LESSON 1 - NERVE CELLS AND NERVE 4.

The Structure of a Neuron

IMPULSES
a. The most distinctive structural feature of neurons is
The Cells of the Nervous System their shape, which varies enormously.
b. Larger neurons contain four major components:
dendrites, cell body, axon, and presynaptic terminal.
A. NEURONS AND GLIA The tiniest neurons may lack axons and well-defined
dendrites.
1. The nervous system is composed of two types of cells:
neurons and glia. Compare the structure of these components in the
following two types of neurons:
a. Neurons: Receive information and transmit it to other
cells. i. A motor neuron: Conducts impulses to muscles
b. Glia: Serve many functions that are difficult to and glands from the spinal cord.
summarize. ii. The cell body of a motor neuron is located in the
c. The human brain contains approximately 86 billion spinal cord.
neurons though that estimate varies from person to iii. A sensory neuron (receptor neurons): Sensitive to
person). certain kinds of stimulation (e.g., light, touch, etc.).

Because of the small size of many neurons and the variation Note that the soma is located on a stalk off the main trunk of the axon.
in cell density from one spot to another, obtaining an
accurate count is difficult.
c. Neuron Components:

i. Dendrites: Branching fibers that extend from the cell

2. Santiago Ramón y Cajal (1852–1934), a Pioneer of
body get narrower at their end. The dendrite’s surface
Neuroscience
is lined with specialized synaptic receptors, at which
a. Used staining techniques to reveal that small gaps the dendrite receives information from other neurons.
separate the tips of one neuron from another. This
discovery showed that the brain—like the rest of the ii. Dendritic spines: Short outgrowths found on some
body—is composed of individual cells. dendritic branches that increase the surface area
available for synapses.

3. The Structure of an Animal Cell iii. Cell body (soma): Contains the nucleus, ribosomes,
mitochondria, and other structures found in most cells.
a. Neurons contain the same basic structures as other
animal cells. iv. Axon: A long, thin fiber (usually longer than dendrites),
b. Membrane (plasma membrane): Composed of two which is the information-sending part of the neuron,
layers of fat molecules, this membrane allows some sending an electrical impulse toward other neurons,
small uncharged chemicals to flow both into and out of glands, or muscles.
the cell. Protein channels allow a few charged ions to
cross the membrane, but most chemicals are unable to v. Myelin sheath: Insulating covering found on some
cross. vertebrate axons.
c. Nucleus: The structure that contains the chromosomes.
d. Mitochondrion: The structure that provides the cell vi. Nodes of Ranvier: Interruptions along the myelin
with energy. It requires fuel and oxygen to function. sheath.
e. Ribosomes: Site of protein synthesis in the cell.
f. Endoplasmic reticulum: A network of thin tubes that vii. Presynaptic terminal (end bulb): Swelling at the tip of
transports newly synthesized proteins to other the axon. The part of the neuron that releases chemicals
locations. Ribosomes may be attached. that cross the junction between one neuron and the next.

The nucleus, membrane, and other structures are

characteristic of most animal cells. The plasma membrane
is the border of the neuron.
d. Neurons may have any number of dendrites but are g. Radial glia: Guides the migration of neurons and the
limited to no more than one axon (which may have growth of axons and dendrites during embryonic
branches). development.

e. Other terms associated with neurons: B. The Blood-Brain Barrier

1) Afferent axons: Brings information into a
structure. 1. The mechanism that keeps most chemicals out of the
2) Efferent axons: Sends information away from a vertebrate brain.
structure.
3) Interneurons (intrinsic neurons): Entirely 2. Why do We Need a Blood-Brain Barrier?
located within a single structure of the nervous a. The blood-brain barrier is needed because the
system. brain lacks the type of immune system present in
the rest of the body.

b. Because neurons generally cannot be replicated and

A. Variations Among Neurons
replaced, the barrier is in place to minimize the risk
a. Neurons vary enormously in size, shape, and function. of irreparable brain damage.
b. A neuron’s function is closely related to its shape.
c. A neuron’s shape is plastic (changeable) as new c. A wall is formed that keeps out most viruses,
experiences can modify the shape of a neuron. bacteria, and harmful chemicals.

d. When viruses do enter, like the rabies virus, it can

5. Glia infect the brain and lead to death.

a. Glia (neuroglia) are the other major nervous system e. A virus that enters your nervous system probably
component. Glia has many different functions, but they do stays with you for life (e.g., chicken pox and
not transmit information like neurons. shingles).
b. Glia are smaller and slightly more numerous than
neurons. Several types of glia exist to perform different
functions. 3. How does the Blood-Brain Barrier Works?

