NERVOUS SYSTEM
The nervous, endocrine and immune systems often cooperate and interact in regulating internal
body functions to maintain homeostasis.
The ability of an organism to survive and maintain homeostasis depends largely on how it
responds to internal and external stimuli.
A stimulus is an agent or a change within the body that can be detected by an organism.
Nerve cells are called neurons. These cells are specialized for transmitting electrical and
chemical signals through a network.
The nervous system consists of this network of neurons and supporting cells.
Neurotransmitters are chemical messengers used by neurons to signal other neurons and that
allows the nerve impulse to be transmitted across a synapse or connection between neurons
and/or receptors.
EVOLUTION AND DIVERSITY OF THE NERVOUS SYSTEM
The nervous system evolved over millions of years. By the time of the Cambrian Explosion 545
million years ago, a nervous system had already evolved.
Poriferans lack nervous system.
Nerve nets and radial nervous systems are characteristic of radially symmetrical animals.
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Nerve nets consist of scattered neurons, impulses may flow in both directions of the
synapse, and the impulse weakens as it spreads from the point of stimulation.
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There is no CNS.
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Found in cnidarians. Some cnidarians have two nerve nets, one for slow for tentacle
movement and another for faster to coordinate swimming.
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Echinoderms have a nerve ring and nerves that extend into various parts of the body.
Bilateral nervous systems are found in bilateral animals.
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Neurons aggregate to form ganglia, nerves, nerve cords and a brain.
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Cephalization includes the clustering of nerve cells at the anterior end of the animal
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Planarians have two ganglia at the anterior end and two parallel nerve cords joined by
transverse nerves.
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Annelids and arthropods have one or two ventral nerve cords that extend the length of
the body. An anterior pair of ganglia dorsally located is needed to respond adequately to
stimuli and to coordinate input.
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Octopuses and squids have the most sophisticated nervous system of invertebrates.
They have large brains, well-developed image-forming eyes and rapid signaling along
giant axons.
FUNCTIONS OF THE NERVOUS SYSTEM
The nervous system has three overlapping functions related to stimuli from within and without
the body.
It is the master controlling and communicating system of the body. It is responsible for behavior,
thought, actions, emotions, and maintaining homeostasis (together with the endocrine system).
Reactions to stimulus depend on four processes:
1. Reception or sensory input: afferent or sensory neurons and sense organs detect the
stimulus.
2. Integration: involves sorting and interpreting information and determining proper
response.
3. Response or motor output: efferent neurons bring the proper message to muscles and
glands.
Neurons that transmit messages to the central nervous system (CNS) are called afferent or
sensory neurons.
Neural messages are transmitted from the CNS by efferent neurons or motor neurons, to
effectors, muscles or glands.
The action by effectors is the response to the stimulus.
Signals are transmitted by nerves. Nerves are bundles of neuron extensions tightly wrapped in
connective tissue.
The nerves that communicate between the body and the CNS form the peripheral nervous
system.
NETWORKS OF NEURONS
1) Neuron structure
Nerve cells are called neurons.
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A typical neuron has cell body, dendrites and an axon.
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Dendrites are short, highly branched cytoplasmic extensions specialized to receive stimuli
and send nerve impulses to the cell body.
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In many brain areas, the finer dendrites have thorny projections called dendrite spines.
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The axon is a long extension sometime more than 1 meter long, and conducts impulses
away from the cell body.
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The conical region of the axon where it joins the cell body is called the axon hillock.
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An insulating layer called the myelin sheath surrounds many axons.
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The axon ends in many terminal branches called axon terminals with a synaptic terminal
or knob at the very end that releases neurotransmitters.
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Axons may branch forming axon collaterals.
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Axons outside the CNS and more than 2 μm in diameter are myelinated.
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The junction between a synaptic terminal and another neuron is called a synapse.
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The transmitting cell is called the presynaptic cell, and the target cell is called the
postsynaptic cell.
A nerve consists of hundreds or thousands of axons wrapped together in connective tissue.
2) The Reflex Arc
A reflex is the simplest nerve circuit. It is called the reflex arc.
A reflex is an automatic response to a stimulus.
Sensory receptor → sensory neuron → interneuron in spinal cord → motor neuron →
effector organ (e. g. muscle)
The cell body of sensory neurons is located in the dorsal root ganglion.
There are many ganglia along the sides of the spinal cord.
A ganglion (pl. ganglia) is a cluster of neuron cell bodies that usually perform a similar function.
If the cluster of cell bodies is located in the brain, it is called a nucleus (pl. nuclei).
3) Supporting cells
Glia are supporting cells. There are several types:
1.
2.
3.
4.
Envelop the neuron to form and insulating sheath around them.
Phagocytes that remove microorganisms and debris.
Lines the cavities of the brain and spinal cord.
Anchor neurons to blood vessels
Until recently, researchers assumed that glia play a supportive role without actually participating
in nerve signaling. Recent studies have suggested that some synaptic interactions do occur
between glia and neurons.
Glial cells are sometimes called collectively neuroglia.
Vertebrates have six types of glial cells.
Four types of glia cells are found in the Central Nervous System, CNS: astrocytes,
oligodendrocytes and ependymal cells.
1. Astrocytes are star-shaped cells that anchor neurons to capillaries, which are the nutrient
supply line. Tight junction anchor these cells to capillaries and contribute to the blood-brain
barrier, which restricts the passage of most substances into the brain.
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They are involved in the exchange between blood and neurons, e. g. take glucose from
the blood and pass it to neurons in the form of lactic acid.
Some are phagocytic.
Some regulate the concentration of K+ in the extracellular fluid of the nervous tissue.
Others recapture or regulate the concentration of released neurotransmitters.
