- Youngest division of biopsychology focusing on neural
Experimental Designs and Confounded Variables:
- Experimental designs involve controlled manipulation
of variables for testing hypotheses.
- Good experimental design is vital to eliminate
confounded variables, ensuring accurate results.
- Quasiexperimental studies resemble experiments but
lack control over all variables, limiting their validity.
- Case studies focus on single cases, offering in-depth
insights but lacking generalizability due to individual
differences.
Pure vs. Applied Research:
- Pure research is driven by curiosity, aiming solely for
knowledge acquisition.
- Applied research seeks direct benefits for humanity,
often translating findings from pure research into
practical applications.
Nobel Prize-Winning Research:
- Several Nobel Prizes have been awarded for
significant contributions related to the nervous
system and behavior.
- Winners include researchers like Pavlov, Golgi, Cajal,
Sperry, Hubel, Wiesel, and others, recognized for
various breakthroughs.
Divisions of Biopsychology:
Physiological Psychology:
- Studies neural mechanisms through direct
manipulation and recording of the brain in controlled
experiments.
- Usually involves animal subjects due to invasive
methods.
Psychopharmacology:
- Focuses on manipulation of neural activity and
behavior using drugs.
- Often applied, aiming to develop therapeutic drugs or
reduce drug abuse.
Neuropsychology:
- Examines psychological effects of brain damage in
human patients.
- Relies on case studies and quasiexperimental studies
due to ethical constraints.
Psychophysiology:
- Studies the relationship between physiological activity
and psychological processes in humans.
- Non-invasive measures include EEG, muscle tension,
and autonomic nervous system indicators.
Limitations of Quasiexperimental Studies:
Participants self-select into groups, introducing
potential confounds.
Factors like education, drug use, or diet might differ
between groups, affecting outcomes.
Cognitive Neuroscience:
bases of cognition (thought, memory, attention).
- Primarily involves human participants and noninvasive
methods, avoiding direct brain manipulation.
- Main method: functional brain imaging, capturing
brain activity during cognitive tasks.
Korsakoff’s Syndrome:
- Severe memory loss condition often observed in
alcoholics.
- Initially attributed to alcohol toxicity, later linked to
thiamine (vitamin B1) deficiency.
- Converging operations approach: Studied through
neuropsychological case studies, quasiexperiments
with humans, and controlled experiments on animals.
Scientific Inference:
- Empirical method in biopsychology, used to study
unobservable processes like neural mechanisms.
- Scientists observe measurable events and infer
unobservable processes logically.
- Example: Studying visual perception of motion
through observing retinal image movements and
inferring brain's role in perception.
Thinking about the Biology of Behavior: From
Dichotomies to Interactions
Introduction
- Dichotomous thinking simplifies complexities in
human existence into mutually exclusive categories.
- Common dichotomies: physiological vs. psychological,
inherited vs. learned.
- Historical origins: Descartes' mind-brain dualism and
nature-nurture debates.
Origins of Dichotomous Thinking
- Historical context: Renaissance led to scientific
observations challenging Church doctrines.
- Descartes' dualism: Physical matter (suitable for
science) vs. the mind (Church's domain).
- Influence on perception: Humans assume a part of
behavior transcends the brain.
Physiological vs. Psychological Dichotomy
- Historical challenge: Julien Offroy de la Mettrie's
pamphlet questions mind-brain dualism.
- Modern evidence: Brain damage and stimulation
demonstrate complex psychological changes.
- Animal studies: Chimpanzees exhibit self-awareness,
blurring the line between human and animal
cognition.
Nature vs. Nurture Dichotomy
- Early perspectives: Nature-centric ethology vs.
nurture-focused experimental psychology.
- Problem with dichotomies: Complex behaviors result
from interactions between genetics and experience.
- Flawed assumptions: Attempts to quantify genetic vs.
experiential contributions oversimplify interactions.
Conclusion
- Dichotomous thinking simplifies intricate biological
processes.
- Modern understanding emphasizes interactions
between genes and experiences.
- Analogies highlight the complexity of nature-nurture
interactions, discouraging reductionist views.
- Call to avoid oversimplified dichotomies, encouraging
a nuanced approach to understanding behavioral
biology.
Evolution and Natural Selection:
 Natural selection is a process that leads to the
evolution of species better adapted to their
environment.
 It was termed "natural selection" by Charles Darwin,
emphasizing its similarity to selective breeding in
domestic animals.
 Darwin's theory asserts that species evolve by
producing offspring with advantageous traits that
enhance survival and reproduction.
Initial Resistance and Understanding:
 The theory of evolution initially faced resistance due
to conflicting beliefs in the 19th century.
 Present-day opposition mainly arises from individuals
unfamiliar with the evidence.
Significance of Evolution:
 Evolution is fundamental in various aspects of human
welfare, medical science, and our understanding of
the world.
 The theory of evolution is supported by abundant
evidence from museums, books, textbooks, and
scientific studies.
Evolution and Behavior:
 Social Dominance: Some species establish social
hierarchies through combative encounters between
males, leading to advantages for dominant males.
 Courtship Displays: Elaborate courtship displays
precede copulation in many species and may
contribute to the evolution of new species.
Course of Human Evolution:
 Evolution from Single-Cell Organisms to Humans:
 Fishes evolved into amphibians around 400 million
years ago.
 Reptiles appeared about 300 million years ago.
 Mammals evolved about 180 million years ago.
 Humans belong to the order "Primate."
 Hominid Evolution:
 Humans belong to the Homo genus, one of the six
genera in the Hominini tribe.
 The early Homo species coexisted with
Australopithecus species for about a half-million
years.
 Modern humans (Homo sapiens) originated in Africa
approximately 275,000 years ago.
 Around 130,000 years ago, modern humans began to
migrate out of Africa.
Misunderstandings about Evolution:
1. Nonlinear Evolution: Evolution does not proceed
in a single linear direction; it's more like a dense bush
than a ladder.
2. Lack of Evolutionary Supremacy: Humans are
just one surviving branch on the evolutionary tree,
and countless species have gone extinct.
3. Non-Teleological Progress: Evolution
doesn't aim for perfection; adaptations occur through
changes in existing development.
4. Not All Traits Are Adaptive: Not all traits or
behaviors are adaptive; some are by-products or
spandrels.
5. Evolution Speed: Evolution can occur suddenly
due to environmental changes or genetic mutations.
6. Survival of Few: Very few species have survived
from the many that have existed over time.
7. Exaptation: Not all characteristics evolved for
their current function; some were co-opted later.
8. Homologous vs. Analogous Structures: Not all
similarities among species imply common ancestry.
Homologous structures share a common origin, while
analogous structures have similarities due to
convergent evolution.
9. Interbreeding with Other Species: Evidence
suggests that Homo sapiens interbred with other
Homo species.
Evolution, Genetics, and Experience
Introduction:
 The text explores the relationship between brain size,
intelligence, and evolutionary development,
challenging the assumption that larger brains equate
to higher intelligence.
Brain Size and Intelligence:
 Varied brain sizes in healthy adult humans (1,000 to
2,000 grams) debunk the notion of a direct link
between brain size and intelligence.
 Larger animals tend to have larger brains due to the
need for more brain tissue to control larger bodies.
 Proposal: Brain weight as a percentage of total body
weight offers a better measure of intellectual capacity,
highlighting humans (2.33%) above elephants (0.20%)
and even shrews (3.33%).
Evolution of the Human Brain:
 Early focus on brain size challenged by the discovery
that humans do not have the largest brains; whales
and elephants surpass human brain sizes.
 Evolutionary shift to comparing different brain
regions; emphasis on the evolution of the cerebrum,
which controls complex adaptive processes.
 Increase in brain size during evolution, with a
significant expansion in the cerebral cortex's surface
area due to increased convolutions.
Evolutionary Psychology: Understanding Mate
Bonding:
 Evolutionary psychology examines human behaviors
through the lens of evolutionary pressures.
 Mating patterns in vertebrates range from
promiscuity to monogamy, driven by reproductive
contributions of males and females.
 Polygyny prevalent due to females' greater
contribution to parenting; selective bonding in
females leads to fierce competition among males.
 Polyandry rare in mammals, occurring in species
where males contribute significantly to parenting.
 Monogamy observed in about 9% of mammalian
species, evolving where undivided male help
enhances female reproductive success.
Fundamental Genetics
Introduction:
 The text delves into the foundational concepts of
genetics, highlighting the contributions of Gregor
Mendel to evolutionary understanding.
Mendelian Genetics:
 Mendel's pioneering work with pea plants revealed
the mechanisms of inheritance.
 Dichotomous traits (distinct characteristics) in true-
breeding lines aided Mendel's experiments.
 Mendel’s crossbreeding experiments demonstrated
dominant and recessive traits, laying the groundwork
for Mendelian inheritance.
Chromosomes, Meiosis, and Genetic Diversity:
 Chromosomes, composed of DNA, carry the genetic
code and come in matched pairs, except for sex
chromosomes (X and Y).
 Meiosis, the process of cell division for gamete
formation, introduces genetic diversity through
random chromosome assortment and crossover.
 Mutations, including those causing genetic disorders
like Down syndrome, contribute to evolution but
often disappear due to reduced fitness.
 DNA replication is a crucial process, ensuring genetic
continuity during cell division; errors and mutations
occur but can contribute to evolution.
Sex Chromosomes and Sex-Linked Traits:
 Sex chromosomes (X and Y) determine an individual’s
sex; most sex-linked traits are on the X chromosome.
 Dominant sex-linked traits occur more frequently in
females, whereas recessive traits manifest in all males
possessing the gene.
 Recessive sex-linked traits, like color blindness,
illustrate gender-based expression patterns due to X
chromosome inheritance.
Genetic Code and Gene Expression:
 Structural genes contain information for protein
synthesis, crucial for cellular activities and structure.
 Gene expression varies among different cell types due
to specific stretches of DNA regulating protein
synthesis.
Epigenetics and Its Implications
Introduction:
 The text explores the emergence of epigenetics, a
field studying inheritance mechanisms beyond genetic
code and expression.
 It highlights the shift from exclusive focus on genes to
understanding the role of epigenetic modifications in
inheritance and development.
Epigenetic Discoveries:
 Epigenetic research challenges the notion of "junk
DNA" by revealing active regions controlling nearby
genes, expanding our understanding of genome
functionality.
 Small RNA molecules have diverse roles, including
regulating gene expression, indicating a complex RNA
landscape.
 Mechanisms like DNA methylation and histone
remodeling control gene expression, showing both
increasing and decreasing effects.
 RNA editing allows alterations in messenger RNA,
while small RNAs and proteins cleave and splice RNA,
shaping genetic expression.
Experience-Induced Epigenetic Changes:
 Epigenetic modifications induced by experiences
(neural activity, hormonal changes, environmental
shifts) can be enduring and impact gene expression.
 Transgenerational epigenetics, observed in mice and
humans, demonstrates the inheritance of experiences
across generations through epigenetic mechanisms.
Selective Breeding of Maze-Bright and Maze-Dull
Rats:
 Maze-Bright Rats: Rats that were selectively bred for
their superior maze-learning abilities.
 Maze-Dull Rats: Rats that were selectively bred for
their poorer maze-learning abilities.
 Tryon's Experiment: Tryon's study demonstrated
that behavioral traits, such as maze-running abilities in
rats, can be selectively bred, challenging the idea that
behavior develops solely through learning.
Phenylketonuria (PKU):
 PKU Gene: A single gene mutation causing a
deficiency in phenylalanine hydroxylase enzyme.
