Clinical Neuroscience Week 9 - Diencephalon, Limbic System, Basal Ganglia

• The diencephalon includes the thalamus, hypothalamus, subthalamus, and epithalamus.

Thalamus

• The thalamus is the largest part of the diencephalon, comprising 80% of its mass.
• It filters and integrates sensory information before sending it to the cortex, controlling information flow and helping the brain prioritize attention.
• It serves as a relay station for incoming motor and sensory information, including hearing, taste, sight, and touch (but not smell).
• All bodily sensory information is processed through the thalamus before reaching the cerebral cortex for interpretation.
• Blood supply comes from the posterior cerebral artery, posterior communicating artery, and basilar communicating artery.
• The thalamus consists of two egg-shaped structures, one in each hemisphere, linked across the third ventricle by the interthalamic adhesion.
• It integrates sensory information, regulates motor activity and consciousness, and plays a role in sleep, wakefulness, learning, memory, and perception.
• Multiple nuclei within the thalamus receive, process, and transmit information to related areas in the cerebral cortex.
• Thalamic nuclei are primarily made of excitatory neurons projecting to the cortex in an organized way.
• The thalamus comprises of various nuclei; each has its unique role ranging from relaying sensory and motor signals to regulating consciousness and alertness.
• The Intramedullary lamina, comprised of white matter, is y-shaped and compartmentalizes the three large cell groups.

Thalamic Nuclei Groups

• Anterior nuclear group
• Medial dorsal nuclear group
• Lateral nuclear group
• The Reticular nucleus, is a sheet that covers the thalamus and influences the other nuclei within.
• Four functional groups of thalamic nuclei: relay, association, nonspecific, and reticular.

Functional Groups - Relay Nuclei

• Anterior Nucleus: Receives input from the mammillothalamic tract, cingulate cortex, and hippocampus, and projects to the prefrontal and cingulate cortex.
• Ventral Anterior (VA): Receives input from the substantia nigra (pars reticulata) and globus pallidus internus, and projects to the prefrontal cortex and frontal eye field.
• Ventral Lateral (VL - anterior part): Receives input from the globus pallidus (internal segment) and cerebellum, and projects to the supplementary motor area.
• Ventral Lateral (VL - posterior part): Receives input from deep cerebellar nuclei, and projects to the motor/pre-motor association cortex.
• Ventral Posterolateral (VPL): Receives input from the medial lemniscus/spinothalamic tract (body) and vestibular pathways, and projects to the somatosensory and vestibular cortex.
• Ventral Posteromedial (VPM): Receives input from the medial lemniscus/spinothalamic tract (head), and projects to the somatosensory cortex (parietal association cortex).
• Medial Geniculate: Receives input from the inferior colliculus, and projects to the primary auditory cortex.
• Lateral Geniculate: Receives input from the optic tract, and projects to the primary visual cortex.

Functional Groups - Association Nuclei

• Lateral Dorsal (LD): Receives input from the pretectum, cingulate cortex, hippocampus, and other thalamic nuclei, and projects to the cingulate cortex.
• Mediodorsal (MD): Receives input from the prefrontal cortex, olfactory areas, amygdala, striatum, superior colliculus and other thalamic nuclei, and projects to the prefrontal and cingulate cortex and frontal eye fields.
• Lateral Posterior (LP)/Pulvinar Complex (PUL): Receives input from other thalamic nuclei/nuclei of the superior colliculus and projects to the Parietal (LP)/ occipital cortex (PUL).

Functional Groups - Non-Specific Nuclei

• Intralaminar: Receives input from the spinothalamic tract, reticular formation, neuromodulatory system, and amygdala, and projects to widespread areas of the cortex and basal ganglia.
• Midline Nuclei: Receives input from the reticular formation, neuromodulatory system, and amygdala, and projects to the cingulate cortex, hippocampus, and corpus striatum.

Functional Groups - Reticular Nuclei

• Reticular Nucleus (RN): Receives input from the cortex, underlying thalamic nuclei, and reticular formation and projects to underlying thalamic nuclei.

