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THE NEUROLOGY OF CONSCIOUSNESS
SECOND EDITION
xii FOREWORD
Observations such as those described above have
also led to a conflation of the phenomenon of attention
with that of consciousness. Some suggest that aware-
ness and perception must interact to provide a
reportable sensory phenomenon or motor action. For
instance, there are those who believe that the defining
characteristic or evidence that can be obtained for a
conscious event is that it is reportable to the self or
others.
The notion of conscious level has proved useful in
clinical medicine, largely because of its prognostic
value. Much recent work with comatose patients has
led to elaboration of schemata of different levels in the
continuum between complete loss of consciousness
and wakefulness. These constructs deal with the state
of awareness in addition to states of access to the con-
tents of consciousness and responses to them. The
interaction between them remains one of the mysteries
of the complex set of brain functions we continue to
call consciousness. These recent advances in classifying
states of impaired consciousness also take into account
the notion of local and global impairments of con-
sciousness. They have used a novel type of report that
itself rests on recordings of brain physiology. The
reproduction of patterns of regionally distributed brain
activity when people imagine complex behavior is
compared to that found in people who cannot commu-
nicate by normal means, when they are invited to
imagine the same behavior. Correspondence has been
found in sufficient cases to suggest that there are con-
scious processes at work in some partially damaged
brains that are capable of integrating various types of
information and relating them to memories. These
observations and their clinical consequences turn con-
sciousness, at least in part, into a social phenomenon
with strong links to communication.
It is becoming clear to the reader that taking the
human neuroscience perspective, normal or pathologi-
cal, the word consciousness remains ambiguous and
so a book that dissects phenomena related to con-
sciousness into individual components is welcome.
What will it require to put the facts about each element
together? It is true there are already models that help
to classify and categorize the different phenomena
associated with consciousness and conscious access
and at least make an attempt to link them together.
The difficulty is that the more the problem of con-
sciousness is investigated, the more one realizes that it
is a difficult problem indeed, involving many func-
tions and many levels of brain organization, each inter-
acting with others in unexplored and unexpected
ways. There is a view—which is not shared by all—
that no global synthesis will be possible without an
understanding of the molecular, cellular, neural, and
physiological principles that govern the various spatial
and temporal levels of brain organization. In the study
of consciousness we may be living at a time analogous
to that which, with the rise of molecular biology,
started the process of understanding how living organ-
isms are constituted some 50 years ago. “Life” was dif-
ficult to define then and “vitalist” theories were still
common. This is no longer the case today. The intro-
duction of a diversity of novel methods and theoretical
paradigms produced new knowledge about bacterial
cells and subsequently about eukaryotic cells too,
despite the continuing absence of a satisfactory a priori
general definition of what “life” is. A similar process
may be in the process of illuminating what conscious-
ness and conscious access are.
The editors and contributors of this excellent text
combine most of the knowledge and data that are cur-
rently available—from normality through disease. For
that they are to be congratulated. Federating this infor-
mation is a prerequisite to integrating it into more gen-
eral processes, which is one of many steps we need to
provide a better understanding of biological mechan-
isms engaged in conscious processing.
Richard Frackowiak
Centre Hospitalier Universitaire Vaudois,
University of Lausanne and Ecole Polytechnique
Fédérale de Lausanne, Lausanne, Switzerland
Jean-Pierre Changeux
Pasteur Institute and Collège de France,
Paris, France
 Preface
Thinking must never submit itself, neither to a dogma, nor
to a party, nor to a passion, nor to an interest, nor to a precon-
ceived idea, nor to anything whatsoever, except to the facts
themselves, because for it to submit to anything else would be
the end of its existence. Henri Poincaré (1854 1912)
‘Truth is sought for its own sake. And those who
are engaged upon the quest for anything for its own
sake are not interested in other things. Finding the
truth is difficult, and the road to it is rough.’ wrote Ibn
al-Haytham (965 1039), a pioneer of the scientific
method. This book addresses one of the biggest chal-
lenges of science; understanding the biological basis of
human consciousness. It does so through observation
and experimentation in neurological patients, formu-
lating hypotheses about the neural correlates of
consciousness and employing an objective and repro-
ducible methodology. This scientific method, as first
proposed by Isaac Newton (1643 1727), has proven
utterly successful in replacing Dark Age ‘magical
thinking’ with an intelligent, rational understanding of
nature. Scientific methodology, however, also requires
imagination and creativity. For instance, methodologi-
cally well-described experiments allowed Louis
Pasteur (1822 1895) to reject the millennia-old
Aristotelian (384 322 BC) view that living organisms
could spontaneously arise from non-living matter.
Pasteur’s observations and genius gave rise to the
germ theory of disease, which would lead to the use of
antiseptics and antibiotics, saving innumerable lives.
The progress of science also largely depends upon
the invention and improvement of technology and
instruments. For example, the big breakthroughs of
Galileo Galilei (1564 1642) were made possible thanks
to eyeglass makers’ improvements in lens-grinding
techniques, which permitted the construction of his
telescopes. Similarly, advances in engineering led to
space observatories such as the Hubble Telescope
shedding light on where we come from. Rigorous sci-
entific measurements permitted to trace back the
beginning of the universe to nearly 14 billion years; the
age of the earth to more than 4.5 billion years; the ori-
gin of life on earth to (very) approximately 3.5 billion
xiii
years; and the apparition of the earth’s first simple ani-
mals to about 600 million years. Natural selection, as
revealed by Charles Darwin (1809 1882) then gave
rise to nervous systems as complex as the human
brain, arguably the most complex object in the uni-
verse. And, somehow, through the interactions among
its 100 billion neurons, connected by trillions of synap-
ses, emerges our conscious experience of the world
and of ourselves.
The study of consciousness has remained within the
scope of philosophy for millennia. Recent empirical
evidence from functional neuroimaging offers a new
way to investigate the mind body conundrum. It also
gives new opportunities to the neurological commu-
nity to improve our understanding and management
of patients with disorders of consciousness. This sec-
ond edition of The Neurology of Consciousness aims at
revising our understanding of the anatomical and
functional underpinnings of human consciousness by
emphasizing a lesion approach through the study of
neurological patients. This second edition seems criti-
cal to us as numerous recent findings and seminal arti-
cles have been published since the first edition of the
book in 2009. The different chapters review the map-
ping of conscious perception and cognition in health
(e.g., wakefulness, sleep, dreaming, sleepwalking and
anaesthesia) and in disease (e.g., post-comatose states,
seizures, split-brains, neglect, amnesia, dementia, and
so on).
‘A genuine glimpse into what consciousness is
would be the scientific achievement, before which all
past achievements would pale’ wrote William James in
1899. Testable hypotheses on consciousness, even if
still far away from solving all problems related to the
neural substrate of consciousness, give us such a
glimpse. In our view, scientific and technological
advances complemented by an adequate theoretical
framework will ultimately lead to an understanding of
the neural substrate of consciousness.
We thank our funding agencies including the
National Institutes of Health, the European Commission,
the McDonnell Foundation, the Mind Science
xiv PREFACE

Foundation Texas, the Belgian National Funds for
Scientific Research (FNRS), the French Speaking
Community Concerted Research Action, the Queen
Elizabeth Medical Foundation, the Belgian American
Education Foundation, the Wallonie-Bruxelles
International, Liège Sart Tilman University Hospital, the
University of Liège and the University of Wisconsin
School of Medicine and Public Health. We learned a lot
while working on this second edition of The Neurology of
Consciousness and we hope you do too while reading it.
Steven Laureys (Liège),
Olivia Gosseries (Liège and Madison),
and Giulio Tononi (Madison)
 List of Contributors
Selma Aybek Laboratory for Behavioral Neurology and
Imaging of Cognition, Department of Neurosciences and
Clinic of Neurology, University Medical Center, Geneva,
Switzerland
Claudio L. Bassetti Department of Neurology, University
Hospital, Bern, Switzerland
Olaf Blanke Laboratory of Cognitive Neuroscience, Brain
Mind Institute, School of Life Sciences, École
Polytechnique Fédérale de Lausanne, Lausanne,
Switzerland; Center for Neuroprosthetics, École
Polytechnique Fédérale de Lausanne, Lausanne,
Switzerland; Department of Neurology, University
Hospital, Geneva, Switzerland
Hal Blumenfeld Departments of Neurology, Neurobiology,
and Neurosurgery, Yale University School of Medicine,
New Haven, CT, USA
Melanie Boly Department of Psychiatry, University of
Wisconsin-Madison, Madison, WI, USA; Department of
Neurology, University of Wisconsin-Madison, Madison,
WI, USA
Marie-Aurélie Bruno Coma Science Group, Neurology
Department and GIGA, University of Liège, Liège,
Belgium
Chris Butler Nuffield Department of Clinical
Neurosciences, University of Oxford, John Radcliffe
Hospital, Oxford, UK
Camille Chatelle Department of Neurology, Massachusetts
General Hospital, Boston, MA, USA; Department of
Physical Medicine and Rehabilitation, Spaulding
Rehabilitation Hospital and Harvard Medical School,
Boston, MA, USA
Athena Demertzi Coma Science Group, Neurology
Department and GIGA, University of Liège, Liège, Belgium
Sebastian Dieguez Laboratory for Cognitive and
Neurological Sciences, Department of Medicine, Hôpital
de Fribourg, Université de Fribourg, Fribourg,
Switzerland
Brian L. Edlow Department of Neurology, Massachusetts
General Hospital, Boston, MA, USA; Athinoula A.
Martinos Center for Biomedical Imaging, Massachusetts
General Hospital, Charlestown, MA, USA
Andreas K. Engel Department of Neurophysiology and
Pathophysiology, University Medical Center Hamburg-
Eppendorf, Hamburg, Germany
xv
Nathan Faivre Laboratory of Cognitive Neuroscience, Brain
Mind Institute, School of Life Sciences, École Polytechnique
Fédérale de Lausanne, Lausanne, Switzerland; Center
for Neuroprosthetics, École Polytechnique Fédérale de
Lausanne, Lausanne, Switzerland
Joseph J. Fins Weill Cornell Medical College and the
Rockefeller University, New York, NY, and Yale Law
School, New Haven, CT, USA
Pascal Fries Ernst Strüngmann Institute for Neuroscience
in Cooperation with Max-Planck-Society, Frankfurt,
Germany; Donders Institute for Brain, Cognition and
Behaviour, Radboud University Nijmegen, Nijmegen, The
Netherlands
Michael S. Gazzaniga Dynamical Neuroscience, University
of California, Santa Barbara, CA, USA; Psychological and
Brain Sciences, University of California, Santa Barbara,
CA, USA
Joseph T. Giacino Department of Physical Medicine and
Rehabilitation, Spaulding Rehabilitation Hospital and
Harvard Medical School, Boston, MA, USA
Olivia Gosseries Department of Psychiatry, University of
Wisconsin-Madison, Madison, WI, USA; Coma Science
Group, Neurology Department and GIGA, University of
Liège, Liège, Belgium
Christof Koch Allen Institute for Brain Science, Seattle,
WA, USA
Andrea Kübler Institute of Psychology, University of
Würzburg, Würzburg, Germany
Ron Kupers Department of Neuroscience and Pharmacology,
University of Copenhagen, Copenhagen, Denmark
Steven Laureys Coma Science Group, Neurology
Department and GIGA, University of Liège, Liège,
Belgium
Nicole L. Marinsek Dynamical Neuroscience, University of
California, Santa Barbara, CA, USA
George A. Mashour Department of Anesthesiology,
Neuroscience Graduate Program, Center for
Consciousness Science, University of Michigan Medical
School, Ann Arbor, MI, USA
Marcello Massimini Department of Biomedical and
Clinical Sciences “Luigi Sacco,” University of Milan,
Milan, Italy
Donatella Mattia Fondazione Santa Lucia, IRCCS, Rome,
Italy
xvi LIST OF CONTRIBUTORS
Michael B. Miller Dynamical Neuroscience, University of
California, Santa Barbara, CA, USA; Psychological and
Brain Sciences, University of California, Santa Barbara,
CA, USA
Lionel Naccache INSERM U1127, Institut du Cerveau et de
la Moelle Epinière, PICNIC Lab, Paris, France, Faculté de
Médecine Pitié-Salpêtrière, Université Pierre et Marie
Curie, Paris, France; Departments of Neurology and of
Clinical Neurophysiology, Assistance Publique Hôpitaux
de Paris, Groupe hospitalier Pitié-Salpêtrière Charles Foix,
Paris, France
Paolo Nichelli Department of Biomedical, Metabolical and
Neural Sciences, University of Modena and Reggio
Emilia, Emilia-Romagna, Italy
Marie-Christine Nizzi Psychology Department, Harvard
University, Cambridge, MA, USA
Adrian M. Owen The Brain and Mind Institute, The
University of Western Ontario, London, ON, Canada
Pietro Pietrini Clinical Psychology Branch, Department of
Neuroscience, University of Pisa Medical School, Pisa,
Italy
Bradley R. Postle Departments of Psychology and
Psychiatry, University of Wisconsin-Madison, Madison,
WI, USA
Maurice Ptito Department of Neuroscience and
Pharmacology, University of Copenhagen, Copenhagen,
Denmark
Geraint Rees Faculty of Life Sciences, School of Life and
Medical Sciences, University College London, London, UK
Mario Rosanova Department of Biomedical and Clinical
Sciences “Luigi Sacco,” University of Milan, Milan, Italy;
Fondazione Europea di Ricerca Biomedica, FERB Onlus,
Milan, Italy
Eric Salmon Cyclotron Research Centre, University of
Liege, Belgium
Nicholas D. Schiff Department of Neurology and
Neuroscience, Weill Medical College of Cornell
University, New York, NY, USA
Caroline Schnakers Department of Psychology and
Neurosurgery, University of California, Los Angeles, CA,
USA
Francesca Siclari Center for Investigation and Research on
Sleep, University Hospital and University of Lausanne,
Switzerland
Giulio Tononi Department of Psychiatry, University of
Wisconsin-Madison, Madison, WI, USA
Naotsugu Tsuchiya School of Psychological Sciences,
Faculty of Biomedical and Psychological Sciences, Monash
University, VIC, Australia
Patrik Vuilleumier Laboratory for Behavioral Neurology
and Imaging of Cognition, Department of Neurosciences
and Clinic of Neurology, University Medical Center,
Geneva, Switzerland
Susan Whitfield-Gabrieli Martinos Imaging Center at
McGovern Institute for Brain Research, Massachusetts
Institute for Technology, Cambridge, MA, USA
Adam Zeman Department of Neurology, University of
Exeter Medical School, Exeter, UK
 C H A P T E R