 Phenylalanine: An amino acid that accumulates in the
body due to PKU, leading to abnormal brain
development.
 PKU Treatment: A phenylalanine-restricted diet,
initiated early in life, can prevent intellectual
disability, demonstrating the interaction between
genetics and environment.
Development of Birdsong:
 Sensory Phase: Young songbirds form memories of
adult songs, guiding their own singing development.
 Sensorimotor Phase: Juvenile males refine subsongs,
gradually resembling adult songs; auditory feedback is
crucial for proper development.
 Age-Limited vs. Open-Ended Learners: Songbird
species exhibit either stable adult songs (age-limited
learners) or can add new songs throughout life (open-
ended learners).
Heritability Estimates: Minnesota Study of Twins
Reared Apart:
 Heritability Estimates: Numerical values
representing the proportion of variability in a trait due
to genetic variation in a specific study.
 Minnesota Study of Twins Reared Apart:
Demonstrated that adult monozygotic twins are more
similar than dizygotic twins, highlighting genetic
influences on behavior.
 Importance of Environmental Variation:
Heritability estimates depend on the environmental
variations present in the studied population.
Epigenetic Effects in Humans:
  Epigenetic Changes: Modifications in gene
expression due to environmental factors, influencing
disease susceptibility.
 Epigenetic Differences in Monozygotic Twins:
Studies reveal epigenetic differences between
monozygotic twins, suggesting the role of experience-
driven changes.
 Disease-Discordant Monozygotic Twin Studies:
Comparing twins with discordant diseases helps
identify disease-related epigenetic differences, aiding
disease understanding.
Twin Studies of the Effects of Experience on
Heritability:
 SES and Intelligence: Low socioeconomic status
affects the heritability of intelligence, emphasizing the
role of environment in realizing genetic potential.
 Implications for Poverty Programs: Understanding
the interaction between genes and environment
supports interventions to reduce poverty, allowing
genetic potential to be realized effectively.
General Layout of the Nervous System
 Central Nervous System (CNS): The division of the
nervous system located within the skull and spine.
 Peripheral Nervous System (PNS): The division of the
nervous system located outside the skull and spine.
Divisions of the Nervous System
 Central Nervous System (CNS): Comprised of the
brain (located in the skull) and the spinal cord (located
in the spine).
 Peripheral Nervous System (PNS): Comprised of the
somatic nervous system and the autonomic nervous
system.
 Somatic Nervous System (SNS): Interacts with the
external environment, with afferent nerves receiving
sensory signals from the skin, muscles, eyes, and
efferent nerves carrying motor signals to skeletal
muscles.
 Autonomic Nervous System (ANS): Regulates the
body's internal environment. Afferent nerves carry
sensory signals from internal organs to the CNS, and
efferent nerves carry motor signals from the CNS to
internal organs.
 Sympathetic Nervous System: Part of the autonomic
nervous system. Its sympathetic nerves project from
the lumbar and thoracic regions of the spinal cord,
involved in the fight-or-flight response.
 Parasympathetic Nervous System: Part of the
autonomic nervous system. Parasympathetic nerves
project from the brain and sacral region of the spinal
cord, primarily responsible for rest and digestion.
 Cranial Nerves: 12 pairs of nerves that project from
the brain. They include purely sensory nerves, such as
the olfactory and optic nerves, and nerves with both
sensory and motor functions. The vagus nerves are
the longest cranial nerves, extending to the gut.
Meninges:
 Protective membranes covering the CNS: dura mater
(outer tough membrane), arachnoid membrane
(spider-web-like membrane), and pia mater (inner
delicate membrane).
 Surround the brain and spinal cord, providing
protection.
 Cerebrospinal Fluid (CSF):
 Fills subarachnoid space, central canal of the spinal
cord, and cerebral ventricles in the brain.
 Supports and cushions the brain, absorbed from the
subarachnoid space into blood-filled spaces.
 Production and absorption processes are complex and
crucial for brain function.
 Blood-Brain Barrier:
 A barrier formed by tightly packed cells of cerebral
blood vessel walls.
 Regulates the passage of molecules into the brain,
allowing certain substances (e.g., glucose) to pass
while impeding large molecules and toxins.
Cells of the Nervous System:
 Neurons:
 Specialized for reception, conduction, and
transmission of electrochemical signals.
 Classified into multipolar (multiple processes),
unipolar (one process), bipolar (two processes), and
interneurons (short or no axon).
 External features include dendrites (receiving signals),
cell body (metabolic center), axon hillock (junction
with axon), and axon (signal transmission).
 Internal features encompass endoplasmic reticulum
(protein synthesis), mitochondria (energy release),
nucleus (DNA-containing structure), and synaptic
vesicles (store neurotransmitters).
 Glia (Glial Cells):
 Supportive cells in the nervous system, approximately
equal in number to neurons.
 Types include oligodendrocytes (form myelin sheaths
in CNS), astrocytes (maintain chemical environment),
microglia (immune defense), and Schwann cells (form
myelin sheaths in PNS).
 Glial Cells:
 Astrocytes: Large star-shaped cells in the nervous
system, involved in various functions such as synapse
regulation and blood-brain barrier maintenance.
 Oligodendrocytes: Cells responsible for myelinating
axons in the central nervous system.
 Schwann Cells: Glial cells that myelinate axons in the
peripheral nervous system.
 Microglia: Small glial cells that respond to injury and
play a role in cell death regulation and synapse
formation.
 Neuroanatomical Techniques:
 Golgi Stain: Staining method revealing neuronal
silhouettes by staining a few neurons entirely black.
 Nissl Stain: Dye-based method used to estimate the
number of neurons in an area by staining neuron cell
bodies.
 Electron Microscopy: Technique providing detailed
images of neuronal structures at the electron level.
 Neuroanatomical Tracing: Methods like anterograde
and retrograde tracing used to trace axonal paths in the
nervous system.
 Directional Coordinates:
 Anterior-Posterior (Rostral-Caudal): Front to back
directional axis in the nervous system.
 Dorsal-Ventral: Top to bottom directional axis, crucial
for understanding brain regions.
 Medial-Lateral: Middle to side directional axis,
indicating positions relative to the body's midline.
 Proximal-Distal: Closeness or farness from the central
nervous system, applicable in the peripheral nervous
system.
 Planes in Brain Sections:
 Horizontal Plane: Cut parallel to the ground, dividing
the brain into upper and lower parts.
 Frontal (Coronal) Plane: Cut from side to side,
creating front and back portions of the brain.
 Sagittal Plane: Cut lengthwise, dividing the brain into
left and right halves.
 Midsagittal Section: Sagittal cut at the midline,
separating the brain into perfectly symmetrical halves.
Anatomy of the Central Nervous System Reviewer
Spinal Cord
 Gray Matter: Inner H-shaped core of the spinal cord,
composed of cell bodies and unmyelinated
interneurons.
 White Matter: Surrounding area of myelinated axons,
giving it a glossy white sheen.
 Dorsal Horns: Two arms of spinal gray matter on the
dorsal side.
 Ventral Horns: Two arms of spinal gray matter on the
ventral side.
 Spinal Nerves: Pairs of nerves attached to the spinal
cord on the left and right sides, divided into dorsal
and ventral roots.
 Dorsal Root Axons: Sensory (afferent) unipolar
neurons, synapsing in dorsal horns.
 Ventral Root Neurons: Motor (efferent) multipolar
neurons, projecting to skeletal muscles or autonomic
ganglia.
Myelencephalon (Medulla)
 Reticular Formation: Complex network of nuclei in
the brain stem, involved in various functions including
arousal, sleep, and attention.
Metencephalon
 Pons: Division of the metencephalon creating a bulge
on the brain stem’s ventral surface.
 Cerebellum: Large structure on the brain stem’s
dorsal surface, vital for sensorimotor control and
cognitive functions.
Mesencephalon (Midbrain)
 Tectum: Dorsal surface of the midbrain, comprising
superior and inferior colliculi for auditory and visual-
motor functions.
 Tegmentum: Ventral to the tectum, housing
structures like the periaqueductal gray, substantia
nigra, and red nucleus.
Diencephalon
 Thalamus: Two-lobed structure atop the brain stem,
comprising sensory relay nuclei for processing and
transmitting sensory signals to the cortex.
 Hypothalamus: Regulates motivated behaviors,
homeostasis, and hormone release from the pituitary
gland.
 Optic Chiasm: Point where optic nerves from each
eye cross.
 Mammillary Bodies: Spherical nuclei involved in
memory and other functions.
Telencephalon
 Cerebral Cortex: Gray matter covering cerebral
hemispheres, involved in voluntary movement,
sensory interpretation, and complex cognitive
processes.
 Neocortex: Six-layered cortex mainly composed of
small, unmyelinated neurons.
 Lobes: Frontal, Parietal, Temporal, Occipital; divisions
on the surface of each hemisphere.
 Fissures: Large furrows dividing lobes; e.g., central
fissure, lateral fissure.
 Gyri: Ridges between fissures and sulci; e.g.,
precentral gyrus, postcentral gyrus.
 Basal Ganglia: Includes amygdala, striatum (caudate
and putamen), and globus pallidus; involved in
voluntary motor responses and decision making.
Limbic System
 Amygdala: Almond-shaped nucleus involved in
emotion, especially fear.
 Hippocampus: Important for certain forms of
memory, particularly spatial location.
 Cingulate Cortex: Large strip of cortex encircling the
dorsal thalamus.
 Fornix: Major tract connecting hippocampus and
mammillary bodies.
 Septum: Midline nucleus located at the anterior tip of
the cingulate cortex.
 Mammillary Bodies: Involved in memory and other
functions; located on the inferior surface of the
hypothalamus.
Synthesis and Packaging of Small-Molecule
Neurotransmitters:
1. Synthesis: Small-molecule neurotransmitters are
synthesized in the cytoplasm of the terminal button.
2. Packaging: They are packaged into synaptic
vesicles by the Golgi complex within the terminal
button.
3. Storage: The vesicles filled with neurotransmitter
are stored in clusters near the presynaptic membrane.
Synthesis and Packaging of Neuropeptides:
1. Synthesis: Neuropeptides are synthesized in the
cell body on ribosomes.
2. Packaging: They are packaged into vesicles by
the Golgi complex in the cell body.
3. Transport: These vesicles are transported along
microtubules to the terminal buttons at a rate of
about 40 centimeters per day.
4. Storage: Neuropeptide-containing vesicles are
usually larger than those containing small-molecule
neurotransmitters and are not as closely packed near
the presynaptic membrane.
Neurotransmitter Release, Receptors, and Synaptic 3. cell bodies - g
Transmission (Cell bodies, also known as somas, contain the
Important Terms and Explanations: nucleus and other organelles of the neuron.)