Thalamic Connections

• All thalamic nuclei, except the reticular nucleus, have reciprocal excitatory connections with the cerebral cortex.
• Sensory inputs are excitatory, and most neurons in the thalamic nuclei are glutamatergic (excitatory projection neurons).
• The reticular nucleus inhibits activity of other thalamic nuclei and does not directly excite the cerebral cortex.
• It acts as a gatekeeper, regulating information flow from the thalamus via inhibitory transmitters (GABAergic interneurons).
• Relay nuclei transmit specific sensory information from different areas of the cortex and project it back to association areas.
• Lateral geniculate is for vision.
• Medial geniculate is for auditory.
• Ventral posterior lateral nuclei is for somatosensory.
• Non-specific nuclei send diffuse projections to large regions of the cortex and are involved in arousal and attention rather than relaying specific sensory data.

Thalamic Nuclei - Ventral Lateral (VL) Nucleus

• The central hub for motor control.
• Receives signals from the cerebellum and basal ganglia.
• Cerebellar input provides information about movement errors.
• Basal ganglia contribute to movement initiation and selection.
• Projects to the primary motor cortex (frontal lobe) along with premotor and supplementary motor cortices.
• Crucial for coordinating and planning movements and learning new movements.
• Anterior Part (VLa): Proximal muscle control.
• Posterior Part (VLp): Distal muscle control.
• Lesions in this area can result in ataxia or tremors.
• It's a target location for deep brain stimulation in patients with Parkinson’s disease.

Thalamic Nuclei - Ventral Posterolateral (VPL) vs. Ventral Posteromedial (VPM) Nuclei

• VPL receives somatosensory input from the body (trunk + limbs) via the dorsal column medial lemniscus (DCML) and spinothalamic tract.
• It projects to the primary somatosensory cortex.
• VPM receives somatosensory input from the head (face + head) via the trigeminal nerve (CN V).
• It projects to the primary somatosensory cortex.
• Damage to the area or the VPM can lead to impaired facial sensation on the contralateral side of the body, impacting touch, pain, and temperature sensations from the face.

Thalamic Nuclei - Medial Geniculate (MGN) vs. Lateral Geniculate (LGN) Nuclei

• MGN serves as the primary auditory relay station, receiving signals from the inferior colliculus and projecting to the auditory cortex (temporal lobe).
• LGN is the primary visual relay station, receiving input from the optic tract and projecting to the primary visual cortex (occipital lobe).

Thalamic Nuclei - Thalamic Reticular Nucleus (TRN)

• It acts as a Gatekeeper.
• It inhibits activity of other thalamic nuclei.
• It regulates the flow of sensory information to the cortex.
• It plays a key role in attention, arousal, and sensory processing by controlling which sensory signals reach the conscious level.

Thalamic Nucleus & Function

• Anterior Nucleus: Connected to the hippocampus, involved in memory.
• Dorsomedial Nucleus: Emotional behavior and memory.
• Ventral Anterior Nucleus (VA): Motor functions.
• Ventrolateral Nucleus (VL): Motor functions.
• Ventral Posterolateral Nucleus (VPL): Relay nuclei to somatosensory cortex, processes somatosensory information.
• Ventral Posteromedial Nucleus (VPM): Relay nuclei to somatosensory cortex, processes somatosensory information.
• Lateral Posterior (Posterolateral) Nucleus: Integrating sensory input and associating it with cognitive functions.
• Medial Geniculate Nucleus: Relay nuclei to auditory cortex; processes auditory info.
• Lateral Geniculate Nucleus: Relay nuclei to visual cortex; processes visual info.
• Pulvinar Nucleus: Processes visual stimuli.
• Reticular Nucleus: Sheet/outer cover of the thalamus; influences activity of other thalamic nuclei.

Clinical Implications - Thalamic Stroke

• Weakness
• Vision changes
• Speech difficulties
• Balance problems
• Pain
• Sensory changes
• Cognitive changes
• Mood changes
• Sleep disturbances
• Blood supply typically arises from small perforating branches from the posterior cerebral artery, posterior communicating artery, and basal communicating artery.

Clinical Implications - Thalamic Pain Syndrome

• Chronic pain results from injury to the thalamus.
• Spontaneous pain occurs on the contralateral side of the thalamic lesion.
• Treatment is difficult and often ineffective.
• Characterized by burning, constrictive pain, allodynia, hyperalgesia, paresthesia, and temperature changes.