2. Oligodendrocytes envelop neurons in the CNS with myelin and insulate them.
3. Schwann cells are found outside the CNS and form an outer cellular sheath around the
axon called neurilemma, and an inner myelin sheath.
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The plasma membrane of the Schwann cell is rich in myelin, a white fatty substance
that acts as an insulator.
Gaps in the myelin sheath are called nodes of Ranvier.
Multiple sclerosis occurs when the myelin sheath around the axons deteriorates and is
replaced by scar tissue.
The damage interferes with the conduction of the nerve impulse.
The cause of MS is a mystery but there is some evidence that indicates that it is an autoimmune
disease.
ION PUMPS AND ION CHANNELS
Ion pumps and ion channels maintain the resting potential of a neuron.
Most animal cells have a difference in electrical charge across the plasma membrane: more
negative on the inside and more positive on the outside of the cell, in the fluid.
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This is called the membrane potential.
The plasma membrane is said to be polarized when one side or pole has a different charge
from the other side.
When this occurs, a potential energy difference exists across the membrane.
If the charges are allowed to come together they have the potential to do work.
Neurons use electrical signals to transmit information.
MEASURING THE MEMBRANE POTENTIAL
Microelectrodes are used to measure the membrane potential of neurons.
The voltmeter registers the membrane potential, the difference in charge across the
membrane.
A resting neuron is the one not transmitting an impulse.
For an impulse to be fired, the plasma membrane of the neuron must maintain a resting
potential. It must be polarized.
The resting potential is the difference in electrical charge across the plasma membrane.
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The inner surface of the membrane is negative.
The interstitial fluid surrounding the neuron is positive.
An electrical potential difference exists across the membrane. It is called the resting or
membrane potential.
The resting potential of a neuron is 70 mV (millivolts).
By convention it is expressed as -70mV because the inner side is negatively charged relative to
the interstitial fluid.
GATED IONS CHANNELS
The resting potential results from the diffusion of K+ and Na+ through ion channels that are
always open; these channels are said to be ungated.
Neurons also have gated ion channels, which open or close in response to one of three kinds
of stimuli.
1. Stretch-gated ion channels: found in cells that sense stretch and open when the
membrane is mechanically deformed.
2. Ligand-gated ion channels are found at synapses and open or close when a specific
chemical signal binds to the channel.
3. Voltage-gated ion channels are found in axons and open or close when the membrane
potential changes.
THE NATURE OF THE NERVE IMPULSE
Action potentials are the signals conducted by axons.
Graded potentials are called graded because their magnitude varies directly with stimulus
strength.
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Hyperpolarization is an increase in the voltage across the membrane.
Depolarization is a decrease in the voltage across the membrane.
The stronger the stimulus, the more the voltage changes due to more channels opening, and
the farther the current flows.
Gated potentials are triggered by some change (stimulus) in the neuron's environment.
Graded potentials are short-lived, local changes in membrane potential that can be either
depolarizations or hyperpolarizations.
These changes cause current flows that decrease in magnitude with distance.
PRODUCTION OF ACTION POTENTIAL
Depolarization of a neuron’s membrane is graded up to a particular voltage called the threshold
voltage.
The nerve impulse is an action potential.
A) Threshold phase:
Electrical, chemical or mechanical stimulus may alter the membrane's permeability to Na+.
The axon contains specific voltage-activated ion channels that open when they detect a change
in the resting potential.
When the change reaches threshold levels, the protein changes shape, the channels open
and Na+ flows into the cell, while the K+ remain closed.
The membrane of a neuron can depolarize by about 15mV without initiating an impulse
The threshold to open the voltage-activated sodium-ion channels is -55mV.
B) Depolarization phase:
Sequence of events:
1. Transient increase in Na+ permeability; K+-gated channels remain closed,
2. Followed by the restoration of Na+ impermeability (repolarization phase),
3. Increase of K+ permeability.
If the depolarization reaches the threshold, it triggers an action potential.
The inside of the cells becomes positive. Polarity reverses due to the influx of Na+.
These causes a momentary reversal of polarity as the membrane depolarizes and overshoots to
+35 mV, creating a spike.
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The spike is an example of positive feedback
After a few milliseconds, the sodium-ion channels close. The closing depends on time rather
than on voltage.
K+ channels also open but more slowly and remain open until the resting potential has been
restored.
Once depolarization occurred in one portion of the membrane, the adjacent areas also become
depolarize and the ion gates open.
Once the depolarization reaches the threshold potential, it triggers a greater depolarization. This
is done by a positive feedback mechanism.
This process is repeated creating a wave of depolarization until the depolarization reaches the
end of the axon.
The magnitude of the action potential is independent of the strength of the stimulus: an all-ornone event.
C) Repolarization phase:
Repolarization occurs in less than one millisecond later when the Na+ channels close and the
membrane becomes impermeable to Na+.
Leakage of K+ out of the cell also occurs and restores the interior of the membrane to its
negative state.
Potassium-gated channels respond slowly and remain open longer. K+ continue to leak out and
this contributes to the temporary hyperpolarization of the membrane (undershoot).
Sodium-potassium pumps begin to function again. This process restores the membrane to the
usual resting potential of -75 mV.
When the membrane is depolarized, it cannot transmit another impulse no matter how great
stimulus is applied because the ion-gated channels are closed and unable to open.
This is called the refractory period, when the membrane is insensitive to stimulus.
D) Propagation of the nerve impulse along the axon.
The action potential is regenerated along the length of the axon.
1. Na+ entering the cell creates an electrical current that depolarizes the next neighboring
region of the membrane.
2. In case of the action potential, the depolarization is strong enough to reach the threshold
in the neighboring regions, re-initiating the action potential there.
3. The membrane is repolarized in the previous region as K+ flow outward.
4. The depolarization-repolarization process is repeated in the next region of the
membrane.