1. Neurotransmitter Release: 4. nondecremental - n
 Exocytosis: The process of neurotransmitter release, (Electrical synapses, facilitated by gap junctions, allow
where synaptic vesicles fuse with the presynaptic nondecremental transmission of signals between
membrane and empty their contents into the synaptic neurons.)
cleft upon stimulation by action potentials. 5. presynaptic facilitation - h
 Voltage-Activated Calcium Channels: Channels on the (Axoaxonic synapses modulate neurotransmitter
presynaptic membrane that open in response to release by enhancing or inhibiting the activity of
action potentials, allowing Ca2+ ions to enter the presynaptic neurons.)
neuron and trigger exocytosis. 6. nondirected synapses - f
2. Receptor Types: (Dendritic spines provide sites for synaptic
 Ionotropic Receptors: Receptors associated with connections and are examples of nondirected
ligand-activated ion channels. Neurotransmitter synapses.)
binding induces immediate postsynaptic potentials by 7. synaptic vesicles - k
opening or closing ion channels. (Synaptic vesicles store neurotransmitters and release
 Metabotropic Receptors: Receptors associated with them into the synaptic cleft during neuronal
signal proteins and G proteins. Neurotransmitter communication.)
binding initiates slower and more varied effects by 8. from cell body to terminal buttons - d
stimulating the synthesis of second messengers or (Axonal conduction of action potentials occurs along
binding to ion channels indirectly. the length of the axon, from the cell body to the
3. Neurotransmitter Binding: terminal buttons.)
 Receptor Subtypes: Different types of receptors to 9. acetylcholinesterase - m
which a neurotransmitter can bind. These subtypes (Acetylcholinesterase is an enzyme that breaks down
enable one neurotransmitter to transmit different acetylcholine, representing enzymatic degradation of
messages to different parts of the brain. neurotransmitters.)
 Autoreceptors: Metabotropic receptors located on 10. short amino acid chains - j
the presynaptic membrane that monitor (Neuropeptides are relatively large neurotransmitters
neurotransmitter levels in the synapse, regulating consisting of short amino acid chains.)
subsequent release. 11. saltatory - a
4. Neurotransmitter Deactivation: (Saltatory conduction is the rapid transmission of
 Reuptake: Common mechanism where action potentials along myelinated axons, "jumping"
neurotransmitters are drawn back into the from one node of Ranvier to another.)
presynaptic buttons by transporter mechanisms after 12. metabotropic receptors - l
release. (Metabotropic receptors are associated with G
 Enzymatic Degradation: Some neurotransmitters are proteins and signal proteins, leading to slower and
broken down by enzymes (e.g., acetylcholinesterase) varied effects in synaptic transmission.)
in the synapse, deactivating their effects. 13. electrical synapses - n
 Recycling: Neurotransmitter molecules, breakdown (Electrical synapses, facilitated by gap junctions, allow
products, and vesicles are recycled in the presynaptic direct electrical communication between neurons.)
neuron, maintaining efficiency. 14. spines - f
5. Glia and Gap Junctions: (Dendritic spines are small protrusions on dendrites,
 Glia: Supportive cells in the nervous system, providing sites for synaptic connections and
particularly astrocytes, involved in releasing chemical representing nondirected synapses.)
transmitters, containing receptors for
neurotransmitters, and coordinating neuronal activity. Neurotransmitter Classes and Drug Actions
 Gap Junctions: Narrow spaces bridged by connexins, Neurotransmitter Classes:
allowing electrical signals and small molecules to pass Amino Acid Neurotransmitters:
between adjacent cells. Facilitates rapid  Glutamate: Most prevalent excitatory
communication. neurotransmitter in the mammalian central nervous
system.
Matching Exercise: Connecting Terms with Definitions  Aspartate: Common excitatory neurotransmitter.
1. fatty - b  Glycine: Inhibitory neurotransmitter, prevalent in the
(Fatty substances make up the myelin sheath, which spinal cord.
facilitates saltatory conduction of action potentials.)  GABA (Gamma-Aminobutyric Acid): Most prevalent
2. sclerosis - a inhibitory neurotransmitter, but can have excitatory
(Sclerosis refers to the hardening or thickening of a effects at certain synapses.
tissue, often used to describe the degeneration of Monoamine Neurotransmitters:
myelin in diseases like multiple sclerosis.)
 Dopamine: Synthesized from tyrosine, involved in
mood regulation and reward pathways.
 Norepinephrine: Also called noradrenaline, involved
in stress response and attention.
 Epinephrine: Also called adrenaline, associated with
the "fight or flight" response.
 Serotonin: Derived from tryptophan, plays a role in
mood, appetite, and sleep regulation.
Indolamines:
 Serotonin: Classified as an indolamine, synthesized
from tryptophan.
Acetylcholine:
 Small-molecule neurotransmitter.
 Found at neuromuscular junctions, autonomic
nervous system synapses, and various central nervous
system synapses.
 Broken down by acetylcholinesterase in the synapse.
Unconventional Neurotransmitters:
 Soluble-Gas Neurotransmitters: Include nitric oxide
and carbon monoxide, diffuse through cell
membranes and stimulate second messengers.
 Endocannabinoids: Similar to THC, produced from
fatty compounds, have effects on presynaptic
neurons.
Neuropeptides:
 About 100 neuropeptides identified, functions
depend on their amino acid sequences.
 Categorized into pituitary peptides, hypothalamic
peptides, brain–gut peptides, opioid peptides, and
miscellaneous peptides.
Drug Actions on Synaptic Transmission:
Agonistic Drug Effects:
 Increase synthesis, storage, or release of
neurotransmitters.
 Bind to autoreceptors and block inhibitory feedback.
 Bind to postsynaptic receptors, either activating them
or enhancing the effects of neurotransmitters.
 Block deactivation processes (e.g., enzymatic
degradation or reuptake).
Antagonistic Drug Effects:
 Block synthesis of neurotransmitters.
 Cause neurotransmitter leakage from vesicles, leading
to degradation.
 Block release of neurotransmitters from terminal
buttons.
 Bind to autoreceptors, inhibiting neurotransmitter
release.
 Bind to postsynaptic receptors without activating
them, blocking access of the usual neurotransmitter.
Influential Research Areas Using Drugs:
Discovery of Receptor Subtypes:
 Different types of acetylcholine receptors (nicotinic
and muscarinic) have distinct distribution and actions,
dispelling the notion of one receptor per
neurotransmitter.
Discovery of Endogenous Opioids:
 Natural opioids like enkephalins and endorphins bind
to specific receptors, explaining the pain-killing effects
of opiates like morphine.
Antipsychotic Drugs and Schizophrenia:
 Effective antipsychotic drugs block dopamine D2
receptors, indicating excessive dopamine activity in
schizophrenia patients.
Understanding neurotransmitter classes and drug
actions is pivotal in comprehending brain function and
designing treatments for neurological disorders.
Neurotransmitter Classes and Drug Actions
Neurotransmitter Classes:
 Amino Acid Neurotransmitters:
 Glutamate: Most prevalent excitatory
neurotransmitter in the mammalian central nervous
system.
 Aspartate: Common excitatory neurotransmitter.
 Glycine: Inhibitory neurotransmitter, prevalent in the
spinal cord.
 GABA (Gamma-Aminobutyric Acid): Most prevalent
inhibitory neurotransmitter, but can have excitatory
effects at certain synapses.
 Monoamine Neurotransmitters:
 Dopamine: Synthesized from tyrosine, involved in
mood regulation and reward pathways.
 Norepinephrine: Also called noradrenaline, involved
in stress response and attention.
 Epinephrine: Also called adrenaline, associated with
the "fight or flight" response.
 Serotonin: Derived from tryptophan, plays a role in
mood, appetite, and sleep regulation.
Indolamines:
 Serotonin: Classified as an indolamine, synthesized
from tryptophan.
Acetylcholine:
 Small-molecule neurotransmitter.
 Found at neuromuscular junctions, autonomic
nervous system synapses, and various central nervous
system synapses.
 Broken down by acetylcholinesterase in the synapse.
Unconventional Neurotransmitters:
 Soluble-Gas Neurotransmitters: Include nitric oxide
and carbon monoxide, diffuse through cell
membranes and stimulate second messengers.
 Endocannabinoids: Similar to THC, produced from
fatty compounds, have effects on presynaptic
neurons.
Neuropeptides:
 About 100 neuropeptides identified, functions
depend on their amino acid sequences.
 Categorized into pituitary peptides, hypothalamic
peptides, brain–gut peptides, opioid peptides, and
miscellaneous peptides.
 distinct distribution and actions, dispelling the notion
of one receptor per neurotransmitter.
Discovery of Endogenous Opioids:
 Natural opioids like enkephalins and endorphins bind
to specific receptors, explaining the pain-killing effects
of opiates like morphine.
Neuroimaging Techniques in Biopsychology
Contrast X-ray Technique: Cerebral Angiography
 Definition: Infusion of radio-opaque dye into cerebral
artery for visualizing cerebral circulatory system
during x-ray photography.
 Application: Localizing vascular damage and detecting
tumor presence via displacement of blood vessels.
 Equipment: X-ray tube, detector rotating around the
patient's head, generating CT scans of horizontal brain
sections.
Radioactivity-Based Techniques: Positron Emission
Tomography (PET)
 Definition: Imaging brain activity using radioactive
fluorodeoxyglucose (FDG) injection, indicating active
neurons through accumulated radioactivity.
 Usage: Identifying brain molecules' distribution,
neurotransmitters, receptors, or transporters.
 Limitation: PET scans represent radioactivity levels,
requiring superimposition on brain images for
interpretation.
Magnetic-Field-Based Techniques: Magnetic
Resonance Imaging (MRI)
 Definition: High-resolution brain imaging using radio-
frequency waves emitted by hydrogen atoms aligning
with a magnetic field.
 Advantages: Provides detailed structural images,
including three-dimensional representations.
 Functional MRI (fMRI): Measures oxygen flow to
active brain areas, correlating neural activity with
cognitive tasks.
 Diffusion Tensor Imaging (DTI): Identifies major brain
tracts based on rapid water diffusion along axon
bundles.
Transcranial Stimulation Techniques:
 Transcranial Magnetic Stimulation (TMS):
Temporarily disrupts brain areas, aiding causal
relationship studies between brain activity and
cognition.
 Transcranial Direct Current Stimulation (tDCS):
Temporarily enhances cortical activity, enabling
controlled assessment of its effects on cognition and
behavior.
Psychophysiological Measures:
 Scalp Electroencephalography (EEG): Records gross
electrical brain activity, useful for diagnosing brain
pathologies and studying states of consciousness.
 Event-Related Potentials (ERPs): Components of EEG
reflecting neural responses to specific stimuli,
measured through signal averaging.
 Eye Movement Recording (EOG): Tracks eye
movements by measuring electrical potential changes
around the eyeball, essential in sleep research and
cognitive tasks.
 Magnetoencephalography (MEG): Records magnetic
fields resulting from neural activity changes, providing
insights into surface brain activity.
Psychophysiological Measures of Somatic Nervous
System Activity:
Muscle Tension:
 Each skeletal muscle composed of muscle fibers
contracts in an all-or-none fashion when activated by
motor neurons.
 Anxious individuals typically display high resting levels
of muscle tension, used as an indicator of
psychological arousal.
Electromyography (EMG):
 Procedure for measuring muscle tension.
 EMG activity recorded between electrodes taped to
skin surface over the muscle of interest.
 Raw EMG signal correlates with muscle contraction;
amplitude reflects the number of muscle fibers
contracting.
 Integrated EMG signal represents total EMG activity
per unit of time, providing a continuous measure of
muscle tension level.
Eye Movement:
 Electrooculography (EOG) records eye movements.
 Electrodes placed around the eyes measure changes
in electrical potential related to eye movement.
 Emotional thoughts and experiences increase skin's
ability to conduct electricity, measured through skin
conductance level (SCL) and skin conductance
response (SCR).
Blood Pressure:
 Blood pressure measured using sphygmomanometer
or more reliable automated methods.
 Blood pressure expressed as systolic over diastolic
pressure in millimeters of mercury (mmHg).
Invasive Physiological Research Methods:
Stereotaxic Surgery:
 Procedure for precisely positioning experimental
devices in the brain.
 Stereotaxic atlas provides directions to target site;
instrument used for precise placement.
Lesion Methods:
Aspiration Lesions:
 Delicately peeling off layers of cortical tissue using
suction through a fine-tipped glass pipette.
