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The Association Areas
Motor, sensory, and language areas account for only about half of the total cerebral cortex
Remaining areas are called association areas:
Prefrontal association cortex
Parietal–temporal–occipital association cortex
Limbic association cortex
Sensory input →
Primary sensory areas (somatosensory, 1o visual, 1o auditory cortices) →
Higher sensory areas →
Association areas →
Higher motor areas →
Primary motor areas →
Motor output
Sensory Input
Relayed from afferent neuronal receptors
Primary Sensory Areas (Somatosensory, 1o Visual, 1o Auditory Cortices)
Initial cortical processing of specific sensory input
Higher Sensory Areas
Further elaboration and processing of specific sensory input
Association Areas
Integration, storage, and use of diverse sensory input for planning of purposeful action
Higher Motor Areas
Programming of sequences of movement in context of diverse information provided
Primary Motor Cortex
Commanding of efferent motor neurons to initiate voluntary movement
Motor Output
Relayed through efferent motor neurons to appropriate skeletal muscles, which carry out desired action
Basal Ganglia
Consist of several nuclei located deep within the cerebral white matter
Primary functions:
Associated with a variety of functions, including motor control, cognition, emotions, and learning
inhibiting muscle tone throughout the body – balance of excitatory and inhibitory inputs to neurons that innervate skeletal muscles
selecting, maintaining purposeful motor activity while suppressing unwanted patterns of movement
coordinates slow, sustained contractions, especially those related to posture and support
Parkinson’s Disease (PD) - Diseases of the Basal Ganglia
Caused by the degeneration of dopaminergic neurons in Substantia Nigra in the midbrain which synapse onto neurons in the basal ganglia:
This circuit is important for smooth movement.
The loss of dopamine results in the characteristic features of PD:
Increased muscle tone, rigidity
Involuntary, useless movements at rest, tremors
Difficulty in initiating and carrying out movement
Huntington’s Disease (HD) - Diseases of the Basal Ganglia
Caused by degeneration of the caudate nucleus
This loss causes some motor circuits to become overactive resulting in the characteristic features of HD:
Chorea - rapid, uncontrolled jerky movements
Memory problems
Thalamus
Deep in the brain near the basal ganglia:
Serves as “relay station” and synaptic integrating center for sensory input
Helps direct attention to stimuli of interest
Capable of crude awareness of sensations but cannot distinguish their location or intensity
Hypothalamus
The integrating centre for homeostatic function
Brain area most involved in directly regulating internal environment
Controls body temperature and food intake
Controls thirst and urine output
Controls anterior pituitary hormone secretion
Produces posterior pituitary hormones
Five functions of the brain stem:
Majority of cranial nerves arise from areas in brain stem
Contains centers that control cardiovascular, respiratory, and digestive function
Regulates postural muscle reflexes
RAS (reticular activating system) controls the overall degree of cortical alertness
Plays a role in the sleep–wake cycle
Cranial Nerves
Optic (vision)
Olfactory (smell)
Oculomotor & trochlear (visual reflexes)
Facial (taste, face muscles)
Vestibulocochlear (hearing and equilibrium)
Glossopharyngeal (taste)
Cerebellum
Important in integration of motor output and sensory perception
Important in balance and planning of movement
Vestibulocerebellum
balance and eye movement
Spinocerebellum
enhances muscle tone and coordinates skilled movements
Cerebrocerebellum
plans and initiates voluntary activity and stores procedural memories
Reticular Activating System (RAS)
Behavioural state system - diffuse modulatory system
Neurons originate in the RAS and project to various areas in the brain
Influences attention, motivation, wakefulness, memory, motor control, mood and metabolic homeostasis
Controls levels of consciousness and sleep-wake cycles (keeps the “conscious brain” awake)
Controls the overall degree of cortical alertness
General anaesthetics depress synaptic transmission in the reticular formation
Blocking ascending pathways between the reticular formation and the cerebral cortex creates a state of unconsciousness.
