Foundational Concepts in Neuroscience A Brain Mind
Odyssey (Norton Series on Interpersonal Neurobiology)
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The Norton Series on Interpersonal Neurobiology
Louis Cozolino, PhD, Series Editor
Allan N. Schore, PhD, Series Editor, 2007-2014
Daniel J. Siegel, MD, Founding Editor
The field of mental health is in a tremendously exciting period
of growth and conceptual reorganization. Independent find-
ings from a variety of scientific endeavors are converging in an
interdisciplinary view of the mind and mental well-being. An
interpersonal neurobiology of human development enables us to
understand that the structure and function of the mind and brain
are shaped by experiences, especially those involving emotional
relationships.
The Norton Series on Interpersonal Neurobiology provides
cutting-edge, multidisciplinary views that further our under-
standing of the complex neurobiology of the human mind. By
drawing on a wide range of traditionally independent fields of
research—such as neurobiology, genetics, memory, attachment,
complex systems, anthropology, and evolutionary psychology—
these texts offer mental health professionals a review and synthe-
sis of scientific findings often inaccessible to clinicians. The books
advance our understanding of human experience by finding the
unity of knowledge, or consilience, that emerges with the transla-
tion of findings from numerous domains of study into a common
language and conceptual framework. The series integrates the
best of modern science with the healing art of psychotherapy.
Dedicated to My Teachers
Contents
List of Figures
Preface: From Molecules to Consciousness
1. Origins
2. Nervous Systems and Brains
3. Chemistry and Life
4. Genes and the History of Molecular Biology
5. How Neurons Generate Signals
6. Synapses, Neurotransmitters, and Receptors
7. Neuroanatomy and Excitability
8. Poison, Medicine, and Pharmacology
9. Psychoactive Drugs
10. Neural Development and Neuroplasticity
11. Sensory Perception
12. Nose and Smell
13. Tongue and Taste
14. Eyes and Vision
15. Ears and Hearing
16. Skin, Touch, and Movement
17. Imaging the Brain
18. Connectivity, Language, and Meaning
19. Memory
20. Rhythms, Sleep, and Dreams
21. Emotion
22. Mind, Consciousness, and Reality
Figure Credits
Notes
References
Acknowledgments
Index
About the Author
List of Figures
1.1. Paleolithic drawings in Chauvet Cave
2.1. Nerve cell and astrocyte glial cell
2.2. Compass jellyfish, Chrysaora hysoscella
2.3. Planarian nervous system
2.4. Basic plan of the vertebrate brain
2.5. Brains ofa fish and a bird
2.6. Brain of amouse
2.7. Human brain: dorsal and lateral views
2.8. Human brain: ventral and medial views
2.9. Open human skull with dura intact and with dura peeled back
2.10. Nerve fibers from brain and spinal cord to body
2.11. Person reacting to fire
2.12. Muscles surrounding the eyeball
2.13. Visual perception and the action of pointing a finger
2.14. Galvani’s experimental apparatus
2.15. Neuron, showing axon and dendrites
2.16. Neuron drawings: human cerebellum and cerebral cortex
3.1. Periodic table of chemical elements
3.2. Diagram of phospholipid bilayer
3.3. Phospholipid bilayer biological membrane
3.4. Myoglobin molecule
4.1. Solvay Conference, Brussels, Belgium, 1927
4.2. Niels Bohr Institute, Copenhagen, Denmark, 1933
4.3. Bacteriophage virus and phage infection
4.4. Gene transcription and translation
5.1. Lipid bilayer membrane with channel proteins
5.2. Relative ion concentrations inside and outside the cell
5.3. Cell membrane with ion-channel proteins
5.4. Hyperpolarization, potassium channels, chloride channels
5.5. Depolarization, sodium channels, calcium channels
5.6. Action potential
5.7. Actions of voltage-gated ion channels during an action
potential
5.8. Location of the axon hillock
5.9. Neuron with myelinated axon
5.10. Electron micrograph of myelinated axon from human cerebral
cortex
6.1. Electrical synapse
6.2. Chemical synapse
6.3. Electron micrograph of chemical synapses
6.4. EPSPs and IPSPs generated in the dendritic field
6.5. Ionotropic receptor
6.6. GPCR and its intracellular cascade
7.1. Muscles in the iris of the eye
7.2. Cholinergic pathways in the human brain
7.3. Serotonergic pathways in the human brain
7.4. Dopamine, norepinephrine, and epinephrine biosynthesis
7.5. Dopaminergic pathways in the human brain
7.6. Noradrenergic pathways in the human brain
8.1. Paracelsus
8.2. Blood vessel cross sections and the blood-brain barrier
8.3. Deadly nightshade, Atropa belladonna