1
Neuroanatomical Basis of Consciousness
Hal Blumenfeld
Departments of Neurology, Neurobiology, and Neurosurgery, Yale University School of Medicine,
New Haven, CT, USA

O U T L I N E

Introduction 3 Cortical Networks and Consciousness 16

The Cortex and Arousal 16
The Consciousness System 5
Attention and Consciousness 16
Subcortical Networks and Consciousness 6 Hemispheric Dominance of Attention 17
The Thalamus and Consciousness 8 Affect, Motivation, and Attention 18
Glutamatergic and Related Arousal Systems 9 The Binding Problem 18
Cholinergic Arousal Systems 9 Top-Down and Bottom-Up Attention Networks 19
GABAergic Arousal Systems 12 Task-Positive and Task-Negative Networks 19
Noradrenergic Arousal Systems 12 Memory Systems and Consciousness 20
Serotoninergic Arousal Systems 13 Volition and Conscious Free Will 21
Dopaminergic Arousal Systems 13 Self-Awareness and Embodiment 22
Histaminergic Arousal Systems 14 Awareness: Conscious Report and
Orexinergic Arousal Systems 15 Contrastive Analysis 22
Adenosine and Arousal 15
Acknowledgments 23
Amygdala and Arousal 15
Attention and Awareness: Roles of Subcortical References 23
Arousal Systems, Tectal Region, Basal Ganglia,
Claustrum, and Cerebellum 15

INTRODUCTION all the various types of information processed by

Consciousness is of great importance to normal
human quality of life. The nature of consciousness and
the best way to understand and define it have long
generated lively debate among scientists, philosophers,
clinicians, and the general public. From a neurological
perspective, consciousness is classically described as
emerging from brain systems that make up the content of
consciousness, regulated by distinct systems that control
the level of consciousness (Plum and Posner, 1982).
The content of consciousness is the substrate upon
which levels of consciousness act. This content includes
hierarchically organized sensory, motor, emotional, and
memory systems in the brain (Figure 1.1). Much of
neuroscience is dedicated to understanding the normal
functioning of these systems. Selective deficits in con-
tents of consciousness, such as loss of a portion of one’s
visual field, or sudden impairment in spoken language,
are also the main subject matter of clinical neurology.
However, level of consciousness can affect all of
these specific functions. The level of consciousness is
controlled by specialized cortical and subcortical sys-
tems that determine the amount of alertness, attention,
and awareness (mnemonic, AAA) (Blumenfeld, 2002).

S. Laureys, O. Gosseries & G. Tononi (Eds) DOI: http://dx.doi.org/10.1016/B978-0-12-800948-2.00001-7

The Neurology of Consciousness, Second edition. 3 © 2016 Elsevier Ltd. All rights reserved.
4 1. NEUROANATOMICAL BASIS OF CONSCIOUSNESS

FIGURE 1.1 The content of consciousness. Parallel interconnected and hierarchically organized sensory and motor systems receive inputs,
generate outputs, and perform internal processing on multiple levels, from relatively simple to highly abstract. Three additional special
systems—mediating memory, emotions and drives, and consciousness itself—act on the other systems in a widely distributed manner,
especially at the highest levels of processing. Source: Modified with permission from Blumenfeld (2010).

Basic alertness (arousal, wakefulness) is necessary for
any meaningful responses to occur. Attention enables
selective or sustained information to be processed.
Finally, awareness is the ability to form experiences
that can potentially be reported later. This chapter will
review the neuroanatomical basis of brain systems that
control the level of consciousness. In analogy with
other cortical-subcortical systems such as the sensory,
motor or limbic systems, the brain networks dedicated
to regulating the level of consciousness can be referred
to as the “consciousness system” (Blumenfeld, 2010,
2012). This chapter begins with an overview of the
main cortical and subcortical structures that constitute
the consciousness system. Next, the major subcortical
networks that regulate level of consciousness are each
discussed in turn, including the thalamus and sub-
cortical arousal nuclei acting through multiple neuro-
transmitters (glutamate, acetylcholine, gamma amino
butyric acid (GABA), norepinephrine, serotonin, dopa-
mine, histamine, orexin) that arise from the upper
brainstem, basal forebrain, and hypothalamus. The
second half of the chapter reviews important cortical
networks for controlling the level of alertness, atten-
tion, and awareness, including systems that select and
encode conscious experiences into memories for sub-
sequent report. This neuroanatomical review of the
cortical and subcortical systems that control level of
consciousness will serve as a general introduction to the
normal functions as well as disorders of consciousness
discussed in the remaining chapters in this book.

THE NEUROLOGY OF CONSCIOUSNESS

 THE CONSCIOUSNESS SYSTEM 5

FIGURE 1.2 The consciousness system. Anatomical structures involved in regulating the level of consciousness, specifically controlling the
level of alertness, attention and awareness. (A) Medial view showing cortical (blue) and subcortical (red) components of the consciousness sys-
tem. (B) Lateral cortical components of the consciousness system. Note that other circuits not pictured here, such as the anterior insula, claus-
trum, basal ganglia, amygdala, and cerebellum, may also play a role in attention and other aspects of consciousness. Source: Reproduced with
permission from Blumenfeld (2010).

THE CONSCIOUSNESS SYSTEM literature (Heilman and Valenstein, 2003; Mesulam,

The specialized brain networks controlling the level
of consciousness can be referred to as the “conscious-
ness system” (Blumenfeld, 2009, 2010) (Figure 1.2).
It has long been recognized through studies based
on human brain disorders (Penfield, 1950; Plum and
Posner, 1972; Von Economo, 1930) as well as experimen-
tal animal models (Bremer, 1955; Moruzzi and Magoun,
1949; Steriade and McCarley, 2010) that the level of
consciousness depends critically on both cortical and
subcortical structures. Here we provide a brief overview
of the cortical and subcortical networks comprising
the consciousness system, which will be discussed in
greater detail in the remaining sections of the chapter.
Cortical components of the consciousness system
include the major regions of the higher-order “heteromo-
dal” (Mesulam, 2000) association cortex (Figure 1.2; see
also Figure 1.11). On the medial brain surface, important
components are the medial frontal, anterior cingulate,
posterior cingulate, and medial parietal (precuneus,
retrosplenial) cortex (Figure 1.2A). On the lateral
surface, major consciousness system networks include
the lateral frontal, anterior insula, orbital frontal, and
lateral temporal-parietal association cortex (Figure 1.2B).
It is important to recognize that individual
components of the higher-order association cortex play
important and well-studied roles in specific cognitive
functions in the dominant and non-dominant hemi-
spheres as described in the behavioral neurology
2000). Recently these same association cortex regions
have also been described as participating in either so-
called task-positive networks based on their activation
during externally oriented attention (Asplund et al.,
2010; Buschman and Miller, 2007; Dosenbach et al.,
2007; Vanhaudenhuyse et al., 2011) or task-negative
networks, also known as the “default mode” based on
activity at rest (Fox et al., 2005; Raichle et al., 2001).
Regardless of the heterogeneous functions of individual
regions or networks, it is the collective activity of wide-
spread areas of bilateral association cortex that deter-
mines the level of consciousness. Taken as a whole, the
higher-order association cortex interacts with subcorti-
cal arousal systems (Steriade and McCarley, 2010) to
exert powerful control over the overall level of arousal,
attention, and awareness.
Subcortical components of the consciousness system
include the upper brainstem activating systems, thala-
mus, hypothalamus, and basal forebrain (Figure 1.2A).
It is likely that other subcortical structures (not shown)
also participate, including portions of the basal gan-
glia, cerebellum, amygdala, and claustrum. Multiple
parallel neurotransmitter systems participate in
subcortical arousal including acetylcholine, glutamate,
gamma amino butyric acid (GABA), norepinephrine,
serotonin, dopamine, histamine, and orexin (Cooper
et al., 2003; Saper et al., 2005; Steriade et al., 1997;
Steriade and McCarley, 2010). Like the diverse cortical
regions already discussed, these subcortical pathways

THE NEUROLOGY OF CONSCIOUSNESS

6 1. NEUROANATOMICAL BASIS OF CONSCIOUSNESS

FIGURE 1.3 Arousal circuits of the pontomesencephalic reticular formation, thalamus, hypothalamus and basal forebrain. (A) Midsagittal
view; (B) coronal view. Widespread projections to the cortex arise from outputs of the pontomesencephalic reticular formation relayed via the
thalamic intralaminar nuclei, basal forebrain, and hypothalamus. Source: Reproduced with permission from Blumenfeld (2010).