c. Astrocytes: A type of glia that absorbs chemicals a. Endothelial cells form walls of the capillaries. These
released by axons and later returns those chemicals to the cells are tightly joined in the brain, blocking most
axon to help synchronize the activity of neurons. Astrocytes molecules from passing. In the rest of the body, the
remove waste products as well, particularly those created endothelial cells are separated by gaps.
after neurons die. Astrocytes also cause blood vessels to b. Small uncharged molecules (e.g., oxygen and carbon
dilate to bring in more nutrients at times of increased brain dioxide) and molecules that can dissolve in the fats of
activity. They act as a recycling system for glutamate the capillary wall can cross passively (without using
released by neurons (absorbing excess glutamate then energy) through the blood-brain barrier.
converting it to glutamine and passing it back into the c. An active transport system (a protein-mediated
neurons). process that uses energy) exists to pump necessary
d. Microglia: Very small cells that remove waste material chemicals, such as glucose, through the blood-brain
as well as viruses, fungi, and other microorganisms. They barrier.
proliferate after brain damage, removing dead or damaged d. The blood-brain barrier is essential for health. For
neurons. example, in Alzheimer’sdisease, the endothelial cells
lining the brain’s blood vessels shrink and harmful
e. Oligodendrocytes: A type of glia that builds the myelin chemicals can enter the brain.
sheaths around certain neurons in the brain and spinal cord. e. The blood-brain barrier poses difficulty in medicine
because it keeps out many useful medications that may
f. Schwann cells: A type of glia that builds the myelin
be used to treat diseases like brain cancer.
sheaths around certain neurons in the periphery of the body.
C. Nourishment of Vertebrate Neurons Sodium-potassium pump: A protein complex found along
the neuron membrane that transports three sodium ions
1. Almost all neurons depend on glucose (a simple sugar)
outside of the cell while also drawing two potassium ions
for their nutrition.
into the cell; this is an active transport mechanism (requires
2. Neurons rely on glucose so heavily because glucose is energy in the form of ATP to function).
practically the only nutrient that crosses the blood-brain
c. When the membrane is at rest, two forces work on
barrier in adults. Ketones can also cross but are in short
sodium ions:
supply.
3. A thiamine (vitamin B1) deficiency leads to an inability i. The electrical gradient: Opposite electrical charges
to use glucose, which could lead to neuron death and a attract, so sodium (which is positively charged) is
condition called Korsakoff’s syndrome (a disorder marked attracted to the negative charge inside the cell.
by severe memory impairment). ii. The concentration gradient (difference in the
distribution of ions between the inside and the
outside of the membrane): Sodium is more
II. The Nerve Impulse concentrated outside the membrane than inside and
is thus more likely to enter the cell than to leave it.
A. The Resting Potential of the Neuron iii. Given that both the electrical and concentration
gradients tend to move sodium into the cell, sodium
1. The membrane of a neuron maintains an electrical
would be expected to quickly enter the cell.
gradient (also called polarization—a difference in
However, when the membrane is at rest, sodium
electrical charge between the inside and outside of the
channels are closed.
cell).
d. Potassium ions are subject to the same two forces;
2. In the absence of any outside disturbance (i.e., at rest),
however, the forces are in opposition to each other.
the membrane maintains an electrical polarization (i.e.,
Potassium ions are positively charged, so the electrical
a difference in electrical charge between two locations)
gradient tends to move potassium in, but since
that is slightly more negative on the inside relative to
potassium is concentrated on the inside of the cell, the
the outside. This difference in electrical potential or
concentration gradient causes potassium to flow out of
voltage is known as the resting potential.
the cell.
3. The resting potential is measured by very thin 5. Why a Resting Potential?
microelectrodes. A typical resting membrane potential
is -70 millivolts (mV). This may vary from one neuron a. The advantage of the resting potential is to allow the
to another. neuron to respond quickly to a stimulus.

4. Forces Acting on Sodium and Potassium Ions:

B. The Action Potential
a. The neuron membrane is selectively permeable, which
allows some molecules to pass freely (e.g., water, 1. Action potential: Messages sent by axons.
carbon dioxide, oxygen) while restricting others. Most 2. Hyperpolarization (increased polarization): Occurs
large molecules and ions cannot cross the membrane. A when the negative charge inside the axon increases
few important ions cross through protein channels. (e.g., 70 mV becomes 80 mV).
3. Depolarization (reduce polarization toward zero):
b. During the resting potential, chloride channels (or Occurs when the negative charge inside the axon
gates) remain open along the membrane, which allows decreases (e.g., 70 mV becomes55 mV).
ions to pass through. Potassium channels are mostly 4. Threshold of excitation: The level that a depolarization
closed, causing potassium to cross the membrane must reach for an action potential to occur. A
slowly. Sodium gates remain closed, restricting the subthreshold stimulation produces a small response
passage of sodium ions. proportional to the amount of current. However, as long
as the stimulation is above the threshold, regardless of
how far beyond, the stimulation produces a big
response.
5. The All-or-None Law C. Propagation of the Action Potential
a. The all-or-none law means that the amplitude and 1. The action potential begins at the axon hillock (a
velocity of an action potential are independent of the swelling located where the axon exits the cell body).
intensity of the stimulus that initiated it.
2. The action potential is regenerated due to sodium ions
b. If the threshold is met or exceeded, an action potential
moving down the axon, depolarizing adjacent areas of the
of a specific magnitude will occur; if the threshold is
membrane.
not met, an action potential will not occur.
3. Propagation of the action potential: Transmission
(movement) of an action potential down an axon. The
6. The Molecular Basis of the Action Potential
action potential moves down the axon by regenerating itself
i. Principles of the action potential: at successive points on the axon.4. The refractory periods
i. At the start, sodium ions are mostly outside the prevent the action potentials from moving in the opposite
neuron and potassium ions are mostly inside. direction (i.e., toward the axon hillock).
ii. When the membrane depolarizes, sodium and
potassium channels in the membrane open.
iii. At the peak of the action potential, the sodium D. The Myelin Sheath and Saltatory Conduction
channels close. 1. Myelin: An insulating material composed of fats and
ii. Voltage-gated channels regulate channels of sodium proteins found on some vertebrate axons. Myelin
and potassium. Their permeability depends on the greatly increases the speed of propagation.
voltage difference across the membrane.
iii. When the action potential reaches its peak, voltage- 2. Myelinated axons: Axons covered with a myelin
activated sodium gates close, and voltage-gated sheath, found only in vertebrates.
potassium channels open, allowing potassium ions to
flow outside of the membrane due to their high 3. Nodes of Ranvier: Short, unmyelinated sections on a
concentration inside the neuron as opposed to outside. myelinated axon.
In addition, the electrical gradient is now pushing the
potassium to flow outward. 4. Saltatory conduction: The “jumping” of the action
iv. A temporary hyperpolarization (membrane potential potential from node to node.
below the resting potential) occurs before the
5. Some diseases, including multiple sclerosis, destroy the
membrane returns to its normal resting potential (this is
myelin along axons; loss of the myelin sheath slows
due to potassium gates opening wider than usual,
down or prevents the propagation of action potentials.
allowing potassium to continue to exist past the resting
potential). 6. The Refractory Period
v. After the action potential, the neuron has more sodium
and fewer potassium ions for a short period (this is soon 1. The refractory period is a period immediately after an
adjusted by the sodium-potassium pumps to the action potential occurs when the neuron will resist the
neuron's original concentration gradient). production of another action potential.
vi. The sodium-potassium pump does not restore the 2. Absolute refractory period: Sodium gates are
resting membrane potential; diffusion does. Less than 1 incapable of opening; hence, an action potential cannot
percent of the total sodium ions present to cross the occur, regardless of the amount of stimulation.
membrane during an action potential. 3. Relative refractory period: Sodium gates are capable
vii. Local anesthetic drugs (e.g., Novocain, Xylocaine, of opening, but potassium channels remain open; a
etc.) block the occurrence of action potentials by stronger than normal stimulus (i.e., exceeding
blocking voltage-activated sodium gates (preventing threshold) will initiate an action potential.
sodium from entering a membrane).
A. Local Neurons
viii. Action potentials only occur in axons; dendrites do not
1. Small neurons with short dendrites and a short (if any)
have voltage-dependent channels. axon. Local neurons do not follow the all-or-none law.
ix. In most neurons, the absolute refractory period is about 2. Graded potentials: Membrane potentials that vary in
1 ms and the relative refractory period is another 2–4 magnitude in proportion to the intensity of the stimulus.
ms. Graded potentials get smaller as they travel.
LESSON 2 - SYNAPSES 7. Spatial summation: Several synaptic inputs originating
from separate locations exert a cumulative effect on a
I. The Concept of the Synapse
postsynaptic neuron.
A. Synapse: Specialized gap between neurons.
a. For example, pinching one point does not produce
B. Properties of Synapses a reflex but pinching two points at once does.
b. This happens because the two points activated
1. Sherrington studied reflexes (automatic muscular separate sensory neurons, whose axons converged
responses to stimuli). onto a neuron in the spinal cord.
2. In a leg flexion reflex, a sensory neuron excites a second
neuron, which in turn excites a motor neuron, which excites
a muscle. II. Inhibitory Synapses