Hypothalamus

• A structure deep within the brain, below the thalamus and above the pituitary gland
• Situated above the brain stem and about the size of an almond; part of the endocrine system.
• Serves as the main link between the endocrine and nervous systems coordinating the autonomic nervous system and pituitary gland activity.
• Maintains body homeostasis by directly influencing the autonomic nervous system and managing hormones.
• Regulates heart rate, blood pressure, and body temperature.
• Releases hormones that direct other hormones or glands to manage bodily functions like sleep, mood, muscle and bone growth, and sex drive.
• Helps to correct for any imbalances.
• Produces dopamine (neurotransmitter released during pleasurable activities, essential for smooth movements) and somatostatin (regulates body functions).

Hypothalamus + Pituitary Gland Functions

• Anterior Pituitary: Hypothalamus-releasing hormones like growth hormone-releasing hormone (GHRH), gonadotropin-releasing hormone (GnRH), corticotropin-releasing hormone (CRH), thyrotropin-releasing hormone (TRH), and dopamine (inhibition).
• Posterior Pituitary: Oxytocin and vasopressin.

Clinical Implications - Hypothalamic Dysfunction

• Hypothalamic-pituitary disorders are: Hypopituitarism, diabetes insipidus, Prader-Willi syndrome, Kallmann syndrome, acromegaly and pituitary gigantism, syndromes of inappropriate antidiuretic hormone, central hypothyroidism, functional hypothalamic amenorrhea, and hyperprolactinemia.
• Signs and Symptoms: HTN or hypotension, water retention or dehydration, weight loss or weight gain, infertility, poor bone health, delayed puberty, muscle loss and weakness, body temperature fluctuations, trouble sleeping (insomnia), frequency need to urinate.
• Dysfunction caused by head injuries, brain infections, potential brain tumors, brain surgery, radiation, birth defects, and inflammatory diseases like multiple sclerosis and neurosarcoidosis.

Subthalamus

• Small region located below the thalamus.
• Plays a crucial role in regulating movement by modulating the activity of the basal ganglia.
• Helps fine-tune and coordinate voluntary muscle movements, specifically initiation and smoothness of movements.
• The main structure is the subthalamic nucleus.
• Dysfunction results in motor dysfunction seen in Parkinson's.
• It is also an area for deep brain stimulation used to treat Parkinson's.
• Hemiballismus presents with sudden violent involuntary movements on one side of the body.

Epithalamus

• Most dorsal, smallest, and oldest part of the diencephalon.
• Lies posterior to the thalamus and forms a posterior part of the roof of the third ventricle.
• Connects the limbic system.
• Three components:
  • Pineal Gland (body): Part of the endocrine system that secretes melatonin, which helps control the circadian sleep cycle and wakefulness cycles.
  • Habenular Nuclei: Receives afferents from the forebrain and projects them to the midbrain; part of the limbic system that regulates emotions and behaviors, and has links to cognitive, emotional, and sensory processing.
  • Stria Medullaris: A bundle of nerve fibers that connects the habenular nuclei with other brain regions (specifically within the limbic system).

Limbic System

• Known as the emotional brain or emotional nervous system.
• Initiates need-directed motor activity for survival based on experience.
• Helps manage emotional life and form memories, influencing certain behaviors.
• Plays a vital role in motivation, learning, and spatial memory.
• Primary components include the amygdala, hippocampus, hypothalamus, thalamus, and cingulate gyrus.
  • Amygdala: Emotional center that processes fear, anger, and pleasure, helps respond to emotional stimuli and forms emotional memories, evaluates the emotional significance of events, and triggers appropriate responses.
  • Hippocampus: Essential for forming new memories, connecting emotions and senses to memories, consolidating short-term memories to long-term storage, involved in spatial navigation.
  • Hypothalamus: Regulates bodily functions (hunger, sleep, body temperature) through hormone regulation, controls the release of hormones from the pituitary gland (influences emotions and stress response), and is the primary regulator of autonomic and endocrine functions.
  • Thalamus: Acts as a relay station, directing sensory and motor signals to the cerebral cortex and plays a role in regulating consciousness, sleep, and alertness.
  • Cingulate Gyrus: Helps interpret situations and guide appropriate behaviors for social interactions; plays a role in empathy and social interactions.