each carry out individual roles, but it is the collective
and parallel actions of all of these systems that
together control the level of consciousness.
Understanding consciousness depends not only on
neuroanatomy but also on neurophysiology. Although
this chapter will focus on the “where” of consciousness,
equally important is “how” these networks interact to
form consciousness. Recent proposed physiological
mechanisms for consciousness include synchronized
oscillations (Buzsaki and Wang, 2012; Llinás and Paré,
1997; Singer, 1998), slow cortical potentials (Li et al.,
2014), connectivity (Boly et al., 2011; Rosanova et al.,
2012; Rubinov and Sporns, 2010), information
integration (Tononi, 2005; Tononi and Koch, 2008), pop-
ulation neuroenergetics (Shulman et al., 2003), and
recurrent or global neuronal processing (Dehaene et al.,
1998; Lamme and Roelfsema, 2000; Sergent and Dehaene,
2004) among others. Much additional work is needed
before the physiological mechanisms of consciousness
are more definitely known. By contrast, when it comes to
neuroanatomy, the past century of research has at least
led to a basic understanding of the most important brain
structures contributing to consciousness. We now turn in
greater detail to these major cortical and subcortical net-
works that constitute the consciousness system.
SUBCORTICAL NETWORKS
AND CONSCIOUSNESS
The main subcortical components of the conscious-
ness system include the midbrain and upper pons, thal-
amus, hypothalamus, and basal forebrain (Figure 1.2).
These structures contribute importantly to maintaining
alertness and arousal. Attention and awareness are also
facilitated by the same midline arousal systems, as well as
by other subcortical networks including the superior colli-
culi, cerebellum, amygdala, basal ganglia, claustrum, and
thalamic reticular nucleus (Crick and Koch, 2005; Dreher
and Grafman, 2002; Krauzlis et al., 2013; O’Halloran et al.,
2012; Zikopoulos and Barbas, 2012).
In terms of alertness and arousal, much has been
learned about the basic anatomy of consciousness by
understanding which brain lesions can cause coma.
Coma is as a state of unarousable unresponsiveness in
which the eyes are closed and no purposeful responses
can be elicited (Fisher, 1969; Plum and Posner, 1972).
Coma occurs either through bilateral damage to wide-
spread cortical areas, or via lesions in a core set of struc-
tures lying in upper brainstem and medial diencephalon.
These critical subcortical arousal structures were initially
identified based on strokes and other localized disorders
in human patients (Penfield, 1950; Plum and Posner,
1972; Von Economo, 1930) as well as lesion, disconnec-
tion, and stimulation experiments performed in animal
models (Bremer, 1955; Moruzzi and Magoun, 1949;
Steriade and McCarley, 2010). In the brainstem, the
subcortical arousal systems begin in the upper pons
and extend to the midbrain. Lesions in this small but
critical region of the upper pons and midbrain pro-
duce profound coma, whereas lesions in the lower
pons or medulla do not typically disrupt conscious-
ness (Figure 1.3). The core brainstem arousal systems
lie in the tegmentum and include a variety of nuclei
embedded within the brainstem reticular formation.
The tegmentum is sandwiched between the more
ventral brainstem basis—containing ascending and
descending white matter pathways; and the more

THE NEUROLOGY OF CONSCIOUSNESS

 SUBCORTICAL NETWORKS AND CONSCIOUSNESS 7
TABLE 1.1 Widespread Projection Systems in the Nervous System
Projection Neurotransmitter
system Location(s) of cell bodies Main target(s) receptor(s)a,b Function(s)c

Reticular Midbrain and rostral pons Thalamic intralaminar nuclei, Unknown Alertness
formation hypothalamus, basal forebrain (glutamate?)d
Intralaminar Thalamic intralaminar nuclei Cortex, striatum (Glutamate?) Alertness
nuclei
Midline Midline thalamic nuclei Cortex (Glutamate?) Alertness
thalamic nuclei
Norepinephrine Pons: locus ceruleus and lateral Entire CNS α1A D, α2A D, β1 3 Alertness, attention, mood
tegmental area elevation
Dopamine Midbrain: substantia nigra Striatum, limbic cortex, D1 5 Movements, initiative,
pars compacta and ventral amygdala, nucleus accumbens, working memory
tegmental area prefrontal cortex

Serotonin Midbrain, pons, and medulla: Entire CNS 5-HT1A F, 5-HT2A C, Alertness, mood elevation,
raphe nuclei 5-HT3 7 breathing control
Histamine Hypothalamus: tubero-mammillary Entire brain H1 4 Alertness
nucleus; midbrain: reticular
formation
Orexin Posterior lateral hypothalamus Entire brain OX1, OX2 Alertness, food intake
(hypocretin)
Acetylcholine Basal forebrain: nucleus basalis, Cerebral cortex including Muscarinic (M1 5), Alertness, memory
medial septal nucleus, and nucleus hippocampus nicotinic subtypes
of diagonal band

Pontomesencephalic region: Thalamus, cerebellum, Muscarinic (M1 5), Alertness, memory

pedunculopontine nucleus and pons, medulla nicotinic subtypes
laterodorsal tegmental nucleus
a
Many of the neurons releasing the neuromodulatory transmitters listed here also release a variety of peptides, which are likely to play a neuromodulatory role as well.
b
Additional receptor subtypes are constantly being added.
c
Functions listed are highly simplified here; see references at the end of this chapter for additional details.
d
Entries in parenthesis with question mark are uncertain.
Source: Modified with permission from Blumenfeld (2010).

dorsal tectum—lying dorsal to the cerebral aqueduct
or fourth ventricle. Lesions outside the brainstem
tegmentum in the basis or tectum do not produce
coma. Bilateral lesions of the thalamus, particularly
in the intralaminar and midline thalamic nuclei, can
also produce profound suppression of arousal.
Subsequent work has revealed that the subcortical
arousal systems consist of multiple parallel neurotrans-
mitter systems and pathways (Figure 1.3; Table 1.1).
Unlike most pathways in the nervous system which
project to relatively narrow target regions, the sub-
cortical arousal systems belong to a set of widespread
projecting systems (Table 1.1) that reach many struc-
tures or even the entire nervous system. Interestingly
the projection systems arising from the upper brain-
stem including the midbrain and upper pons (ponto-
mesencephalic reticular formation; Figure 1.3) tend
to project upward to the cortex, diencephalon, and basal
forebrain while those in the lower pons and med-
ulla project downward to the brainstem, cerebellum,
and spinal cord. The upward projecting systems were
originally called the “ascending reticular activating sys-
tem” (ARAS) (Moruzzi and Magoun, 1949) recognizing
their important role in arousal. Although the term
“ARAS” is still occasionally used, in reality these
arousal systems arise from a variety of specific nuclei
(Table 1.1) rather than from what was formerly consid-
ered a single diffusely organized system.
The subcortical arousal systems in the midbrain and
upper pons have three main targets (Figure 1.3):
(i) Putative glutamatergic neurons from the reticular
formation and cholinergic neurons from the pedun-
culopontine tegmental nucleus and laterodorsal
tegmental nucleus (LDT) project mainly to the thala-
mus, particularly to the thalamic intralaminar nuclei
which, in turn, increase cortical arousal. (ii) Other
neurons project to the nucleus basalis and hypothala-
mus, which again relay arousal influences to the cortex.
(iii) Finally, the monaminergic neurotransmitter
systems (norepinephrine, dopamine, serotonin; not
shown in Figure 1.3) project directly to the entire fore-
brain including the cortex and subcortical structures.

THE NEUROLOGY OF CONSCIOUSNESS

8 1. NEUROANATOMICAL BASIS OF CONSCIOUSNESS
FIGURE 1.4 Major inputs to the pontomesencephalic reticular
formation and related structures. Source: Reproduced with permission
from Blumenfeld (2010).
In addition to these ascending connections, the subcor-
tical arousal systems are also highly interconnected
and strongly influence each other’s function through
multiple connections within and between the brain-
stem, thalamus, hypothalamus, and basal forebrain.
The upper brainstem arousal systems are influenced
by a variety of inputs including numerous regions of the
association cortex and limbic cortex, as well as sensory
pathways such as the anterolateral pain transmission
pathways (Figure 1.4). Inhibitory influences arise from
the thalamic reticular nucleus (not to be confused with
the reticular formation) as well as other GABAergic
inputs (Parent and Steriade, 1981, 1984; Ropert and
Steriade, 1981). The arousal systems are also strongly
regulated by brainstem and hypothalamic circuits con-
trolling circadian sleep rhythms (Saper et al., 2005, 2010).
The subcortical arousal systems will now each be
discussed in greater detail to more fully appreciate
the functional anatomy of these complex parallel
arousal systems and their contributions to conscious-
ness. It should be noted that, unlike gross lesions of
the brainstem-diencephalic arousal systems, lesions
or pharmacological blockade of the individual pro-
jecting neurotransmitter systems do not cause coma.
Blockade of some neurotransmitters, especially ace-
tylcholine or histamine, produces severe confusion
and drowsiness, but not coma. Thus, the normal
awake, conscious state does not depend on a single
projection system, but rather on the parallel action of
multiple anatomical and neurotransmitter systems
acting together (Table 1.1). After discussing each of
the major subcortical arousal systems, emphasizing
their role in maintaining the alert state, we will then
briefly discuss their role together with other subcorti-
cal structures (tectal region, basal ganglia, claustrum,
cerebellum) in attention and awareness.
THE NEUROLOGY OF CONSCIOUSNESS
The Thalamus and Consciousness
Nearly all information destined for the cortex first
reaches the thalamus. The thalamus transmits this
information and then receives an even greater number
of reciprocal connections back from the cortex.
Therefore the thalamus plays a key role in all aspects
of forebrain function including consciousness. The
thalamus relays the content of consciousness, and also
controls the level of consciousness through specialized
circuits that regulate the level of arousal and are
crucial for selective attention.
Organization of the thalamus can be described
based on regions or based on projections. The regional
organization of the thalamus divides the thalamic sub-
nuclei proceeding from lateral to medial (Figure 1.5)
into the thalamic reticular nucleus located most
laterally, followed by the lateral nuclear group which
contains the largest number of thalamic relay nuclei
(Table 1.2). Continuing medially, next comes the
Y-shaped white matter internal medullary lamina
which separates the lateral, anterior, and medial
nuclear groups from each other (Figure 1.5). Embedded
within the internal medullary lamina lie the intra-
laminar nuclei. Finally a thin layer of midline thalamic
nuclei are located most medially, adjacent to the third
ventricle (Figure 1.5).
Projection patterns can also be used to classify the
thalamic subnuclei (Table 1.2). Some, such as the
ventral posterior lateral nucleus, a somatosensory
relay in the lateral thalamus, project to a relatively
localized region of cortex, and are referred to as
specific relay nuclei. Others, such as the thalamic intra-
laminar nuclei, have more widespread projections to
many cortical areas, and are called diffusely or widely
projecting (“nonspecific”) nuclei.
The specific thalamic relay nuclei communicate with
the cortex regarding each sensory and motor function,
and are therefore responsible for all the individual
contents of consciousness. On the other hand, the
widely projecting thalamic nuclei influence the overall
level of cortical arousal, and therefore control the
level of consciousness. The rostral intralaminar nuclei
(central lateral, paracentral, central medial nuclei;
Table 1.2) and midline thalamic nuclei are thought to
be particularly important for activating the cortex
(Figure 1.3). As was already discussed, the intralaminar
thalamus plays a key role in transmitting arousal influ-
ences from the midbrain and upper pontine cholinergic
and glutamatergic systems to the cortex.
The thalamic reticular nucleus forms a thin shell of
predominantly GABAergic inhibitory neurons on the
lateral thalamus (Figure 1.5). As axons traverse this
nucleus traveling from thalamus to cortex or from
cortex back to thalamus they give off collateral
 SUBCORTICAL NETWORKS AND CONSCIOUSNESS 9

FIGURE 1.5 The thalamus. Main nuclear divisions and nuclei are shown (see also Table 1.2). The posterior portion of the reticular nucleus
has been removed. Source: Reproduced with permission from Blumenfeld (2010).