3. Reflex arc: The circuit from sensory neuron to muscle a. The interneuron sends a message to block activity of
response. motor neurons to the extensor muscles in the same leg and
the flexor muscles of the other three legs of a dog.
4. Sherrington discovered that:
b. Inhibitory postsynaptic potential (IPSP): A temporary
a. Reflexes are slower than conduction along an axon. hyperpolarization of a postsynaptic cell (this occurs when
b. Several weak stimuli presented at slightly different K+ leaves the cell or Cl enters the cell after it is stimulated).
times or locations produce a stronger reflex than a
single stimulus does.
c. Excitation of one set of muscles leads to a relaxation
A. Relationship Among EPSP, IPSP, and Action Potentials
of others.
1. The more synapses that a response travels through, the
5. Speed of a Reflex and Delayed Transmission at the
longer the response takes.
Synapse
2. The probability of an action potential on a given neuron
a. Sherrington measured reflex speeds using dogs. He
depends on the ratio of EPSPs to IPSPS at a given moment.
pinched a dog’s foot and observed that the dog
flexed the leg after a short delay. He then measured 3. Spontaneous firing rate: The ability to produce action
the total distance that the impulse traveled, from the potentials without synaptic input (EPSPs and IPSPs
skin receptor to the spinal cord then from the spinal increase or decrease the likelihood of firing action
cord back down the leg. potentials).
b. Sherrington observed a rate of 15 second/s.
c. Previous research indicated potential never
velocities of 40 m/s. III. Chemical Events at the Synapse
d. Sherrington concluded that the delay must be
caused by one nerve communicating with the other. A. The Discovery of Chemical Transmission at Synapses

6. Temporal summation: Repeated stimuli within a brief 1. T.R. Elliott (1905): Suggested that the sympathetic
time having a cumulative effect. nerves stimulate muscle by releasing adrenaline or a
similar chemical. However, the evidence was not
a. Presynaptic neuron: The neuron that delivers the convincing, and most scientists continued to belief that
synaptic transmission. synapses communicated electrically.
b. Postsynaptic neuron: The neuron that receives the 2. Otto Loewi (1960): Fluid from a stimulated frog heart
message. was transferred to another heart. The fluid caused the
c. Graded potential: Either depolarization new heart to react as if stimulated.
(excitatory) or hyperpolarization(inhibitory) of the 3. Lead to the conclusion that synaptic transmission
postsynaptic neuron. depends on chemical rather than electrical stimulation.
d. A graded depolarization is known as an excitatory 4. This discovery revolutionized our understanding and
postsynaptic potential (EPSP) and occurs when led to research developing psychiatric drugs.
Na+ ions enter the postsynaptic neuron. EPSPs are
not action potentials: The EPSP’s magnitude
decreases as it moves along the membrane.
B. The Sequence of Chemical Events at a Synapse e. It is possible for a neuron to accumulate excessive
levels of a neurotransmitter. MAO (monoamine
1. The major events at a synapse:
oxidase) breaks down transmitters into inactive
a. The neuron synthesizes chemicals that serve as chemicals.
neurotransmitters. It synthesizes the smaller f. Catecholamines (dopamine, epinephrine, and
neurotransmitters in the axon terminals and norepinephrine): Three closely related.
synthesizes neuropeptides in the cell body.
b. Action potentials travel down the axon. At the
presynaptic terminal, an action potential enables 5. Release and Diffusion of Transmitters
calcium to enter the cell. Calcium releases
a. When an action potential reaches the axon terminal, the
neurotransmitters from the terminals and into the
depolarization causes voltage-dependent calcium gates
synaptic cleft, the space between the presynaptic
to open. As calcium flows into the terminal, the neuron
and postsynaptic neurons.
releases a neurotransmitter into the synaptic cleft within
c. The released molecules diffuse across the narrow
1 2 ms. This process of neurotransmitter release is
cleft, attach to receptors, and alter the activity of the
called exocytosis.
postsynaptic neuron. Mechanisms vary for altering
b. After being released by the presynaptic neuron, the
that activity.
neurotransmitter diffuses across the synaptic cleft to the
d. The neurotransmitters will separate from their
postsynaptic membrane, where it will attach to
receptors and (in some cases) are converted into
receptors. This takes no more than 0.01 ms across a
inactive chemicals.
cleft that is 20–30 nanometers wide.
e. The neurotransmitter molecules may be taken back
c. The brain uses dozens of neurotransmitters, but no
into the presynaptic neuron for recycling, or they
single neuron releases them all.
may diffuse away.
d. Most neurons release a combination of
f. Some postsynaptic cells send reverse messages to
neurotransmitters and neuropeptides, which carry more
control the further release of neurotransmitters by
complex messages.
the presynaptic cells.
e. A neuron may receive and respond to many
neurotransmitters at different synapses.
2. Synthesis of Transmitters
a. Neurons synthesize neurotransmitters from 5. Activating Receptors of the Postsynaptic Cell
precursors derived originally from food. a. A neurotransmitter can have two types of effects
b. Compounds containing a catechol and an amine when it attaches to the active site of the receptor:
group. ionotropic or metabotropic effects.
c. Choline is the precursor for acetylcholine. Choline
7. Ionotropic effects
is obtained from certain foods.
d. The amino acids phenylalanine and tyrosine are a. At one type of receptor, neurotransmitters exert
precursors for the catecholamines. ionotropic effects, corresponding to the brief on/off
e. The amino acid tryptophan is the precursor for effects.
serotonin. The amount of tryptophan in the diet b. Sodium and potassium channels along an axon are
controls the levels of serotonin. transmitter-gated orligand-gated channels. When
the neurotransmitter attaches, it opens a channel.