8.4. Henbane, Hyoscyamus niger, and jimson weed, Datura
stromonium
8.5. Tea plant, Camellia sinensis
9.1. Tobacco plant, Nicotiana tabacum
9.2. Opium poppy, Papaver somniferum
9.3. Coca plant, Erythroxylum coca
10.1. Transcription factors and DNA transcription
10.2. Drawing of ahuman embryo at three weeks after conception
10.3. Neuron growth cone drawings by Ramon y Cajal
10.4. Internal cytoskeleton of an axon’s growth cone
10.5. Microfilaments and microtubules
10.6. Sperry’s frog vision experiment
10.7. Hippocampus
11.1. Bacterial flagella
11.2. Phototropism in a Phycomyces sporangiophore
11.3. Café wall illusion
11.4. Electromagnetic spectrum
11.5. Flower patterns in visible and ultraviolet light
11.6. Skylight polarization pattern
11.7. Earth’s magnetic field
12.1. Olfactory receptor cells
12.2. Alembic (alchemical still)
12.3. Black pepper, Piper nigrum
12.4. Durians for sale
13.1. Taste bud
13.2. Neural taste pathways in human brain
13.3. Chili plant, Capsicum annuum
14.1. Cross section of the human eyeball
14.2. Response of human rod and cone cells to visible light
wavelengths
14.3. Distribution of cones and rods in the human retina
14.4. Diagram of rod cell and cone cell
14.5. Diagram of a rhodopsin molecule in a lipid bilayer membrane
14.6. Cell layers in vertebrate retina
14.7. Location of the lateral geniculate nuclei
14.8. Visual areas in the posterior cerebral cortex
15.1. Bell vibrating after a clapper strikes it
15.2. Sound shown as a sine wave
15.3. Sine waves with lower and higher frequencies
15.4. Sine waves with higher and lower amplitudes
15.5. Complex sinusoidal waveform
15.6. Sound waveforms from various sources
15.7. Jew’s harp
15.8. Fourier decomposition of sound produced by a Jew’s harp
15.9. Fourier analysis of different notes played on the piano
15.10. Human ear
15.11. Bony labyrinth of the human inner ear
15.12. Inner-ear hair cell
15.13. Side view of the human skull
16.1. Somatosensory neuron endings in skin
16.2. Dorsal root ganglion neuron
16.3. Primary somatosensory cortex
16.4. Somatosensory map in anterior parietal lobe
16.5. Representation of a person’s phantom hand
16.6. Golgi-stained Purkinje neuron in the human cerebellum
17.1. Spinning toy top
17.2. MRI of human brain
17.3. Cap with sixty-four electrodes for EEG recording
17.4. Human EEG recorded simultaneously from thirty-one
electrodes
18.1. Cell layers in the human cerebral cortex
18.2. Cortical language areas
18.3. Visual information in a split-brain patient’s brain
18.4. Drawings by split-brain patient
18.5. Electron micrograph of neuropil from rat hippocampus
19.1. Piis the ratio of circumference (C) to diameter (D)
19.2. Hippocampus and adjacent areas
19.3. Seahare, Aplysia californica
19.4. Synaptic connections in Aplysia californica
20.1. Progression through NREM and REM stages during sleep
21.1. Chimpanzee facial expression
21.2. Muscles of the human face
21.3. Molecular structure of oxytocin
21.4. Oxytocinergic and vasopressinergic projections from
hypothalamus
21.5. Dopaminergic reward pathways
Preface: From Molecules to Consciousness
Neuroscience—the science of brain and behavior—is one of the most
exciting fields in the landscape of contemporary science. Its rapid
growth over the last several decades has spawned many discoveries
and a large number of popular books. Contemporary news is filled
with stories about the brain, brain chemistry, and behavior. Photos
and drawings of brains and nerve cells grace the pages of newspapers
and magazines. Neuroplasticity—the capacity of neural connections
to change and reorganize—has become a buzzword. The notion that
your mind can change your brain is pervasive.
We read that drugs used to treat mental conditions such as
depression, anxiety, and psychosis are among the best-selling phar-
maceuticals in history. Ads for these drugs depict neurons and neu-
rotransmitter molecules. One hears that brain science is being used
increasingly in the courtroom—that brain structure and functional
activity are used in arguing for guilt or innocence in accusations of
criminal behavior. It is said that advertising agencies are spending
millions of dollars to study the neural activity in people’s brains when
they look at ads or use products. What is going on with all of this?
How are we to understand fact from fiction?
My intention in this book is to provide the backstory—a description
of how nervous systems work and how the workings of our nervous
system relate to our mind and our behavior. My description is concise
yet thorough, rigorous yet easy to follow (and, I hope, fun to read).