The Limbic System
Surrounds the brain stem and is not a separate structure
It is an interconnected ring of forebrain structures:
Includes portions of cerebral lobes, the basal nuclei, the thalamus, and the hypothalamus
Utilizes the neurotransmitters norepinephrine, dopamine, and serotonin
Depression is associated with defects in limbic system neurotransmitters
Complex interacting network is associated with emotions, basic survival, sociosexual behavior, motivation and learning
Limbic system functions:
Emotion: subjective feelings and moods and the physical responses associated with these feelings (amygdala)
Basic behavioural patterns: aimed at survival and perpetuation of the species
Motivation: directing behaviour toward goals
Learning: acquiring knowledge or skills as a result of experience and/or instruction
Memory: storage of acquired knowledge for later use
Medial Prefrontal Cortex
Executive function: decision making, control of emotion & impulses
Cingulate Cortex
Motivation, drive, mood: decreased activity correlates with depression
Medial Temporal Lobes
Episodic memory formation of recent event sequences
Subcortical Structures of Limbic System
Hippocampus (within medial temporal lobe)
Episodic memory formation, context & location
Hypothalamus
Homeostasis & basic drives: food, water, sex, aggression
Amygdala
Involved in fear & affective learning.
Activation causes anxiety, (ablation eliminates fear recognition & learning)
Learning
Is the acquisition of knowledge or skills as a consequence of experience, instruction, or both
It is widely believed that rewards and punishments are integral parts of many types of learning
Memory
The storage of acquired knowledge for later recall:
Short-term: seconds to hours
Long-term: days to years
Working memory: temporarily holds and interrelates various pieces of information relevant to a current mental task
Memory Trace
A neural change responsible for retention or storage of knowledge. These traces are present across multiple regions of the brain (sites of original auditory, visual, etc. experience)
Declarative memory (“what” memories of people, places etc):
Important for remembering facts and events – involves Hippocampus
Procedural memory (“how to” memories):
The learning of new motor skills – involves Cerebellum
Consolidation of short-term declarative memory into long term memory:
The function of the medial temporal lobe (hippocampus and adjacent areas of temporal lobe), occurs largely during sleep
Requires gene activation leading to protein synthesis and synaptic changes
Short-term Memory
Involves transient changes in synaptic activity.
Habituation: decreased responsiveness to a repetitive and indifferent stimulus
Sensitization: increased responsiveness to mild stimuli following a noxious stimuli
Both involve changes in ion channels and currents
Long-term Memory
Involves formation of new, permanent synaptic connections.
Activation of specific genes that control synthesis of proteins needed for lasting structural or functional changes in pre- or postsynaptic membranes
Long term potentiation (LTP) is important for initial storage into long term memory
Working Memory
Concept that working memory temporarily holds and interrelates various pieces of information that are relevant to a current mental task (7 +/- 2 objects)
Hold and process data for immediate use - can include newly acquired and previously stored knowledge that is transiently accessed
Critical for ability to reason, plan and make judgments
By comparing and manipulating new and old information within your working memory you can, for example, find your way home, carry on a conversation etc
Takes place within prefrontal cortex
Alzheimer’s Disease
Characterized by short-term memory loss in early stages
Followed eventually by loss of long-term memory - eg, recognition of family members
Confusion, disorientation, personality changes
Loss of ability to read, write, calculate
Language ability and speech also impaired
Brain alterations - neurofibrillary tangles, amyloid plaques
Characteristic loss of cholinergic neurons in the basal forebrain - cells that project to hippocampus
Neuronal loss in hippocampus & temporal lobes → cortex (cholinergic neurons – neurons that use acetylcholine as their neurotransmitter.)
Language
Localized in one hemisphere – left hemisphere (95% of right-handers, 60-70% of left-handers)
Broca’s and Wernicke’s area
Broca’s Area
Speaking ability:
Damage results in an inability to send the proper commands to the motor cortex to form the words.