Limbic Lobe

• Usually includes the orbital frontal cortex, cingulate gyrus, parahippocampal gyrus, and uncus.
• Close connections inlcude temporal pole, the hypothalamus and reticular formation, and nucleus accumbens.
• The nucleus accumbens (in the basal ganglia) is involved in processing rewarding experiences, motivation, and translating emotional responses into motor activity.
• It is a key component of the brain’s pleasure center.
• Damage to the limbic system can result in anxiety, addiction, obsessive-compulsive disorder, depression, and post-traumatic stress disorder.

Limbic Network Function - MOVE

• M = Memory & Motivation: Drive; memory, attention and retrieval, declarative learning; motivation, desire to learn, try, or benefit from the external environment.
• O = Olfaction: Only sensory system that does not pass through the thalamus as a second-order synapse in the sensory pathway before reaching the cerebral cortex.
• V = Visceral: Drive, thirst, hunger, and temperature regulation; endocrine functions; sympathetic and parasympathetic reactions; hypothalamic regulation over the autoimmune system and peripheral autonomic nervous system (ANS) responses that reflect limbic function.
• E = Emotion: Feelings and attitude; self-concept and worth; emotional body image; tonal responses of motor system affected by limbic descending pathways; attitude, social skills, opinions.

Basal Ganglia

• Consists of the striatum (caudate nucleus, putamen, and nucleus accumbens), globus pallidus (external and internal segments), subthalamic nucleus, and substantia nigra (pars reticulata and pars compacta).
• Plays a crucial role in movement and learning new motor skills by regulating voluntary movement and filtering out unnecessary motor signals.
• Structures form a hub of interconnected pathways that integrate information from various brain areas to control movement, cognition, and emotion.
• Acts as a gatekeeper to initiate movement.
• Plays a role in cognitive functioning, decision-making, reward processing, and learning through neural circuits that project to various brain areas.
• Information flows from the basal ganglia in complex loops, with input primarily from the cerebral cortex and output back to the cortex via the thalamus.
• The basal ganglia are not a primary component of the limbic system, but they have an influence on cognitive functioning in terms of decision making, reward processing, motivation, and learning.
• They have connections with the limbic system and are best known for their role in motor control but are critical for emotion and behavioral inhibition.
• Basal Ganglia and Neighboring Structures
• Located deep within the brain, playing a significant role in motor control, learning, and various cognitive functions.

Basal Ganglia - Striatum

• Caudate Nucleus: C-shaped structure divided into a head, body, and tail (tail connects to the amygdala); receives input from various areas of the cortex and sends signals to the thalamus.
• Putamen: Receives inputs from various areas of the cortex including sensory information (pain and touch) and sends signals to other brain structures like the substantia nigra pars reticulata.
• Modulation of activity of motor neurons and involved in motor control, cognitions, and reward processing through reinforcement learning.
• The Putamen has touch on processing positive and negative emotions, regulating emotional responses.
• Nucleus Accumbens: Located near the front of the brain, below the anterior commissure. Is a key part of reward processing, motivation, and emotional regulation.
• Dysfunction can result in disorders like addiction and depression.
• It consists of a cluster of interconnected nuclei and is the largest structure of the subcortical Basal Ganglia.

Striatum Function

• Control movements, reward and decision making, learning, habit formation, and social interactions.
• Degeneration of striatal neurons can lead to movement disorders such as Parkinson’s and Huntington’s disease.

Basal Ganglia Inputs

• Two inputs to the striatum: cortex and substantia nigra.
  • Cortical projections release glutamate to the striatum, which sends excitatory signals to the striatum.
  • Substantia nigra projections release dopamine to the striatum, which can either be excitatory or inhibitory.
    • D1 expressed receptors are excitatory and D2 expressed receptors are inhibitory.
• About 90% of the striatum is GABAergic neurons, which are inhibitory projections neurons.

Basal Ganglia - Globus Pallidus

• Situated medial to the putamen.
• With the putamen, globus pallidus form the lentiform nucleus.
• The globus pallidus receives input from the striatum and sends inhibitory outputs to the thalamus.
  • Globus pallidus external is the intrinsic nucleus that relays information to the globus pallidus internal.
  • Globus pallidus internal is the output nucleus from the basal ganglia that exerts control over the thalamus.
• Palatal outputs are inhibitory to the subthalamic nucleus in the thalamus.
• It consists of an External (lateral) segment (GPe) and Internal (medial) segment (GPi).

