Neurophysiology

What is the nervous system?

-it is a complex network of neurons and other associated cells that are responsible for transmitting signals
between the brain and the body

The nervous system

• nerve tissue within the brain and spinal cord = the central nervous system
• nerve tissue outside the brain and spinal cord = the peripheral nervous system

Neurons
• the primary functional unit of the CNS and the PNS
• communicate with each other in networks
• communicate via electrical and chemical signals
• can be over a meter in length!

Dendrite

. 3
rodeevier
o

Axon
terminal

schwann cell

Sheath
Myein
 How do neurons differ from other cells of the body?
Similarities Differences

They have axons and dendrites which are

They have a cell membrane specialized structures

They communicate via electrical signals

They have a nucleus and organelles

They have similar cellular processes (protein

They make special chemicals (called
synthesis, energy production)
neurotransmitters)
Neurons receive and pass on signals
• neurons carry signals from the body to the brain
• neurons carry signals from the brain to the body to execute a response

Synapses
• allow neurons to communicate
• the presynaptic neuron releases a chemical into the synaptic cleft
• this chemical binds to, and modulates the activity of postsynaptic neurons
• those chemicals can be categorised into one at 3 types

1. Neurotransmitters - shortrange
2. Neuromodulators - medium range
3. Neurohormones - long range (released into the blood)
• 86 billion neurons in the human brain
• Action potentials
• an electrical current is the flow of electric charge. This can be in the forms of electrons, ions and
electrolytes
• in the spaces between neurons, there are lots of sodium ions I sodium that has last an electron and
become positively charged
• the inside of the cell is slightly negatively charged compared to the outside L resting potential at a
neuron is - 70mv.
• sodium ions want to move into the inside of the cell
• the flow of ions across the cell membrane is an electrical current
• this electric current can be propagated through a neuronal cell bod
• This how of electric current in a neuron is called an action potential.

Cell membranes are impermeable to these charged ions, so they cannot move backwards and forwards
across the membranes therefore Cell membranes have "gates" or "channels" that allow the movement of
specific ions. These ion gates/ion channels open in response to various stimuli, including
neurotransmitters, and electric currents
* lons can move in or out of the cell to raise or lower the potential difference between the inside and
outside of the cell
* If the inside becomes more positively charged this is called depolarisation
* If the inside becomes more negatively charged this is called hyperpolarisation
* When a neurotransmitter is released from a pre-synaptic neuron, this can initiate a small number of ion
channels opening on the post-synaptic neuron to allow a few sodium ions to cross the cell membrane
* If the resting potential rises to -55mV an action potential is initiated
HOW DOES MOVEMENT OF IONS ACROSS THE CELL MEMBRANE RESULT
& IN AN "ACTION POTENTIAL"?
* In response to a signal (a neurotransmitter, for example) a small number of ion channels open to allow a
few Nations to cross the cell membrane
* This results in a rise in the membrane potential from -70mV
* If the threshold potential (-55mV) is reached a huge number of ion channels open allowing many Na+ ions
to cross the cell membrane
* When the membrane reaches +40mV potassium (K*) channels open
* Potassium channels open later than sodium channels because they open slowly
* Potassium channels are slow to open BUT ALSO slow to close
* Because of this, potassium continues to leave the cell, and the cell hyperpolarizes (-90mV)
• Cell has hyperpolarized
• Gone past resting potential of -70mV
• During the refractory period, no more action potentials can be initiated

Nerve impulses
* Really important to realise that what we have just described only relates to a small section of the cell
membrane
* How is this signal "propagated" from one end of the neuron to the other?
* Myelin plays an important role
* Insulates, and allows signal to propagate to axon terminal
* "Saltatory conductance"
• An influx of sodium ions causes localized depolarization on a section of membrane
• If the signal is strong enough, the influx of sodium ions will depolarize the adjacent piece of membrane
to the threshold potential
• This sequential depolarization of adjacent pieces of membrane allows the signal to be propagated in one
direction
• NB - Refractory period means the signal can't move backwards!
1. A signal (usually neurotransmitter) reaches a neuron dendrite
2. Small number of Nat channels open - Nat moves into the cell
3. If enough Nat moves into the cell and threshold potential is reached - action potential initiated
4.Lots of Nat channels open, and a huge number of Na* ions move into the cell
5. Inside of the cell becomes positive due to influx of Na+ (+40mV)
6. K+ channels are slower to open, and open at +40mV
7. At this membrane potential Na+ channels close and no more Nat enters the cell
8. K+ moves out of the cell
9. The cell moves back towards its resting potential as a result of +ve K+ ions moving out
10. K+ continues to leave the cell causing hyperpolarisation (refractory period)

