excitatory and inhibitory signals project to inhibitory spinal inter-neurons, which in turn project onto the motor neurons
Reflexes
no subject to volitional control, they are automatic in response to sensory stimuli, usually involve multiple neurons that form a reflex arc
Reflexes are...
For the most part, they do learn, they emerge without prior experience. It is possible however, to acquire new reflexes through learning
How are Reflexes Strengthened?
The strength of many reflexes can be adjusted through trial and error learning to compensate for the changes in the body or the external environment
Central Patten Generator
vegetative processes and bodily movements that are subject to sensory modulation but do not require sensory inputs to generate the rhythmic activity
Locomotor rhythms are created within the central nervous system independently of sensory inputs
Half Centered Oscillators:
consist of two neurons that reciprocally inhibit each other. Some CPGs contain half-center oscillators, but most contain more than two sets of inhibitory neurons as well as several types of excitatory interneurons
· Tonic excitation is required to keep the network going
CPG for walking in mammals
Can be modulated by external inputs
Modulation of CPG for walking
Entrained by leg movements when decerebrate cats are placed on a moving treadmill where they can adjust their speed based on that of the treadmill
Walking CPG
Can be modulated by descending input from the midbrain locomotor area
Stimulation of midbrain locomotor area
1. Walking CPG becomes active
2. With increasing stimulation intensity, its rhythm accelerates
Human spinal cords do contain CPG but this CPG is more difficult to activate in humans than in other vertebrates
Motor Programs
movements that we have to think consciously about like picking up a water bottle
Emotional or actional movements
Primary motor cortex activity codes for movement which means it does not react to it
Motor Cortex
Located in the posterior portion of the frontal lobe, anterior to the central sulcus
Primary motor cortex (M1)
Thin strip of tissue in the most posterior part of the Motor Cortex
Corticospinal neurons are most concentrated here
Most project to the intermediate zone and spinal cord interneurons
Corticobulbar projections
Project to the medulla where they target control respiration, urination, and other vegetative functions as well as regions
Premotor cortex
Most anteriorly in the Motor Cortex
Involved in muscle recruitment and body positioning (goal oriented actions)
Supplementary motor area
Most medially in the Motor Cortex
(33%) goal oriented that fire no matter how the joystick was held (monkeys)
(50%) were muscle contraction oriented which only fired when the joystick was held in the way that it was trained
Motor Cortex
Most voluntary movements, many unconscious and involuntary reflexes including the muscle stretch reflexes, involve long loop circuits through the motor cortex
Motor Cortex Lesions in Humans
Homunculus
A way to show that adjacent regions in the motor cortex tend to control adjacent body parts
Motor cortex
The head and hand can be evoked from disproportionately large regions (face and hand overrepresented)
Controls movements rather than muscles
Population code
Movements are encoded in the pattern of activity across a population of neurons
Neurons of M1 have broad tuning curves which is why population code is needed
Cortical movement control involves the concerted activity of many different neurons and individual neurons are activated during many different movements
Premotor cortex
Contains multiple subdivisions and its neurons exhibit remarkably complex activity patterns
Most neurons increase their firing rates shortly before and during arm, hand, lip, or tongue movements
Codes for the intention of others and goals of motor acts
Mirror neurons
Increase their firing rate when the organism observes the movement that they select for when they perform the movement themselves
Code for the intention of others
Constraint-induced movement theory
Requires patients who have lost the function of one arm to put their good arm in a sling for several hours at a time and use the impaired arm to perform various tasks
The idea is that forcing subjects to use their impaired arm will increase plasticity within the brain regions controlling that impaired arm
Cerebellar granule cells
Lie just beneath the Purkinje cell layer, 1st major source of input to Purkinje cells
Axons are called parallel fibers because they split into left and right branches that extend in straight lines parallel to one another and to the cerebellar surface
Purkinje cells respond to granule cell activity only when thousands of those cells fire simultaneously. Even then, granule cell activity elicits only simple spikes in the Purkinje cells
Inferior Olive
Lies in the medulla and is a second major source of input to Purkinje cells
Each neuron sends an axon to just a single Purkinje cell but forms thousands of synapses in that cell (climbing fibers)
Whenever an inferior olive neuron fires an action potential the Purkinje cell to which that neuron projects also fires an unusually large and long action potential called a complex spike
All cerebellar Purkinje cells use GABA as their main neurotransmitter and therefore inhibit their target cells
Cerebellum functions using adaptive feedforward control
Adjusted through learning, feedforward means before learning adjusts the commands so that the target state is reached with only minimal error
Adaptive Purkinje cells adapt through trial-and-error learning allowing a controller to predict the error that would occur if there had been no previous learning and to correct for it before the error arises
Cerebellar circuits
Help organisms discriminate sensations that are caused by their own movements from sensations that are caused by external factors
Frontostriatal system
Overarching function includes the selection of goals, of actions to attain those goals, and of specific movements to perform the actions
Frontostriatal system
Includes parts of the neocortex as well as several subcortical structures which are collectively referred to as the basil ganglia
Dorsal Striatum
Involved in learning and control of stereotyped movements
Receives inputs from the posterior regions of the frontal lobe, including the motor and premotor cortices
Outputs to the dorsal pallidum which projects to the thalamus
Ventral Striatum
Generally involved in generating cravings for food, water, sex, and drugs
Receives inputs from the orbital prefrontal cortex which occupies the inferior surface of the frontal love
Outputs to the ventral pallidum which projects to the thalamus
Thalamus
Areas project to the frontal lobe: the parts receiving input from the ventral pallidum project to the anterior prefrontal cortex and the parts receiving input from the dorsal pallidum project to more posterior portions of the frontal lobe
Ventral Membrane
Made up of the substantia nigra, compact division (SNc) and the ventral tegmental area (VTA)
Direct pathway through the striatum
Contains medium spiny neurons that project directly to the pallidum