GPe + GPi + Putamen = Lenticular (Lentiform) Nucleus

Basal Ganglia Outputs

• The primary output region of the basal ganglia is the internal segment of the globus pallidus.
• This region sends inhibitory GABAergic projections to the nuclei in the thalamus.
• The thalamus then projects back out to the cerebral cortex primarily to the motor areas.
• Outputs from the basal ganglia leave through the internal segment of the globus pallidus.
• Inhibitory projections release GABA onto the thalamus, which then sends glutamate projections to the cortex to communicate with the cerebral cortex.

Basal Ganglia - Substantia Nigra

• Two parts:
  • Pars Reticulata (SNr): Sends inhibitory signals to the GPi.
  • Pars Compacta (SNc): Produces dopamine.

Substantia Nigra Function

• The substantia nigra is essential for controlling body movements, particularly by producing dopamine, also has a role in the chemical signaling in the brain that affects learning, mood, judgment, decision, decision making and other processes.
• There are also two parts of each substantia nigra
  • Pars Reticulata hold a chemical called GABA inhibits activity in the brain cells and used to stop signals. Has an important job in regulating motor control. Receives input from the striatum and subthalamic nucleus.
• Pars Compacta cells produce serves mainly as a projections of the basal ganglia circuit in that it supplies the striatum with dopamine and dopamine modulates basal ganglia activity.
• The globus pallidus, internal and external, taper into the midbrain as a reticular part of the substantia nigra. Meaning its continuously active or firing at a sustained rate
• Substantia nigra (Pars Compacta) is tonically active. And tonically has functions as such: releasing dopamine which helps us to focus and retain information and contributes to feelings of well being and happiness.
• Substantia nigra are inhibitory gabergic neurons that project to the thalamus and brainstem nuclei and whose activity is modified by the direct and indirect pathways which favors activity in the direct pathway.

Substantia Nigra Degeneration

• Degeneration of neurons in the substantia nigra is a primary feature of Parkinson’s disease – leading to motor symptoms like tremors and rigidity, balance and coordination problems, muscle weakness, and spasms, shakiness and tremors, trouble focusing, thinking or problem solving and difficulty controlling eye movements that result in vision impairment. Degeneration also results in lower levels of dopamine which are associated with Parkinson’s disease.

Basal Ganglia - Subthalamic Nucleus

• The subthalamic nucleus composes the majority of the subthalamus.
• It is an oval or lens-shaped group of neurons.
• Thought to play a role in motor integration and body movement accuracy and has a touch on the limbic associated functions and is important for inhibition of unwanted movements.
• It sends excitatory signals to the internal globus pallidus, which in turn inhibits the thalamus.
• Subthalamic nuclei are the only glutamatergic neurons in the basal ganglia and is excitatory to the globus pallidus internal and is also part of the indirect pathway that controls motor function.

Clinical Implications – Damage to Subthalamic Nucleus

• Lesions can cause decreased stimulation of the globus pallidus internus within the indirect pathway, with consequent disinhibition of the thalamus with the resulting hyperkinetic movements.
• Can lead to movement disorders such as hemibolism, which is uncontrolled violent movement on one side of the body.

Basal Ganglia Connections

• Cerebral Cortex: Basal ganglia (BG) receives inputs from the cerebral cortex (areas involved in motor control, cognition, and sensory processing), inputs directed to the striatum (caudate nucleus + putamen), and BG processes/outputs back out to the cortex via the thalamus.
• Thalamus: Serves as a “Relay station” / “air traffic control”, transmits processed signals from the BG and projects them back out to the cortex “Feedback Loop”.
• Cerebellum: Works alongside the BG to ensure fluid and accurate motor execution. Cerebellum is involved in timing and precision of movements, BG focus is initiation and regulation of movement patterns
• Limbic System: BG is connected to the Limbic System but is not a primary component of it, the BG + Limbic System collaboration helps to integrate emotional and motivational states with motor actions Impacts behaviors and habits
• Pre-Frontal Cortex: Communicates with the BG to guide and support goal-directed behaviors and the formation of habits.