5. Because of the refractory period, the wave of depolarization cannot move backwards
towards the cell body. It can move only in the forward direction.
Continuous conduction occurs in unmyelinated axons.
In unmyelinated neurons, the speed of transmission is proportional to the diameter of the axon.
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Axons with larger diameter transmit more rapidly.
Squids and other invertebrates have large, unmyelinated axons.
Resistance to the flow of electrical current is inversely proportional to the cross-sectional area
of the a conductor.
Many vertebrate axons are surrounded by a myelin sheath.
Myelin increases the conducting speed of action potentials by insulating the axon membrane.
The voltage-activated ion Na+ and K+ channels are concentrated at the nodes called nodes
of Ranvier where the membrane is in contact with the interstitial fluid.
In myelinated axons, depolarization (action potential) jumps from one node of Ranvier to the
next.
This mode of conduction is called saltatory conduction (saltare = to leap in Latin).
It is fifty times faster than continuous conduction: 120 meters/sec.
COMMUNICATION BETWEEN CELLS AT SYNAPSES
A synapse is the junction between two neurons or between a neuron and an effector
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Neuromuscular junction or motor end plate is the synapse between a muscle and a
neuron.
Presynaptic neuron and postsynaptic neuron.
Signals across the synapse can electrical or chemical.
Electrical synapses contain protein channels (gap junctions) that connect the cytoplasm of two
neurons and allow ions to flow directly from one neuron to the next.
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It allows the passage of ions from one neuron to the next and the impulse is directly
transmitted.
They occur between axons and cell body, cell body to cell body, dendrites and
axons, axons and axons, dendrites and dendrites.
For quick communication and coordination between many neurons.
Chemical synapses are separated by the synaptic cleft.
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Most synapses are chemical.
Chemical messengers or neurotransmitters conduct the message.
When depolarization reaches the end of the axon it cannot jump across the cleft.
The electrical signal is converted to a chemical one.
Neurotransmitters are the chemicals that conduct the signal across the synapse and bind to
chemically activated ion channels in the membrane of the postsynaptic neuron.
Synaptic vesicles at the tip of the axon contain neurotransmitters.
1. Arrival of the depolarization wave opens calcium channels and allows Ca2+ influx into the
axon terminal.
2. Ca2+ act as messengers that cause the vesicles to fuse with the axon's membrane and
release the neurotransmitters into the cleft by exocytosis.
3. Ca2+ are quickly removed from the cytosol, either taken up by mitochondria or ejected by
calcium pumps.
4. Neurotransmitters diffuse across the synaptic cleft and binds reversibly to specific protein
receptors clustered on the postsynaptic membrane.
5. Binding of neurotransmitters causes the protein receptors to change shape and open ion
channels that initiate a depolarization wave in the postsynaptic neuron.
6. Neurotransmitters are quickly destroyed by enzymes or reabsorbed by the presynaptic
neuron.
7. Removal of neurotransmitters closes the ion channels and terminates the synaptic
response.
DIRECT SYNAPTIC TRANSMISSION
Binding of neurotransmitters causes the protein receptors to change shape and open ion
channels that initiate a depolarization wave in the postsynaptic neuron. This is called direct
synaptic transmission.
Each neuron may synapse with hundreds of other neurons. Some of these synapses are
excitatory and other inhibitory.
Presynaptic knobs may cover as much as 40% of the postsynaptic neurons' dendrites and cell
body.
In excitatory synapses, the binding of neurotransmitters open sodium channels and bring the
membrane closer to the threshold making it easier to depolarize it.
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This is called the excitatory postsynaptic potential or EPSP.
In inhibitory synapses, the neurotransmitters open potassium ion channels making it easier for
the K+ to leak out and for Cl- and hyperpolarizing the membrane making it more difficult to reach
the threshold.
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This is called inhibitory postsynaptic potential or IPSP.
Both EPSP and IPSP are graded potentials that depend on the number of neurotransmitters
that bind to the receptors of the postsynaptic neuron.
Several synaptic terminals acting on one postsynaptic neuron have a cumulative impact on the
membrane potential at the hillock bringing it closer to the threshold.
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This additive effect is called summation.
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In temporal summation the presynaptic neuron(s) transmit to the postsynaptic
neuron in rapid succession.
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In spatial summation, several different synaptic terminals stimulate a postsynaptic
cell at the same time and have an additive effect.
Summation is also applicable to IPSP.
INDIRECT SYNAPTIC TRANSMISSION
In indirect synaptic transmission, a neurotransmitter binds to a receptor that is not part of an ion
channel.
This activates a signal transduction pathway involving a second messenger in the postsynaptic
cell.
Cyclic AMP, cAMP, is one of those second messengers.
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Neurotransmitter (e. g. norepinephrine) activates a G protein
G protein activates adenylyl cyclase , which converts ATP to cAMP
cAMP activates a protein kinase, which phosphorylates specific channel proteins in the
postsynaptic membrane, causing them to open or close.
NEUROTRANSMITTERS
More than 40 different chemicals are known or suspected to function as neurotransmitters.
Each type of neuron is thought to release one type of neurotransmitter.
A postsynaptic neuron may have more than one type of receptors for neurotransmitters.
1. Acetylcholine is important in muscle contraction, and in the autonomic nervous system.
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It can be inhibitory or excitatory.
It is released from motor neurons called cholinergic neurons, and from the CNS.
2. Biogenic amines are neurotransmitters derived from amino acids.
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Norepinephrine, epinephrine and dopamine are biogenic amine or catecholamine
derived from tyrosine.
Biogenic amines mostly function within the CNS.
Dopamine and serotonin affect mood and have been linked to depression, attention
deficit disorder; Parkinson's and schizophrenia result from lack of dopamine.