4. Storage of Transmitters
8. Metabotropic Effects and Second Messenger Systems
a. Most neurotransmitters, such as acetylcholine, are
synthesized in the presynaptic terminal. a. At other receptors, neurotransmitters exert
b. Vesicles: Tiny nearly spherical packets that store metabotropic effects by initiating a sequence of
high concentrations of neurotransmitters near the metabolic reactions that are slower and longer
presynaptic terminal. lasting than ionotropic effects.
c. Nitric oxide is an exception to this rule, as neurons c. Specifically, when the neurotransmitter attaches to
do not store nitric oxide for future use. the receptor, it alters the configuration of the rest of
d. There are also substantial amounts of the receptor protein, enabling a portion of the
neurotransmitters outside the vesicles. protein inside the neuron to react with other
 molecules. Activation of the receptor by the c. Nicotine: Compound found in tobacco.
neurotransmitter leads to activation of G-proteins,
i. Stimulates the nicotinic receptor (a type of
which are attached to the receptor.
acetylcholine receptor) in both the central nervous
d. G-proteins: A protein coupled to the energy-
system and the neuromuscular junction of skeletal
storing molecule guanosinetriphosphate (GTP).
muscles.
e. Second messenger: Chemicals that carry a
ii. Can also increase dopamine release by attaching to
message to different areas within a postsynaptic
neurons that release dopamine in the nucleus
cell; the activation of a G-protein inside a cell
accumbens.
increases the amount of the second messenger.
d. Opiate drugs: Derived from, or chemically similar
to those derived from, the opium poppy.
9. Neuropeptides
i. Familiar opiates include morphine, heroin,
a. Neuropeptides are often referred to as and methadone.
neuromodulators. ii. Opiate drugs exert their effects by binding to
b. Whereas the neuron synthesizes most other the same receptors as endorphins.
neurotransmitters in the presynaptic terminal, it
12. Inactivation and Reuptake of Neurotransmitters
synthesizes neuropeptides in the cell body and then
slowly transports them to other parts of the cell. a. Neurotransmitters need to be inactivated because
c. Whereas other neurotransmitters are released at the they may continue to excite or inhibit the receptor.
axon terminal, the neuropeptides are released Neurotransmitters are inactivated in different ways.
mainly by dendrites, and also by the cell body and b. Acetylcholine is broken down by the enzyme
the sides of the axon. acetylcholinesterase (AChE);the two components
are reabsorbed and recycled.
c. Serotonin and the catecholamines (i.e., dopamine
10. Variation in Receptors and norepinephrine) are taken up by the
presynaptic neuron. This process is called
a. The brain has a great variety of receptors, including
reuptake; it occurs through specialized proteins
at least 26 types of GABA receptors and at least
called transporters.
seven families of serotonin receptors, differing in
d. Some serotonin and catecholamine molecules are
their structure.
converted into inactive chemicals by enzymes such
b. Receptors differ in their chemical properties,
as COMT (converts catecholamines) and MAO
responses to drugs, and roles in behavior.
(converts both catecholamines and serotonin).
e. Stimulant drugs, including amphetamine and
cocaine, inhibit the transporters for dopamine, thus
11. Drugs that Act by Binding to Receptors decreasing reuptake and prolonging dopamine’s
a. A drug that chemically resembles a effects.
neurotransmitter can bind to its receptor. i. Amphetamine also blocks the serotonin and
b. Hallucinogenic drugs: Drugs that distort norepinephrine transporters.
perception.
ii. Methamphetamine’s effects are like those of
i. Many hallucinogenic drugs like lysergic acid amphetamine, but stronger.
diethylamide (LSD) resemble serotonin and
iii. Most antidepressant drugs also block the
bind to serotonin type 2A (5-HT2A)
dopamine transporter, but weaklier than
receptors.
amphetamine and cocaine do.
ii. These drugs provide stimulation at
inappropriate times or for longer than usual iv. When stimulant drugs increase the accumulation
durations. of dopamine in the synaptic cleft, COMT breaks
down the excess dopamine faster than the
presynaptic cell can replace it. A few hours after
taking a stimulant drug, a user has a deficit of
dopamine and enters a withdrawal state, marked by
 reduced energy, reduced motivation, and mild 4. Protein hormones and peptide hormones are
depression. composed of chains of amino acids.
a. Proteins are longer chains and peptides are shorter.
b. Protein and peptide hormones bind to membrane
f. Methylphenidate (Ritalin): Stimulant drug often receptors and activate a second messenger within
prescribed for attention-deficit/hyperactivity disorder. the cell—exactly the same process as at a
metabotropic synapse.
i. Works like cocaine by blocking the reuptake of
dopamine at presynaptic terminals. 5. Hormones secreted by the brain control the secretion of
other hormones.
ii. The effects of methylphenidate are much longer
lasting and less intense as compared to cocaine, yet also a. The pituitary gland is attached to the hypothalamus and
come with a risk of addiction. consists of two distinct glands, the anterior pituitary and
the posterior pituitary, which each produce different
13. Negative Feedback from the Postsynaptic Cell hormones.
a. Autoreceptors: Presynaptic receptors sensitive to b. The posterior pituitary is composed of neural tissue like
the same neurotransmitter they release. Detect the the hypothalamus. Two hormones, oxytocin and
amount of transmitter released and inhibit further vasopressin (also known as antidiuretic hormone) are
synthesis and release (that is, they provide negative released from the posterior pituitary. However, both of these
feedback). hormones are synthesized in the hypothalamus.
b. Postsynaptic neurons can respond to stimulation by
releasing special chemicals that travel back to the c. The anterior pituitary, composed of glandular tissue,
presynaptic terminal, where they inhibit further synthesizes six hormones.
release of transmitter. Nitric oxide, anandamide,
d. The hypothalamus controls the release of these six
and 2-AG (sn-2 arachidoyl glycerol) are three such
hormones by secreting releasing hormones that stimulate
chemicals.
or inhibit the release of other hormones.
c. Cannabinoids: The active chemicals in marijuana,
bind to anandamide or 2- AG receptors on
presynaptic neurons or GABA.