For more than twenty years I have taught a class on introductory
neuroscience at the University of California, Berkeley. Hundreds of
students take this class every year. Many are freshmen in their first
year of college. Some are seniors just before graduation. Students
having every sort of disciplinary interest take the class—biological
and physical sciences, social sciences, engineering, economics, busi-
ness, arts, and humanities. Some come to the class already having a
passion for science. Others are afraid of science and sometimes have
particularly strong aversive reactions to chemistry. Occasionally a few
high school students or a few graduate students take the class. And
there are generally a few retired professionals and other folks from the
Berkeley community sitting in on lectures.
My aim as instructor is to provide a comprehensive overview of
brain structure and function—beginning with atoms and molecules,
building up to membranes and cells, progressing to neurons, neural
circuits, and sensory organs, and then on to perception, memory, lan-
guage, emotion, sleep, and dreams. A theme throughout the semester
is the mystery of how mental experience (which is all we ever truly
are able to know) is related to the workings of our brain and body—the
deep mystery of mind and consciousness.
That class is the framework for this book. As such, the book is
comprehensive enough to use as a text in an introductory course on
neuroscience. It gives quite a bit of detail about the structure and
function—largely at the molecular and cellular levels—of the human
nervous system. Most of the material in this book is basic enough that
it will not readily be outdated. Textbooks that endeavor to keep up
with the very latest developments need to be rewritten and updated
frequently. An instructor using this book for a class can always keep
things current by supplementing with additional material drawn
from contemporary neuroscience research. This book provides the
foundation.
I have taught this subject not only in university classrooms but
also in a variety of continuing-education workshop settings for in-
terested individuals of all kinds. I have taught various components
of this material in workshops held in the unique setting of the Esalen
Institute on the California coast, and I have been privileged to teach
neuroscience to Tibetan Buddhist monks and nuns in India as part of
a science-education program initiated by the Dalai Lama. In all these
diverse settings, this approach to the subject—exacting yet affable—
has been well received.
Not just for use in the classroom, this book is also meant for anyone
interested in learning about how the brain works. It is for those of you
who wish to understand just where that news article on brain scans
is coming from, or how to appreciate a story about antidepressant
drugs. I develop everything as much as possible from the ground up,
so that no specific technical background is required to read this book.
For those not familiar with molecular structure diagrams, I hope you
will quickly come to appreciate their simplicity, beauty, and utility. My
intention is to make even the most complex material easy and fun to
understand, at least in its essential features.
A goal is for readers to come away with some intuition about how
the brain works, to appreciate the beauty and power of molecular and
cellular explanations, and at the same time to appreciate that the un-
fathomable complexity of living systems places substantial limits on
any sort of seemingly simple explanation. This last point is frequently
forgotten.
Contemporary physical, chemical, and biological sciences have
enjoyed awesome success over the last several centuries in accounting
for a great deal of what we observe. From the farthest reaches of the
universe, billions of light years away, to the inner structure of atomic
nuclei, physical theories provide a stunningly powerful and coherent
explanatory framework. Living systems, even with their extraordi-
nary complexity, appear to be falling nicely within this remarkable
explanatory framework. However, scientific understanding of the
human mind—and how our capacity for conscious awareness relates
not only to the brain but also to everything else we believe we know
about the physical universe—is considered to be among the greatest
mysteries in modern science, perhaps the greatest mystery. It is my
hope that this book will provide the foundation from which to more
deeply appreciate this awesome subject.
FOUNDATIONAL CONCEPTS
IN NEUROSCIENCE
A Brain-Mind Odyssey
CHAPTER 1
Origins
Early in the winter of 1994, a trio of speleologists made an amazing
find: a limestone cavern in the south of France containing some of the
most spectacular examples of prehistoric art ever seen. The charcoal,
ochre, and carved wall paintings of Chauvet Cave date from more than
thirty thousand years ago and are among the oldest currently known
examples of Paleolithic cave art. Before its modern discovery in De-
cember 1994, it is likely no one had entered this cave for more than
twenty thousand years.
A number of other caves in this region of southwestern Europe
are also filled with wall paintings dating from ten thousand or more
years ago. Often these drawings are deep inside the caves, far removed
from the entrances and completely impervious to light from outside
—places of deep darkness. The Paleolithic humans who made these
drawings carried torches and stone lamps and built small fires to
illuminate their artistic undertakings. It is also likely, given the op-
portunity afforded by these nether regions so isolated from outside
light, that these ancient humans used the darkness as an aid to inner
exploration.
In darkness is the absence of the compelling visual stimuli that fill