branches to the thalamic reticular neurons. The
thalamic reticular neurons, in turn, project to the thala-
mus and inhibit the specific thalamocortical neurons
corresponding to individual corticothalamic loops.
The reciprocal connections between thalamic relay
nuclei and the thalamic reticular nucleus are thought
to play an important role in generating corticotha-
lamic rhythms during normal sleep and waking
activity, as well as in pathological rhythms such as
epilepsy (McCormick, 2002; McCormick and Bal, 1997;
McCormick and Contreras, 2001; Steriade et al., 1993b).
These physiological rhythms are crucial for regulating
the level of consciousness. In addition, the thalamic
reticular nucleus influences arousal through long-range
inhibitory projections to the pontomesencephalic retic-
ular formation (Parent and Steriade, 1984). Selective
attention may also be mediated through the particular
arrangement of reticular thalamic neurons and their
directed inhibitory projections to the thalamus, which
are capable of generating an inhibitory surround
around a “searchlight” of focused attention in a narrow
band of thalamocortical channels (Crick, 1984; Mayo,
2009; Pinault, 2004).
Glutamatergic and Related Arousal Systems
Glutamate is the most prevalent excitatory neuro-
transmitter of the central nervous system. For many of
the arousal systems the most likely neurotransmitter is
glutamate, although it has not been identified with
certainty (Steriade and McCarley, 2010). Arousal
system pathways probably mediated by glutamate
include those arising from the midbrain and upper
pontine reticular formation that project to the thalamus
and basal forebrain (Steriade, 2004; Steriade et al.,
1993a), as well as the widespread projections from the
thalamic intralaminar nuclei to the cortex (Figure 1.3).
It is not known whether other excitatory amino acid
neurotransmitters such as aspartate might also play a
significant role in arousal.
Cholinergic Arousal Systems
Acetylcholine is the major neurotransmitter of the
peripheral nervous system, but in the central nervous sys-
tem it has a more neuromodulatory function, where its
role in arousal has been studied extensively. The two
main sources of cholinergic projections neurons in the
central nervous system lie in the brainstem pontomesen-
cephalic reticular formation and in the basal forebrain
(Figure 1.6; Table 1.1). At the junction of the midbrain and
pons, the pedunculopontine nucleus is located in the lat-
eral reticular formation, while the laterodorsal tegmental
nucleus lies in the periaqueductal gray (Mesulam et al.,
1983). The pedunculopontine nucleus stretches from the
caudal midbrain substantia nigra pars reticulata into the
rostral pons towards the superior cerebellar peduncle
(Mena-Segovia et al., 2009; Rye et al., 1987). The nucleus
has a gradient of increasing cholinergic and decreasing
GABAergic neurons as it extends caudally, and also con-
tains glutamatergic neurons (Wang and Morales, 2009).
Cholinergic neurons from these brainstem nuclei project
to the thalamus, including the intralaminar nuclei, play-
ing an important role in arousal. Brainstem cholinergic

THE NEUROLOGY OF CONSCIOUSNESS

TABLE 1.2 Major Thalamic Nuclei
Nucleia
RELAY NUCLEI
Lateral nuclear group
Ventral posterior lateral
nucleus
Ventral posteromedial
nucleus
Lateral geniculate nucleus
Medial geniculate nucleus
Ventral lateral nucleus
Ventral anterior nucleus
Pulvinar
Lateral dorsal nucleus
Lateral posterior nucleus
Ventral medial nucleus
Medial nuclear group
Mediodorsal nucleus (MD)
Anterior nuclear group
Anterior nucleus
Midline thalamic nuclei
Paraventricular, parataenial,
interanteromedial, inter-
mediodorsal, rhomboid,
reuniens (medial ventral)
INTRALAMINAR NUCLEI
Rostral intralaminar nuclei
Central medial nucleus
Paracentral nucleus
Central lateral nucleus
Caudal intralaminar nuclei
Centromedian nucleus
Parafascicular nucleus
RETICULAR NUCLEUS
Reticular nucleus
a
Some additional, smaller nuclei have not been included here.
b
In addition to the inputs listed, all thalamic nuclei receive reciprocal inputs from the cortex and from the thalamic reticular nucleus. Modulatory cholinergic, noradrenergic, serotonergic,
and histaminergic inputs also reach most thalamic nuclei.
c
1 represents least diffuse (specific thalamic relay nuclei);11 represents moderately diffuse; 11 1 represents most diffuse.
Source: Modified with permission from Blumenfeld (2010).
Diffuseness of
projections to
Main inputsb Main outputs cortexc Proposed functions
Medial lemniscus, Somatosensory cortex 1 Relays somatosensory spinal
spinothalamic tract inputs to cortex
Trigeminal lemniscus, Somatosensory and taste 1 Relays somatosensory cranial
trigeminothalamic tract, cortex nerve inputs and taste to
taste inputs cortex
Retina Primary visual cortex 1 Relays visual inputs to cortex
Inferior colliculus Primary auditory cortex 1 Relays auditory inputs to
cortex
Internal globus pallidus, deep Motor, premotor, and 1 Relays basal ganglia and
cerebellar nuclei, substantia supplementary motor cerebellar inputs to cortex
nigra pars reticulata cortex
Substantia nigra pars reticulata, Widespread to frontal 11 1 Relays basal ganglia and
internal globus pallidus, deep lobe, including cerebellar inputs to cortex
cerebellar nuclei prefrontal, premotor,
and supplementary
motor cortex
Tectum (extrageniculate visual Parietotemporo-occipital 11 Behavioral orientation toward
pathway), other sensory inputs association cortex relevant visual and other
stimuli
See anterior nucleus 11 Functions with anterior nuclei
See pulvinar 11 Functions with pulvinar
Midbrain reticular formation Widespread to cortex 11 1 May help maintain alert,
conscious state
Amygdala, olfactory cortex, Frontal cortex 11 Limbic pathways, major relay
limbic basal ganglia to frontal cortex
Mammillary body, hippocampal Cingulate gyrus 1 Limbic pathways
formation
Hypothalamus, basal forebrain, Amygdala, hippocampus, 11 Limbic pathways
amygdala, hippocampus limbic cortex
Deep cerebellar nuclei, globus Cerebral cortex, striatum 11 1 Maintain alert consciousness;
pallidus, brainstem ascending motor relay for basal ganglia
reticular activating systems and cerebellum
(ARAS), sensory pathways
Globus pallidus, ARAS, sensory Striatum, cerebral cortex 11 1 Motor relay for basal ganglia
pathways
Cerebral cortex, thalamic relay Thalamic relay and None Regulates state of other
and intralaminar nuclei, ARAS intralaminar nuclei, ARAS thalamic nuclei
 SUBCORTICAL NETWORKS AND CONSCIOUSNESS 11

FIGURE 1.6 Cholinergic projection systems. (A) Overview and inset showing axial section through caudal midbrain. (B) Coronal section
through basal forebrain. See also Table 1.1. Source: Reproduced with permission from Blumenfeld (2010).

arousal is thought to act synergistically with non-
cholinergic putative glutamatergic pontomesencephalic
neurons that project to intralaminar thalamus and basal
forebrain (Figure 1.3) (Rasmusson et al., 1994, 1996;
Steriade, 2004; Steriade et al., 1993a). In sleep, pontogen-
iculate waves arise from cholinergic brainstem neurons
projecting to thalamocortical neurons in the lateral genic-
ulate nucleus (Steriade et al., 1989, 1990). The pedunculo-
pontine nucleus also has numerous ascending and
descending motor projections and is involved in
controlling locomotion (Inglis and Winn, 1995).
Interestingly, the brainstem has very few direct
cholinergic projections to the cortex and nearly all
facilitatory effects of the brainstem cholinergic systems
on cortex are mediated via the thalamus (Beninato and
Spencer, 1987; Cornwall et al., 1990; Hallanger et al.,
1987; Hallanger and Wainer, 1988; Heckers et al., 1992;
Jones and Webster, 1988; Satoh and Fibiger, 1986). The
major source of cholinergic input to the cortex is the
basal forebrain (Figure 1.6; Table 1.1). Cholinergic
neurons in the nucleus basalis of Meynert and
surrounding regions (substantia innominata, globus
pallidus, and preoptic magnocellular nucleus) not only
project to almost the entire neocortex (Mesulam et al.,
1983; Rye et al., 1984) but also innervate some nuclei in
the thalamus (reticular thalamic, mediodorsal, antero-
ventral/anteromedial, and ventromedial nuclei)
(Heckers et al., 1992; Parent et al., 1988; Steriade et al.,
1987). The hippocampal archicortex, however,
receives cholinergic inputs mainly from the medial
septal nuclei and nucleus of the diagonal band of
Broca (Rye et al., 1984). Additional cholinergic neu-
rons lie in the medial habenula, and short-range cho-
linergic neurons are present in the striatum and to a
more limited extent within the cortex itself. Like the
brainstem cholinergic nuclei, the basal forebrain con-
tains both cholinergic and non-cholinergic neurons,
including GABA and glutamate as transmitters
among others (Brashear et al., 1986; Alvaro Duque
et al., 2007).
The brainstem and basal forebrain cholinergic
systems work together to abolish cortical slow wave