Radio-Frequency Lesions:
 Passing high-frequency current through
stereotaxically positioned electrode to create small
subcortical lesions.
Knife Cuts:
 Tiny, well-placed cuts used to eliminate conduction in
specific nerves or tracts.
Reversible Lesions:
 Temporarily eliminating activity in brain area through
cooling or injecting anesthetic.
Electrical Stimulation:
 Electrical brain stimulation achieved through bipolar
electrodes, eliciting behavioral responses depending
on location, current parameters, and test
environment.
Invasive Electrophysiological Recording Methods:
Intracellular Unit Recording:
 Records graded fluctuations in neuron's membrane
potential.
Extracellular Unit Recording:
 Records action potentials of neurons through
extracellular microelectrode.
Multiple-Unit Recording:
 Records total number of action potentials from
multiple nearby neurons.
Invasive EEG Recording:
 Records EEG signals through implanted electrodes in
laboratory animals.
Measuring Chemical Activity of the Brain:
 2-Deoxyglucose Technique:
- Radioactive 2-DG injected into animal, absorbed by
active neurons, brain slices subjected to
autoradiography to visualize brain activity areas.
 Cerebral Dialysis:
- Measures extracellular concentration of specific
neurochemicals in behaving animals, involving
implantation of tube with semipermeable section in
the brain for chemical analysis.
Locating Neurotransmitters and Receptors in the
Brain:
Immunocytochemistry:
 Uses antibodies labeled with dye or radioactive
element to locate neuroproteins in the brain.
 Can locate neurotransmitters by binding to their
enzymes.
 In Situ Hybridization:
 Utilizes hybrid RNA strands complementary to target
neuroprotein mRNA to mark location of neurons
releasing the neuroprotein.
Routes of Drug Administration:
 Drugs administered through feeding, intragastrically,
intraperitoneally, intramuscularly, subcutaneously,
intravenously, or stereotaxically implanted cannula for
direct brain delivery.
 Cannula used to overcome blood-brain barrier for
drug delivery to brain.
Selective Chemical Lesions:
 Selective neurotoxins like kainic acid, ibotenic acid, or
6-hydroxydopamine injected to make specific neuron-
destroying lesions.
In Situ Hybridization (Figure 5.19):
 Technique: In situ hybridization is a method used to
visualize the localization of specific mRNA molecules
in a tissue section.
 Description: A color-coded frontal section through a
rat brain shows high mRNA expression for an
endorphin in the striatum (in red and yellow).
 Significance: Reveals gene expression patterns in
specific brain regions.
Melanopsin Knockout Mice:
 Technique: Gene knockout involves deleting a specific
gene from an organism's genome.
 Description: Melanopsin knockout mice lack the gene
responsible for synthesizing melanopsin, a protein in
retinal neurons.
 Result: Knockout impaired but didn't eliminate the
mice's ability to adjust circadian rhythms in response
to light-dark cycles.
 Insight: Demonstrates melanopsin's role in regulating
circadian rhythms but highlights the complexity of
behavior genetics.
Gene Replacement Techniques:
 Technique: Gene replacement involves replacing a
defective gene with a functional one.
 Description: Defective human genes associated with
disorders like schizophrenia can be replaced in
animals like mice, creating transgenic mice.
 Potential: Offers possibilities for understanding
diseases and exploring gene therapies, although
challenges remain in practical application.
 Green Fluorescent Protein (GFP) and Brainbow:
 Technique: GFP fluoresces green when exposed to
blue light, aiding visualization of specific cells.
 Application: GFP can be inserted into target cells or
expressed universally, aiding researchers in
visualizing specific neurons.
 Brainbow: Technique utilizing mutated fluorescent
proteins of different colors to label neurons distinctly,
enabling tracing neural pathways.
 Opsins and Optogenetics:
 Technique: Opsins are light-sensitive ion channels
that, when illuminated, open and allow ions to enter or
exit the cell.
 Optogenetics: Involves inserting opsin genes into
specific neurons, enabling control of neuron activity
with light.
 Application: Used in living animals, optogenetics
allows precise modulation of neuronal activity, aiding
neuroscience research.
 Abbreviations:
 CT: Computed Tomography
 MRI: Magnetic Resonance Imaging
 PET: Positron Emission Tomography
 2-DG: 2-Deoxyglucose
 fMRI: Functional Magnetic Resonance Imaging
 MEG: Magnetoencephalography
 TMS: Transcranial Magnetic Stimulation
 EEG: Electroencephalography
 ERP: Event-Related Potential
 AEP: Auditory Evoked Potential
 EMG: Electromyography
 EOG: Electrooculography
 SCL: Skin Conductance Level
 SCR: Skin Conductance Response
 ECG: Electrocardiography
 EKG: Electrocardiogram
 IP: Intraperitoneal
 IM: Intramuscular
 IV: Intravenous
 SC: Subcutaneous
 6-OHDA: 6-Hydroxydopamine
 GFP: Green Fluorescent Protein
 Open-Field Test:
 Definition: A behavioral paradigm used to assess
species-common behaviors in animals. The subject is
placed in a large, barren chamber, and its activity,
including movement, exploration, and interaction
with the environment, is recorded.
 Indicators of Fearfulness: Low activity scores, high
bolus counts (excrement pieces), thigmotaxis
 (tendency to stay close to the walls), reduced
exploration.
 Application: Often used to measure anxiety and fear
responses in laboratory animals, especially rodents.
 Tests of Aggressive and Defensive Behavior:
 Definition: Behavioral paradigms designed to study
species-common aggressive and defensive behaviors.
 Indicators of Aggressive Behavior: Aggressive
postures, vocalizations, physical attacks, dominance
displays.
 Indicators of Defensive Behavior: Freezing, fleeing,
defensive postures, vocalizations.
 Application: Used to investigate social hierarchies,
territorial behaviors, and responses to threats or
challenges within animal populations.
 Tests of Sexual Behavior:
 Definition: Behavioral paradigms used to observe and
measure species-common sexual behaviors in
animals, especially mammals.
 Indicators of Sexual Behavior: Mounting,
intromission, copulatory behaviors, courtship rituals,
mate selection.
 Application: Applied in studies related to
reproductive behaviors, mate preferences, and the
effects of hormones or neural manipulations on
sexual responses.
Important Biopsychological Concepts and Terms
1. Aggressive and Defensive Behavior:
 Definition: Patterns of aggressive and defensive
behavior observed and measured during encounters
between dominant and intruder rats.
 Behaviors: Dominant male exhibits aggression
(sideways movement, hair erect, biting), intruder
exhibits defensiveness (rearing, pushing away, rolling
over).
2. Sexual Behavior in Rats:
 Definition: Study of physiological bases of rat sexual
behavior, focusing on copulatory acts and responses.
 Behaviors: Male mounts female, intromission occurs,
followed by ejaculation. Measures include mounts for
intromission, intromissions for ejaculation, and post-
ejaculation interval.
3. Learning Paradigms:
Pavlovian Conditioning Paradigm:
 Definition: Pairing a neutral stimulus with an
unconditional stimulus to elicit a conditional
response.
 Example: Tone (conditional stimulus) paired with
meat powder (unconditional stimulus) leads to
salivation (conditional response).
Operant Conditioning Paradigm:
 Definition: Response rate modification through
reinforcement (increases behavior) or punishment
(decreases behavior).
 Example: Animals press lever for self-stimulation;
reinforcing certain behaviors increases their
occurrence.
4. Seminatural Animal Learning Paradigms:
Conditioned Taste Aversion:
 Definition: Avoidance response to tastes of food
followed by illness, promoting survival by preventing
ingestion of harmful substances.
 Example: Rats associating a particular taste with
illness avoid consuming it.
Radial Arm Maze:
 Definition: Maze with arms radiating from a central
area; tests rodents' spatial memory and navigation
abilities.
 Example: Rats learn to visit baited arms and avoid
unbaited ones, relying on spatial cues for orientation.
Morris Water Maze:
 Definition: Circular pool with invisible escape
platform; assesses rats' spatial learning and memory
as they find the hidden platform.
 Example: Rats swim, locate the hidden platform, and
learn to navigate based on spatial cues.
 Conditioned Defensive Burying:
 Definition: Defensive behavior in response to aversive
stimuli, involving digging and burying the stimulus.
 Example: Rats bury aversive stimuli (e.g., shock
probe), exhibiting defensive responses for protection.
5. Significance of Learning Paradigms in
Biopsychological Research:
 Importance: Learning paradigms help psychologists
understand behavior, control animal responses, and
infer cognitive states.
 Relevance: Studying learning processes aids in
comprehending sensory, motor, motivational, and
cognitive aspects of animals.
Tests of Aggressive and Defensive Behavior:
 Colony-Intruder Paradigm:
 Definition: A method to observe and measure
aggressive and defensive behavior in rats during
encounters between dominant and intruder males.
 Behaviors: Dominant male displays aggression
(sideways movement, erect hair, biting), intruder
displays defensiveness (freezing, boxing, rolling over).
 Defensive Behavior Assessment:
 Definition: Tests assess rat's reactivity to the
experimenter, e.g., resistance to being picked up.
 Measurement: Scored categories range from no
resistance (lowest) to biting (highest).
 Elevated Plus Maze:
 Definition: A maze elevated above the floor with
closed and open arms, used to measure rat's
defensiveness/anxiety.
 Measurement: Proportion of time spent in protected
closed arms indicates defensiveness.
Tests of Sexual Behavior:
 Male Rat Sexual Behavior:
 Measures: Number of mounts for intromission,
intromissions for ejaculation, interval between
ejaculation and reinitiation of mounting.
 Female Rat Sexual Behavior:
 Measure: Lordosis quotient, indicating the proportion
of mounts that elicit lordosis (receptive posture).
Traditional Conditioning Paradigms:
 Pavlovian Conditioning Paradigm:
 Definition: Neutral stimulus (conditional stimulus)
paired with unconditional stimulus, leading to a
conditional response.
 Operant Conditioning Paradigm:
 Definition: Response (e.g., lever press) is reinforced or
punished, influencing its future occurrence.
 Example: Self-stimulation paradigm, where animals
stimulate brain pleasure centers by pressing a lever.
Seminatural Animal Learning Paradigms:
 Conditioned Taste Aversion:
 Definition: Avoidance response to food tastes
associated with illness.
 Importance: Enhances survival by preventing
consumption of potentially harmful substances.
 Radial Arm Maze:
 Definition: Maze with arms radiating from a central
area; tests spatial memory and foraging behavior in
rodents.
 Morris Water Maze:
 Definition: Circular pool where rats swim to find an
invisible platform; assesses spatial learning and
memory.
 Conditioned Defensive Burying:
 Definition: Rats bury an object associated with
aversive stimuli; used to study anxiety-related
behavior and drug effects.
 Retina Structure:
 Receptors: Specialized cells that detect light and
convert it into neural signals.
 Horizontal Cells: Neurons that facilitate lateral
communication between receptors and bipolar cells.
 Bipolar Cells: Neurons that relay signals from
receptors to ganglion cells.
 Amacrine Cells: Neurons that contribute to lateral
communication among bipolar, horizontal, and
ganglion cells.
 Retinal Ganglion Cells: Neurons whose axons form
the optic nerve and transmit visual information to the
brain.
 Binocular Disparity:
 Definition: The difference in the position of the same
image on the two retinas.
 Importance: Greater binocular disparity for close
objects helps the brain construct three-dimensional
perceptions from two-dimensional retinal images.
