me when he has no one at home to bear
him company. I allow him——’
‘I thought his daughter had come to keep him company.’
‘Poor Cara! she is a sweet child; but, at seventeen, what can she know of
his troubles?’
‘Softly, softly,’ said Mr. Sommerville; ‘one plea is enough at a time. If
Mr. Beresford is without a companion, it does not matter that his daughter is
only seventeen; and whatever her age may be, if she is there he cannot be
without companionship. My dear lady, be reasonable. If he has a child
grown up, or nearly so, he should stay at home. A great many of us have not
even that inducement,’ said the old man, who was an old bachelor; ‘but no
kind lady opens her doors to us.’ He looked at her sharply with his keen
eyes; and she felt, with intense annoyance, that she was getting agitated and
excited in spite of herself.
‘Mr. Sommerville,’ she said, with some dignity, ‘if anyone has been
misrepresenting my friendship for Mr. Beresford, I cannot help that. It is
wicked as well as unkind; for I think I have been of use to him. I think I
have helped him to see that he cannot abandon his life. I don’t mean to
defend myself. I have not done anything to be found fault with; friendship
——’
‘Is a delusion,’ said the old man. ‘Friendship between a man and a
woman! There is no sense in it. I don’t believe a word of it. Meaning no
harm to you, my dear lady. You don’t mean any harm; but if you talk to me
of friendship!’
‘Then I had better say nothing,’ she answered quickly. ‘My husband’s
representative—if you call yourself so—has no right to treat me with
rudeness. I have nothing more to say.’
‘My dear lady,’ said old Mr. Sommerville, ‘if I have appeared rude I am
unpardonable. But you’ll forgive me? I mean nothing but your good. And
all I want is a little prudence—the ordinary precautions.’
‘I will none of them!’ she said, with a flush of indignation. ‘I have
nothing to be afraid of, and I will not pretend to be prudent, as you call it.
Let the world think or say what it pleases—it is nothing to me.’
Then there was a pause, and Mrs. Meredith betook herself to her work—
a woman’s safety-valve, and laboured as if for a wager, while the old
plenipotentiary sat opposite to her, confounded and abashed, as she thought.
But Mr. Sommerville was too old and experienced to be much abashed by
anything. He sat silent, collecting his forces for a renewed attack. That was
all. He had a sincere friendship for her in his way, and was as anxious to
prevent scandal as any father could have been; and now it occurred to him
that he had begun at the wrong end, as he said. Women were kittle cattle.
He had failed when he dwelt upon the danger to herself. Perhaps he might
succeed better if he represented the danger to him.
‘I have made a mistake,’ said the hypocritical old man. ‘It can do no
harm to you, all that has come and gone. I was thinking of my own selfish
kind that give most weight to what affects themselves, and I am rightly
punished. A lady sans reproche like yourself may well be sans peur. But
that is not the whole question, my dear madam. There is the man to be
considered.’
When he said this she raised her eyes, which had been fixed on her
work, and looked at him with some anxiety, which was so much gained.
‘You will not doubt my word when I say there’s a great difference
between men and women,’ said the old diplomatist. ‘What is innocent for
one is often very dangerous for the other, and vice versâ: you will not deny
that.’
Then he made a pause, and looking at her for reply, received a sign of
assent to his vague proposition, which indeed was safe enough.
‘How can you tell that Mr. Beresford receives as pure benevolence all
the kindness you show him? It is very unusual kindness. You are kind to
everybody, madam, above the ordinary level; and human creatures are
curious—they think it is their merit that makes you good to them, not your
own bounty.’
She did not make any reply, but continued to look at him. Her attention
at least was secured.
‘If I were to tell you the instances of this that have come under my own
observation! I have known a poor creature who got much kindness in a
house on account of his defects and deficiencies, and because everybody
was sorry for him; who gave it out, if you’ll believe me, and really thought,
that what his kind friends wanted was to marry him to the daughter of the
house! It’s not uncommon, and I dare say, without going further, that you
can remember things—which perhaps you have laughed at——’
‘All this has nothing to do with Mr. Beresford,’ she said, quietly, but
with a flush of rising offence.
‘No, no.’ He made a hesitating answer and looked at her. Mrs. Meredith
fell into the snare.