14. Electrical Synapses

a. A few special-purpose synapses operate electrically
because it is faster than chemical transmission
(somewhat confirming Sherrington’s hypothesis).
b. Gap junction: In an electrical synapse, the
membrane of one neuron comes into direct contact
with the membrane of another.

C. Hormones
1. A hormone is a chemical that is secreted primarily by
glands but also by other cells and is conveyed by blood
to other organs whose activity it influences.
2. Unlike neurotransmitters, which are released directly to
another neuron, hormones convey messages to any
organ that can receive them.
3. Endocrine glands produce hormones.
 LESSON 4 - GENETICS, EVOLUTION, 9. Sex-Linked and Sex-Limited Genes
DEVELOPMENT, AND PLASTICITY a. Genes located on sex chromosomes are known as
sex-linked genes. All other chromosomes are
I. Genetics and Evolution of Behavior
referred to as autosomal chromosomes. Genes
A. Mendelian Genetics located on autosomal chromosomes are autosomal
genes.
1. During the nineteenth century, Gregor Mendel b. A female mammal has two X chromosomes; a male
demonstrated that inheritance occurs through genes has one X and one Y chromosome.
(units of heredity that maintain their identity from one c. When biologists speak of sex-linked genes, they
generation to the next). usually mean X-linked genes.
2. As a rule, genes come in pairs, as they are aligned along d. Sex-limited genes: Genes present in both sexes but
chromosomes (strands of genes). One exception to this active mainly in one sex.
rule is male sex chromosomes, which do not come in
pairs. 10. Genetic Changes
3. A gene is a portion of a chromosome, which is a. Genes change in several ways.
composed of a double-stranded chemical called b. Mutation: A heritable change in a DNA molecule.
deoxyribonucleic acid (DNA). Changing one base in DNAto any of the other three
4. A strand of DNA serves as a template (model) for the types means that the mutant gene will code for a protein
synthesis of ribonucleic acid (RNA). with a different amino acid at one location in the
a. RNA is a single-stranded chemical: One type of molecule.
RNA molecule—messengerRNA— serves as a i. Mutations are rarely advantageous. One rare
template for the synthesis of protein molecules. example: FOXP2 gene, which facilitates language
b. DNA contains four “bases”—adenine, guanine, development.
cytosine, and thymine. ii. ii. Other types of mutations: duplication or
c. The order of those bases determines the order of deletion; microduplication or microdeletion.
corresponding bases along an RNA molecule— Scientists believe microduplications and
adenine, guanine, cytosine, and uracil. microdeletions maybe responsible for some cases
of schizophrenia.
5. Proteins can be part of the structure of the body or serve
as enzymes (biological catalysts that regulate chemical 11. Epigenetics: Changes in gene expression without
reactions in the body). modification of the DNA sequence (e.g., brain changes
If individuals have an identical pair of genes on the two resulting from drug addiction).
chromosomes, they are homozygous for that gene; if they
instead have an unmatched pair of genes, they are a. How could an experience modify gene expression?
heterozygous for that gene (e.g., a gene for black hair on i. Proteins called histones bind DNA into a shape that
one chromosome and a gene for brown hair on the other). is more like a string wound around a ball.
ii. To activate a gene, the DNA must partially unwind
6. Genes are dominant, recessive, or intermediate. from the histones.
d. Dominant genes show a strong effect in either iii. The result of an experience—maternal deprivation,
homozygous or heterozygous conditions. cocaine exposure, new learning, or whatever—in
e. Recessive genes show their effects only in some way alters the chemical environment within a
homozygous conditions (e.g., a carrier for both a cell.
dominant black hair gene and a recessive brown iv. In some cases, the outcome adds acetyl groups
hair gene will have black hair). (COCH3) to the histone tails near a gene, causing
f. An intermediate gene occurs in a phenotype where the histones to loosen their grip on the DNA, and
there is incomplete dominance in the heterozygous facilitating the expression of that gene.
condition. v. Another possibility is to add or remove methyl
groups from DNA, usually at the promoter regions
7. Often, more than one single gene contributes to the at the beginning of a gene.
appearance of certain characteristics (e.g., at least 10
genes contribute to variations in eye color). b. Experiences act by altering the activity of genes.
c.
8. Partial expression of a gene is also possible. d.
B. Heredity and Environment
1. Most variations in behavior depending on the combined
b. Have humans stopped evolving?
influence of many genes and environmental influences.
c. Does “evolution” mean “improvement”? The flaw
2. If the variations in some characteristics depend largely
in this argument is that evolution depends on
on genetic differences, the characteristic has high
reproduction, not just survival. Evolution improves
heritability.
fitness, which is operationally defined as the
3. Heritability in humans is studied in three ways:
number of copies of one’s genes that endure in later
a. Comparing monozygotic (identical) twins and
generations. The genes of the current generation
dizygotic (fraternal) twins.
evolved because they were fit for previous
b. Studying adopted children and their biological
generations. They may or may not be adaptive in
parents.
the future.
c. Identifying specific genes linked to some behavior.
d. Does evolution benefit the individual or the
4. Environmental Modification species? Neither: It benefits the genes!