 Accommodation:
 Definition: The adjustment of the lens shape to focus
on objects at varying distances.
 Process: Controlled by ciliary muscles, allowing the
lens to flatten for distant objects and assume a curved
shape for close objects.
 Pupil and Iris:
 Pupil: The hole in the iris through which light enters
the eye.
 Iris: Donut-shaped bands of contractile tissue
controlling the pupil size.
 Function: Regulates the amount of light entering the
eye, balancing sensitivity and acuity.
 Light and Wavelength:
 Light Definition: Electromagnetic energy perceived
by the visual system.
 Wavelength: Distance between wave peaks;
determines color perception.
 Visible Spectrum: Wavelengths between 380 and 760
nanometers detectable by the human visual system.
 Binocularity and Vision: Front-facing eyes provide
depth perception by combining images from both
eyes, enhancing visual understanding.
 Retinal Processing:
 Neural Pathway: Light passes through retinal layers
before reaching receptors; signals are transmitted back
through layers to ganglion cells.
 Inside-Out Structure: Receptors activated after light
passes through other retinal layers.
 Blind Spot: Gap in the receptor layer where ganglion
cell axons exit the eye, creating a visual blind spot.
 Visual Illusions:
 Fortification Illusions: Visual disturbances preceding
migraine headaches, characterized by expanding gray
areas and flickering lines.
 Significance: Studied to understand visual processing
mechanisms; demonstrates perceptual phenomena
related to neurological conditions.
Evolution of Eye Position:
 Side-by-Side Eyes: Provides a larger field of vision,
beneficial for prey animals to detect predators from
various directions.
 Front-Facing Eyes: Offers depth perception by
combining images from both eyes, advantageous for
predators to accurately assess distances.
Binocular Disparity in Depth Perception:
 Demonstration: Compare views with each eye
closed; notice greater disparity for closer objects.
 Importance: Brain uses binocular disparity to
construct three-dimensional perceptions from two
retinal images, enhancing depth perception.
 Retinal Neurons:
 Receptors: Specialized cells detecting light, initiating
neural signals.
 Horizontal Cells: Facilitate lateral communication
between receptors, bipolar cells.
 Bipolar Cells: Relay signals from receptors to
ganglion cells.
 Amacrine Cells: Specialized for lateral
communication, enhancing neural coordination.
 Retinal Ganglion Cells: Axons form optic nerve,
transmit visual data to the brain.
 Variety of Subtypes: Over 60 retinal neuron types
identified, including 30 retinal ganglion cell subtypes
(Cepko, 2015; Seung & Sümbül, 2014).
Communication in the Retina:
 Chemical and Electrical Signaling: Neurons
communicate chemically via synapses and electrically
via gap junctions (Pereda, 2014).
 Lateral Communication: Horizontal and amacrine
cells specialized for communication across major
sensory input channels.
Retinal Layer Arrangement:
 Inside-Out Structure: Light passes through four
layers before reaching receptors, initiating neural
transmission.
  Signal Transmission: Activated receptors transmit
neural messages back through retinal layers to
ganglion cells.
 Visual Challenges: Incoming light distorted by retinal
tissue; gap in receptor layer creates the blind spot.
 Visual Completion:
 Filling in Blind Spots: Visual system fills in missing
information, creating continuous perceptions.
 Continuous Bar Phenomenon: Brain completes
visual input, demonstrating the visual system's
complexity.
 Role in Normal Vision: Completion vital in normal
vision, creating perception of entire objects from
partial information.
 Surface Interpolation: Extracts edge information,
inferring appearance of large unpatterned surfaces.
 Fovea: Specialized for high-acuity vision, reduces
distortion of incoming light, minimizing visual
challenges.
 Creative Solution to Blind Spot: Brain creatively
fills blind spot gaps using surrounding receptor
information.
Cone and Rod Vision:
 Duplexity Theory: Cones (photopic vision) for high-
acuity color vision in good lighting; rods (scotopic
vision) for dim illumination, lack detail and color.
 Convergence Differences: Scotopic system has high
convergence (less acuity), rods converge on a single
retinal ganglion cell; photopic system has low
convergence (higher acuity), fewer cones converge on
each retinal ganglion cell.
 Distribution on Retina: Fovea contains cones,
specialized for high-acuity vision; rods more prevalent
in peripheral retina, less detail but increased
sensitivity.
Spectral Sensitivity and Purkinje Effect:
 Spectral Sensitivity Curves: Photopic (cone-
mediated) and scotopic (rod-mediated) sensitivity
curves vary, affecting perception of different
wavelengths.
 Purkinje Effect: Shift in relative brightness perceived
during transition from photopic to scotopic vision due
to differences in spectral sensitivity curves.
Eye Movement and Fixational Eye Movements:
 Involuntary Eye Movements: Tremor, drifts,
saccades; critical for visual function, prevent fading of
visual objects during fixation.
 Role of Eye Movements: Images need to move on the
retina; stabilized retinal images cause objects to fade
and reappear, demonstrating the importance of eye
movements in vision.
 Visual Transduction:
 Definition: Visual transduction is the conversion of
light to neural signals by visual receptors.
 Discovery of Rhodopsin: Breakthrough in 1876 with
extraction of rhodopsin, red pigment in rods.
 Rhodopsin Properties: Rhodopsin bleached and lost
light-absorbing capacity in continuous intense light,
regained function in the dark.
 The process of visual perception is a complex
interplay of various specialized cells and mechanisms
within the eye and the brain. To summarize the key
points discussed in the text:
Retinal Neurons:
 Types of Neurons: Receptors, horizontal cells,
bipolar cells, amacrine cells, and retinal ganglion cells
form the neural network in the retina.
 Subtypes: Over 60 different types of retinal neurons
identified, including about 30 retinal ganglion cell
subtypes.
 Communication: Neurons communicate chemically
via synapses and electrically via gap junctions.
Horizontal and amacrine cells specialize in lateral
communication.
Visual Completion and Perception:
 Filling in Blind Spots: Visual system fills in missing
information, creating continuous perceptions even in
blind spots.
 Continuous Bar Phenomenon: Brain completes
visual input, showcasing the visual system's ability to
construct a whole perception from partial information.
 Role in Normal Vision: Completion is essential for
normal vision, allowing perception of entire objects
from limited information.
 Surface Interpolation: Extracts edge information to
infer appearance of large surfaces, emphasizing the
brain's role in perception.
 Cone and Rod Vision:
 Duplexity Theory: Cones mediate high-acuity color
vision in good lighting; rods dominate in dim
illumination but lack detail and color.
 Convergence Differences: Scotopic system (rods) has
high convergence, sacrificing acuity for sensitivity.
Photopic system (cones) has low convergence,
preserving acuity.
 Distribution on Retina: Fovea contains cones
specialized for high-acuity vision; rods more prevalent
in peripheral retina, enhancing sensitivity in low light.
Spectral Sensitivity and Purkinje Effect:
 Spectral Sensitivity Curves: Photopic and scotopic
spectral sensitivity curves differ, influencing
perception of different wavelengths.
 Purkinje Effect: Shift in relative brightness during
transition from photopic to scotopic vision due to
differences in spectral sensitivity curves.
Eye Movements and Fixational Eye Movements:
 Involuntary Movements: Tremor, drifts, saccades;
crucial for maintaining visual perception during
fixation.
 Role of Eye Movements: Prevent fading of visual
objects by keeping retinal images in motion,
highlighting their vital role in vision.
 Visual Transduction:
 Definition: Visual transduction is the conversion of
light to neural signals by visual receptors.
 Rhodopsin Discovery: Rhodopsin, a red pigment in
rods, undergoes bleaching in continuous intense light
but regains function in the dark, playing a central role
in visual transduction.
Rhodopsin and Rod-Mediated Vision:
 Rhodopsin Absorption: Rhodopsin's light absorption
and bleaching are fundamental in rod-mediated vision.
 Predicting Human Vision: Rhodopsin's absorption
predicts human vision under various conditions,
especially scotopic (low-light) conditions.
 Relationship with Scotopic Sensitivity: Rhodopsin's
absorption of different wavelengths corresponds to
 human and animal ability to detect various
wavelengths under scotopic conditions.
 Rhodopsin as a G-Protein–Coupled Receptor:
Rhodopsin is a G-protein–coupled receptor responding
to light, initiating intracellular chemical events.
Retina-Geniculate-Striate System:
 Predominant Pathways: About 90% of retinal
ganglion cell axons are part of retina-geniculate-striate
pathways.
 Retinotopic Organization: Each level of the system
maps the retina; adjacent stimuli on the retina excite
adjacent neurons in the system.
 Primary Visual Cortex (Striate Cortex): Receives
signals from lateral geniculate nuclei, processes retinal
information, disproportionate representation of the
fovea.
M and P Channels:
 Parallel Channels: Lateral geniculate nuclei transmit
signals through two parallel channels—parvocellular
(P) and magnocellular (M) layers.
 P (Parvocellular) Channels: Responsive to color,
fine pattern details, stationary or slow-moving objects;
majority input from cones.
 M (Magnocellular) Channels: Responsive to
movement; majority input from rods.
 Projection to Visual Cortex: M and P areas in lower
layer IV project to different areas in visual cortex.
 Edge Perception:
 Significance of Edges: Edges define object extents
and positions, crucial for visual perception.
 Visual Edge Definition: Visual edge is the junction of
two different areas in a visual image.
 Species' Edge Perception: Many species excel in
edge perception due to evolutionary pressure.
Quiz Questions (Fill in the Blanks):
4. Iris are donut-shaped bands regulating
light reaching the retina.
5. Prey animals have sideways eyes,
enabling a larger field of vision against predators.
6. Binocular disparity is the difference in
the position of the same image on two retinas.
7. Amacrine cells and horizontal cells are
responsible for lateral communication.
8. Surface interpolation is the process of
perceiving large surfaces by extracting edge
information and making inferences about shape.
9. In scotopic vision, hundreds of rods
converge on a single retinal ganglion cell.
10. The majority of visual neurons respond
only to changing images.
11. Rhodopsin is a G-protein–coupled
receptor that does not respond to neurotransmitter
molecules.
12. The path that all signals take to reach the
right primary visual cortex from the left visual field is
either ipsilateral or contralateral.
Lateral Inhibition and Contrast Enhancement:
 Mach Bands: Illusory bands, Mach bands, enhance
contrast at edges, making them appear more distinct.
 Contrast Enhancement Mechanisms: Contrast
enhancement occurs in our nervous systems, making
perceived edges sharper than actual physical
properties of light.
 Physiological Basis in Horseshoe Crabs: Study on
horseshoe crabs demonstrated lateral inhibition in
ommatidia, where firing rates and inhibition levels
affect contrast enhancement.
Methods of Hubel and Wiesel:
 Mapping Receptive Fields: Hubel and Wiesel
mapped receptive fields using microelectrodes,
recording responses to various stimuli.
 Receptive Fields Characteristics: Receptive fields in
the foveal area are smaller, circular, and monocular at
all levels (retinal ganglion cells, lateral geniculate
neurons, and lower layer IV neurons).
 Receptive Fields in Retina-Geniculate-Striate
System:
 Excitatory and Inhibitory Areas: Neurons in the
retina-geniculate-striate system have receptive fields
with excitatory and inhibitory areas, with responses
varying based on the location of light within these
areas.
 On-Center and Off-Center Cells: Neurons can be
classified as on-center or off-center cells. On-center
cells respond with "on" firing to central light and "off"
firing to peripheral light. Off-center cells show the
opposite response pattern.
 Contrast Sensitivity: On-center and off-center cells
respond best to contrast, where maximizing contrast
between center and periphery of the receptive field
influences firing rates.