‘If he has misunderstood my sympathy for his troubles, if he has
ventured to suppose——’
‘Cara has gone out with her aunt,’ said Edward, coming in hastily; ‘but
there is surely something wrong in the house. Mr. Beresford called me into
his room, looking very much distressed. He told me to tell you that he
thought of leaving home directly; then changed his mind, and said I was not
to tell you.’
‘Why do you tell me then?’ cried his mother, with impatience. ‘What is
it to me where he is going? Am I always to be worried with other people’s
troubles? I think I have plenty of my own without that.’
Edward looked at her with great surprise. Such outbreaks of impatience
from his gentle mother were almost unknown to him. ‘He looks very ill,’ he
said: ‘very much disturbed: something must have happened. Why should
not I tell you? Are you not interested in our old friend? Then something
very extraordinary has happened, I suppose?’
‘Oh, my boy,’ cried Mrs. Meredith, in her excitement, ‘that is what Mr.
Sommerville has come about. He says poor James Beresford comes too
often here. He says I am too kind to him, and that people will talk, and he
himself thinks—— Ah!’ she cried suddenly, ‘what am I saying to the boy?’
Edward went up to her hurriedly and put his arm round her, and thus
standing looked round defiant at the meddler. Oswald, too, entered the room
at this moment. The hour for luncheon approached, and naturally called
these young men, still in the first bloom of their fine natural appetites, from
all corners of the house. ‘What’s the matter?’ he said. But he had another
verse of his poem in his head which he was in great haste to write down,
and he crossed over to the writing-table in the back drawing-room, and did
not wait for any reply. Edward, on the contrary, put the white shield of his
own youthfulness at once in front of his mother, and indignant met the foe.
‘People have talked a long time, I suppose,’ said Edward, ‘that there was
nobody so kind as my mother; and I suppose because you have trained us,
mamma, we don’t understand what it means to be too kind. You do, sir?’
cried the young man with generous impertinence; ‘you think it is possible to
be too innocent—too good?’
‘Yes, you young idiot!’ cried the old man, jumping up in a momentary
fury. Then he cooled down and reseated himself with a laugh. ‘There is the
bell for lunch,’ he said; ‘and I don’t mean to be cheated out of the luncheon,
which, of course, you will give me, by the freaks of these puppies of yours,
madam. But Oswald is a philosopher; he takes it easy,’ he added, looking
keenly at the placid indifference of the elder son.
‘Oswald takes everything easy,’ said Mrs. Meredith, with a sigh. And
they went downstairs to luncheon, and no man could have been more
cheerful, more agreeable than the old Indian. He told them a hundred
stories, and paid Mrs. Meredith at least a score of compliments. ‘This
indulgence will put it out of my power to be at your levée this afternoon,’
he said; ‘but there will be plenty of worshippers without me. I think the
neglected women in this town—and no doubt there’s many—should bring a
prosecution against ladies like you, Mrs. Meredith, that charm more than
your share; and both sexes alike, men and women. I hear but one chorus,
“There’s nobody so delightful as Mrs. Meredith,” wherever I go.’
‘We are all proud of your approbation,’ said Oswald, with much
solemnity: he was always light-hearted, and had no desire to inquire
particularly into the commotion of which he had been a witness. But
Edward kept his eyes upon his mother, who was pale with the excitement
she had come through. What that excitement meant the young man had very
little idea. Something had disturbed her, which was enough for her son; and,
curiously enough, something had disturbed the neighbours too, whom
Edward accepted without criticism as we accept people whom we have
known all our lives. He was curious, and rather anxious, wondering what it
might be.
But as for Mrs. Meredith, the idea of communicating to her sons even
the suggestion that she could be spoken of with levity, or criticised as a
woman, appalled her when she thought of it. She had cried out, appealing to
the boys in her agitation, but the moment after felt that she could bear
anything rather than make them aware that anyone had ventured upon a
word to her on such subjects. She exerted herself to be as vivacious as her
visitor; and as vivacity was not in her way, the little forced gaiety of her
manner attracted the attention of her sons more than the greatest seriousness
would have done. Even Oswald was roused to observe this curious change.