THE NEUROLOGY OF CONSCIOUSNESS

12 1. NEUROANATOMICAL BASIS OF CONSCIOUSNESS
FIGURE 1.7 Noradrenergic projection systems. See also Table 1.1. Source: Reproduced with permission from Blumenfeld (2010).
activity and promote an alert state (Dringenberg and
Olmstead, 2003; Steriade, 2004). Cholinergic arousal in
the central nervous system is mediated predominantly
by muscarinic acetylcholine receptors, although nico-
tinic receptors may also play an important role in
arousal and attention (Bloem et al., 2014). As would be
expected from the connections of cholinergic neurons
described here, pharmacological blockade of central
cholinergic neurotransmission produces an acute state
of delirium and memory loss. However, despite the
importance of acetylcholine in consciousness, selective
damage to cholinergic neurotransmission does not pro-
duce coma (Blanco-Centurion et al., 2006, 2007; Buzsaki
et al., 1988; Fuller et al., 2011), presumably because
of the multiple parallel neurotransmitter systems
participating in consciousness as already discussed.
GABAergic Arousal Systems
Found in local inhibitory interneurons throughout
the cortex and subcortical structures, GABA is the
most prevalent inhibitory neurotransmitter in the cen-
tral nervous system and plays a major role in regulat-
ing arousal. Several long-range GABAergic projection
systems also contribute to controlling arousal. Some
GABAergic neurons in the basal forebrain are thought
to promote arousal because these inhibitory neurons in
turn project to cortical inhibitory interneurons (Freund
and Meskenaite, 1992). However, the overall effects of
basal forebrain GABAergic neurons on arousal may
be heterogeneous because of variable firing patterns
with respect to cortical activation and sleep-wake
cycles (Hassani et al., 2009; Jones, 2004; Manns et al.,
2000); and because parvalbumin-containing GABAergic
neurons are related to cortical desynchrony whereas
neuropeptide Y-containing neurons may have the
opposite effect (Alvaro Duque et al., 2000, 2007).
Other important long-range GABAergic projections
mainly inhibit arousal. These include neurons in the
THE NEUROLOGY OF CONSCIOUSNESS
ventral lateral preoptic nucleus, which have wide-
spread inhibitory projections to virtually all subcortical
arousal systems (Saper et al., 2010); lateral septal
GABAergic neurons thought to inhibit the basal fore-
brain and hypothalamus (Mesulam and Mufson, 1984;
Varoqueaux and Poulain, 1999); and the thalamic
reticular nucleus which contains GABAergic neurons
projecting both to the remainder of the thalamus and
to the brainstem reticular formation (Parent and
Steriade, 1984; Steriade et al., 1984). In addition,
GABAergic neurons in the globus pallidus internal
segment inhibit regions of the thalamus including the
intralaminar nuclei. It has been proposed that paradox-
ical arousal effects of GABA agonists such as zolpidem
in minimally conscious state, or benzodiazepines in
catatonia may occur when these agents inhibit the glo-
bus pallidus, thereby removing tonic inhibition of the
intralaminar thalamus (Brown et al., 2010; Giacino
et al., 2014). Activation of these multiple GABAergic
inhibitory projections converging on the subcortical
arousal systems has also been proposed as the mecha-
nism for loss of consciousness in partial seizures
(Blumenfeld, 2012; Englot and Blumenfeld, 2009).
Noradrenergic Arousal Systems
Norepinephrine (noradrenaline)-containing neurons
are located in the locus ceruleus in the rostral pons
adjacent to the fourth ventricle, as well as in the nearby
lateral tegmental area extending into the more caudal
pons and medulla (Figure 1.7; Table 1.1). Ascending
noradrenergic projections reach the cortex, thalamus
and hypothalamus (Foote et al., 1983; Morrison et al.,
1981; Pickel et al., 1974) to regulate sleep-wake cycles,
attention, and mood, while descending projections to
the brainstem, cerebellum, and spinal cord modulate
autonomic function and gating of pain. Many attention-
enhancing drugs such as amphetamines augment
noradrenergic function. Norepinephrine is thought to
 SUBCORTICAL NETWORKS AND CONSCIOUSNESS
FIGURE 1.8 Serotonergic projections systems. See also Table 1.1. Source: Reproduced with permission from Blumenfeld (2010).
play an important role in promoting arousal (Berridge,
2008; Berridge et al., 2012; Constantinople and Bruno,
2011). For example, selective α-2 agonists such as
clonidine or the anesthetic agent dexmedetomidine
markedly depress arousal possibly by inhibiting locus
ceruleus neurons (Correa-Sales et al., 1992; De Sarro
et al., 1987; Scheinin and Schwinn, 1992). However,
selective removal or blockade of noradrenergic neurons
affects arousal but does not produce coma (Berridge
et al., 1993; Blanco-Centurion et al., 2004, 2007; Cirelli
and Tononi, 2004; Hunsley and Palmiter, 2003) again
reinforcing the notion of multiple parallel systems
promoting consciousness.
Serotoninergic Arousal Systems
Serotonergic neurons are found predominantly in the
midline raphe nuclei of the midbrain, pons, and medulla
(Figure 1.8; Table 1.1). The more rostral serotonergic
neurons in the midbrain and upper pontine raphe
nuclei project to the entire forebrain, participating in
sleep-wake regulation; dysfunction of serotonergic
systems is thought to play a role in a number of
psychiatric disorders including depression, anxiety,
obsessive-compulsive disorder, aggressive behavior, and
eating disorders. More caudal serotonergic neurons in the
pons and medulla are important for modulation of
breathing, pain, temperature control, cardiovascular, and
motor function.
The most important rostral raphe nuclei participating
in arousal are the dorsal raphe and median raphe
(Jacobs and Azmitia, 1992; Wiklund et al., 1981). The role
of serotonergic neurons in arousal is complex, possibly
because the large diversity of serotonin receptors
(Hannon and Hoyer, 2008) leads to effects that either
THE NEUROLOGY OF CONSCIOUSNESS
13
promote or inhibit arousal in different brain regions
(Dugovic et al., 1989; Dzoljic et al., 1992; Kumar et al.,
2007; Lemoine et al., 2007; Leonard and Llinás, 1994;
Luebke et al., 1992; Monckton and McCormick, 2002;
Muraki et al., 2004; Rogawski and Aghajanian, 1980).
Rostral brainstem serotonergic neurons have been pro-
posed to promote arousal in response to hypoventilation
and increased carbon dioxide levels, perhaps playing an
important role in preventing sudden infant death syn-
drome and sudden unexplained death in epilepsy
(Buchanan and Richerson, 2010; Kinney et al., 2009;
Richerson and Buchanan, 2011; Sowers et al., 2013).
Dopaminergic Arousal Systems
Most dopaminergic neurons are located in the
ventral midbrain, either in the substantia nigra pars
compacta or in the adjacent ventral tegmental area
(Figure 1.9; Table 1.1). These mesencephalic nuclei give
rise to the following three ascending dopaminergic
projection systems: (i) the mesostriatal (nigrostriatal)
pathway projects from the substantia nigra to the
caudate and putamen (Matsuda et al., 2009); (ii) the
mesolimbic pathway arises mainly from the ventral
tegmental area and projects to limbic structures
including the medial temporal lobe, amygdala, cingu-
late gyrus, septal nuclei, and nucleus accumbens
(Fallon, 1981; Oades and Halliday, 1987); (iii) the
mesocortical pathway arises mainly from the ventral
tegmental area as well as scattered neurons in the
vicinity of the substantia nigra and ventral periaque-
ductal gray, projecting to the prefrontal cortex
(Figure 1.9) as well as to the thalamus (Garcia-Cabezas
et al., 2009; Groenewegen, 1988; Lu et al., 2006;
Sanchez-Gonzalez et al., 2005). Dopaminergic neurons
14 1. NEUROANATOMICAL BASIS OF CONSCIOUSNESS

FIGURE 1.9 Dopaminergic projections systems. See also Table 1.1. Source: Reproduced with permission from Blumenfeld (2010).

in the ventral tegmental area also project caudally to
various brainstem nuclei and to the spinal cord (Oades
and Halliday, 1987).
Dopamine may contribute to maintaining the
waking state at least in part through effects on other
subcortical arousal circuits (Deutch et al., 1986;
Lu et al., 2006; Neylan et al., 1992; Qu et al., 2008;
Volkow et al., 2009). Effects of dopamine on the
thalamus and cortex can be either excitatory or inhibi-
tory (Bandyopadhyay and Hablitz, 2007; Govindaiah
et al., 2010; Lavin and Grace, 1998; Penit-Soria et al.,
1987). Impaired dopaminergic transmission to the pre-
frontal cortex has been proposed to be important for
the apathetic negative symptoms of schizophrenia, and
may also contribute to states of markedly reduced
motivation, initiative and action/intention seen in
frontal lobe disorders, abulia, and akinetic mutism
(Combarros et al., 2000; Kim et al., 2007; Yang et al.,
2007). Amantadine improves arousal in chronic disor-
ders of consciousness, although it is unclear whether
the mechanism is through enhanced dopaminergic
neurotransmission or effects of this medication on
other neurotransmitter systems (Giacino et al., 2012).
Histaminergic Arousal Systems
Histamine-containing neurons are found mainly in
the tuberomamillary nucleus (Panula et al., 1984) of
the posterior hypothalamus (Figure 1.10; Table 1.1),
although a few scattered histaminergic neurons are
also found in the midbrain reticular formation.
Widespread ascending projections of histaminergic
neurons from the tuberomamillary nucleus reach
nearly the entire forebrain including cortex and thala-
mus, while descending projections target the brainstem
and spinal cord (Brown et al., 2001; Hong and Lee,
2011; Lin et al., 1996; Panula et al., 1989).
Anti-histamine medications are intended to act on
peripheral histamine release from mast cells, but are
well-known to induce drowsiness presumably through
central actions (White and Rumbold, 1988). Histamine
can produce arousal effects on cortex (Dringenberg