* Demyelinating diseases slow the conduction of action potentials

* Neurotoxins - can block ion channels
* Anaesthetics can also work in this way (procaine)
* Hypo/hyperkalemia - blood potassium dysregulation results in the threshold potential being greater or
smaller than normal - results in action potentials being instigated more or less frequently than normal

Neuronal connectivity (nerve tracts and nervous tissue organisation)

• On their own, neuronal axons are quite vulnerable (very thin and sometimes have to traverse large
distances).
• Evolution has bundled these neuronal axons together to make them tougher (strength in numbers!)
• In the PNS they are called nerves
• In the CNS they are called nerve tracts
• Sciatic nerve is the largest
• Sometimes the cell bodies of neurons within nerves cluster together - these are referred to as ganglia
• Ganglia can be thought of as relay stations
 Types of neurons
• Can be classified based on structure, function or directionality
Afferent (sensory) neurons - carry information from organs/tissues to the CNS
Efferent (motor) neurons - carry information from the CNS to target cells
Interneurons - connect afferent and efferent neurons within the CNS

Afferent (sensory) neurons are critically important for converting internal and external stimuli into action
potentials which can be sent to the brain for interpretation, and it necessary to instigate a response
Sensory neurons are activated by various stimuli
vision/light
sound
taste
smells
temperature
pain
spatial awareness
chemicals in blood
Sensory receptors located in the sensory neuron dendrites (or non neural cells that synapse onto
sensory neurons) receive the signals
Can be classified based on their location:
Exteroceptors occur at or near the surface of the skin; include those for tactile sensations, such as
touch, pain, and temperature, as well as those for vision, hearing, smell, and taste
Interoceptors (visceroceptors) respond to stimuli occurring in the body from visceral organs and blood
vessels
Proprioceptors respond to stimuli occurring in skeletal muscles, tendons, ligaments, and joints. These
receptors collect information concerning body position
* Can also be classified by the stimulus detected:
Mechanoreceptors (physical force such as pressure and stretch)
Photoreceptors (light)
Thermoreceptors (temperature)
Chemoreceptors (taste, smell, body chemistry)
Nociceptors (tissue damage/pain)

Spinal nerves and cranial nerves

• Sensory neurons in the PNS connect to the CNS through spinal and cranial nerves
• Spinal nerves lie throughout your body and connect to the CNS via the spinal cord
• Cranial nerves innervate the head and connect to the CNS via the brain

Cranial nerves
* Largely responsible for relaying information between the brain, and the head and neck
* Some afferent, some efferent, some

Spinal nerves
• Spine is composed of multiple bones joined by sutures
* Vertebrae are composed of individual vertebra connected via disks of connective tissue, that allow
movement
* 31 pairs of spinal nerves enter and leave the spinal column through notches between the stacked vertebrae
Spinal cord
• 8 pairs cervical nerves
• 12 pairs thoracic nerves
• 5 pairs lumbar nerves
• 5 pairs sacral nerves
• 1 coccygeal segment
 ·
White matter:
Posterior (dorsal)
gray matter :

Spinal cord
* Sensory neurons (within spinal nerves) carrying information to the CNS enter the spinal cord through
the dorsal horn (blue lines)
* Motor neurons efferent nerves exit the CNS at the ventral horn (red lines)
* The cell bodies of sensory neurons are located within the dorsal root ganglia
The nervous system tracts
These sensory and motor nerve pathways are arranged into "tracts"
* ascending tracts (sensory)
* descending tracts (motor)
Medial lemniscus pathway - 2 tracts
- fine touch and positional sense
Spinocerebellar tract
- body positioning (muscles/tendons)
Anterolateral (spinothalamic tract)
- info from skin
- lateral spinothalamic (pain and temperature)
- anterior spinothalamic (touch)
Spinoreticular tract (anterolateral)
- response to pain
Ascending tract neurons
• Spinothalamic tracts and medial lemniscus tracts consist of 3 neurons with decussation
• Spinocerebellar tracts - 2 neurons with no decussation
- Lateral spinothalamic tract
* Spinoreticular tract
- 4 neurons with decussation