a. Even a trait with high heritability can be modified 5. Brain Evolution

by environmental interventions.
a. Humans have bigger and better brains (at least we
b. Phenylketonuria, or PKU: A genetic inability to
think so) than other species.
metabolize the amino acid phenylalanine. If PKU
is not treated, phenylalanine accumulates to toxic b. Humans may have been able to evolve such big
levels impairing brain development. and leaving a brains without sacrificing other functions, because of
child mentally retarded, restless, and irritable. our diet.
Although PKU is hereditary. condition,
c. We devote more energy to our brains and less to
environmental interventions can modify it.
physical strength.
5. How Genes Affect Behavior
.
a. If we speak of a “gene for alcoholism,” we should
6. Evolutionary psychology: Concerns about how
not imagine that the gene itself causes. alcoholism.
behaviors evolved.
Rather, it produces a protein that under certain
circumstances increases the probability of a. The emphasis is on evolutionary and functional
alcoholism. explanations (how behavior may be useful to a
b. Exactly how a gene increases the probability of a population and why natural selection would favor it).
given behavior is a complex issue.
b. Some proposed evolutionary explanations are
speculative and controversial.
C. The Evolution of Behavior
c. Altruistic behavior: Behaviors that benefit others
1. Evolution: Change over generations in the frequencies rather than the individual committing the behavior. This
of various genes in a population. is in contrast to the belief that genes evolve for their
2. Offspring generally resemble their parents, but because own benefit.
of mutations and recombination, new inheritable
variations in the gene pool can occur. i. Perhaps a better explanation is a kin selection—
3. Artificial selection: Breeding animals for desirable selection for a gene that benefits the individual’s
individual characteristics (this causes changes in relatives. A gene spreads if it causes you to risk
various genetic frequencies in a population). your life to protect your children, who share many
4. Common Misunderstandings about Evolution of your genes, including perhaps a gene for
altruism.
a. Does the use or disuse of some structure or ii. Another explanation is reciprocal altruism, the
behavior cause an evolutionary increase or idea that individuals help those who will return the
decrease in that feature? This idea is a carryover of favor.
biologist Jean-Baptiste Lamarck’s theory of iii. A more controversial hypothesis is group selection.
evolution through the inheritance of acquired According to this idea, altruistic groups thrive
characteristics, known as Lamarckian evolution. better than less cooperative ones.
 c. This phenomenon is also seen in animals, like
songbirds, who lose neurons in areas necessary for
II. The Development of the Brain
singing in the fall and winter, only to regain
A. Maturation of the Vertebrate Brain neurons in those areas in the spring.
d. In general, new neurons do not form in other parts of
1. The human central nervous system begins to form when the adult mammalian brain. This is evidenced by the
the embryo is about two weeks old. age of a radioactive isotope of carbon in one’sbrain and
2. A neural tube forms around a fluid-filled cavity; this heart cells.
structure eventually sinks under the skin surface and
develops into the hindbrain, midbrain, and forebrain.
The fluid-filled cavity becomes the central canal and B. Pathfinding by Axons
the four ventricles.
1. Axons travel over long distances to precise locations.
3. The human brain weighs approximately 350 grams at
How do they find their way?
birth and around 1000 grams at one year of age. The
average adult brain weighs between 1200 and 1400 2. Chemical Pathfinding by Axons
grams.
i. Weiss (1924) grafted an extra leg to a salamander
4. Growth and Development of Neurons
and eventually axons grew into it so that the leg
Five Steps of Neuron Development: moved in sync with the salamander’s other legs.
Weiss suggested that the nerves attached to muscles
a. Proliferation: Production of new cells; cells along
at random and then sent a variety of messages, each
the ventricles of the brain divide. Some cells
one tuned to a different muscle.
remain where they are as stem cells, continuing to
divide. Others become primitive neurons and glia b. Specificity of Axon Connections
that migrate to other locations.
i. Evidence suggests Weiss was wrong—sensory
b. Early in development, the primitive neurons begin
axons find their way to their correct targets.
to migrate (move). Chemicals known as
immunoglobins, and chemokines guide the new ii. Sperry (1943) discovered that severed optic nerve
cells to their eventual destination in the brain. axons will grow back to their original targets in the
c. At first, a primitive neuron looks like any other cell. tectum. He showed that this process was dependent on
Gradually, the neuron differentiates, forming its chemical gradients in the \target cells by severing the
axon and dendrites; the axon grows before the optic nerve and rotating the eye by 180°.
dendrites.
d. A later and slower stage of neuronal development c. Chemical Gradients
is myelination, the process by which glia produce i. A growing axon follows a path of cell-surface
the insulating fatty sheaths that accelerate molecules, attracted by some chemicals, and
transmission in many vertebrate axons. Myelin repelled by others, in a process that steers the axon
forms first in the spinal cord and then in the in the correct direction.
hindbrain, midbrain, and forebrain. ii. Eventually, axons sort themselves over the surface
e. The final stage is synaptogenesis, or the formation of their target area by following a gradient of
of synapses. Although this process begins before chemicals.
birth, it continues throughout life, as neurons form
new synapses and discard old ones. 3. Competition Among Axons as a General Principle
a. Postsynaptic cells strengthen the synapses of some
5. New Neurons Later in Life
cells and weaken synapses with others.
b. Neural Darwinism: During development of the
a. The traditional belief was that adult vertebrate
nervous system, synapses form randomly before a