‘What has happened?’ he said to his brother. He thought the Spy had been
finding fault with the expenditure of the household, and thought with alarm
of his own bills, which had a way of coming upon him as a surprise when
he least expected them. It was almost the only thing that could have roused
him to interest, for Oswald felt the things that affected Oswald to be of
more importance than anything else could be. As for Edward, he awaited
somewhat tremulously the disclosure which he expected after Mr.
Sommerville’s departure. But Mrs. Meredith avoided both of them in the
commotion of her feelings. She shut herself up in her own room to ponder
the question, and, as was natural, her proud impulse of resistance yielded to
reflection. Her heart ached a good deal for poor Beresford, a little for
herself. She, too, would miss something. Something would be gone out of
her life which was good and pleasant. Her heart gave a little sob, a sudden
ache came into her being. Was there harm in it? she asked herself, aghast.
Altogether the day was not a pleasant one for Mrs. Meredith. It seemed to
plunge her back into those agitations of youth from which surely middle
age ought to deliver a woman. It wronged her in her own eyes, making even
her generous temper a shame to her. Had she been too good: as he said—too
kind? an accusation which is hurtful, and means something like insult to a
woman, though to no other creature. Too kind! No expression of contempt,
no insinuated slander can be more stinging than this imputation of having
been too kind. Had she been too kind to her sorrowful neighbour? had she
led him to believe that her kindness was something more than kindness?
She, whose special distinction it was to be kind, whose daily court was
established on no other foundation, whose kindness was the breath of her
nostrils; was this quality, of which she had come to be modestly conscious,
and of which, perhaps, she was a little proud, to be the instrument of her
humiliation? She was not a happy wife, nor indeed a wife at all, except in
distant and not very pleasant recollection, and in the fact that she had a
watchful husband, at the end of the world, keeping guard over her. Was it
possible that she had given occasion for his interference, laid herself open
to his scorn? It seemed to the poor woman as if heaven and earth had
leagued against her. Too kind; suspected by the jealous man who watched
her, despised by the ungrateful man by whom her tender generosity had
been misinterpreted! She sent down a message to Cara that she was not
going out. She sent word to her visitors that she had a headache. She saw
nobody all day long. Too kind! The accusation stung in the tenderest point,
and was more than she could bear.
CHAPTER XXV.
AN IDEALIST.
IN THE ‘HOUSE.’
Roger had not renewed his visit to Cara for some weeks. He had been too
much cast down and discouraged by that first Sunday for which he had
prepared so elaborately, and looked forward to with so much eagerness. But
discouragement, like everything else, wears out, and when he had gone
round the circle from anger to disapproval, from disapproval to contempt,
from contempt to pity, Roger found himself with some surprise back at his
original point, longing to see Cara, and ready to believe that anything that
had come between them had been accidental. The two Merediths would not
be there for ever, and Cara no doubt, poor girl, must be pining for someone
from her old home, and would be glad to see him, and hear all that
everybody was doing. He was sorry he had said a word to his mother about
what happened in the Square; indeed he had done nothing but regret ever
since the indiscretion which tempted him to complain; for Mrs. Burchell
was one of those inconvenient persons who never forget the indignant
criticisms of injured feeling, but continue to repeat and harp upon it long
after that feeling has sunk into oblivion or changed into contempt. Very
soon the softening influences of his early love, and the longing he had after
the object of it, made Roger forgive Cara all her imagined sins against him;
but his mother could not forget that he had been slighted, and punished his
betrayal of his wound by incessant reference to the evils in the Square. This
of itself helped on his recovery, since to find fault yourself with those to
whom you are attached is a very different thing from hearing them assailed
by others. The process ended by a serious quarrel with Mrs. Burchell, who
would not give up this favourite subject, and taunted her son with his want
of proper pride, and inclination to put up with anything, when she heard of
his intention to go back. ‘If I had been so treated anywhere, I would never
go near them again. I would not invite people to trample upon me,’ cried
the Rector’s wife. ‘I might forgive, but I should never forget.’ ‘My dear,’
the Rector had said, ‘Roger has himself to look to: we are not able to do
very much for him; and Cara will be a kind of heiress. I should not mind
any trifle of that sort, if he has serious views.’ ‘What do you call serious
views?’ cried Roger, ashamed and wretched, and he plunged out of the
house without waiting for an answer, and betook himself to those wintry
woods of which Agnes was thinking at the ‘House,’ and which even in
winter were sweet. Roger had no sordid intentions, which was what his
father meant by ‘serious’ views; and though he was well enough satisfied
with his daily work, and not, like Agnes, troubled by any ideal, yet he felt,
like his sister, the wretched downfall of existence into misery and
meanness, between his mother’s prolonged and exaggerated resentment and
his father’s serious worldliness. That boyish love of his was the highest
thing in the young man’s mind. If nothing else that was visionary existed in
his nature, his semi-adoration of Cara, which had lasted as long as he could
recollect, was visionary, a touch of poetry amid his prose, and to hear it
opposed, or to hear it sordidly encouraged alike shocked and revolted him.