THE NEUROLOGY OF CONSCIOUSNESS

 SUBCORTICAL NETWORKS AND CONSCIOUSNESS
FIGURE 1.10 Histaminergic projections systems. See also
Table 1.1. Source: Reproduced with permission from Blumenfeld (2010).
and Kuo, 2003; McCormick and Williamson, 1989) and
thalamus (McCormick and Williamson, 1991). In addi-
tion to the cortex and thalamus, arousal actions of
histamine may be mediated by activation of other sub-
cortical arousal systems including other hypothalamic
nuclei (Brown et al., 2001; Lin et al., 1994), the basal
forebrain (Dringenberg and Kuo, 2003; Khateb et al.,
1995; Zant et al., 2012), brainstem cholinergic (Khateb
et al., 1990; Lin et al., 1996), and noradrenergic nuclei
(Korotkova et al., 2005). Effects of histamine are
receptor-dependent as activation of H1 receptors pro-
motes wakefulness, whereas H3-receptors appear to
have the opposite role (Huang et al., 2006; Khateb
et al., 1990; Lin et al., 1990, 1996; Valjakka et al., 1996).
Orexinergic Arousal Systems
Orexin (hypocretin) is a peptide produced in
neurons of the perifornical, lateral, and posterior
hypothalamus (Peyron et al., 1998; Sakurai et al., 1998),
which project to both cortex and virtually all sub-
cortical arousal systems (Chemelli et al., 1999; Peyron
et al., 1998) to promote the awake state. The arousal
effects of orexin likely arise from both cortical and
subcortical mechanisms (Bourgin et al., 2000; Eriksson
et al., 2001; España et al., 2001; Hagan et al., 1999;
Horvath et al., 1999; Kiyashchenko et al., 2002;
Tsunematsu et al., 2011; van den Pol et al., 2002).
Abnormalities of the orexin system are thought
to play a role in narcolepsy, a disorder characterized
by excessive daytime sleepiness and pathological
transitions into rapid eye movement sleep (Anaclet et al.,
2009; Chemelli et al., 1999; Gerashchenko et al.,
2003; Hara et al., 2001; Lin et al., 1999; Nishino et al., 2000;
THE NEUROLOGY OF CONSCIOUSNESS
15
Peyron et al., 2000; Thannickal et al., 2000). The beneficial
effects of modafenil in preventing the symptoms of
narcolepsy may in part be through activation of orexin
neurons (Chemelli et al., 1999).
Adenosine and Arousal
Although the neuroanatomical sources of adenosine
are not well characterized, this neuromodulator may
be important in mechanisms of conscious arousal
(Huang et al., 2014; Liu and Gao, 2007). The effects of
adenosine on arousal are generally inhibitory, and cir-
cadian fluctuations in adenosine levels peak just prior
to the initiation of sleep. Adenosine receptors are
found in both cortex and thalamus, where they have
an overall inhibitory function on arousal. Caffeine
blocks adenosine receptors and this may be one impor-
tant mechanism whereby coffee promotes alertness
(Huang et al., 2011; Lazarus et al., 2011).
Amygdala and Arousal
Because the amygdala has widespread and reciprocal
cortical-subcortical connections that contribute to aro-
usal particularly in response to emotions, it is appro-
priate to include this complex of nuclei located in the
anteromedial temporal lobe as an important subcortical
component of the consciousness system (Steriade and
McCarley, 2010). The main components of the amygda-
loid nuclear complex are the corticomedial, basolateral,
and central nuclei, as well as the bed nucleus of the
stria terminalis. The basolateral nucleus is largest in
humans and has widespread direct and indirect connec-
tions to the cortex, basal forebrain, and medial thalamus
(Aggleton, 2000; LeDoux, 2007). The smaller cortico-
medial nucleus participates in appetitive states via the
hypothalamus, as well as in olfaction. The central
nucleus, although smallest, has important connections
with the hypothalamus and brainstem participating in
arousal and autonomic control (LeDoux, 2007).
Attention and Awareness: Roles of Subcortical
Arousal Systems, Tectal Region, Basal Ganglia,
Claustrum, and Cerebellum
To complete our discussion of subcortical networks
regulating the level of consciousness, it is important to
again emphasize the functions of the consciousness
system in controlling alertness, attention, and aware-
ness, and to briefly mention several additional sub-
cortical structures participating in these functions. As
we have already discussed, the thalamus and other
multiple parallel subcortical arousal systems in the
upper brainstem, hypothalamus, and basal forebrain
(Table 1.1) are essential for maintaining the alert state.
16 1. NEUROANATOMICAL BASIS OF CONSCIOUSNESS
These same systems also play a key role in controlling
attention and awareness not only in a permissive sense
(e.g., being in a coma is not compatible with attention
and awareness), but also by facilitating the addi-
tional processing in cortical and subcortical networks
necessary for attention and for awareness.
Several additional subcortical structures also play
a role. Components of the tectal region, specifically
the superior colliculi and pretectal area form an impor-
tant circuit along with the pulvinar of the thalamus
to direct saccadic eye movements towards salient
stimuli, and the same circuits also participate in
directed attention (Krauzlis et al., 2013). The basal
ganglia have major reciprocal connections with the
thalamic intralaminar nuclei and this circuit as well as
other basal ganglia connections may contribute to
arousal and attention functions (Dreher and Grafman,
2002; Hager et al., 1998; Ring and Serra-Mestres, 2002).
The claustrum is a thin layer of neurons located in the
white matter between the putamen and insula, with
widespread cortical connections that have been pro-
posed to play an important role in the attention and
awareness aspects of consciousness (Crick and Koch,
2005). Finally, the cerebellum has major reciprocal
connections with the prefrontal cortex and has also
been proposed to participate in attention, although
this remains somewhat controversial (Baumann and
Mattingley, 2014; Bischoff-Grethe et al., 2002; Dreher
and Grafman, 2002; O’Halloran et al., 2012).
CORTICAL NETWORKS
AND CONSCIOUSNESS
The cortical components of the consciousness system
include widespread regions of association cortex in
the bilateral cerebral hemispheres, particularly in the
lateral frontal, anterior insula, lateral parietal (and adja-
cent temporal-occipital cortex), medial frontal, medial
parietal (precuneus) and cingulate cortex (Figure 1.2).
As has already been discussed, individual cortical
components of the consciousness system have specific
well-studied functions in behavioral neurology which
contribute to the various contents of consciousness along
with specific primary and secondary sensorimotor and
limbic cortical areas (Figure 1.11). However, it is the
collective action of widespread bilateral association
cortex regions that gives rise to regulation of the level of
alertness, attention, and conscious awareness.
In this section we will review the contributions of
the cerebral cortex to arousal and the generation of an
alert, awake state. We will next discuss attention in
somewhat greater detail, describing several formula-
tions of cortical systems that control different aspects
of attention. The relationship between memory
THE NEUROLOGY OF CONSCIOUSNESS
systems and consciousness will then be considered, as
well as the role of cortical networks in self-awareness,
planning voluntary action and free will. Finally, we
will review cortical networks revealed by contrastive
analysis (perceived vs non-perceived stimuli) to play
an important role in conscious awareness.
The Cortex and Arousal
The most important input to subcortical arousal
systems, including the thalamus, hypothalamus,
basal forebrain, and the multiple brainstem arousal
systems is the cerebral cortex itself. It has long been
known that stimulation of the higher-order heteromo-
dal frontoparietal association cortex increases arousal
(Figure 1.11) (Segundo et al., 1955). Conversely, abla-
tion of these same regions of the higher-order associa-
tion cortex markedly decreases arousal (Ropert and
Steriade, 1981; Steriade and McCarley, 2010; Watson,
et al., 1977), although the subcortical arousal systems
also receive inputs from primary sensorimotor cortices
(Catsman-Berrevoets and Kuypers, 1981; Ropert and
Steriade, 1981; Rossi and Brodal, 1956). In further sup-
port of the importance of the cerebral cortex in main-
taining consciousness it was recognized by clinicians
that unilateral cortical lesions usually do not markedly
depress level of consciousness, but bilateral lesions of
the association cortex can produce coma (Plum and
Posner, 1972; Posner et al., 2007). The parietal cortex of
the non-dominant (usually right) hemisphere appears
to play a particularly important role in arousal where
large lesions—although not producing coma—do often
produce a markedly drowsy clinical state with forced
eye closure. Thus, in addition to its important role in
producing the specific individual contents of con-
sciousness, the cerebral cortex is also a major driver in
regulating the overall level of conscious arousal.
Attention and Consciousness
There has been recent debate on the relationship
between attention and consciousness. Some view atten-
tion and consciousness as orthogonal functions that
can be fully dissociated and operate independently
(Koch and Tsuchiya, 2007). Others consider attention
and consciousness to be essentially identical, constitut-
ing different names for the same set of functions
(Prinz, 2000, 2012). Still others posit that attention is
necessary for but not identical to consciousness
because additional functions are needed for conscious
awareness (Dehaene et al., 2006; Kouider and Dehaene,
2007). These very different understandings of the rela-
tionship between attention and consciousness may
arise at least in part from different models for defining
and understanding attention. There are a large number
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 Chalender had not moved, did not suspect. He was wounded; his
fever was high. He might not live.
Perhaps he had been in a delirium. Perhaps Patty had been
merely trying to quiet him. But she had been saying, “How wicked we
are!” as if cheaply absolving herself of sin by confession.
Suppose RoBards charged her with disloyalty and she denied it.
What proof had he? He was the only witness. He could not divorce
her for merely kissing a wounded visitor.
Divorce! How loathsome! Nobody had yet forgotten old Aaron
Burr’s brief marriage to old Betty Jumel or the recent noisome
lawsuit that followed, in which Burr flattered her with four
corespondents to her one for him.
As a lawyer RoBards had many divorce trials brought to him and
he abominated them. He had never had a nightmare so vile as the
thought that he might have to choose between clamorous divorce
and smothered disgrace.
He wanted to die now rather than make the choice. To kill
Chalender would seem almost a lesser horror. But that also meant
exposure to the public. The burial of Chalender would but throw
open his own home like a broken grave. It was only a detail that
Chalender had saved his life the night of the fire when RoBards
could not climb back to the wharf and no one else heard or heeded
him.
To butcher a wounded man, guilty so ever; to strip a woman stark
before the mob, evil so ever; to brand his children, to blotch his
home with scandal—pure infamy! But, on the other hand, to spare a
slimy reptile; to be the cheap victim of a woman’s duplicity; to leave
his children to her foul ideals; to make his home a whited sepulcher
—infamy again. He felt that the children must be first to be
considered. But which way was their welfare to be sought?
Then the children themselves ran in upon his swooning mind,
Imogene and Keith. He felt their tendril fingers wrap about his inert
hands. He heard their piping cries of welcome. He fell back from the
door and was so weak, so sick that they easily pulled him to his
knees and clambered on his back and beat him, commanding
“Giddap!” and “Whoa, Dobbin!”
The very attitude was a degradation. He was actually crawling, a
brute beast on all fours with his young on his back. When he flung
them off Keith bumped his head and began to cry, Immy to howl and
boo-hoo! And they ran to their mother protesting that their Papa was
mean, and hurted them. They turned to Chalender for protection.
And this was Chalender’s first warning that RoBards had come home
—home! what a dirtied word it was now—“home!”
RoBards scrambled to his feet and dashed out of the intolerable
place.
Only the old tulip tree had dignity now. With a Brahmanic majesty
it waved its long-sleeved arms above him and warned him that he
must not let life drive him mad. His decision one way or the other did
not matter much. Nothing he did or left undone mattered much. The
leaves would come and go and come anew. The farmer was still
striding along after his plough in a silhouette cut out against a scarlet
west.
Just one thing seemed important: the house pleaded with him not
to dishonor it. It was older than he. It had cradled him. It had cradled
his children. It wanted to cradle their children’s children. The
lengthening shadow of the chimney had crept along the grass now
till it lay like a soft coverlet on the beds of the little twain that had
been lent him for a while. The very chimney had a soul of its own,
and a good name. It seemed to implore him not to brand it as a place
of evil resort.
His knees gave way and he dropped to the ground, rendered idiot
by the contradiction of his impulses. He saw old negro Cuff staring at
him. The farmer’s wife paused at the back door to wonder. At an
upper window Patty’s Teen leaned out to fix on him the white stare of
her black face.
Then someone came stepping toward him as timidly as a rabbit in
dew-chilled grass. Someone sank down by him with a puff of floating
silk and a drift of perfume across his nostrils. And then his wife
spoke in the coldest calmest voice he had ever heard from her, as if
his discovery of her had discovered her to herself and had aged her
in a moment.
“Mist’ RoBards!” she pleaded, “Mist’ RoBards if it will save you any
trouble, I’ll kill myself. I’ll fling myself down the well, or let you kill me
if you would like that better. Some day you were bound to catch us
together, Harry and me. I’m almost glad you did at last. I’ve been bad
enough to destroy my own soul, but don’t let me break your heart or
ruin your life. I’m not worth your grieving for, Mist’ RoBards. I’ve been
as wicked as I could be and for a long while, and now you’ve found
me out—and I’m glad. Even if you kill me, I’m glad.”
But he was not glad. Suspicion had burned and hurt, but
knowledge was a knife through the heart; it was mortal. It killed
something in him. One soul of his many souls was slain. His other
souls were in a panic about its deathbed, as Patty went on, her voice
queerly beautiful for all the hideous things it told:
“Harry doesn’t know that you saw him—us. Nobody does. He isn’t
in his right mind. He is weak and sick and I made myself pretty just
to make him quit laughing at me. And if he dies, it will be my fault.
“And that would be funny—for such a worthless little fool as me to
cause so much trouble for two men, two such fine men. He is fine, in
spite of all his wickedness, and he’s doing a great work that must go
on. Let me go away and disappear somewhere. I’ll drown myself in
your river, if I can find a place deep enough. And Harry need never
know why. I don’t want him to know that you saw us. I couldn’t stand
that. It’s of you I’m thinking. I don’t want him to know that you know
about this terrible thing. It isn’t so bad, if he doesn’t know you know.
For then you’d have to kill him, I suppose.
“But please don’t kill him, for then they’ll try you and send you to
prison or hang you and choke you to death before all the people. Oh,
don’t let that happen, David. You couldn’t be so cruel to me as to let
them kill you and hurt you and bury you in the Potter’s Field on my
account—don’t do that to me, Davie. I’ve loved you. In my way, I’ve
loved you. I’m not good enough for you, but—if any harm should
come to you, I’d die. Don’t look like that, Mist’ RoBards! Oh, don’t
look so helpless and heartbroken and so unhappy. Don’t torture me
to death that way!”
And then it was he that sobbed and not she. He could feel her
clutching at him and lifting him from the grass reeking with his tears.
She drew his head into her soft arms and into her lap and set her lips
against his cheek, but dared not kiss him, though her tears beat on
his clenched eyelids like the first big drops of a long rain.
One little mercy was vouchsafed him and that was the sinking of
the sun behind the hill; the blessed twilight came with its infinite
suavity and the impalpable veils it draws across the harsh edges of
things and thoughts.
He saw the tide of the evening wind where it eddied along the
grass and overflowed his hands and his face. He heard the farmer
go up the dusty lane that muffled the tread of the tired horses, but
not the little clink-clink of the harness rings. He supposed that the
farmer was staring and wondering at him as he himself stared inside
his own eyelids at the world within him, and wondered at that.
It grew cold. His wife’s hands chilled as they clenched his. He
could feel her shiver. He could just hear her whisper through her
chattering teeth:
“Please come in, Mist’ RoBards.”
He put away her arms and got to his feet. Then his dignity took on
the look of mere sulkiness. When he saw Patty unable to rise, and
huddled in a dismal heap, he bent and lifted her to her feet. She
seemed unable to stand or walk; so his arm of its own volition or
habit went round her to hold her up.
And at that she threw her arms about him and buried her face in
his breast and sobbed. He looked through blurred eyes at the
ambiguous sky where stars were thrilling in the rosy afterglow. In the
dark house someone was lighting lamps. The lamps and the stars
were tenderly beautiful, but they came only when all else was black.
From the hall door a rug of warm yellow ran across the porch and
down the steps into the path. The children began to call, “Mamma!
Papa! where are you?”
 The house yearned toward him with its deep bosom. Something
with the arms of a spirit reached out from it and drew him in.
It was wrong to yield, but he had an utter need of peace for a
while. He was wounded worse than Chalender, and needed more
care.
All that night it was as if Indians prowled about the house, savages
that longed to drag forth the people within, to howl slanders and
truths about them, to fasten them to stakes and dance a torture
dance about them, cut off their eyelids and blind them with ruthless
light. There were no Indians to fear now, save the stealthy reporters
and the more merciless newspapers.
But the house baffled them; it was a strong stockade. They should
not have its children yet a while. It had won another day in its long
battle against the invading strangers.
 CHAPTER XIV