3. Amino acids and peptides function as neurotransmitters in the CNS
GABA (gamma aminobutyric acid) is an amino acid that inhibits neurons in the brain and
spinal cord, and produces IPSP.
Glycine, glutamate and aspartate are amino acids that function as neurotransmitters.
Some neurotransmitters are small molecules that act rapidly.
Others are neuropeptides, larger molecules that modulate the effects of the small-molecule
neurotransmitters.
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Substance P is an excitatory neurotransmitters involved in the perception of pain.
Endorphins are neuropeptides that decrease the perception of pain. They also produce
euphoria and other emotional states.
4. Gaseous signals of the nervous system
Nitric oxide, NO, and carbon monoxide, CO, are local regulators.
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NO is involved in the production of erection in males during sexual arousal by relaxing the
smooth muscles in the blood vessels in erectile tissue of the penis allowing the influx of
blood.
Viagra inhibits the enzyme that slows the muscle-relaxing effects of NO.
Gaseous neurotransmitters are synthesized on demand and are not stored in vesicles.
VERTEBRATE NERVOUS SYSTEMS
The vertebrate nervous system is regionally specialized.
In all vertebrates, the nervous system shows cephalization and distinct CNS and PNS
components.
Brain (ventricles)
Central
Spinal cord (central canal)
Vertebrate
Nervous
Sensory
Sensing external and internal environment
Peripheral
Somatic
Motor
Autonomic
Sympathetic
Parasympathetic
Enteric Division
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The somatic nervous system has receptors and efferent nerves from the CNS to muscles.
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The autonomic nervous system has receptors and efferent nerves from the CNS to organs.
Axons within the CNS are located in well-defined bundles or tracts, whose myelin sheaths give
them a whitish appearance, the white matter.
The gray matter consists mainly of dendrite, unmyelinated axons, and clusters of nerve-cell
bodies, or nuclei.
The peripheral nervous system (PNS) consists of paired nerves and their associated ganglia.
Cranial nerves originate in the brain and extend to the head and neck, and one pair, the vagus,
into the thorax and abdomen.
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There are 12 pairs of cranial nerves.
The spinal nerves arise from the spinal cord and supply all parts of the body except the head
and some areas of the neck.
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There are 31 pairs of spinal nerves.
Sympathetic NS permits the body to respond to stressful situations; it is involved in arousal
and energy generation.
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Preganglionic neurons transmit message from CNS to the paravertebral sympathetic
ganglion chain, and postganglionic neurons transmit the message to the effector.
Nerves emerge from the middle region of the spinal cord.
Most pathways consist of a chain of two neurons.
Synapse is inside the ganglion.
Sympathetic neurons release norepinephrine at their target organs.
Parasympathetic NS restores the body to resting state, and actively maintains normal body
functions.
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Parasympathetic preganglionic neurons synapse with postganglionic neurons in
ganglia near or within the walls of the effector organs.
Parasympathetic neurons release acetilcholine at their target organs.
Enteric Division consists of networks of neurons in the digestive tract, pancreas, and
gallbladder.
Controls the organs’ secretions and peristalsis.
The enteric division can function independently but it is normally under the control of the
sympathetic and parasympathetic nervous system.
EMBRYONIC DEVELOPMENT OF THE BRAIN
The embryonic neural tube differentiates into three regions:
1. Presencephalon or forebrain → telencephalon → cerebrum.
diencephalon → thalamus, hypothalamus.
2. Mesencephalon or midbrain → mesencephalon → midbrain (part of brain stem)
3. Rhombencephalon or hindbrain → metencephalon → pons, cerebellum
myelencephalon → medulla oblongata
These primary divisions in turn differentiate to give rise to specific structures of the adult brain.
THE BRAINSTEM
The medulla oblongata, the pons, and the midbrain are derived from the embryonic hindbrain.
They function in homeostasis, coordination of movement, and conduction of information to
higher brain centers.
The medulla and pons
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The brain stem sends axons to the cerebellum and cerebral cortex by releasing
norepinephrine, dopamine, serotonin and acetylcholine.
These signals cause changes in attention, alertness, appetite and motivation.
The medulla contains centers that control visceral functions, including breathing, heart and
blood vessel activity, swallowing, vomiting, and digestion.
The pons contributes to some of these activities, e. g. regulation of breathing.
All sensory and motor messages from the higher brain regions most pass through the brain
stem to the rest of the body. Conduction of information is one main function of the brainstem.
Coordinates large body movements such as walking.
Messages from the right side cross to the left side of the body and vice versa in the medulla.
Arousal And Sleep
Sleep and arousal are controlled by several centers in the brainstem and cerebrum.
The reticular formation of the brainstem contains over 90 separate nuclei.
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Maintains consciousness and determines the degree of alertness.
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Receives messages from sensory receptors through the spinal cord and communicates with
the cerebral cortex.
The more input the cortex receives, the more alert the person is.
It filters out familiar and repetitive information that may cause a brain overload.
The pons and medulla have centers that cause sleep when stimulated, and midbrain has a
center that causes arousal.
Serotonin may be the neurotransmitter of the sleep-producing centers.
Melatonin, produced by the pineal body, seems to have an influence in the sleep-wake cycle.
The function of sleep is not understood.
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One hypothesis is that sleep is involved in the consolidation of learning and memory.
THE CEREBELLUM
The cerebellum develops from the metencephalon, and is located dorsal to the pons and
medulla.
It processes and integrates information from the cerebral motor cortex and from receptors
(visual, equilibrium, etc.), and provides instructions to cerebral motor cortex about the precise
timing and appropriate patterns of skeletal muscle contraction, for smooth, coordinate
movements, agility and posture.
The coordination occurs unconsciously. It checks for errors during motor, perceptual, and
cognitive functions.
Cognitive functions learning, decision making, consciousness, and an integrated sensory
awareness of the surroundings.
