Eye movements that are automatic responses to stimuli
Reflex eye movements?.??.?.?
Repeatable: the same stimulus always evokes the same response (not influenced by internal states)
Mediated by spinal circuits (circuits located in the spinal cord)
Mediated by reflex arcs: mechanisms are basically chains of neurons (pathways) between sensory receptors and muscles
Reflex eye movements will be used to address the following three misconceptions regarding the nature of reflexes
Why move the eyes?
To keep the images on the retinas relatively still
To position the images of things of interest onto the foveas (the most sensitive regions of the retinas)
If retinal images move on the retina, it is impossible to see clearly => it is vital to keep the images steady
When we stand or sit still, our heads are typically fairly still as well
When we walk or run or drive on a bumpy road, our heads move up and down (bob), side to side (roll and weave) and sometimes forwards and backwards (nod): a lot of this movement cannot be prevented
If the eyes jiggle around with an angular motion of 1° amplitude it is impossible to see clearly
Vestibular-ocular reflex (VOR)
A reflex that involves the vestibular organs, keeping the eyes fixated on an object of interest as the head moves
Without the VOR, every time the head jiggles, the eyes would jiggle and clear vision would be impossible
Just living jiggles your head – breathing jiggles it, blood pulsing through the arteries jiggles it
If the vestibular organs are lost, balance is quickly recovered (not as good as before, but plenty good enough for everyday living)
The VOR keeps your eyes fixated on an object of interest as your head moves
The vestibulo-ocular reflex
Eyes counter-rotate so that they remain pointing in the same direction
Operates in all three dimensions: horizontal, vertical and torsional components
The VOR is mediated by circuits which do not involve the spinal cord at all
We also have reflexes that keep our heads stable – meaning that the VOR doesn't have to do all the work – the vestibulocollic and cervicocollic reflexes
Most invertebrates and some amphibians cannot make eye movements, birds have only a very limited ability to do so
Types of eye movement
Saccadic eye movements (very rapid shifts or jumps from one position in the orbit to another)
Slow, smooth movements that gradually shift an eye's position in its orbit
Saccadic movements
Shifts in the direction of gaze from one thing to another, made using rapid eye movements
Conjugate eye movements
Eyes move together in the same direction, through the same angle
Disconjugate eye movements: eyes move together in opposite directions
Horizontal saccades
Simultaneous contraction of left eye lateral rectus + right eye medial rectus
Horizontal saccades
To make a leftward saccade, activating signals are sent to the left abducens nucleus, which excites the motor neurons and also interneurons that carry the signal to the right oculomotor nucleus
Muscles involved in horizontal saccades
Medial and lateral recti
Left saccade: simultaneous contraction of left eye lateral rectus + right eye medial rectus
Horizontal saccades
1. Excitatory signals sent to left abducens
2. Excite motor neurons and interneurons that carry signal to right oculomotor nucleus
Contraction of muscles on left sides is not sufficient, muscles on right sides need to relax
How right side muscles relax
By shutting off (inhibiting) excitation of these muscles
Paramedian pontine reticular formation (PPRF)
Generates a burst of excitation that causes the eyes to move rapidly to a new position (the saccadic movement itself)
Nucleus prepositus hypoglossi (NPH)
Contains loops that feedback the output back into the nucleus, so it cannot be described in terms of chains (arcs)
Pathway for reflex horizontal saccades
1. Retina(s)
2. Superior colliculi
3. PPRF
Pathway for voluntary horizontal saccades
1. Frontal cortex
2. Superior colliculi
3. PPRF
When making a voluntary saccade
You often also turn the head
Turning the head
Elicits the VOR response that tries to keep the eyes looking in the same direction
Vestibular-ocular reflex (VOR)
Reflex response that tries to keep the eyes looking in the same direction when the head turns
To look in a new direction, you need to turn the VOR off when making a voluntary head turn
Whether a VOR response occurs depends on your intent
6 Common Misconceptions about reflex behaviour
Simple responses
Stereotyped responses
Mediated by spinal circuits
Mediated by reflex arcs
Repeatable
Not acquired or modified by learning and experience
Questions on saccade circuits should focus on what the circuit does and what the different parts do, not on the names of nuclei
If the pathway transmitting signals to the left lateral rectus is severed
The eye cannot be pulled to the left
If the nucleus paragiganto-cellularis is removed
The inhibition on the muscles moving the eye to the right is removed, opposing the leftward movement