He resolved never to mention Cara’s name again, nor to make any reference
to the Square, to shut up his sentiments about her in his own bosom,
whether these were sentiments of admiration or of offence. Supposing she
was cold to him—and it would be very natural that she should be cold, as
he had never gone back to her, nor visited her but once—he would bear it
and make no sign; never again would he subject her name to comments
such as these. Fathers and mothers do badly by their children when they
force them to such a resolution. Roger kept his word all through the weary
Sunday, and did not say even that he would not return home for the next;
but he made his arrangements all the same.
When the next Sunday came the heart of the aunt at Notting Hill was
once more gladdened by the sight of him; and in the afternoon he duly set
out for the Square. Perhaps his dress was not so elaborate nor his necktie so
remarkable as when he first went there. He had sworn to himself that he
would form no special expectations and make no grand preparations, and on
the whole he was happier on his second visit. Miss Cherry, whom he found
at the Square, was very glad to see him, and Mr. Beresford spoke to him
kindly enough, and Cara was sweet and friendly. But they treated his visit
as a call only; they did not ask him to dinner, which was a disappointment.
They offered him a cup of tea, which Roger did not care for, being scarcely
fashionable enough to like five o’clock tea, and let him go when they went
to dinner, forlorn enough, turning him out as it were upon the streets full of
people. To be sure Roger had his aunt at Netting Hill, who was very glad to
see him, who would give him supper and make him very comfortable. Still,
as he had hoped perhaps to be asked to stay, to spend the evening with Cara,
it gave him a very forlorn sensation, when they bade him cheerfully good-
by at the sound of the dinner-bell. He went out into the evening streets,
where many people were going to church, and many coming back from
their afternoon walk, going home to their families in twos and threes.
Scarcely anyone seemed to be alone but himself. Still he said to himself he
had no right to grumble, for they had been kind—and next Sunday he
would go again; and with this melancholy yet courageous resolution he
made a little pause at the corner of the street, asking himself where he
should go now? His aunt would have taken tea and gone to evening church
before he could get to Netting Hill. So he changed his direction and went
manfully the other way, to the ‘House,’ to visit his sister, arguing his
disappointment down. Why should they have asked him to dinner? Besides,
he did not go for dinner, which would have been mercenary, but for Cara—
and he had seen Cara, without those Merediths thrusting themselves into his
way; and she had been very kind, and Miss Cherry had been kind, and there
was no reason why he should not go again next Sunday afternoon. So why
should he be discouraged? There was Agnes, whom he had not seen since
she had gone into this ‘House,’ as they called it. It was only right that a man
should go and look after his own sister, even if he did not approve of her. So
Roger employed his undesired hour of leisure in the way of duty, and went
to see Agnes, gradually calming himself down out of his disappointment on
the way.
The Burchells were not what is called a family devoted to each other.
They were good enough friends, and took a proper brotherly and sisterly
interest in what happened to each other, especially as every new piece of
family news brought a certain amount of enlivenment and variety and a new
subject for conversation into the monotonous family life; but they were
prosaic, and Agnes was the one among them whom the others did not
understand much, and not understanding, set down bluntly as fantastic and
incomprehensible. Had she fallen in love with somebody or had a
‘disappointment,’ they would have entered to a certain degree into her
feelings, and even now Roger could not quite divest himself of the thought,
that, though he knew nothing of it, something of this kind must be at the
root of her withdrawal from home. An ideal life, what was that? Neither
Roger nor any of the rest understood what she could mean, or really