THE DIENCEPHALON - EPITHALAMUS, THALAMUS AND HYPOTHALAMUS
The forebrain gives rise to the thalamus, hypothalamus and cerebrum.
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Telencephalon develops into the cerebrum.
Diencephalon gives rise to the epithalamus, thalamus and hypothalamus.
The epithalamus is the most dorsal portion of the diencephalon and forms the roof of the third
ventricle.
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It includes the choroid plexus, a cluster of capillaries that produce the cerebrospinal fluid.
The pineal body is tiny projection of the epithalamus.
The thalamus incoming information from all the sense is sorted out in the thalamus and sent on
to the appropriate higher brain centers for further interpretation in integration. It is the gateway
to the cerebral centers.
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It contains about 12 nuclei that receive information from a specific region of the cerebral
cortex.
Impulses related to emotions, visceral functions, funnel through the thalamic nuclei.
The hypothalamus forms the floor of the third ventricle and receives olfactory messages and
regulates the function of internal organs, maintains homeostasis (temperature, respiration,
regulation of pituitary gland, appetite, etc.)
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The hypothalamus contains the body thermostat that regulates hunger, thirst and
temperature
It regulates the sexual and mating behavior, fight-or-flight response, rage and pleasure.
The hypothalamus and circadian rhythms.
Animals exhibit regulatory rhythmic behavior, e. g. seasonal reproduction, migration, sleeparousal, etc.
Circadian rhythms is a physiological cycle of about 24 hours that is present in all eukaryotic
organisms and that persists even in the absence of external cues.
The biological clock is a component of the circadian rhythm.
In mammals, a pair of structures called the suprachiasmatic nuclei (SCN) in the hypothalamus
functions as a biological clock.
Cells of the SCN produce specific proteins under the influence of light.
The rhythm does no exactly match events in the environment, and external cues are necessary
to keep cycles timed to the outside world.
Visual information received through the eye keep the rhythm synchronized with the daily light
and dark cycle.
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The human circadian rhythm is of 24 hours and 11 minutes.
THE CEREBRUM
The cerebrum develops from the embryonic telencephalon.
In most vertebrates, the cerebrum is divided into right and left hemispheres.
Each hemisphere consists of an outer covering of gray matter, the cerebral cortex, and internal
white matter and a cluster of nuclei called basal nuclei deep within the white matter.
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White matter is made of myelinated axons
Gray matter (cerebral cortex) is made of cell bodies and dendrites.
The cerebrum is the largest and most prominent part of the human brain.
In humans, the largest and most complex part of the brain is the cerebral cortex.
Sensory information analyzed, motor commands are issued, and language is generated in
the cerebral cortex.
Characteristic to mammals is the neocortex, an additional outer layer of cortex consisting of six
sheets of neurons running tangential to the brain surface.
The neocortex is associated with greater cognitive ability and more sophisticated behavior.
The human neocortex amounts to about 80% of the brain mass.
Cerebral cortex is made of gray matter arranged into sulci.
White matter lies beneath the cerebral cortex.
Corpus callosum is made of axons and connects right and left hemispheres.
Axons are arranged into bundles (tracts).
The basal ganglia, a cluster of nuclei within the white matter, are important centers of motor
function.
Regions of the brain
Regions of the brain are specialized for different functions.
Sensory areas receive information from senses and receptors.
Motor areas control the movement of voluntary muscles.
Association areas are the site of intellect, learning, memory, language, and emotion; interprets
sensory information.
The cortex has been mapped into areas responsible for certain functions:
Occipital lobe: visual centers.
Temporal lobes: auditory centers.
Parietal lobes receive information about heat, touch and pressure.
Other areas are involved in complex integrative activities.
The size of the motor area in the brain for any given part of the body is proportional to the
complexity of movement involved and not to the amount of muscle.
FUNCTIONS OF THE CEREBRAL CORTEX
The cerebral cortex controls voluntary movement and cognitive functions.
The major increase in the size of the neocortex that occurred during mammalian evolution was
mostly an expansion of the association areas that integrate higher cognitive functions and make
more complex behavior and learning possible.
Information Processing In The Cerebral Cortex
Sensory information is sent through the thalamus to the appropriate lobe, from here the
information is sent to adjacent association areas, which associate the input with different types
of senses, and assess the significance of the overall sensory input. The signals are then
transmitted to still more association areas. The association areas then compose an appropriate
motor response, which is used by the motor cortex to direct the movement of skeletal muscles.
The primary motor cortex and the somatosensory cortex are located and the rear of the frontal
lobe.
The somatosensory cortex sends information to the motor cortex which in turn sends
commands down the thalamus and spinal cord to skeletal muscles for a particular movement.
In both the motor cortex and the somatosensory cortex, the neurons are distributed in an orderly
fashion according tot eh part of the body that generates the sensory input of receives the motor
commands. See fig. 48.28
Lateralization of cortical function
During early development brain functions are placed into opposite cerebral hemispheres, and a
division of labor is created. This is called lateralization.
Each hemisphere has unique abilities not shared by its partner.
The left hemisphere is normally specialized for high-speed serial information processing
essential to language and logic operations, e. g. language, mathematics, logic operations, and
the processing of serial sequences of information.
The right hemisphere is stronger at pattern recognition, nonverbal ideation, and emotional
processing, e. g. face recognition, spatial relations, nonverbal ideation, intuition, artistic skills,
creative endeavors, music, emotional processing in general.
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Typical right-cerebral dominant people are left-handed and male.
The two hemispheres have instantaneous communication with one another via connecting fiber
tracts, as well as complete functional integration. Both hemispheres work together and never
work alone at any task.
In some cases, lateralization results in cerebral confusion and learning disabilities.
Language and speech
Language is a process that involves multiple areas of the brain.