THAT night RoBards slept apart from his wife—in the spare room.
He owed that much to his wrongs and she dared not try to wheedle
him into the dangerous neighborhood of her beauty.
But first they heard the children’s prayers together. It was bitter to
hear their sleepy voices asking forgiveness for their tiny sins and
murmuring, “God bless Mamma and Papa and Mister Chalender!
Amen!” Then the wet little good-night kisses scalded the cheeks of
the divided parents who leaned across the cradles as across coffins
and waited till sleep carried their babes away to the huge nursery of
night. Then they parted without a word, without the challenge of a
look.
He slept, too. All night he slept, better than ever. His strength had
been shattered in a moment as if a bolt of lightning had riven him. He
was a dead man until the morning brought resurrection and the
problems of the daylight.
The first thing he heard was a loud shout:
“Jump her, boys! jump her! No water! There’s no water! We’ve got
to get some gunpowder! Up she goes! Down she comes!”
It was Harry Chalender in a delirium fighting the Great Fire again.
His frenzy gave him the horrible sanctity of the insane.
The doctor came over after breakfast. He shook his head. The
wound was dangerous: the pick-blade had made an ugly gouge and
gangrene might set in. There was pus in the wound. There was
fever, of course, high, racking fever that fried his flesh till the very
skin seemed to crackle.
RoBards had not expected to go back to town for several days. He
had needed the cool remoteness of his farm. But now the solitude
was like that uttermost calm into which the angels fell and made it
Pandemonium. Now the place was crowded with invisible devils
gibbering at him, shaking their horned heads over him in hilarious
contempt, tempting him to everything desperate.
He made an excuse to Patty that he had to return to the city. He
spoke to her with the coldest formality. She made no effort to detain
him, but this was plainly not from indifference, for she answered like
a condemned prisoner in the dock.
“All right, Mist’ RoBards. I understand.”
It broke his heart to see her meek. All the fire of pride was gone
out of her. She was a whipped cur thing, and he could not put out his
hand to caress her.
Something in him, a god or a fiend, tried to persuade him that she
was not to blame, that she had been the prey of currents stronger
than herself. But whether the god or the fiend whispered him this, the
other of the two spirits denied it as a contemptible folly.
According to the law, women, as soon as they married, lost all
rights to their souls, wills, properties, and destinies: yet if men were
to forgive their wives for infidelities no home would be safe. This
new-fangled mawkishness toward the wicked must have a limit
somewhere.
He had to go into his library for a lawbook that he had brought with
him on an earlier visit to his home—“visit” seemed the nice, exact
word, for he was only a visitor now. Harry Chalender was the master
of the house.
RoBards expected to find the usurper in a delirium. But Chalender
was out of the cloud for the moment. With a singularly fresh and
boyish cheer, he sang out:
“Hello, David! How’s my old crony? Don’t let me keep you out of
your shop. Go ahead and work and don’t mind me. I’m pretty sick, I
suppose, or I’d take myself out of your way. Forgive me, won’t you?”
He asked forgiveness for a possible inconvenience, but kept in his
black heart the supposed secret of his treachery! Yet something
compelled RoBards to laugh and say that he was to make himself at
home and feel right welcome. The dishonest glance he cast toward
Chalender was met with a look of smiling honesty that reminded
David of some lines he had heard the English actor Kean deliver at
the theatre:

“My tables—meet it is I set it down,

That one may smile and smile and be a villain.”

Yet he smiled himself, and felt that many a villain was more the
hero than he. He hurried out of the room, fleeing from the helpless
sick man who smiled and had no conscience to trouble him.
He found that his horse had gone lame and could not take him all
the way to New York. He drove the limping nag only so far as White
Plains, and sent Cuff back with him. He waited in front of Purdy’s
Store until the Red Bird coach was ready to start. He saw Dr.
Chirnside waiting for the same stage, and he dreaded the ordeal of
the old preacher’s garrulity. But there was no escape. The parson
had come up to look over the churches in the Bedford Circuit and he
was pretty sure to indulge in one of his long tirades against the evils
of the times—especially the appalling atheism of the country, an
inheritance from the Revolutionary sentiments. The colleges were
full of it—of atheism, drunkenness, gambling—but Dr. Chirnside
seemed to dread atheism far less than he dreaded the other sects of
his own faith. Methodists, Baptists, Presbyterians were gaining a
foothold in the countryside and he almost choked when he referred
to the Catholics.
All the way down to New York Dr. Chirnside’s tongue kept pace
with the galloping horses. He began with the stage itself. He
remembered when even carriages were almost unknown in the rural
districts. Gentlemen rode horses and carried their necessaries in
valises swung from the saddle; ladies rode on pillions. Then light
wagons came in, and carioles next, gigs, chaises, and chairs. And
now stages with their luxury and their speed of nearly ten miles an
hour!
As if that were not enough, a steam railroad was to ruin the peace
of the country. Had Mr. RoBards ridden behind one of the engines
that now drew the railway cars from the City Hall all the way to
Harlem? No? He had been fortunate in his abstemiousness.
“The speed of these trains is only another instance of our mad
passion for hurry. After a time people will return to their sanity, and
the stage coaches will drive the fire-breathing monsters back to the
oblivion they came from.
“Another evil of the railroad is that it will bring more and more of
the wicked city element into the country. The aqueduct has
practically ruined an entire region. Have you seen the hollow
Chinese wall they are building for the Croton water? Ah, yes! Indeed!
Most impressive, but if man’s work destroys God’s beautiful country
where will be the profit?
“The Continental Sabbath will soon destroy the rural peace as it
has already destroyed New York’s good name. The chains are no
longer drawn across Broadway before the church services and any
Tom, Dick, or Harry may now drive his rattletrap past the sacred
edifice on his way to some pagan holiday.
“In the good old days even taking a walk on Sunday was
recognized as a disrespect to the Lord. Nowadays men go driving!
And not always without fair companions of the most frivolous sort. In
my day a gentleman, passing his most intimate friend on the way to
church, would greet him with a cold and formal nod. Nowadays
people smile and laugh on Sunday as if it were merely a day like
another! Where will it end? I tremble to think of it.
“I have just witnessed an example of the extent to which the new
lawlessness is carrying us. Fortunately I was able to deal with it
sternly.”
He told how some of the aqueduct laborers had spent their
Sunday off, not in pious meditation and fasting, but in sauntering
about the country. Their paganism had gone so far that when they
came upon a patch of wild whortleberries growing by the roadside,
they brazenly began to pick and eat them and gather others to take
to their camp.
 “Driving home from the service I chanced to see them, and I
determined to put a stop at once to this violation of the laws of God
and man. I ordered the county sheriff to arrest the culprits. They
were fined a shilling each for the sacrilege.
“Unfortunately, this was not the end of it. The depths of human
depravity were disclosed in the behavior of these gross men. Only
last Sabbath, instead of going to church, they hung about the village.
Most unluckily, the sheriff’s daughter carelessly went into the garden
and picked a few currants for the midday dinner. Whereupon the
laborers called on her father and demanded that he arrest his own
daughter. He had to do it, too, and pay her fine of a shilling. It will be
a lesson to the wicked girl, but it rather undoes the good I was able
to impress on the laborers.”
Dr. Chirnside was aghast at such levity, such contempt for sacred
things, but RoBards took no comfort in the thought that, since man’s
quenchless thirst for horrors could be slaked with such trivial
atrocities, his own tragedy was only one example more.
He felt an almost irresistible impulse to seize the clergyman by the
sleeve and cry:
“What would you say if I told you of what has been going on in my
own home? My wife is a member of your congregation; she has
been brought up with every warning against immodesty of thought or
action, and yet—and yet——”
He could not frame the story even in thought. He could not tell it.
Yet if he did not tell, the secret would gnaw his heart away like a rat
caged within.
Dr. Chirnside could hardly have found appropriate gloom for this
disaster since he was already in such despair over the habits of the
modern women, that he had no superlatives left for their dishonor.
As the stage swung down into the city, lurching through mudholes
that occasionally compelled it to take to the sidewalk and scatter the
pedestrians like chickens, he pointed out a girl strolling along with a
greyhound on the leash of a blue silk ribbon.
 “See how our girls walk abroad unattended,” he gasped. “That
young female has at least a dog to protect her, but it is appalling how
careless parents are. No wonder our foreign critics are aghast at the
license we allow our ladies. They go about without a father or a
husband to guard them from the insolence of bystanders. It is the
custom, too, to permit couples who have been formally betrothed to
be alone together without any guardian. In most of the homes sofas
have been imported for them to sit upon. No wonder that New
England people say that their old custom of bundling was less
immodest. The very word sofa implies an Oriental luxury.
“The dress of our women, too, is absolutely disgusting. When I
was young there was an outcry against a new fashion of shortening
the skirts in the rear so that the heels were visible. People frankly
cried ‘Shame!’ at the sight of them. Nowadays ankles are openly
exposed. Look at that pretty creature stepping across the gutter. She
is actually lifting her petticoats out of the mud. No wonder those men
all crane their necks to ogle! And her satin shoes are hardly more
than cobwebs!
“Their immodesty does not stop at the ankles. The bare bosom is
seen! Really! I blush to mention what young females of excellent
family do not blush to reveal.
“It is perilous to health, too. You see our ladies gadding about in
the bitterest weather with their necks uncovered, while gentlemen
shiver under their great coats with five or six capes and heavy stocks
and mufflers besides.
“But the men are hardly more modest. This new fashion of—dare I
refer to it?—of buttoning the pantaloons down the front instead of on
the sides! It is astounding. One or two sermons have already been
preached against it and I think I shall refer to it myself next Sabbath.
Pardon me!”
There was a respite while he took out his pocketbook and made a
note of this urgent matter. RoBards remembered his own
memorandum that a man may smile and be a rake as well. He could
hardly keep from plucking at the parson’s sleeve and confessing:
 “When you are in your pulpit, cry out also that one of the town’s
pets, the popular Harry Chalender, has ruined the good name of my
wife and our children and stained the old RoBards mansion with the
wreckage of the Seventh Thou-shalt-not!”
But Dr. Chirnside was putting up his pencil and putting forth his
lean, cold hand for a farewell clasp. The stage was nearing City Hall
Park and he must get out his fare and get down at his parsonage.
And a little further below was the Astor House, which RoBards
must call home henceforth.
Dr. Chirnside had referred for his “thirteenthly” to the barbaric
luxury of the new hotel, and to the evil influence of such hostelries on
home-life. It had a bathtub on every floor! What Oriental luxury would
come next? In many of the more religious states bathing was a
misdemeanor, but in New York every crime flourished—and every
slothfulness. The modern woman, unlike her mother, was too
shiftless to care for her own household or even to oversee her
servants: she preferred to live in a hotel and have more time and
convenience for her idle mischiefs.
But RoBards mused dismally that his home had gone to wrack and
ruin first, and that the hotel was his only refuge.
 CHAPTER XV