Processing
* The signal is transmitted to the relevant area of the brain
* For example, somatosensory information is transmitted to the somatosensory cortex of the postcentral
gyrus (responsible for conveying information regarding location and type of sensation/touch)
* Visual information is transmitted to the visual cortex etc etc
* Further processing allows sensory information to be incorporated into memory, but also initiates a
response
* This can be voluntary or involuntary
* Motor functions are largely controlled by the frontal lobe
* Primary motor cortex (also called the somatomotor cortex is located in the frontal lobe
* The primary motor cortex stimulates spinal cord neurons to stimulate skeletal muscle movement
* The primary motor cortex receives input from multiple regions that aid in the planning and subsequent
execution of movement
* Frontal lobe is also responsible for higher cognitive functioning
* Famous case is Phineas Gage (1848)

Motor tracts
* Descending tracts (motor neurons) can be autonomic or somatic
* There are 3 major types of descending tract
* Corticospinal (pyramidal) tracts (voluntary control of skeletal muscles). These nerves exit at the spinal
cord. Decussates
* Corticobulbar (pyramidal) tracts (voluntary control of skeletal muscles). These nerves exit at the brainstem.
No decussation
* Subconscious (extrapyramidal) tracts (involuntary movement, balance, body positioning, muscle tone)
Motor tracts - somatic (voluntary)
* The corticospinal and corticobulbar tracts (pyramidal tracts) descend from the motor cortex
* 2 neuron system. Upper motor neuron sends a signal from the motor cortex
* The upper motor neuron synapses with a lower motor neuron in the brainstem (corticobulbar) or spine
(corticospinal) and the lower motor neuron exits the CNS at this point
* Corticospinal tract decussates. Corticobulbar does not.
* This decussation occurs in the medulla pyramids

Motor tracts- extrapyramidal (involuntary)

• Extreapytimidal tracts - involuntary movement ( posture, locomotion, autonomic functions,
balance)
• There are a number of these tracts, and they do not pass through the meulla pyramids
• They regulate lower motor neurons and result in involuntary skeletal muscle movement

So far we have learned

• neurons communicate via action otentials
• Action potentials are initieted via the release of nurotransmitters from a presynaptic cell, binding
to a postsynaptic cell
• Neuronal axons are bundled together into "tracts"
• These tracts are ascending or descending, and take sensory information from the periphery to
the CNS, or from the CNS to the periphery to instigate a response
• So far we have discussed voluntary movement
PNS
• The peripheral nervous system( nervous tissue outside the CNS) can be subdivided into nervous
tissue that controls voluntary movements ( somatic system ) and involuntary system ( autonomic
system)
• Somatic nerve signals are transmitted to and from the brain (cerebral cortex) via the cranial and
spinal nerves
• Autonomic system is regulated by the hypothalamus, medulla and the pons ( regulates breathing,
heart, glands, digestion etc.)

Autonomic system
• general visceral afferent fibres transmit sensory information from internal organs, glands etc to the
CNS ( medulla, hypothalamus and pons)
• Motor neurons emanate from both the brain stem and the spinal cord

PNS STRUCTURE
* Autonomic system differs from somatic system
* In the somatic system, the motor neuron cell bodies lie within the CNS
* In the autonomic system, a preganglionic neuron residing in the CNS, synapses with a second
postganglionic motor neuron at an autonomic ganglion that resides outside the CNS
* The postganglionic neurons are clustered into various nerves which extend to target organs
Come from different regions of the CNS
* Sympathetic-from the thoracolumbar region
* Parasympathetic-from the craniosacral region
Differing locations of ganglia
*Sympathetic - close to spinal cord in a chain
* Parasympathetic - close to target organs
Differing lengths of postganglionic fibers
* Sympathetic - Long
* Parasympathetic - Short
Postganglionic branching
• Sympathetic - lots, so that multiple organs can be mobilized at once
• Parasympathetic - very little branching

Neurotransmitters
* Both pre- and post-ganglionic neurons release neurotransmitters
* In most neurons, electrical signals (action potentials) result in neurotransmitter release into the
synaptic cleft and stimulate or inhibit an action potential in a post-synaptic cell