Characteristics of Simple and Complex Cells:
 Simple Cells: Receptive fields of simple cells have
antagonistic "on" and "off" regions, respond to bars of
light or dark edges, and have straight-line borders.
They are monocular and exhibit orientation specificity.
 Complex Cells: Receptive fields of complex cells are
larger, respond to straight-line stimuli in specific
orientations, and do not have static "on" and "off"
regions. They are also monocular but can be binocular,
with ocular dominance and sensitivity to retinal
disparity for depth perception.
Organization of Primary Visual Cortex:
 Vertical Columns: Primary visual cortex is organized
into functional vertical columns, each responding to
stimuli from the same retinal area, dominated by the
same eye, and preferring the same straight-line angles.
 Clustering and Preferences: Functional columns
analyzing input from one retinal area cluster together,
with half receiving input from the left eye and the
other half from the right eye. Each cluster contains
neurons with various orientation preferences for
straight-line stimuli.
Excitatory and Inhibitory Areas:
 Excitatory Area: Part of a neuron's receptive field
where stimulation leads to an increase in firing rate.
 Inhibitory Area: Part of a neuron's receptive field
where stimulation leads to a decrease in firing rate.
On-Center and Off-Center Cells:
 On-Center Cells: Neurons that respond with increased
firing ("on" firing) when light is shone in the central
 region of their receptive fields and decreased firing
("off" firing) in response to light in the periphery.
 Off-Center Cells: Neurons that respond with
decreased firing ("off" firing) when light is shone in
the central region of their receptive fields and
increased firing ("on" firing) in response to light in the
periphery.
Contrast Sensitivity:
 Contrast Sensitivity: The ability of neurons, especially
on-center and off-center cells, to respond best to the
contrast between the center and periphery of their
receptive fields. Maximizing this contrast influences
firing rates.
Simple and Complex Cells:
 Simple Cells: Neurons in the primary visual cortex
with receptive fields having antagonistic "on" and
"off" regions. They respond to bars of light or dark
edges and have straight-line borders. Simple cells are
monocular and exhibit orientation specificity.
 Complex Cells: Neurons in the primary visual cortex
with larger receptive fields that respond to straight-
line stimuli in specific orientations. They lack static
"on" and "off" regions. Complex cells can be
monocular or binocular, have ocular dominance, and
are sensitive to retinal disparity for depth perception.
Vertical Columns:
 Vertical Columns: Functional columns in the primary
visual cortex where neurons respond to stimuli from
the same retinal area, are dominated by the same
eye, and prefer the same straight-line angles.
Cortical Mechanisms of Vision and Conscious
Awareness: Visual Cortex Organization Visual Cortex
Classes:
 Primary Visual Cortex: Receives input from thalamic
visual relay nuclei (lateral geniculate nuclei); located
in the posterior occipital lobes.
 Secondary Visual Cortex: Receives input from primary
visual cortex.
 Visual Association Cortex: Receives input from
secondary visual cortex and other sensory systems'
secondary areas.
 Locations in the Brain:
 Prestriate Cortex: Surrounds the primary visual
cortex; part of secondary visual cortex.
 Inferotemporal Cortex: Located in the inferior
temporal lobe; part of secondary visual cortex.
 Posterior Parietal Cortex: Largest area of association
cortex receiving visual input.
 Flow of Visual Information:
 Primary to Secondary Visual Cortex: Visual
information flows from primary visual cortex to
various secondary visual cortex areas.
 Hierarchy in Visual Processing: Neurons in higher
visual areas have larger receptive fields and respond
to more specific and complex stimuli as compared to
lower areas.
Term Connections from Exercise: Scan Your Brain
1. Ommatidia: Large receptors (c) in the lower layer
IV.
2. Color Constancy: Contextual influences (d)
ensuring perceived color stability despite varying
illumination.
3. Simple Cortical Cells: Firing on the periphery of
receptive field (f) for contrast enhancement.
4. Striate Cortex: Primary visual cortex, main
entrance point for visual signals.
5. Perception of Edges: Contrast-enhancement (g)
for clear edge detection.
6. Trichromatic: Component theory (e), which
involves three types of cones.
7. Stimulation Outside Receptor Field: Contextual
influences (d) affecting neuron responses beyond
receptive fields.
8. Off-Center Cells: Unresponsive to diffuse light
(h), providing specific responses to light in the center
of the receptive field.
9. Cytochrome Oxidase: Mitochondrial enzyme (b)
used to visualize peglike columns rich in these
enzymes.
Damage to Primary Visual Cortex: Scotomas and
Completion
 Scotoma: Area of blindness in the contralateral visual
field of both eyes caused by damage to the primary
visual cortex.
 Perimetry Test: Used to map scotomas; patient
focuses on a fixation point, and areas of blindness are
identified when the patient fails to see a flashed light.
 Completion Phenomenon: Patients with scotomas
may complete images even in blind areas, leading to
conscious unawareness of deficits. Example: seeing a
complete face when focusing on a nose despite half
the face being in the scotoma.
 Blindsight: Ability to respond to visual stimuli in a
scotoma without conscious awareness. Patients might
accurately reach for objects they claim not to see.
Functional Aspects of Blindsight:
 Blindsight in Scotoma Patients: Patients can respond
to visual stimuli in their blind field without conscious
awareness.
 Example: D.B. Case: Patient with blindsight couldn't
consciously see objects in his blind field but could
accurately reach and differentiate objects when
forced to guess.
Functional Areas of Secondary and Association Visual
Cortex
 Functional Areas: More than 30 different functional
areas of visual cortex in macaque monkey, specialized
for various visual analyses.
 Interconnections: Over 300 interconnecting pathways
identified between these areas, often reciprocal
connections.
 Human Visual Cortex: About a dozen functional areas
identified in human visual cortex using PET, fMRI, and
evoked potentials.
Dorsal and Ventral Streams (LO 6.22):
  Dorsal Stream: From primary visual cortex to dorsal
prestriate cortex to posterior parietal cortex;
processes spatial stimuli, like object locations and
movement.
 Ventral Stream: From primary visual cortex to ventral
prestriate cortex to inferotemporal cortex; processes
object characteristics, such as color and shape.
 Specificity in Ventral Stream: Neuronal clusters
respond to specific object classes (e.g., faces,
animals), indicating different visual functions within
the ventral stream.
Processing Auditory Information:
 Cochlear Sensitivity:
 Humans can detect differences in pure tones differing
in frequency by as little as 0.2 percent.
 Cochlear coding principle: Different frequencies
stimulate hair cells at different points along the
basilar membrane.
 Complex sounds activate multiple sites, carried by
various auditory neurons.
 Tonotopic Organization:
 Similar to the retinotopic organization in the visual
system.
 Auditory system is organized tonotopically, where
different frequencies are processed in specific regions
of the cochlea.
Auditory Processing Challenge:
 Auditory system must process multiple frequencies
simultaneously in complex environments (like
parties).
 Auditory system manages to separate and combine
frequency messages from different sources.
 The mechanism behind this ability remains a mystery.
Pathways from Ear to Primary Auditory Cortex:
 No single major auditory pathway comparable to the
retina-geniculate-striate pathway in the visual system.
 Complex network of auditory pathways.
 Axons from auditory nerves synapse in cochlear
nuclei, leading to superior olives, inferior colliculi,
medial geniculate nuclei of the thalamus, and finally,
the primary auditory cortex.
 Auditory signals from each ear are combined at a low
level (superior olives) before transmission to both
ipsilateral and contralateral auditory cortex.
Sound Localization Mechanisms:
 Mediated by lateral and medial superior olives in
different ways.
 Neurons in medial superior olives respond to
differences in the time of arrival of signals from both
ears, aiding in sound localization.
 Neurons in lateral superior olives respond to
differences in sound amplitude from both ears.
 Both medial and lateral superior olives project to the
superior colliculus, mapping auditory space, indicating
the location of sensory inputs in space.
Studying Sound Localization in Barn Owls:
 Barn owls are exceptional at locating sources of
sounds, especially in the dark.
 Auditory neurons in the barn owl's superior colliculus
are finely tuned, responding to specific locations
within the owl's hearing range.
Organization of Auditory Cortex:
 Primary Auditory Cortex (Core Region):
 Receives majority input from the medial geniculate
nucleus.
 Located in the temporal lobe, hidden within the
lateral fissure.
 Comprises three adjacent areas: A1, R, and RT.
Secondary Auditory Cortex:
 Surrounds the core region, forming the belt.
 Areas outside the belt are parabelt areas.
 Approximately 20 separate areas of auditory cortex
exist in primates.
Auditory Pathways:
 Anterior Auditory Pathway (What) and Posterior
Auditory Pathway (Where):
 Anterior pathway is involved in identifying sounds.
 Posterior pathway is involved in locating sounds.
 Posterior pathway may also be related to action
preparation.
Auditory-Visual Interactions:
 Posterior Parietal Cortex:
 Some neurons have visual, auditory, or both receptive
fields, covering the same location in the environment.
 Functional brain imaging shows sensory interactions
occur not only in association cortex but also at the
primary sensory cortex level.
Perception of Pitch:
 Perception of pitch occurs in a specific area anterior
to the primary auditory cortex.
 This area responds to pitch rather than frequency,
suggesting it's where sound frequencies are
converted into the perception of pitch.
Effects of Auditory System Damage:
 Auditory Cortex Damage:
 Studying human auditory cortex damage is
challenging due to its location.
 Lesions in nonhumans have been studied.
 Large lesions involving core, belt, and parabelt areas
surprisingly do not result in severe permanent
deficits, indicating the complexity of subcortical
circuits in auditory functions.
Effects of Auditory Cortex Lesions:
 Similarity Across Species (Humans and Monkeys):
 Following bilateral lesions, there is temporary
complete hearing loss, which recovers in subsequent
weeks.
 Permanent effects include the loss of sound
localization ability and impaired frequency
discrimination.
Unilateral Auditory Cortex Lesions:
 Disrupts the ability to localize sounds contralateral to
the lesion but has lesser effects on ipsilateral sounds.
 Unilateral lesions lead to minor deficits in
 distinguishing contralateral sounds.
 Deafness in Humans:
 Prevalence and Causes:
 Deafness affects around 360 million people globally.
 Total deafness is rare, occurring in only 1% of hearing-
impaired individuals.
 Severe hearing issues commonly result from damage
to the inner ear, middle ear, or the nerves connected
to them.
Types of Hearing Impairments:
 Conductive deafness: Associated with ossicle damage.
 Nerve deafness: Related to cochlear or auditory nerve
damage, often due to hair cell receptor loss.
Specific Frequency Deafness:
 Partial cochlear damage can result in specific
frequency nerve deafness.
 Elderly individuals often have difficulty hearing high
frequencies, affecting speech understanding.
Tinnitus and Deafness:
 Hearing loss can be associated with tinnitus (ringing
of the ears).
 Tinnitus persists even if the affected ear's nerve is cut,
indicating central auditory system changes as the
cause.
Cochlear Implants:
 Cochlear implants can benefit some individuals with
nerve deafness.
 Implants convert sounds into electrical signals,
stimulating the auditory nerve.
 Early implantation is crucial to maximize benefits, as
disuse can lead to alterations in auditory neural
pathways.
Knowledge Review Questions:
1. The area of the sensory cortex receiving input
directly from thalamic relay nuclei is the primary
sensory cortex.
2. Sensation is the process of detecting the
presence of stimuli.
3. Simultaneous analysis of a signal in different
ways by multiple pathways of a neural network is
referred to as parallel processing.