A large continuous region for language comprehension and articulation is located on the left
hemispheres; portions of the frontal and temporal lobes are involved.
The different areas of this region of the left hemisphere are:
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Wernicke's area is involved in sounding out unfamiliar words.
Broca's area is for speech production.
A region to the anterior of the lobe is involved in language comprehension and word analysis.
Most of the lateral and ventral parts of the temporal lobe coordinate the auditory and visual
aspects of language as when naming objects or reading.
Emotions
The limbic system within the brain is involved in human emotions and memory.
It consists of a group of structures located in each hemisphere, and surrounding the upper part
of the brain stem.
Parts of the diencephalon, the hypothalamus and the thalamus, and inner parts of the cerebral
cortex including the amygdala and the hippocampus make the limbic system.
The main limbic structures are the hypothalamus and the anterior nucleus of the thalamus.
Sensory areas of the neocortex and other brain centers, the limbic system generates the
feelings we generally call emotions.
Limbic system is central to some behaviors like the nurturing of infants and bonding between
individuals, and some primary emotions like laughing and crying.
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We are born with brain circuits prepare to recognize caretakers, human faces, and to
express fear, distress and anger.
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Later on, learning and memory build a history of what actions work in obtaining food and
warmth.
And learning right from wrong by observing the approving or disapproving faces of the
caretakers.
The hippocampus and amygdala of the limbic system are involved in emotional memory.
The limbic system interacts with the prefrontal lobes. There is intimate relationship between our
feelings (mediated by the emotional brain, limbic system), and our thoughts (mediated by our
cognitive brain, prefrontal lobes).
As a result we react emotionally to things that are happening.
Communication between the limbic system and the cortex explains why emotions sometimes
override logic and, conversely, why reason can stop us from expressing our emotions in
inappropriate situations.
Memory and learning
Information is held for a time in short-term memory in areas in the frontal lobes and then
forgotten if unnecessary.
If the information is relevant, a long-term memory process is activated that requires the
hippocampus.
The capacity of the short-term memory is limited to seven or eight pieces of information like the
digits in a telephone number or a short sentence.
The capacity of long-term memory is limitless. The ability to commit information to the long-term
memory declines with age.
The transfer of information from short-term memory to long-term memory is affected by many
factors and is mediated by the amygdala, including:
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Emotional state: we learn better when we are alert, motivated, surprised and aroused.
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The neurotransmitter norepinephrine is involved in memory processing of emotionally
charged events, and is released when we are excited or stressed out.
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Rehearsal: repetitive actions. "Practice makes perfect."
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Association of new information to old information.
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Automatic memory: not all memories are consciously formed. We may remember an
irrelevant fact of a happening, e. g. "the speaker was bold-headed."
Nerve cells can make new connections when learning motor skills, e. g. learning how to ride a
bicycle, how to tie your shoes.
Motor skills are learned by repetition that does not involve the recall of the individual steps or
remembering specific information.
Cellular mechanisms of learning
Functional changes in the synapses of the hippocampus and the amygdala are directly related
to memory storage and emotional conditioning.
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Long-term conditioning (LTD) happens when there is a decrease in responsiveness to
an action potential by a postsynaptic cell. It is induced by weak repetitive stimulation.
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Long-term potentiation (LTP) is an enhanced responsiveness to action potentials by a
postsynaptic cell. A single action potential from the presynaptic cell has a much greater
effect at the synapse than before.
Short-term memory involves brief changes in neurotransmitter receptors and second
messengers are linked to ion channels in postsynaptic neurons.
Specific protein kinases are activated by the secondary messenger cyclic AMP.
These kinases phosphorylate and affect specific ion channels.
In long term memory, the changes in the postsynaptic neuron are slower but longer lasting.
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Cyclic AMP activates protein kinases which enter the nucleus and lead to gene
activation.
The kinases phosphorylate a regulatory protein known as CREB.
CREB turns on the transcription of certain genes.
Some types of learning are related to permanent changes in presynaptic terminals and
postsynaptic neurons.
LTP is associated with the release of the neurotransmitter glutamate by the presynaptic cell.
Glutamate opens gated-channels that allow Ca2+ to enter the postsynaptic cell making it more
responsive to stimulation by initiating a cascade of enzyme activity.
Consciousness
Brain studies show changes in neuronal activity that is correlated with conscious perception,
unconscious versus conscious processing, and retrieval of memories.
Consciousness involves memories and the comprehension of external reality, as well as the
person's emotional state.
It involves many parts of the cerebral cortex.
Consciousness varies from minute to minute: alertness, drowsiness, sleep, stupor, coma.
CNS INJURIES AND DISEASES
CNS injuries and diseases are the focus of much research.
Nerve cell development
Signal molecules released by cells along the way direct an axon’s growth by binding to
receptors on the plasma membrane of the growth cone.
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The leading edge of a growing axon is called the growth cone.
The genes and basic events involved in axon guidance are similar in invertebrates and
vertebrates.
Knowledge of these events may be applied one day to stimulate axonal regrowth following CNS
damage.
Neural stem cells
The adult human brain contains stem cells that can differentiate into mature neurons.
Mature neurons with their many connections are not able to undergo cell division.
The induction of stem cell differentiation and transplantation of cultured stem cells are potential
methods for replacing neurons lost to trauma or disease.
DISEASES AND DISORDERS OF THE NERVOUS SYSTEM
Schizophrenia is characterized by hallucinations, delusions, blunted emotions, and other
symptoms. The patient loses the ability to distinguish reality.
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The cause is unknown.
It has both environmental and genetic factors.
It probably involves multiple genes.
Depression includes bipolar disorder, characterized by manic and depressive phases, and
major depression, in which patients have a persistent low mood.
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It has It has both environmental and genetic factors.
Stress during childhood may be an important factor.