PNS Neurotransmittes
* The 2 main neurotransmitters in the PNS are:
- acetylcholine (ACh; cholinergic neurons)
- norepinephrine (noradrenaline; adrenergic neurons)
* All ANS preganglionic fibres are cholinergic, as are almost all parasympathetic postganglionic fibres
• Nicotinic receptors are found on the cell bodies of all postganglionic neurons, both sympathetic and
parasympathetic
* Acetylcholine released from the preganglionic neurons binds to these nicotinic receptors
* This causes depolarization and excitation - preganglionic cholinergic neurons are therefore always
excitatory
* Acetylcholine released by all parasympathetic postganglionic neurons bind to muscarinic receptors on
target cells.
* This can be inhibitory or excitatory
• Postganglionic sympathetic neurons secrete norepinephrine, which binds to adrenergic receptors
* There are many subclasses of adrenergic receptor - a1, a2, B1, B2
* alpha one most common on effector tissues and is excitatory
* a2 is inhibitory
* B1 and B2 can be excitatory or inhibitory depending on cell type

PNS neurotransmitters removal

* Excess neurotransmitter
- diffuses away
- broken down
- transported back into the presynaptic cell (or other)
* Following release ACh is removed from the synaptic cleft via degradation (acetylcholinesterase)
* Noradrenaline is removed by preganglionic cell reuptake then degradation (monoamine oxidase)
REFLEX ARC
* Nerve pathway involved in a reflex
* Can occur independently of the brain (i.e. at the level of the spine)
* These are "spinal reflexes"
1. Sensory receptor activated
2. Sensory (afferent) neurons carry these signals to the spinal cord or brain stem
3. These sensory neurons synapse with an interneuron in the spinal cord or brain stem
4. Motor (efferent) neurons carry the return signal to the muscles
5. The motor fibres innervating the muscle cause a muscle response
• Usually involve 3 types of neuron - sensory neuron, interneuron (relay neuron), and motor
neuron
-monosynaptic reflex arcs only utilise sensory and motor neurons
-polysynaptic reflex arcs utilise interneurons in addition to sensory and motor
* Reflex arcs can also be long or short
- long reflex arcs involve the CNS
- short reflex arcs occur independent of the CNS
* Reflex arcs are either:
- autonomic (affect organs/viscera)
- somatic (affect muscles)

Enteric nervous system

* Refers to the nervous tissue of the gut
* Can act independently of the CNS
* Intrinsic neurons are completely located within the gut wall
* Autonomic neurons of the CNS that innervate the gut are called extrinsic neurons
* Has long reflex arcs (CNS integrating centre) and short reflex arcs (self-contained; no CNS
communication)
* Short reflex arcs are responsible for glandular/cellular secretion, gut motility etc
* Long reflex arcs (transmitted from the CNS)
- sight, smell, thought of food stimulate gut activity
- emotional responses (butterflies in stomach, stress induced diarrhoea)

THE BRAIN
• Weighs approximately 1-1.5kg (2% of body weight)
* Utilises glucose as an energy source (20% of glucose-derived energy)
* The human brain is much larger in comparison to similarly sized animals
* The reasons for this are unclear — ecological/social pressures, language????
• brain in men is slightly larger in volume , but this does not correspond to IQ
•
• Women more likely to suffer depression
• Men more likely to abuse alcohol/drugs, and much more likely to suffer from dyslexia, and autism
• Men have better hand-eye-coordination, dealing with multiple inputs to working memory, tracking
moving objects etc
• Women have better verbal comprehension, writing ability, long-term memory retrieval etc
• Many studies have demonstrated similar IQ's between sexes, but woman have higher EQ's.....

THE CEREBRUM
• The cerebrum is divided into 4 lobes
• Frontal lobe- reasoning/planning/decision making, but also skeletal muscle movement (motor cortex,
and taste (gustatory cortex)
•
•
Occipital lobe- sight (visual cortex), and also the identification of visual stimuli
Parietal lobe- sense of touch (somatosensory cortex), and language processing
G
• Temporal lobe- memory, interpreting visual information, auditory processing (auditory cortex), smell
(olfactory cortex)

THE BRAINSTEM
• The brainstem connects the brain to the spinal cord and consists of the midbrain, the medulla
(oblongata), and the pons
• The cranial nerves connect to the CNS at the level of the brainstem
• The sensory tracts and the motor tracts pass through the brainstem
O
• Fundamentally important in breathing, sleeping heart function, arousal etc