brains gain all their neurons during early
selection process keeps some and rejects others
development and could only lose neurons later in
(this is only partly accurate since synapse
life.
formation is also influenced by chemical guidance
b. The differentiation of stem cells (undifferentiated
and neurotrophic factors).
cells) is an exception to the traditional belief. For
example, stems cells in the nose remain
undifferentiated throughout life, and periodically
divide to replace a dying olfactory receptor.
 5. Social influences also affect the developing brain.
Children of impoverished or abused mothers have
C. Determinants of Neuronal Survival
increased problems in both academic and social
1. While working on the sympathetic ganglion, Rita Levi- functioning.
Montalcini discovered that muscles that synapse with the
axons from the ganglia don’t determine how many neurons
are produced but which synapses survive. E. Differentiation of the Cortex
2. Levi-Montalcini discovered that muscles produce and 1. Neurons differ in shape and chemistry.
release nerve growth factor (NGF), which promotes the 2. Ultimate shape of neurons and functions of regions
survival and growth of axons. depend on input received.
3. Immature neurons experimentally transplanted from
3. Axons that don’t receive enough NGF degenerate and
one part of the developing cortex to another develop the
their cell bodies die. All neurons are born with this suicide
properties characteristic of their new location. Neurons
program and will automatically die if the right synaptic
transplanted at a later stage develop some
connection is not made. This programmed cell death is
characteristics of the new location while retaining
called apoptosis.
others of the initial location.
4. Neurotrophin: A chemical (like NGF) that promotes the 4. In immature ferrets, researchers rerouted the optic
survival and activity of neurons. In addition to NGF, the nerve on one side of the brain away from its normal
brain also uses brain-derived neurotrophic factor (BDNF) thalamic target onto a thalamic target that usually gets
as a neurotrophin. BDNF is the most abundant neurotrophin input from the ears. They found that the parts of the
in the adult mammalian cortex. thalamus and cortex that formerly received auditory
information reorganized to process visual information.
5. Initially, all areas of the developing nervous system
produce far more neurons than will survive into adulthood.
This loss of cells is a natural part of development.
F. Fine-Tuning by Experience
6. After maturity, the apoptotic process becomes dormant,
1. Because of the unpredictability of life, we have evolved
and neurons do not need neurotrophins to survive.
the ability to redesign our brain (within limits) in response
Neurotrophins are used in adult brains to increase the
to experience.
branching of axons and dendrites throughout life.
Deficiencies of neurotrophins lead to cortical shrinking and 2. Experience and Dendritic Branching
are linked to several brain diseases.
a. Environmental enrichment leads to a thicker cortex,
more dendritic branching, and improved
performance on learning tasks in rats.
D. The Vulnerable Developing Brain
b. Much of the benefit of enriched environments in rats
1. The earliest stages of development are remarkably is simply due to activity. Increased size expansion of
similar across species. A series of genes known as neurons has also been demonstrated in humans as a
homeobox genes—found in vertebrates, insects, plants, function of physical activity.
even fungi, and yeast— regulate the expression of other c. Enriched environments enhance sprouting of axons
genes and control the start of anatomical development. and dendrites in a wide variety of species including
2. During early development, the brain is highly humans. Some believe this is evidence of the
vulnerable to malnutrition, toxic chemicals, and psychological term “far transfer,” which suggests
infections that would produce only mild problems at enhanced capacity in one task leads to enhanced
later ages. capacity of other tasks.
3. Fetal alcohol syndrome (FAS): Caused by alcoholic d. However, enriching environments produce
consumption during pregnancy. Symptoms include enhanced capacity of only the tasks that are relevant
decreased alertness, hyperactivity, facial abnormalities, in those environments. One researcher looked at a
mental retardation, motor problems, and heart defects. computer program designed to “train your brain.”
4. Infant's brains are especially sensitive to alcohol After six weeks of using the program several times
because it suppressed the release of glutamate, the a week, much of the 11,000 who participated saw
brain’s main excitatory transmitter. Thus, neurons substantial improvements in their ability to
receive less excitation and undergo apoptosis. complete the computer task. Yet this improvement
does not extend to other tasks.
 e. One of the best ways to maintain intellectual vigor However, in extreme cases the reorganization creates
is physical activity. problems.
ii. Focal hand dystonia (musician’s cramp): this happens
in musicians who practice extensively when the
1. Effects of Special Experiences expanded representation of each finger overlaps its
neighbor. The fingers become clumsy, fatigue easily,
i. Extensive practice of a particular skill makes a person and make involuntary movements that interfere with
more adept at that skill. In a few cases, researchers have the desired task. A similar condition called “writer’s
identified brain changes that are associated with cramp” can happen to people who spend all day writing.
increased expertise at a particular skill.
ii. Brain Adaptations in People Blind Since Infancy
iii. People blind from birth are better at discriminating
G. Brain Development and Behavioral Development
between objects by touch and have increased activation
in their occipital cortex (visual cortex) while 1. Adolescence
performing touch tasks.
a. Adolescents are widely regarded as impulsive and
iv. Further research using magnetic stimulation to
prone to seek immediate pleasure.
inactivate brain areas demonstrated that blind people
b. Research shows adolescents are able to make
use the occipital cortex to discriminate between tactile