4. The mathematical procedure for breaking down
complex waves into component sine waves is called
Fourier analysis.
5. Sine wave vibrations are also called pure tones.
6. The three auditory ossicles are malleus, incus,
and stapes.
7. The layout of the auditory system tends to be
tonotopic, organized by frequency.
8. The axons of auditory nerves synapse in the
ipsilateral cochlear nuclei.
9. One function of the superior olives is sound
localization.
10. The thalamus is made up of a fine sheet of
neurons located just underneath the neocortex,
toward the middle of the brain.
11. The oval window is the membrane transferring
vibrations from the ossicles to the fluid of the cochlea.
12. Many studies of auditory-visual interactions have
focused on association cortex in the posterior parietal
cortex.
Somatosensory System
 Somatosensations: Sensations of touch and pain from
the body are known as somatosensations.
 Somatosensory System Divisions:
 Exteroceptive System: Senses external stimuli applied
to the skin.
 Proprioceptive System: Monitors body position
information from muscles, joints, and balance organs.
 Interoceptive System: Provides general information
about internal body conditions (e.g., temperature,
blood pressure).
Cutaneous Receptors:
 Variety of Receptors: Skin contains various receptors,
including free nerve endings, Pacinian corpuscles,
Merkel's disks, and Ruffini endings.
 Receptor Adaptation: Receptors adapt either rapidly
(e.g., Pacinian corpuscles) or slowly (e.g., Merkel's
disks) to stimuli, influencing tactile sensations.
 Functional Significance: Fast and slow adaptation
provides information about dynamic and static
qualities of tactual stimuli.
Dermatomes:
 Definition: Areas of the body innervated by left and
right dorsal roots of a specific segment of the spinal
cord.
 Overlap: Significant overlap between adjacent
dermatomes reduces somatosensory loss if a single
dorsal root is damaged.
Somatosensory Pathways:
 Dorsal-Column Medial-Lemniscus System:
 Functions: Carries touch and proprioception
information.
 Pathway: Sensory neurons enter spinal cord, ascend
ipsilaterally in dorsal columns, synapse in medullary
dorsal column nuclei, decussate, and reach
contralateral ventral posterior thalamus.
 Projection Areas: Primary somatosensory cortex (SI),
secondary somatosensory cortex (SII), and posterior
parietal cortex.
Anterolateral System:
 Functions: Carries pain and temperature information.
 Pathway: Most neurons synapse upon entry into
spinal cord, then decussate (or ascend ipsilaterally)
and travel to the brain in contralateral anterolateral
spinal cord tracts.
 Tracts: Includes spinothalamic tract, spinoreticular
tract, and spinotectal tract.
 Projection Areas: Somatosensory cortex and other
brain regions receive thalamic pain and temperature
information.
 Functional Implications: Complete transection of
both ascending somatosensory paths results in the
loss of body sensation below the injury site.
Cortical Areas of Somatosensation:
Primary Somatosensory Cortex (SI):
 Somatotopic Organization: Mapped by Penfield and
colleagues in 1937.
  Mapping Method: Electrical stimulation of cortical
surface in conscious patients.
 Result: Patients reported somatosensory sensations
in various body parts.
 Somatotopic Map: Organized according to a map of
the body surface.
 Somatosensory Homunculus: Represents
somatotopic organization; distorted, with more
representation for body parts used in tactile
discriminations (e.g., hands, lips, tongue).
 SI Sensitivity: Most sensitive to tactile input from
body parts involved in discrimination.
 Adjacent Area: Secondary somatosensory cortex (SII)
located ventral to SI in the postcentral gyrus and
extending into the lateral fissure.
SII Features:
 Input: Receives input from SI and is considered
secondary somatosensory cortex.
 Contralateral and Ipsilateral Input: Receives
substantial input from both sides of the body.
 Output: Projects to the association cortex of the
posterior parietal lobe.
 Columnar Organization: Similar to visual and auditory
cortex, primary somatosensory cortex exhibits
columnar organization.
 Functional Strips: Four functional strips within SI,
each with specific somatotopic organization and
sensitivity to different somatosensory inputs (e.g.,
light touch, pressure).
Functional Implications and Deficiencies:
 Tactile Discrimination: Different areas of primary
somatosensory cortex prefer specific types of tactile
stimulation (e.g., light touch or heat) on the same
body part.
 Discrimination Difficulties: Limited capacity of the
back's representation in SI results in difficulty
discriminating objects touching the back.
 Demonstration: Test tactile discrimination abilities on
a friend's back with varying numbers of simultaneous
touches, noticing the challenges in discrimination
unless touches are considerably apart.
 Object Perception: Dorsal stream from SI to posterior
parietal cortex contributes to multisensory integration
and attention direction. Ventral stream projects to SII
and participates in object shape perception.
Effects of Damage to Primary Somatosensory Cortex:
 Mild Effects: Damage effects are often mild due to
parallel pathways in the somatosensory system.
 Study: Corkin, Milner, and Rasmussen (1970) assessed
the effects of damage to the primary somatosensory
cortex.
Somatosensory System and Association Cortex:
Processing Somatosensory Signals in Association
Cortex:
 Destination: Somatosensory signals are sent to
association cortex in prefrontal and posterior parietal
cortex.
 Posterior Parietal Cortex: Contains bimodal neurons
responding to both somatosensory and visual stimuli.
 Spatial Relationship: Visual and somatosensory
receptive fields of bimodal neurons are spatially
related; movements of the hand affect visual
receptive fields.
 Case Study - W.M.: Stroke-induced left hemianopsia;
improved visual detection in left visual field when left
hand was extended, further improved with a tennis
racket, demonstrating somatosensory influence on
visual perception.
Somatosensory Agnosias:
 Types: Two major types of somatosensory agnosia:
1. Astereognosia: Inability to recognize
objects by touch.
2. Asomatognosia: Failure to recognize
parts of one's own body, usually unilateral, associated
with extensive damage to right temporal and
posterior parietal lobe.
 Association with Anosognosia: Often accompanies
anosognosia, where patients fail to recognize their
own symptoms; common in neurological disorders.
 Relation to Contralateral Neglect: Asomatognosia is a
component of contralateral neglect, the tendency not
to respond to stimuli contralateral to a right-
hemisphere injury.
Rubber-Hand Illusion:
 Phenomenon: Feeling an extraneous object (e.g.,
rubber hand) is part of one's own body.
 Induction Method: Synchronous stroking of hidden
hand and visible rubber hand creates illusion.
 Neural Mechanisms: Involves association cortex in
posterior parietal and frontal lobes; bimodal neurons
with visual and somatosensory fields are crucial.
Perception of Pain:
 Adaptiveness of Pain: Paradoxical aspect; warns
against harmful activities or signals need for
treatment.
 No Clear Cortical Representation: Painful stimuli
activate various cortical areas (thalamus, SI, SII, insula,
anterior cingulate cortex), but no specific cortical area
is necessary for pain perception.
 Cortical Areas: Anterior cingulate cortex involved in
expectation, emotional reaction, and adaptive
responses to pain, not in perception itself.
 Descending Pain Control: Cognitive and emotional
factors, as well as analgesic drugs, can effectively
suppress pain.
 Descending Analgesia Circuit: Periaqueductal gray
(PAG) stimulation has analgesic effects; specialized
receptors for opioid analgesic drugs found in PAG;
endogenous opioid analgesics like endorphins block
pain signals through descending circuit.
 Model: Descending analgesia circuit involves PAG
stimulating serotonergic neurons of raphé nuclei,
which excite interneurons in spinal cord dorsal horn,
blocking incoming pain signals.
Neuropathic Pain:
 Definition: Neuropathic pain is severe chronic pain
without a recognizable pain stimulus, often triggered
by innocuous stimuli.
 Causes: Caused by pathological changes in the
nervous system induced by an original injury;
involvement of aberrant microglia triggering
hyperactivity in neural pain pathways.
 Ineffectiveness of Treatments: Neuropathic pain
persists even after cutting nerves in perceived pain
location; existing medications for injury-related pain
are usually ineffective.
Chemical Senses: Smell and Taste:
 Olfaction (Smell) and Gustation (Taste): Chemical
senses monitoring airborne and solution-based
chemicals respectively in the environment.
 Adaptive Roles:
1. Evaluation of Potential Foods:
Encourages consumption of energy and nutrient
sources, avoiding toxins, especially in natural
environments.
2. Social Interactions: Regulates social
interactions in many species; pheromones influence
physiology and behavior of conspecifics; example of
hamsters' aggressive and sexual behavior controlled
by pheromones.
 Human Pheromones: Research on human
pheromones suggests their presence (e.g., menstrual
synchronization, sex prediction), but direct evidence
as sex attractants remains inconclusive.
Olfactory System:
 Receptor Cells: Olfactory receptor cells in nasal
mucosa contain receptor protein molecules
stimulated by airborne chemicals.
 Receptor Diversity: Rats and mice have about 1,000
receptor types; humans have about 300; each
olfactory receptor cell contains one type of receptor
protein.
 Encoding Odors: Each odor encoded by pattern of
activity across receptor types; axons terminate in
olfactory glomeruli, discrete clusters near olfactory
bulbs' surface.
 Topographic Organization: Glomeruli sensitive to
specific odors systematically arrayed on olfactory
bulbs; mirror symmetry between left and right bulbs;
similar layout in related species.
 Neurogenesis: New olfactory receptor cells
continuously created, replacing deteriorated ones;
axons' targeting mechanism remains mysterious.
 Projection Pathways: Olfactory bulbs project to
medial temporal lobe structures, including amygdala
and piriform cortex; limbic pathway mediates
emotional response; thalamic-orbitofrontal pathway
mediates conscious perception of odors.
Gustatory System:
 Taste Receptor Cells: Found on the tongue and oral
cavity, occurring in clusters called taste buds; each
taste bud has various receptor cell types.
 Taste Bud Composition: Each taste bud comprises 50
to 100 receptor cells; one presynaptic cell
communicates with neurons carrying signals away
from the bud; taste bud cells replaced every few
weeks.
 Primary Tastes: Initially assumed to be sweet, sour,
bitter, salty, and umami; discovery of various g-
protein-linked receptor proteins and mechanisms.
 Taste Pathways: Gustatory afferent neurons exit
mouth through facial (VII), glossopharyngeal (IX), and
vagus (X) cranial nerves; terminate in solitary nucleus
of medulla, project to ventral posterior nucleus of
thalamus, and then to primary gustatory cortex in
insula; secondary gustatory cortex in orbitofrontal
cortex.
 Broad vs. Narrow Tuning: Each receptor cell narrowly
tuned to one taste or very few tastes; broader tuning
observed in presynaptic cells and cortical neurons;
mechanisms translating receptor protein responses
into specific tastes not fully understood.
Brain Damage and the Chemical Senses:
 Anosmia: Inability to smell; caused by displacement
of brain due to head trauma, shearing olfactory
nerves; also linked to various neurological disorders
(e.g., Alzheimer’s, epilepsy).
 Ageusia: Rare; damage to ear on one side can cause
partial ageusia limited to anterior two-thirds of the
tongue; gustatory information from this area passes
through facial nerve (VII) via middle ear.
 Coding in Cortex: Primary gustatory cortex
chemotopically organized; responses to odor, texture,
and temperature in food; secondary gustatory cortex
neurons signal food pleasantness and are influenced
by satiety.
 arietal cortex) and bottom-up (sensory circuitry)
mechanisms; attention strengthens responses to
attended aspects and weakens responses to
unattended aspects, termed the push-pull
mechanism.