THE CEREBELLUM
• The cerebellum receives input from the motor cortex regarding movement
• Input from peripheral sensory neurons and inner ear (proprioceptive)
• Very important in the co-ordination/timing/precision/ execution of movement/posture
• Disorders result in ataxia
O
 O
THE DIENCEPHALON
• The diencephalon contains a number of structures including the thalamus, the hypothalamus, and the
pineal gland
• The thalamus is a relay station through which sensory and motor tracts pass on their way to the
cerebrum
• Hypothalamus is a centre for homeostasis, and has multiple sensory inputs
• Pineal gland secretes melatonin
E
THE BASAL GANGLIA
• Important in voluntary movement and behaviour, particularly learning and emotion
• Control (amongst other things) the fluidity of movement, by forming a loop with the motor cortex, and
the thalamus
• Disorders include Parkinson's, OCD, addiction

THE LIMBIC SYSTEM

• Located below the cerebrum
S
• The limbic system supports learning and memory
• Also controls the visceral autonomic (involuntary ) system
• Several regions of the brain are interconnected to regulate these processes, including the amygdala
(emotional processing), the hippocampus (memory consolidation), and the nucleus accumbens
(pleasure, reward)
• Reduced hippocampal volume associated with depression and anxiety
CNS PROTECTION
• skull/vertebrae
• CSF
• Meninges
• Blood brain barrier (except a limited number of areas, inclucing hypothalamus, and vomiting centre
• Brain is protected by connective tissue and the skull
• Connective tissues is called the meninges

MENINGITIS
• inflammation of the meningis- fever headaches
• 9 million people over 400000 deaths
• Viruses, bacteria, fungus ( Neisseria meningitedes causes petechial rash)

CEREBROSPINAL FLUID (CSF)

• Cerebrispinal fluid is a salty soltion secreted by the ventricles that bathes the brain
• Very important for - boyancy/support, removal of waste products, distribution of substances around
the brain
• Is analogous to blood plasma, but with low protein and very few cells
• Synthesised in the ventricles of the brain 2x lateral and a 3rd and 4th ventricle
• The CFS is primarily synthesised by ependymal cells located within regions called the choroid plexus in
each of the ventricles
• The CFS drains from the lateral ventricles into the 3rd then 4th ventricle
• Is then "pumped" around the brain in the subarachnoid space, and also down the central canal of the
spinal cord
• Blockages = hydrocephaly
• Finally leaves the subarachnoid space by entering the sinuses
• Major sinus is the superior sagittal sinus
BLOOD BRAIN BARRIER
• The blood-brain barrier regulates what has access to the CNS
- toxins
- infectious agents
- proteins and neuroactive agents
• Brain capillaries are made from endothelial cells
• In brain there are "tight junctions"
• No blood-brain barrier
- medulla oblongata (vomiting centre)
- hypothalamus (receives various hormonal signals)
• Oxygen can move freely across the blood-brain-barrier
• Some larger molecules can move across but only by "active transport" (includes glucose)
• Lipid soluble compounds can move freely across
• Infectious agents are usually stopped

SPECIAL SENSES
• Special senses have specialized organs devoted to them
- eyes
- ears (hearing and vestibular)
- nose
- tongue
• Touch is not considered a special sense
• In the special sense organs a stimulus is converted into a signal
• This stimulus acts on sensory receptors found on sensory neurons (or cells associated
with sensory neurons)
• The receptors for smell are neurons, but the other special senses use non-neural cells
that synapse onto sensory neurons
• Can be further classified by the stimulus detected:
- Mechanoreceptors
- Photoreceptors
- Thermoreceptors
- Chemoreceptors
- Nociceptors
• Stimuli initiates an action potential if it is of sufficient strength

OLFACTORY SYSTEM
• Large nasal cavity contains 20 million olfactory sensory neurons
• Dendrites of these neurons located in the olfactory epithelium (3cm2)
• Axons pass through the cribiform plate
• These neurons contain olfactory receptors capable of binding odorant molecules, and stimulating an
action potential
• The primary neuron synapses with the secondary neuron in the olfactory bulb
• The secondary neuron projects along the olfactory tract to the olfactory cortex (not through
thalamus)
• The olfactory bulb is linked to the amygdala and hippocampus (limbic system)
• As a result certain smells can elicit very strong memories and emotional responses

GUSTATORY SYSTEM (TASTE)