reasonable, mature decisions when they have had time
stimuli and Braille symbols, but sighted people do not.
to consider the options carefully. However, they are
Similar results are also found using verbal stimuli.
impulsive when making quick decisions, especially in
the face of peer pressure.
Music Training
2. Old Age
i. The auditory cortex response to pure tones is twice as
a. On average, older people’s memory and reasoning
large for professional musicians as for non-musicians.
begin to fade. Many neurons lose some of their
Moreover, a part of the temporal cortex was found to be
synapses, and the remaining synapses change more
30 percent larger in professional musicians.
slowly.
ii. Violin players have a larger area devoted to the left
b. The frontal cortex begins thinning at age 30. However,
fingers in the postcentral gyrus than nonmusicians.
there is great variance in the level of deterioration in
different people.
c. The volume of the hippocampus also gradually
f. Special Training in Adulthood
declines.
i. Might adult experiences modify brain anatomy also? In d. In old age, even the blood contains chemicals that
a sense, the answer is, “Yes, of course.” impair cognitive function.
ii. The issue is whether an adult experience produces a big
enough effect that we might observe it with MRI or
similar technology. II. Plasticity After Brain Damage
iii. Many studies have in fact reported changes in adult
A. Brain Damage and Short-Term Recovery
brain anatomy from tasks such as learning to juggle
three balls, 16 hours of playing a complex video game, 1. Brain damage can result from a number of causes,
or 40 hours of playing golf for the first time. However, including tumors, infections, exposure to radiation or
skeptics raise serious objections. toxic substances, and degenerative conditions such as
iv. The only replicated finding is that physical exercise Parkinson’s and Alzheimer’s disease.
expands part of the hippocampus in older people.
v. In short, we should reserve judgment about most of the 2. Closed head injury: A sharp blow to the head that does
reported effects of brief experiences on the adult brain. not actually puncture the brain. The most common
cause of brain damage in young people. Closed head
injuries damage the brain because of rotational forces
g. When Brain Reorganization Goes Too Far that drive the brain tissue against the inside of the skull.
i. Typically expanded cortical representation of
3. Reducing the Harm from a Stroke
personally important information is beneficial.
 a. Stroke (cerebrovascular accident): A temporary 4. Denervation Supersensitivity
loss of blood flow to the brain. This is a common
a. If a certain set of synapses becomes inactive—
cause of brain damage, especially in the elderly.
perhaps because of damage elsewhere in the
b. Ischemia: The most common type of stroke; loss
brain—the remaining synapses become more
of blood flow caused bya blood clot orother
responsive, more easily stimulated. This process of
obstruction of an artery.
enhanced response is known as denervation
c. Hemorrhage: A less common type of stroke;
supersensitivity or receptor supersensitivity.
bleeding due to the rupture of an artery.
b. Denervation supersensitivity is a way of
d. Ischemia and hemorrhage lead to common
compensating for decreased input. However, the
problems including edema (fluid accumulation),
increased sensitivity can lead to intense responses
increased potassium levels due to dysfunctional
in normal inputs, which can result in prolonged
sodium-potassium pumps, and increased release of
pain.
glutamate.
e. Immediate Treatments
i. Decreasing cell death after a stroke can be 5. Reorganized Sensory Representations and the Phantom
accomplished by administering tissue Limb
plasminogen activator (tPA) clot-busting
drugs, which restore blood flow following a. If a brain area loses a set of incoming axons, we can
ischemia, or by using drugs that antagonize expect some combination of increased response
glutamate activity. (denervation supersensitivity) by the surviving
ii. The most effective known method of axons and collateral sprouting by axons that
preventing brain damage after strokes in ordinarily attach to some other target.
laboratory animals is to cool the brain. Cooling b. Physicians have long noted that most people with
protects the brain after ischemia by reducing amputations experience a phantom limb, a
overstimulation, apoptosis, and inflammation. continuing sensation of an amputated body part.
iii. Exposure to cannabinoids (the chemicals found
in marijuana) minimizes the damage caused by i. Modern methods show that phantom limbs
strokes in laboratory animals. develop when the relevant portion of the
somatosensory cortex reorganizes and
becomes responsive to alternative inputs.
B. Later Mechanisms of Recovery
6. Learned Adjustments in Behavior
1. Increased Brain Stimulation
a. Much recovery from brain damage depends on
a. Diaschisis: Decreased activity of surviving
learning to make better use of the abilities that were
neurons after other neurons are destroyed.
spared. A brain-damaged person or animal may
Behavioral deficits due to diaschisis can sometimes
also learn to use abilities that at first appear lost, but
be improved with the use of stimulant drugs.
are only impaired.
2. Regrowth of Axons i. For example: Monkeys with a
deafferented (loss of sensory or afferent
a. Under certain circumstances, damaged axons can
input) limb fail to use it because walking
grow back. However, regeneration is minimal in
on three limbs is apparently easier than
the mature mammalian central nervous system,
trying to move the impaired limb.
possibly because of a large amount of scar tissue or
However, if forced, they can learn to use
the secretion of growth-inhibiting chemicals.
the deafferented limb.
3. Axon Sprouting
b. One treatment for people recovering from a stroke
a. Ordinarily, the surface of dendrites and cell bodies is to force them to use the weaker limb by
is covered with synapses, and a vacant spot doesn’t preventing them from using the normal limb.
stay vacant for long.
b. After a cell loses input from an axon, it secretes
neurotrophins that induce other axons to form new
branches, or collateral sprouts, which take over
the vacant synapses.