Basal Ganglia Pathologies

• Parkinson's disease: Progressive neurodegenerative disease involving the loss of dopamine. Symptoms include tremors, stiffness, bradykinesia, and balance issues. Symptom management involves meds/supportive therapies; DBS
• Hemiballismus: Damage to the Substantia Nigra resulting in sudden, violent, large-amplitude movements. Symptoms management via meds/supportive therapies; DBS.
• Huntington’s disease: Hereditary neurodegenerative disorder (mutation of HTT gene) "chorea" behavioral changes, and eventually a progressive dementia. Management: Symptom management via meds/ supportive therapies; DBA (severe cases). Life expectancy limited secondary to complications from the disease.

Basal Ganglia Circuits - Direct vs. Indirect Pathways

• Direct - “Go” yields movement initiation.
• Indirect - “No-go” yields inhibited movement.

Direct Pathway

• Begins in the striatum, which sends inhibitory projections to the internal segment of the globus pallidus; the globus pallidus then sends inhibitory output to the thalamus.
• The cortex sends excitatory glutamate projections that activates the striatum.
• With the reduced inhibition of the internal segment of the globus pallidus, the thalamus can more effectively activate the cortex.
• The overall effect is to promote movement by reducing inhibition on the thalamus to allows it to activate the cortex more effectively.
• When input from either the cortex or substantia nigra increases in intensity, the direct pathway is activated
• Neurons in the striatum involved in the direct pathway express D1 dopamine receptor, therefore projections from both the cortex and the substantia nigra activate the neurons in the striatum

Indirect Pathway

• Like the direct pathway, input into the basal ganglia arises from the cortex and substantia nigra.
• The cortex sends glutamate or excitatory projections to the striatum, which activates it.
• The substantia nigra sends inhibitory dopamine projections to the striatum, acting on D2 receptors in the neurons involved in the indirect pathway
• Therefore, this pathway is part of the indirect pathway of the basal ganglia which inhibits movement.
• The overall effect is to increase inhibition on the thalamus, reducing its ability to activate the cortex, and thus suppressing movement

Indirect Pathway Activation

• It is activated by excitatory cortical input, activating the inhibitory striatal neurons.
• This leads to inhibitions of the external segment of the globus pallidus neurons, resulting in disinhibition of the excitatory neurons in the subthalamic nucleus
• It increases inhibition of the thalamus, therefore leading to decreased thalamic output to the cortex, leading to suppression of movement

Inhibition of the Indirect Pathway

• Can be inhibited by dopamine release from the substantia nigra, which acts on the D2 receptors in the striatum.
• The activation of these receptors by dopamine is inhibitory, which reduces the activity he of the striatal neurons
• Is the release of the inhibition of the external segment of the globus pallidus neurons.
• • more active the external segment globus pallidus neurons are, they send a stronger inhibitory or GABA signal to the subthalamic nucleus, reducing its activity, so The final result = the inhibition of the indirect pathway by dopamine leads to increased activity in the thalamus, which in turn increase excitatory outputs to the cortex, which facilitates movement by reducing the inhibitory effects of the indirect pathway.

Summary of Internal Processing

• Cortical activation of the direct pathway leads to increased thalamic output.
• Cortical activation of the indirect pathway leads to decreased thalamic output.
• Substantia nigra activation (via D1) of the direct pathway leads to increased thalamic output.
• Substantia nigra inhibition (via D2) of the indirect pathway leads to increased thalamic output.
• Input Direct pathway > Cortex (glutamate), it has the Activation (increased thalamic output), The Substantia nigra (dopamine) its Activation (D1) (increased thalamic output),
• Input Indirect pathway > Cortex (glutamate) activation (decreased thalamic output), the the Substantia nigra (Inhibition (D2) (increased thalamic output).

Basal Ganglia Loops

• Motor loop
• Cognitive loop
• Limbic loop
• Oculomotor loop
• The loops through the basal ganglia have different functions but follow the same general circuit.

Cortex Inputs/ Outputs from the Pallidum

• The cortex inputs to the striatum, internal processing through the basal ganglia circuits occurs, then the outputs from the pallidum projects to the thalamus, which sends outputs to the cortex
• Basic Circuits: Motor Loops: concerned with voluntary and learned movements, Cognitive Loop is concerned with planning a motor intentions. Oculomotor Loop:Plays a role in voluntary eye movements.
  • plays a role in executive functions like behavioral inhibition (preventing impulsive behaviors), planning and problem solving, and mediating socially appropriate behaviors, The Limbic Loop plays a role in the processing of emotion and reward is Concerned with emotional aspects of movement.