• 5 tastes - sweet (sugars: sucrose etc), sour (H*), salty (alkali metal ions: Nat etc), bitter
(denatonium!!), and umami (flavour enhancer; receptors respond to glutamate/ MSG)
• Within the papillae, there are lots of taste buds, consisting of taste cells that stimulate primary
gustatory neurons
• Taste cells are epithelial
• Only the apical tips project through a taste pore into the mouth
• Salty" and "sour" cells synapse with primary gustatory neurons (H+ and Na+ enter through ion
channels, cause depolarization, and release of serotonin
• Bitter", "umami" and "sweet" tastes stimulate taste cells to release ATP which stimulates the
gustatory neuron
• Gustatory neuron axons enter the brain through cranial nerves 7, 9, and 10 to the medulla where
they synapse
• The information is then passed through the thalamus to the gustatory cortex

HEARING
• The ear can be divided into outer, middle and inner sections
• The inner ear is responsible for vestibular functions (equilibrium)
• The rest is for hearing
• The inner ear consists of 2 major structures
- cochlea (sensory receptors for hearing)
- vestibular apparatus (with semicircular canals) - equilibrium/balance
• Sound is the interpretation of the frequency, amplitude, and duration of sound waves that reach our
ears
• Does a tree that falls in a forest make a noise if no one is there?!?
1. Sound waves
2. Converted to mechanical vibrations (ear drum)
3. Converted to fluid waves in the cochlea
4. Fluid waves activate hair cells (sensory receptor cells)
5. Hair cells release neurotransmitter onto the primary auditory neuron
6. Neurotransmitter triggers action potential in primary auditory neuron
• Sound waves are directed down the ear canal until they hit the tympanic membrane
• The membrane vibrates causing the malleus, then incus, then stapes to vibrate
• As the stapes vibrates, it pushes and pulls on a thin piece of tissue (the oval window)
• The oval window vibration result in fluid waves in the liquid filled cochlea
• As the waves move through the cochlea they bend sensory hair cells in the duct
• Movement of hair cells opens ion channels creating electrical signals
• This results in neurotransmitter release and stimulation.
• This neurotransmitter release initiates action potentials in primary auditory neurons (cochlear
nerves)
• The auditory nerve is a branch of cranial nerve 8
• The neurons run to the cochlear nuclei in the medulla
• The secondary sensory neurons then project to the pons
• The pathway then runs through the thalamus to the auditory cortex

EQUILIBRIUM
• Equilibrium is maintained via hairs cells lining the vestibular apparatus of the inner ear
• These hair cells function just like the cochlear hair cells, but rather than sound waves it is gravity
and acceleration that move the cilia
• Activation of vestibular hair cells results in neurotransmitter release and a resulting action potential
on the primary sensory neurons of the vestibular nerve (a branch of cranial nerve 8)
• These neurons can synapse in the medulla or run straight to the cerebellum (equilibrium processing
centre) through the thalamus

VISION
• The eye focuses light through a lens and an aperture (pupil) onto a light sensitive surface (the retina)
• The photoreceptors on the retina convert light energy to electrical energy
• Neural pathways transmit the signal from the retina to the brain
• Photoreceptors of the retina convert light to electrical energy
• There are different types of neuron in the retina
- photoreceptors (rods and cones)
- bipolar neurons
- ganglion neurons
• Phototransduction is the process by which light is converted to electrical energy.
•
• Light sensitive pigment is bound to the discs in the rod and cone cells
• This visual pigment in rods is called rhodopsin
• There are 3 visual pigments in cone cells that are related to rhodopsin
• All of these pigments are stimulated by different wavelengths (colours) of visible light
• Rhodopsin - visual pigment made from a protein called opsin, and a retinal (vitamin A derivative)
• In the dark rhodopsin is inactive, and the rod/cones cells constantly release the neurotransmitter
glutamate
• Activation of rhodopsin by a photon of light releases retinal, and results in inhibition of glutamate
release
• This glutamate is normally released onto bipolar cells
• Bipolar cells can be "light-on" or "light-off"
• Light-on bipolar cells are inhibited by glutamate release in the dark
• Light-off bipolar cells are excited by glutamate release in the dark
• Bipolar cells then make inhibitory or excitatory synapses with ganglion cells
• The on- or off-state of bipolar cells allows for contrast, colour and sharpness
• The activated ganglion cell bodies transmit the signal along axons through the optic nerve, then
synapse in the thalamus before the signal is finally transmitted to the visual cortex