Simultanagnosia:
 Definition: A disorder of attention characterized by an
inability to perceive more than one object at a time,
even though individual objects can be identified when
presented individually.
 Associated Brain Region: Simultanagnosia is
associated with bilateral damage to the posterior
parietal cortex, specifically the dorsal stream
responsible for visually localizing objects in space.
Principles of Sensorimotor Function:
1. Hierarchical Organization:
 Definition: Commands cascade down through
hierarchical levels from association cortex to muscles;
higher levels specify general goals, not specific
actions.
 Parallel Structure: Both sensorimotor system and
companies have parallel hierarchical structures;
signals flow through multiple paths, allowing diverse
control methods.
  Functional Segregation: Each level of the hierarchy
comprises different units (neural structures or
departments) with distinct functions.
2. Motor Output Guided by Sensory Input:
 Importance of Sensory Input: Sensory feedback,
including visual, somatosensory, and vestibular inputs,
guides motor responses by providing information
about the body's state.
 Continuous Monitoring: Similar to efficient
companies, the sensorimotor system continuously
monitors its activities and uses sensory feedback to
refine and adjust movements.
 Ballistic Movements: Brief, high-speed movements
like swatting a fly are exceptions and do not rely on
sensory feedback.
3. Learning Changes in Sensorimotor Control:
 Conscious Control vs. Unconscious Regulation: Initial
motor responses are consciously controlled; with
practice, responses are organized into integrated
sequences controlled unconsciously by lower levels of
the CNS.
 Similarity to Company Development: Similar to a
company's growth, sensorimotor learning involves the
coordination of individual actions into prescribed
procedures performed at lower hierarchical levels.
General Model of Sensorimotor System:
 Hierarchy and Functional Segregation: Illustrated in
Figure 8.1, the model shows hierarchical structure,
functional segregation, parallel connections between
levels, and various feedback pathways.
 Focus of the Chapter: The chapter delves into neural
structures controlling voluntary behavior, tracing
major motor signals from association cortex down the
sensorimotor hierarchy to skeletal muscles, explaining
the principles of sensorimotor system organization.
Sensorimotor Association Cortex:
 Hierarchy's Peak: Association cortex resides atop the
sensorimotor hierarchy, divided into posterior parietal
association cortex and dorsolateral prefrontal
association cortex.
Posterior Parietal Association Cortex:
 Role in Movement Initiation: Provides necessary
information for movement initiation, integrating body
part positions and external object locations.
 Integration of Sensory Information: Receives inputs
from visual, auditory, and somatosensory systems,
crucial for spatial localization and attention.
 Output and Specialization: Sends outputs to various
motor cortex areas, each specialized for different
movements of eyes, head, arms, or hands.
 Studies on Stimulation and Control: Electrical
stimulation at specific posterior parietal areas induced
intentions or feelings of action in conscious patients.
Neural population activity studies revealed control of
neurons via imagined actions, decoded into motor
imagery.
Consequences of Damage:
 Apraxia: Impairs voluntary movements out of context,
often remedied in natural situations; caused by
unilateral damage to left posterior parietal cortex.
 Contralateral Neglect: Impairs response to stimuli on
the side opposite the brain lesion, particularly
egocentric left; often linked with large lesions of the
right posterior parietal cortex.
 Object-Based Contralateral Neglect: Some patients
fail to respond to the left side of objects, indicating a
nuanced deficit.
 Unconscious Perception: Patients with contralateral
neglect may unconsciously perceive stimuli, as
evidenced by consistent gaze patterns and improved
identification of fragmented drawings.
Dorsolateral Prefrontal Association Cortex:
 Role in Sensorimotor Function: Receives inputs from
posterior parietal cortex, sending projections to
secondary motor cortex areas, primary motor cortex,
and frontal eye field.
 Neuronal Activity Patterns: Neurons exhibit diverse
response properties, with some responsive to object
characteristics, others to object locations, and some
to a combination of both. Some neurons respond to
the impending response itself and tend to fire earlier
than neurons in other motor areas.
 Decision Making: Likely involved in the decision-
making process for initiating voluntary movements.
Interactions with posterior parietal cortex and other
frontal areas are critical for these decisions.
Secondary Motor Cortex:
 Complexity Beyond Two Areas: Originally thought to
consist of two areas (supplementary motor area and
premotor cortex), recent research indicates at least
eight areas in each hemisphere, with subdivisions like
supplementary eye field and cingulate motor areas.
 Programming Specific Movements: Involved in
programming specific movement patterns after
receiving general instructions from dorsolateral
prefrontal cortex. Imaging studies demonstrate brain
activity patterns during imagined or planned
movements.
 Functional Variations: Efforts to differentiate
functions among these areas continue. Earlier
research focused on distinctions between
supplementary motor area and premotor cortex, but
consistent theories have not emerged yet.
Mirror Neurons:
 Discovery and Significance: Discovered in the early
1990s, mirror neurons are active both when an
individual performs a goal-directed movement and
when observing the same movement by others. Their
discovery is significant as they provide a potential
mechanism for social cognition, allowing mapping of
others' actions onto one's own repertoire, facilitating
social understanding, cooperation, and imitation.
Mirror Neurons:
 Role in Social Cognition: Mirror neurons respond to
the understanding of the purpose of an action rather
than superficial characteristics. They fire not only
 when observing an action but also when the subject
hears the associated sound or mentally represents the
action.
 Localization and Studies: Mirror neurons have been
found in various areas of the macaque monkey cortex.
In humans, research primarily relies on functional MRI
studies, showing activation in motor cortex areas
during action performance, observation, or
imagination. However, individual mirror neurons in
humans are rarely described due to limited
opportunities for direct neuronal recording during
behavioral tests.
Primary Motor Cortex:
 Conventional View: Penfield and Boldrey's 1937
mapping revealed somatotopic organization: specific
cortical sites corresponded to contralateral muscle
movements. Earlier studies suggested neurons
encode movement directions. The conventional belief
was that each neuron fired maximally for movements
in a particular direction.
 Challenges to Conventional View: Recent studies
challenged the directional tuning concept. Neurons
were recorded in freely moving monkeys, showing
activity related to movement endpoints rather than
movement directions. Additionally, prolonged bursts
of stimulation produced complex, species-typical
movements involving multiple body parts, challenging
the idea of individual muscle control.
 Current View: New stimulation techniques, using
longer bursts of current, revealed complex natural
movements instead of individual muscle contractions.
Stimulation of a site led to coordinated movements
involving multiple body parts. The response was not
dependent on the initial position, indicating a flexible
sensorimotor system. This suggests the presence of
an action map in addition to the traditional
somatotopic map in the primary motor cortex.
Sensorimotor Circuits and Brain Structures:
1. Sensory signals that are produced by a response
and guide the continuation of the response are
referred to as efference copies.
2. Contralateral neglect is the disturbance of a
patient’s ability to respond to a stimulus on the side
of the body opposite to the side of a brain lesion.
3. Patients who have great difficulty performing
movements when asked to do so out of context but
can readily perform them spontaneously in natural
situations are experiencing apraxia.
4. Mirror neurons fire when individuals perform a
particular goal-directed hand movement or when
they observe the same goal-directed movement
performed by another.
5. Stereognosis refers to the process of identifying
objects by touch.
6. The primary motor cortex is the main point of
departure of motor signals from the cerebral cortex
to lower levels of the sensorimotor hierarchy.
7. The foot area of the motor homunculus is in the
central fissure.
8. Although the cerebellum constitutes only 10
percent of the mass of the brain, it contains more
than half of the brain’s neurons.
9. The basal ganglia are part of neural loops that
receive input from various cortical areas and
transmit it back to the cortex via the thalamus.
10. Damage to either the cerebellum or the basal
ganglia can also produce cognitive changes, in
addition to their primary motor functions.
Sensory and Motor Systems
Muscle Contraction Types: Isometric vs. Dynamic
Contraction To understand muscles, it's crucial to
recognize their elasticity. Muscle contraction can be
isometric, increasing tension without shortening, or
dynamic, pulling muscles together by shortening.
Muscle tension can increase through motor neuron
firing or firing rate changes.
Receptor Organs: Golgi Tendon Organs and Muscle
Spindles Muscle activity is monitored by Golgi tendon
organs (detect muscle tension) and muscle spindles
(sense muscle length changes). They play roles in
providing CNS with information about muscle tension
and preventing muscle damage during extreme
contraction.
Muscle-Spindle Feedback Circuit Muscle spindles have
intrafusal muscles and motor neurons. Intrafusal motor
neurons prevent muscle spindles from becoming slack
during extrafusal muscle contraction. They maintain
spindle responsiveness to slight extrafusal muscle
length changes.
Stretch Reflex Stretch reflex, like the patellar tendon
reflex, prevents external forces from altering the body's
position. When a muscle is suddenly stretched, spindle
afferent neurons send signals to the spinal cord,
triggering a compensatory muscle contraction,
maintaining stability.
Reciprocal Innervation Antagonistic muscles are
innervated to allow smooth motor response: when one
contracts, the other relaxes. In the withdrawal reflex,
excitatory interneurons excite the elbow flexor, while
inhibitory interneurons inhibit the elbow extensor,
coordinating muscle activities.
Recurrent Collateral Inhibition Recurrent collateral
inhibition involves inhibitory interneurons (Renshaw
cells) inhibiting motor neurons that innervate the same
muscle. This inhibition, ensuring motor neurons do not
overwork, helps in smooth muscle function.
Walking: A Complex Sensorimotor Reflex Walking is
controlled by complex sensorimotor reflexes involving
visual, somatosensory, and balance information.
Remarkably, spinal circuits alone can control walking, as
evidenced by studies involving animals with severed
spinal cords.
Exercise: Scan Your Brain
1. f: Contraction of antagonistic muscles
2. k: Bulges
3. a: Axons innervate neurons
4. g: Local feedback circuits
5. b: Control movements of limbs
 6. n: Extensor
7. c: Smallest units of motor activity
8. d: Act to bend
9. m: Muscle spindles
10. e: Smooth unimpeded motor response
11. l: Stretch reflex
12. h: Two synapses
13. j: Small inhibitory interneurons
14. i: Complex reflex
Chapter 8: Central Sensorimotor Programs and
Learning
A Hierarchy of Central Sensorimotor Programs The
sensorimotor system functions as a hierarchy of
central sensorimotor programs. Higher levels issue
commands to lower levels, which, in turn, control
movements of the muscles. The cerebellum and basal
ganglia coordinate these programs, allowing smooth
and efficient motor responses. Once activated, each
level operates based on sensory feedback without
direct cortical control, demonstrating the system's
inherent plasticity.
Characteristics of Central Sensorimotor Programs
1. Motor Equivalence: The system can accomplish
tasks in various ways, exhibiting motor equivalence.
Specific central sensorimotor programs are adapted
to different situations, emphasizing the system's
flexibility.
2. Sensory Information and Consciousness: Central
sensorimotor programs can operate without
conscious awareness, highlighting the separation of
conscious perception and sensory control of behavior.
3. Development and Practice: Central sensorimotor
programs for species-typical behaviors can develop
without practice, while practice can modify or create
new programs. Response chunking and shifting
control to lower levels are key processes in
sensorimotor learning.
Functional Brain Imaging of Sensorimotor Learning
Functional brain imaging studies, like Jenkins and
colleagues' PET study, have explored neural correlates
of sensorimotor learning. These studies identify brain
regions involved in learning new motor sequences.
Changes in these regions occur as tasks are practiced,
indicating the brain's adaptability during learning.
However, the nature of these changes remains a topic
for future research.