brain
Introduction to Brain and Behavior
Authors: Bryan Kolb, Ian Q. Whishaw, G. Campbell Teskey
Focuses on how neurons communicate and adapt, covering:
A chemical message
Varieties of neurotransmitters and receptors
Neurotransmitter systems and behavior
Adaptive role of synapses in learning and memory
A Chemical Message
Otto Loewi (1921): Conducted an experiment with frog hearts
First isolation of a chemical messenger
Demonstrated the role of the vagus nerve and neurotransmitter acetylcholine (ACh) in slowing heart rate
Acetylcholine (ACh):
First neurotransmitter discovered in both the peripheral nervous system and central nervous system (CNS)
Activates skeletal muscles in the somatic nervous system (SNS)
May excite or inhibit internal organs in the autonomic nervous system (ANS)
Research Focus 5-1: The Basis of Neural Communication in a Heartbeat
Heart rate adjusts to match energy expenditure and body’s nutrient and oxygen needs
Loewi's demonstration showed that the frog’s vagus nerve contains a chemical that signals the heart to slow down.
Later, Loewi identified two messenger chemicals:
Excitatory message: signals a speed up
Inhibitory message: signals a slow down
Experiment 5-1: How Does a Neuron Pass on a Message?
Loewi’s complementary experiments led to discoveries regarding other neurotransmitters:
Epinephrine (EP, or adrenaline):
Acts as a hormone to prepare the body for fight or flight during stress
Also serves as a neurotransmitter in the CNS
Norepinephrine (NE, or noradrenaline):
Neurotransmitter found in the brain and in the parasympathetic division of the autonomic nervous system
Accelerates heart rate in mammals
Neurotransmitters
Neurotransmitter:
Chemical released by a neuron onto a target, exerting an excitatory or inhibitory effect
Outside the CNS, many neurotransmitters circulate in the bloodstream as hormones
Slower action than that of neurotransmitters
Loewi's work sparked extensive research into the number of neurotransmitters
Actual number of transmitters: 60 confirmed; 200 posited
Structure of Synapses
Discovery of synaptic structure aided by electron microscopy in the 1950s
Neurotransmitters are packaged in vesicles at the end terminal of axons
Chemical synapse:
Junction where messenger molecules (neurotransmitters) are released from one neuron to excite or inhibit another
Most synapses in mammals are chemical
Chemical Synapse
Depicted in illustrations to demonstrate synaptic interactions
Clinical Focus 5-2: Parkinson Disease
James Parkinson initially described the condition; named Parkinson's disease by Jean-Martin Charcot
Key findings in understanding its neural basis:
Degeneration of the substantia nigra opposite the symptomatic side
Symptoms manifest when dopamine (DA) levels drop below 10% of normal in the basal ganglia
Experiments in rats indicated that selectively destroying dopamine-containing neurons produced Parkinsonian symptoms, affirming dopamine's role in the substantia nigra to basal ganglia pathway
Terms to Learn: Structure of Chemical Synapses
Presynaptic membrane: Axon terminal
Postsynaptic membrane: Dendritic spine
Synaptic cleft: Space between presynaptic and postsynaptic membranes
Tripartite synapse: Involves pre- and postsynaptic neurons and supporting glial cells
Synaptic vesicle: Contains neurotransmitters, located in presynaptic area
Storage granule: Presynaptic structure for holding neurotransmitters
Postsynaptic receptor: Receives neurotransmitters, located on postsynaptic neuron
Anterograde synaptic transmission: From presynaptic to postsynaptic
Transporter: Protein that moves neurotransmitters in the neuron
Neurotransmission in Five Steps
Steps of anterograde synaptic transmission:
Neurotransmitter synthesis occurs within the neuron
Neurotransmitter is packaged and stored in vesicles at the axon terminal
During an action potential, neurotransmitter is released into the synaptic cleft
Neurotransmitter binds to receptors on the postsynaptic membrane
Neurotransmitter is degraded or removed to stop its action
Steps 1 and 2: Neurotransmitter Synthesis, Packaging, and Storage
Neurotransmitters derive from two main pathways:
Synthesized in axon terminal: Using dietary building blocks via transporters
Synthesized in cell body: Following DNA instructions (peptide transmitters), then transported to axon terminal
Origins of neurotransmitters include:
Peptide
Lipid
Gaseous
Ion
Step 3: Neurotransmitter Release
At the axon terminal, action potential opens voltage-sensitive calcium (Ca²⁺) channels