Alzheimer’s disease is an age-related dementia including memory loss and confusion.
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It is caused by neurofibrillary tangles are bundles of degenerated neurons and glial cells.
Plaques are deposits of an insoluble peptide, β-amyloid, produced by the neuron
membrane.
It is a progressive disease.
Neurons die in large numbers leading to shrinkage of the brain.
Parkinson’s disease is a motor disorder caused by the death of dopamine-secreting neurons in
the substantia nigra located in the midbrain region.
Early Embryology
CLEAVAGE
The zygote is totipotent.
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It has the potential to produce all the cell types of the new individual.
Cleavage is a series of rapid mitotic divisions. These cells are called blastomeres.
At about 32-cell stage the embryo is a solid ball called morula.
BLASTULATION
Eventually a hollow ball of cells form the blastula with a fluid filled blastocoel.
Cells at these stages are capable of ameboid movement.
Surface proteins allow cells to recognize one another and determine which ones to adhere to form tissues.
Cytoplasm is not homogeneous and helps determine the course of development.
The amount of yolk determines the pattern of cleavage.
Yolk is a mixture of proteins, phospholipids and fats.
Yolk is evenly distributed in the eggs of most invertebrates and simple chordates: isolecithal eggs.
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Entire egg divides producing cell of even size (holoblastic cleavage).
Cleavage may be radial (deuterostomes) or spiral (protostomes).
Many vertebrate eggs have the concentrated at one end of the cell, known as the vegetal pole: telolecithal
eggs. The other pole is more metabolically active and is called the animal pole.
GASTRULATION
Gastrulation is the embryological process by which the blastula becomes a three-layered gastrula.
Zygote  morula  blastula  gastrula.
The three germ layers are the endoderm, mesoderm and ectoderm.
Each layer gives rise to specific structures.
There are different patterns of gastrulation.
In amphioxus and sea star, cells from the blastula wall invaginate and eventually meet the opposite wall.
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The cavity formed is called the archenteron.
This is the developing gut and the opening is called the blastopore.
In amphibians, the yolk-laden cells obstruct invagination.
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Cells from the animal pole move down over the yolk rich cells and invaginate forming the
dorsal lip of the blastopore.
In birds, epiblast cells migrate toward the midline to form a thickened cellular region, known as the
primitive streak.
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There is narrow furrow at its center, the primitive groove.
Cells migrate in from the epiblast, sink down and move out laterally and anteriorly in the
interior. These cells form the notochord.
ORGANOGENESIS
Organogenesis is the process of organ formation.
Brain, notochord and spinal cord are among the first organs to develop
First the notochord, in all chordate embryos, develops as a cylindrical rod of cells on the dorsal side
The developing notochord induces the overlying ectoderm to thicken and form the neural plate.
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Induction: certain cells stimulate or influence the differentiation of neighboring cells.
Cells from the neural plate move downward and form the neural groove flanked by the neural folds.
The ridges of the neural folds increase and eventually meet forming the neural tube.
The neural tube is formed beneath the surface. Its anterior portion will form the brain and the rest will
differentiate into the spinal cord.
The neural crest consists of cells that lie near the neural tube and will differentiate into sensory neurons
(dorsal root ganglia and postganglionic sympathetic neurons.
The digestive track is formed as a separate foregut and hindgut.
These two simple tubes are lined with endoderm.
Outpocketings give rise to the liver, pancreas and trachea.
The trachea as it grows downward gives rise to the lung buds.
A series of small outpocketings of the pharynx are the pharyngeal pouches. There are corresponding
inpocketings from the ectoderm, the branchial grooves.
EXTRAEMBRYONIC MEMBRANES
Terrestrial vertebrates have four extraembryonic membranes: chorion, allantois, yolk sac and amnion.
Chorion and amnion enclose the entire embryo.
The amniotic cavity is the space between the amnion and the embryo. It becomes filled with the amniotic
fluid.
In humans the allantois is nonfunctional except that its blood vessels form part of the umbilical cord.
In reptiles and birds, the allantois stores nitrogenous wastes.
The yolk sac in higher vertebrates contains little or no yolk.
In the early mammalian embryo, the yolk sac absorbs nutrients and oxygen from the
mother and then degenerates.
MAMMALS
Mammals fall into three groups that differ in the way they use the amniotic egg.
Monotremes
These primitive mammals produce a shelled egg like their reptilian ancestors. Only three species exist
today: two species of spiny anteater (echidna) and the duckbill platypus. [View (48K)]
Marsupials
Marsupials do not produce a shelled egg. The egg, which is poorly supplied with yolk, is retained for a
time within the reproductive tract of the mother. The embryo penetrates the wall of the uterus. The yolk
sac provides a rudimentary connection to the mother's blood supply from which it receives food, oxygen,
and other essentials. However, this interface between the tissues of the uterus and the extraembryonic
membranes never becomes elaborately developed, and the young are born in a very immature state.
Despite their tiny size, they are able to crawl into a pouch on the mother's abdomen, attach themselves to
nipples, and drink milk from her mammary glands.
Marsupials are still abundant in Australia, but only the opossum is found in North America.
Placental mammals
In placental mammals, the extraembryonic membranes form a placenta and umbilical cord, which
connect the embryo to the mother's uterus in a more elaborate and efficient way. The blood supply of the
developing fetus is continuous with that of the placenta. The placenta extracts food and oxygen from the
uterus. Carbon dioxide and other wastes (e.g., urea) are transferred to the mother for disposal by her
excretory organs.
Humans are placental mammals.
http://users.rcn.com/jkimball.ma.ultranet/BiologyPages/E/ExtraembryonicMembranes.html
SUMMARY OF ANIMAL CLASSES
CLASS MYXINI Hagfishes
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Have a partial cranium (skull) but no vertebrae, and so they are not truly vertebrates.