Wernicke’s Area
Language comprehension:
Damage results in an inability to attach meaning to words or choose the appropriate words
Language Disorders
Aphasia – is a defect in language processing caused by dysfunction of the dominant cerebral hemisphere (i.e. stroke)(Broca’s aphasia, Wernicke’s aphasia)
Speech impediments – defects in the mechanical aspect of speech
Dyslexia – difficulty in learning to read because of inappropriate interpretation of words. Due to developmental abnormalities in connections between the visual and language areas of the cortex. (independent of intelligence)
Cortical Pathway for Speaking a Written Word or Naming a Visual Object
To speak about something seen, the brain transfers the visual information from the primary visual cortex to the angular gyrus of the parietal-temporal-occipital association cortex, which integrates inputs such as sight, sound, and touch
To speak about something heard, the brain transfers the auditory information from the primary auditory cortex to the angular gyrus
The information is transferred to Wernicke’s area, where the choice and sequence of words to be spoken are formulated
This language command is then transmitted to Broca’s area, which translates the message into a programmed sound pattern
This sound program is conveyed to the precise areas of the primary motor cortex that activate the appropriate facial and tongue muscles for causing the desired words to be spoken
Electrical activity patterns define arousal states:
Electrical activity in neurons can be measured by EEGs (Electroencephalograms)
Surface electrodes placed on the scalp can detect depolarizations of the cortical neurons in the region under the electrode
Represents Excitatory Post Synaptic Potentials (EPSPs) and Inhibitory Post Synaptic Potentials (IPSPs) in cell bodies of the cortical layers
Patterns of activity can be detected
In awake states - neurons are firing but not always in a coordinated fashion
This desynchronization may be a result of ascending signals coming from the Reticular Activating System (RAS)
What is the value of the EEG?
Often used as a clinical tool in diagnosis of cerebral dysfunction
Epilepsy – shows distinctively abnormal traces
Seizures occur when collections of neurons undergo synchronous action potentials that produce stereotypical, involuntary spasms and alterations in behaviour
Neuronal excitability coupled with compromised inhibitory activity or prolonged activation of excitatory transmitters
Legal determination of death
Electrocerebral silence – flat EEG
Needs to be coupled with other stringent criteria
Also used to distinguish various stages of sleep
Alpha & Beta Waves
Waking brain activity
The alpha rhythm in the EEG is 8-13 Hz: index of cortical inactivity
Present in adult who is awake but relaxed with eyes closed
Amplitude is negatively correlated with cortical activity
Beta rhythm in the EEG is 13-30 Hz: decreased synchronization with cortical activity
In individuals who are alert and attentive to external stimuli or exert specific mental effort
Maximum Alertness
depends on sensory input that stimulates the RAS and subsequently the activity levels of the CNS as a whole
Sleep-wake Cycle
normal cyclic variation in awareness
brain’s overall activity is not reduced during sleep (sleep is an active process)
Two types of sleep characterized by different EEG patterns and different behaviours:
Slow-wave sleep – delta-wave; non-rapid eye movement, NREM
Paradoxical sleep – rapid eye movement, REM
Stages of Sleep
Slow wave sleep occurs in 4 stages – each displaying a lower frequency, but higher amplitude EEG wave
At onset of sleep, you move through the stages, from light sleep (1) to deep sleep (4) in a 35-45 min period
Then it reverses through the same stages in same amount of time
A short period of REM sleep occurs at the end of each slow-wave sleep cycle
Paradoxical sleep – because the EEG wave form resembles that of an awake alert individual
90 minutes/cycle
Time in REM stage increases whereas time in stages 3 and 4 decreases over the cycles
Slow-Wave Sleep
Displays slow waves
Considerable muscle tone; frequent shifting
Minor reductions in heart rate, respiratory rate, and blood pressure
Dreaming is rare
Sleeper is easily awakened
80% of sleeping time
Has 4 stages
Paradoxical Sleep
Similar to the EEG of an alert, awake person
Abrupt inhibition of muscle tone; no movement
Heart rate, respiratory rate, and blood pressure are irregular
Dreaming is common
Sleeper is hard to arouse but apt to wake up spontaneously
Rapid eye movements occur
The best-supported evidence suggests that main functions of sleep include:
Conservation of energy, defense from predation
Restorative function for the brain and body: healing, growth, immune function
An important role in the cerebral changes that underlie consolidation of long-term memory in cortex