Ca²⁺ enters and binds to calmodulin forming a complex that causes vesicles to release neurotransmitter contents
Exocytosis: Process of transmitting neurotransmitters into the synaptic cleft
Step 4: Receptor-Site Activation
After neurotransmitter release, it diffuses across the synaptic cleft to activate postsynaptic receptors
Types of transmitter-activated receptors:
Ionotropic receptors: Directly control ion channels
Metabotropic receptors: Indirectly influence cellular activity through second messengers
Autoreceptors: Regulate neurotransmitter release
Quantum: The quantal release of neurotransmitters
Step 5: Neurotransmitter Inactivation
After signaling, neurotransmitters are removed from receptors and synaptic cleft through:
Diffusion: Moving away from receptor sites
Degradation: Enzymatic breakdown
Reuptake: Neurotransmitter is reabsorbed into presynaptic neuron
Astrocyte uptake: Supported by glial cells
The Versatile Synapse
Synapses exhibit considerable variation in type, location, structure, and function
This diversity allows synapses to be a flexible biological communication system
Transmitters can influence neuron actions by connecting to dendrites, cell body, or axon
Electrical Synapses
Gap junction: Contains connexin proteins connecting adjacent cell membranes to form hemichannels
Allows ions to pass directly between neurons
Regulated gates can open or close, facilitating rapid communication
Can enable dual chemical and electrical transmission (mixed synapse)
Excitatory and Inhibitory Messages
Excitatory synapse characteristics:
Typically located on dendrites
Round vesicles with dense material on membranes
Wider synaptic clefts
Larger active zones
Inhibitory synapse characteristics:
Found mainly on cell body
Flat vesicles with less dense material on membranes
Narrow synaptic clefts
Smaller active zones
Research Focus 5-3: Dendritic Spines; Small but Mighty
Dendritic spines: Key structures in learning and memory
Can act independently and perform structural changes
Impairments can lead to cognitive disabilities and Alzheimer’s disease
Evolution of Complex Neurotransmission Systems
Chemical communication may have originated from single-celled organisms’ feeding behaviors
Juices secreted via exocytosis to capture prey (similar to neurotransmitter release)
Quiz Questions
The neurotransmitter that accelerates the heart rate in mammals is called:
a. epinephrine
b. norepinephrine (correct)
c. acetylcholine
d. bradycardia
A chemical with an excitatory or inhibitory effect when released by a neuron onto a target is called:
a. molecule
b. neurotransmitter (correct)
c. impulse
d. messenger
Synaptic structure was first revealed in the 1950s, using:
a. electron microscopy (correct)
b. digital X-rays
c. chemotherapy
d. gamma radiation
Varieties of Neurotransmitters and Receptors
Extensive diversity in neurotransmitters and receptors allowing versatile interactions
Some neurotransmitters can have both excitatory and inhibitory effects depending on their site of action
Interaction among neurotransmitters at a single synapse may enhance overall potency
Criteria for Identifying Neurotransmitters
Identification of New CNS Neurotransmitters
Staining techniques: Identify specific chemicals in living tissues
Stimulating: Use of microelectrodes on neuronal targets
Collecting: Preserve tissue in saline to detect neuronal communication
Renshaw Loop
All motor-neuron axons from the spinal cord use acetylcholine to communicate
Each may have an axon collateral within the spinal cord synapsing on an interneuron, which then synapses back on the motor neuron's cell body
Toxin example: Strychnine blocks the Renshaw loop leading to increased motor neuron activity, risking respiratory failure
Broadening the Term Neurotransmitter
Expanded definition includes chemicals that:
Transmit messages affecting postsynaptic voltage
Alter synaptic structure
Communicate retrograde signals to influence presynaptic activity
Classes of Neurotransmitters
Small-molecule transmitters: Quick-acting, synthesized in axon terminals
Peptide transmitters
Lipid transmitters
Gaseous transmitters
Ion transmitters
Small-Molecule Transmitters
Fast-acting neurotransmitters synthesized in axon terminals from nutrient sources
Table of Best-Known Small-Molecule Neurotransmitters:
Acetylcholine (ACh)
Amines:
Dopamine (DA)
Norepinephrine (NE)
Epinephrine (EP)
Serotonin (5-HT)
Amino Acids:
Glutamate (Glu)
Gamma-aminobutyric acid (GABA)
Glycine (Gly)
Histamine (H)
Purines:
Adenosine
Adenosine triphosphate (ATP)