Their skeleton is composed of cartilage, and lacks vertebrae.
They have a notochord made of cartilage.
Hagfish also lack jaws.
They are considered vertebrates only if vertebrates are equated with the Craniata:
neural crest, paired sensory organs of the head (cephalization), brain and skull
(cranium).
There are about 30 species all of them marine, bottom-dwelling scavengers.
CLASS CEPHALASPIDOMORPHA Lampreys
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There are about 35 species of lamprey living in fresh and salt water.
Lampreys are anadromous.
Have a complete cranium or braincase.
Their skeleton is entirely of cartilage.
The notochord persists in the adult and is surrounded by cartilage.
Pairs of cartilaginous projections extend upwards and partially surround the nerve cord.
These are considered
rudimentary true vertebrae.
Have a single "nostril" on the dorsal side of the
They lack paired appendages.
The larva is a suspension feeder.
Some species are parasitic on fish.
CLASS CHONDRICHTHYES Cartilaginous fishes
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Probably originated in salt water.
Skin with placoid scales; their structure is similar to that of teeth.
Endoskeleton entirely cartilaginous. This is a derived characteristic.
Notochord replaced by vertebrae.
Two pairs of fins: pectoral and pelvic pairs.
Two-chambered heart.
5 to 7 pairs of gills with separate and exposed gill slits.
No swim bladder or lung.
Lateral line sensitive to changes in water pressure detects vibrations.
Internal fertilization and separate sexes.
Pelvic fins modified into claspers in males to act as an intromittent organ
Oviparous, eggs are released and the young are born outside the mother;
Ovoviviparous, fertilized eggs are kept in the uterus where they hatch;
Viviparous, you develop in the uterus and are fed by the mother through a placenta.
About 750 species: sharks, rays and chimeras
CLASS OSTEICHTHYES Bony fishes
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Probably originated in fresh water.
Ossified skeleton made of calcium phosphate.
Skin usually with overlapping dermal scales, some fish without scales.
Skeleton with many vertebrae replace the notochord.
Median and paired fins supported by fin rays of cartilage or bone.
Jawed mouth terminal; most have many teeth.
Gills supported by bony gill arches and covered by a common bony flap, the
operculum.
Two-chambered heart.
Swim bladder usually present.
Sexes separate and fertilization is usually external; some species have internal
fertilization
Development mostly oviparous.
About 30,000 species.
CLASS AMPHIBIA Frogs, toads, salamanders, newts, caecilians.
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Aquatic larva undergoes metamorphosis into a terrestrial adult in many species.
Amphibian means "two lives."
Skin smooth and moist with many glands, some of which may be poison glands; no
scales; involved in gas exchange.
Respiration by lungs, skin and gills, either separately or in combination; external gills in
the larval form and may persist throughout life in some.
Three-chambered heart: two atria and one ventricle.
Systemic and pulmonary circulation.
Ectothermal.
Separate sexes; fertilization usually external in frogs and toads and mostly internal in
salamanders and caecilians.
Eggs without shell.
Most species return to water to reproduce.
About 4,800 species.
CLASS REPTILIA Snakes, turtles, lizards, crocodilians.
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Leathery skin containing keratin that prevents desiccation; they don't breath through the
skin like amphibians.
Better lungs than amphibians; shift from buccal pumping by aspiration.
A 3-chambered heart with an almost complete septum separating the ventricle into
two chambers.
Internal fertilization.
Behavior helps regulate body temperature.
The amniotic egg.
CLASS AVES Birds
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All birds have feathers.
All birds have toothless beaks.
All birds have 2 legs and 2 wings.
Most birds have 4 toes. A few birds that live only on the ground, such as the Ostrich,
have only 3 toes.
All birds are endothermic.
Excrete solid waste in the form of uric acid.
Lack diaphragm.
Sexes separate.
Fertilization internal; eggs with much yolk and calcareous shells.
Vocal calls and complex songs.
Complex behavior, e.g. courtship, migration.
Birds are adapted to fly.
There are about 8,600 species of living birds in the world.
CLASS MAMMALIA
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Body covered with hair.
Skin with sweat, scent, sebaceous and mammary glands.
Mouth with differentiated teeth.
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Four limbs in most, adapted for many forms of locomotion.
Four-chambered heart.
Respiration system with lungs and larynx.
Muscular diaphragm present.
Highly developed nervous system and brain.
Three ear ossicles.
Endothermic.
Internal fertilization; mostly viviparous; fetal membranes (amnion, chorion, allantois).
Females nourish the young with milk from mammary glands.
There are about 4,500 species of mammals extant.
OTHER:
CHARACTERISTICS OF PRIMATES
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Ability to move the four limbs in various directions.
Grasping power of hands and feet.
Slip-resistant cutaneous ridges and the ventral side of hands and feet with tactile
sensitive organs, e. g. Meissner corpuscles.
Retention of the clavicle to support the pectoral girdle.
Flexible vertebral column that allows twisting and turning.
Anthropoid primates also have a...
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Highly developed brain.
Long postnatal growth period with parental care and long learning period.
Not all primates have all the characteristics e. g. humans do not have grasping feet.
MAJOR FEATURES OF HUMAN EVOLUTION
Brain size: hominoids 6 million ago had brains averaging 400-450 cm3; modern humans
average 1,300 cm3
Jaw shape: hominoid ancestors have longer jaws, prognathism, while modern humans have
shorter jaws and pronounced chins.
Bipedal posture: hominoid ancestors were quadrupeds; modern humans are bipedal.
Reduced size difference between the sexes: hominoids show a greater difference between
males and females than modern humans, where males and females are more alike in body
size.
Family structure: modern humans are monogamous, the newborn is exceptionally dependent
on the mother with extended parental care of the young.