THE sumptuousness of the Astor House only emphasized RoBards’

exile. From his window he would look down upon the seething
throngs along Broadway, the tall beavers of the men and the poke
bonnets of the women bobbing along as on a stream.
He would seek escape from solitude overlooking the multitude by
retreating to the inner court and the fountain flaunting its crystal
plumes in the turfed garden. But there was that quadrangle of many-
windowed walls about him, and he felt Argus eyes upon him
everywhere. Behind every curtain somebody seemed to be watching
him. The expense of the luxurious hotel was heavy—a dollar a day it
cost him, but he could not face boarding-house inquisitiveness.
In his office he would sit and brood across his pine table with its
green baize cover, and stare at the pine boxes that held his books
and the files of his cases tied with red tape. He would dip his quill
into the inkstand of gray stone and make careless scratches on the
paper before him. When he looked at them afterward they made him
wonder if he were going mad. These crazy designs would serve as
evidence for his commitment to any asylum.
On the margins of his briefs he would wake to find that he had
been making crude contours of Patty’s scoop hat, her big eyes, or
the nape of her neck. He would blot her out in a fury of rage, and
attack his work.
The case of Jessamine vs. the City of New York was still hanging
fire. Many of the claims of people who were forced to sell their lands
for the aqueduct were still unsettled though their farms were covered
with stone and trenched with ditches.
Yet now RoBards felt that the city had its justice. He had fought for
the country and the country had betrayed him. Vile wickedness had
found shelter and prosperity in the gentlest seclusions.
 It was a mockery that he should be the counsel for old Jessamine.
What did he owe the dotard except hatred for bringing into the world
so pretty a perjurer? The father had made Tuliptree Farm almost
untenable by his whimpering stupidity and the daughter had driven
him into exile by her ruthless frivolity.
From his law office and his hotel RoBards would flee to a club. He
had joined the fashionable Union Club just formed, but the members
always asked him about his wife, and he had to speak of her with
affection and respect.
The affection was still in his heart, but the respect—he marveled at
his ability to adore one whom he despised, to hang his whole life on
the broken reed of a little woman’s wavering fancy.
He frequented the theatre but he found discomfort there, since
almost all the stories dealt with tragic or comic flirtations. He liked to
go to the Bowery Theatre, but it was always burning down. Mary
Taylor, “Our Mary” as they called her, puzzled him because she had
a reputation for private morality and yet she was a convincing
actress of spicy rôles. Patty was not an actress at all—she was
positively imbecile in the drawing-room plays she had taken part in;
yet her private life proved that her home was but a stage to her.
Behind the private life of people there was so often another private
life. And he had never been admitted until now to the green room of
his own domestic theatre. Patty was a convincing actress of
Innocence.
Moods of retaliation were frequent. There were opportunities
enough. It amazed him now that he was alone in the city to see how
many chances were offered him to make some other husband a fool.
Young girls of fifteen or sixteen, who had not yet been married, or
were only betrothed, dazed him by the black wisdom in their eyes.
They scampered and made pretenses of terror before him like
kittens or puppies begging him to pursue them.
Others were to be had of a more public character. It was estimated
that there were ten thousand downright wicked women in the town.
The streets at night were so crowded with them that innocent young
girls, poor seamstresses or polite damsels whom some emergency
forced to be abroad, were not only ogled and bespoken, but
sometimes seized and kissed by the loiterers.
The haunts of evil were well known, some of them foul dens, but
others mansions. Yet the very sense that Patty had absolved him of
obligation to her; that she herself had severed their contract,
annulled the temptation. What excitement could he find it taking
sneakingly what nobody could prevent his taking openly?
Besides, as a lawyer he knew that the traps of blackmail lay all
about the town—springes to catch woodcocks. The heads of many
families were paying perennial taxes on such indiscretions. He knew
of one banker who had been mulcted of thirty thousand dollars just
because he chanced to be in the house where Helen Jewett was
murdered. The trial of the young clerk charged with the crime was
enormously exploited by the noisy newspapers.
That clerk was ruined for life, and he might well have wished that
he had been employed by Mr. Tappan, the abolitionist silk merchant
who compelled each of his clerks to sign a pledge never to visit a
theatre or make acquaintance with actor, actress, or other person of
evil life, never to be out of his boarding house after ten o’clock at
night, never to miss the two prayer meetings a week, or the two
Sunday services, or fail to report of a Monday morning the church,
the preacher, and the text of the day before.
A final check upon any recklessness in RoBards’ lonely humors
was the feeling that if he also sinned he would be robbed of his
precious indignation against Patty. He was no prig, no prude. He had
lived. But just now the one food of his soul was the sense of being
cruelly wronged. It was gall, but it sustained him somehow.
In the eyes of the law a husband’s infidelity was almost negligible,
but RoBards felt that if he were to break his vows he would acquit
Patty of blame for being false to hers. There were families in town,
according to gossip-mongers and the gossip papers, where husband
and wife were mutually and commonly disloyal. But he could think of
nothing more hideous than such households.
 He was Saint Anthony in a lonely cavern, but only one devil tried
his soul and that was the bewitching spirit of his pretty wife. Patty
drifted through his dreams like a wind-driven moth. She poised and
flitted and opened her arms like a moth’s wings. And it seemed
impossible that he should long resist her.
One morning he read in the Herald (whose editor Mr. Bennett had
recently had a knockdown fight with General Webb of the Courier) a
statement that Mr. Henry Chalender had recovered from his wound
and was once more active in the completion of his section of the
aqueduct. The Herald added that this news would give relief and
pleasure to the numberless admirers of the popular idol.
This paragraph filled RoBards with mixed emotions. During his
long indecision, his Hamlet-like soliloquies and postponements,
nature had healed the wound in Chalender’s flesh, and, though he
would not admit it, had nearly healed the wound in RoBards’ soul.
There was a relief of tension at least. The world was going on.
Chalender was well and busy—perhaps he was renewing his amour
with Patty. Perhaps, deserted and lonely, she would yield again. That
would be a double damnation. Anyone might sin and recover, but to
slip back again was to be lost forever.
Yet who was to uphold her in the hour of weakness? Who was to
drive the wolf away from the ewe?
Insidiously the temptations RoBards had denounced as
complacency, servility, wanton desire took on now the aspect of duty.
It was his duty to go home and take up wedlock again, to save the
little silly beauty he had married from becoming a monster of iniquity.
Now that his house was freed of the intruder, homesickness came
over him like a fever. He yearned for the hills of Westchester, those
earthen breakers foaming with trees, and carrying on their crests
houses like ships anchored on waves that never moved.
His long sojourn in New York began to attract open comment,
particularly as the heat was so vicious that it looked curious for
anyone to remain who could get out. There was nobody in town now
but nobodies.
 What excuse had he to linger? He had to rise and go back. He had
not slain Chalender. This abstention in itself had amounted to an
acquittal. If he were not going to punish Chalender, why should he
punish himself? If his aim were to escape gossip, why encourage it?
He went home. Patty was in the yard playing a game with the
children. They seemed to have grown amazingly since he left. They
ran to him screaming welcome. It was bliss to feel their warm hands
clutching him.
He could see that Patty was afraid to move either toward him or
away. She had never written to him, but he had felt that this was
meekness rather than neglect. She waited now struggling between a
cry of joy and a fit of tears.
He pretended that it was for the children’s sake that he called out:
“Hello, Patty!”
“Hello, David!” she murmured. Suddenly her eyes were gleaming
with tears.
 CHAPTER XVI

THE old Jessamines stared at him, but summed up their curiosity

and their resentment in a “Well! So you’re back?”
“Yes,” he said, the answer sufficient to the question.
He was embarrassed to find that a cousin of his wife’s was visiting
the farm and the spare room was filled. He had to go back with Patty.
But they were like two enemies in the same cell.
Sometimes he would wake suddenly in the night from a hell of
self-contempt. He would both sweat and shiver with remorse for the
shame of having let Chalender live.
In his half-insanity it seemed a belated duty to go out and
assassinate the villain. To shoot him down openly would be too noble
a punishment—like shooting a spy. To garrote him, string him up
squirming from a tree limb would be best. Major André had wept
pleading to be shot, but they had hanged him—not far from Tuliptree
Farm. And only recently people had dug up his grave and found the
tiny roots of a tree all grown about his curly hair.
Chalender had sneaked into RoBards’ home and Patty had played
the Benedict Arnold to surrender the citadel to the enemy. He
deserved to be put out of the way like a poisoned dog, a sheep-killer,
a lamb-worrier.
Sitting up in his bed with night all about him RoBards would enact
some grisly murder, often while Patty slept at his side unheeding the
furies that lashed her husband and mocked him for his forgiveness
like Christ’s of the woman brought before him.
In the restored innocence of sleep, Patty’s face was like a little
girl’s with its embroidery of her curls, one shoulder curved up, a
round white arm flung back above her head, her bosom slowly lifting
and falling with her soft breath.
 Sometimes as he gazed at her, his heart welled with pity for her; at
other times he was frantic to commit murder because of her.
But the big tree at the window would try to quiet him like an old
nurse; it would go “Hush, hush!” The house would seem to sigh, to
creak as if its bones complained. And it, too, would counsel him, “No!
no!”
The ferocity of such debates would wear him out more than a
prolonged contest in court, and he would sink back and draw sleep
over him as a black blanket of respite from thought.
At other times when Patty was gracious and full of laughter, when
she was in a mood to be a child with her children and play with them,
there would be a heavenliness in life that made RoBards cry aloud
within himself, “Thank God I kept the secret.”
By and by there was a child again at Patty’s little breast—the fifth
in number, the third alive. She had resigned herself to motherhood
now. She nursed the babe and took all the care of it without
complaint. She met RoBards at night when he came up from town,
with stories of the wonderful things the new son had achieved or the
older babes had said.
It pleased him quaintly to find his wild, restless Patty becoming a
subdued and comfortable matron, telling unimportant anecdotes
importantly. She kept her grace and her beauty and she could never
grow slattern; but she was maternal now to her marrow.
Regarding the deep peace of his country family, RoBards was
profoundly glad that he had forgone the swift passionate delights of
revenge. If he had slain Chalender or published the scandal in the
courts, Patty would not have been his now. That child whom she had
named after himself, David Junior, would have been doomed to an
unhonored name. This house would have been pointed to as a
monument of scandal. It would be neglected, empty, haunted.
The neighbors never dreamed of the hidden shame. They said:
“Nothin’ ever happens up your way. You’re one lucky man.”
Nearly every other dwelling had some scandal hung upon it like a
signboard from a tavern. Not many miles up the road was a house of
a strange memory. A widow lived there—she called herself a widow,
but the neighbors called her “a queer un.” They told how a negro
preacher freed by his master had settled up in New Hampshire fifty
years before and been so much respected that he had married a
white woman and had many children; and these children had had
children. And one of them had married this woman when she was
young and high-stepping. The first she knew of her husband’s grand-
parentage was when a gossip twitted her with it. She said nothing,
but made an excuse for a trip to the Bermudas with her husband. As
soon as she got him there, she sold him at a good price into slavery,
and came home calling herself a widow.
For that matter, one of the presidents of the United States had
been sued in the open courts by a negress for the support of their
child; it was said that he sold many of his mulatto children, and that
his only indulgence was that when any of his own escaped he would
not hunt them with hounds, but laughed and let the rascals escape if
they could.
The present president of the United States had been in the divorce
court and had turned Washington inside out with domestic
bickerings.
Nearly all the founders of the republic had been plastered with
scandal. Many of them were infidels and Dr. Chirnside was always
bewailing the decay of religion under the republic.
So RoBards reasoned that if there were scandal in his home, it
was only what every other home had. The good thing was that his
shame was hidden. His house was looked upon as a place of honor.
It was unsullied. It must be kept of good repute. There was a certain
kind of hypocrisy that was wholesome and decent and necessary to
good citizenship.