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Thalamus
Yash Bhandari (03)
BPT 1st Year
Introduction
 Situated at the rostral end of the brainstem
 Function : Important relay & Integrative station for information passing to all
areas of the cerebral cortex, the basal ganglia, the hypothalamus, and the
brainstem
General Appearances
 Large, egg shaped mass of grey matter
 Forms major part of the diencephalon
 2 thalami situated on each side of the third ventricle
 ANTERIOR END- is narrow and rounder
- forms the posterior boundary of the interventricular foramen
 POSTERIOR END- expanded to form Pulvinar, which overhangs the Superior colliculus
 INFERIOR SURFACE- continuous with the tegmentum of the midbrain
 MEDIAL SURFACE- forms part of lateral wall of the third ventricle & it usually connected to the
opposite thalamus by a band of grey matter called the Interthalamic connection
Subdivision
 Covered on its- superior surface by a thin layer of white matter, called the Stratum zonale &
lateral surface by another layer, the External medullary lamina
 Grey matter of thalamus is divided by a vertical sheet of white matter, the Internal medullary lamina,
into medial & lateral halves
 Internal medullary lamina consists of nerve fibers that pass from one thalamic nucleus to another
 Anterosuperiorly, the internal medullary lamina splits, resembling a Y shape
 Anterior part lies between the limbs of the Y
 Medial & Lateral lie on the sides of the stem of the Y
 Each of the three parts of the thalamus contains a group of thalamic nuclei
 Smaller nuclear groups are situated within the internal medullary lamina & some are on medial & lateral
side
Thalamus
Anterior
part
Medial
Part
Lateral
Part
Anterior Part
 Contains the anterior thalamic nuclei
 Receives mammillothalamic tract from the mammillary nuclei
 Also receives reciprocal connections with the cingulate gyrus & hypothalamus
 Function: it is closely associated with that of the limbic system & is concerned with emotional tone &
the mechanism of recent memory
Medial Part
 Contains the large Dorsomedial nucleus & several smaller nuclei
 It has two connections with the whole prefrontal cortex of frontal lobe of cerebral hemisphere
 It also has connections with the hypothalamic nuclei
 It is interconnected with all other groups of thalamic nuclei
 Function: Integration of a large variety of sensory information, including somatic, visceral, and olfactory
information, including somatic, visceral, and olfactory information, and the relation of this information
to one’s emotional feelings & subjective states
Lateral Part
Lateral Part
Dorsal tier of
Nuclei
Ventral Tier of
Nuclei
1) Ventral
anterior
nucleus
2) Ventral
lateral nucleus
3) Ventral
posterior
nucleus
Ventral
Posteromedial
nucleus (VPM)
Ventral
Posterolateral
nucleus (VPL)
A) Dorsal Tier of the Nuclei
 Includes the Lateral dorsal nucleus, the lateral posterior nucleus and the
pulvinar
 Nuclei detail is not clear
 They are know, However, to have interconnections with other thalamus nuclei
and with the parietal lobe, cingulate gyrus, and occipital & temporal lobes
B) Ventral Tier of the Nuclei
 In Craniocaudal sequence:
 1) Ventral anterior nucleus
 Connected to the reticular formation, the substantia nigra, the corpus
striatum, and the premotor cortex and with many of the other nuclei
 Since this nucleus lies on the pathway between the corpus striatum & the
motor areas of the frontal cortex, it probably influences the activities of the
motor cortex
 2) Ventral lateral nucleus
 Has connection with Reticular formation, substantia nigra, corpus striatum, premotor cortex, in addition, has
a major input from the cerebellum & a minor input from the red nucleus
 Main projections pass to the motor & premotor regions of cerebral cortex
 This thalamic nucleus probably influences motor activity
 3) Ventral posterior nucleus
 Ventral Posteromedial nucleus (VPM)
 Receives the ascending trigeminal & gustatory pathways
 Ventral Posterolateral nucleus (VPL)
 Receives the important ascending sensory tracts, the medial & lateral lemnisci
 The thalamocortical projections from these important nuclei pass through the posterior limb of the internal
capsule & corona radiate to the primary somatic sensory areas of the cerebral cortex in the postcentral gyrus
(areas 3,1,2)
Other Nuclei of the Thalamus
 Intralaminar nuclei
• Small collection of nerve cells within the internal medullary lamina
• Receives afferent fibers from the reticular formation, spinothalamic & trigeminothalamic tract; they send
efferent fibers to other thalamic nuclei, which in turn project to the cerebral cortex, & fibers to the corpus
striatum
• Nucleus influence the levels of consciousness and alertless
 Midline nuclei
• Groups of nerve cells adjacent to the third ventricle and in the interthalamic connection
• Receives afferent fibers from the reticular formation
• Precise functions are unknown
 Reticular nucleus
• It is a thin layer of nerve cells sandwiched between the external medullary lamina & posterior limb of the
internal capsule
• Afferent fibers converge on this nucleus from the cerebral cortex and the reticular formation, and its output
is mainly to other thalamic nuclei
• Function: it may be concerned with a mechanism by which the cerebral cortex regulates thalamic activity
 Medial Geniculate body
• Form part of the auditory pathway & is a swelling on the posterior surface of the thalamus beneath the
pulvinar
• Afferent fibers to the medial geniculate body form the inferior brachium & come from the inferior colliculus
• Inferior colliculus receives the termination of the fibers of the lateral lemniscus
• Medial geniculate body receives auditory information from both ears but predominantly from the opposite ear
• Efferent fibers leave the medial geniculate body to form the auditory radiation, which passes to the auditory
cortex of the superior temporal gyrus
 Lateral Geniculate body
• Forms part of the visual pathway & is a swelling on the undersurface of the pulvinar of the thalamus
• Nucleus consists to six layers of nerve cells & is the terminus of all but a few fibers of the optic tract (except
the fibers passing to the pretectal nucleus)
• Fibers are the axons of the ganglion cell layer of the retina & come from the temporal half of the ipsilateral
eye & from the nasal half of the contralateral eye, the latter fibers crossing the midline in the optic chiasma
• Receives visual information from the opposite field of vision
• Efferent fibers leave the lateral geniculate body to form the visual radiation, which passes to the visual
cortex of the occipital lobe
Connections
Thalamic Radiations
 Thalamic radiation is the collection of nerve fibers connecting thalamus & cerebral cortex
 It contains both thalamocortical & corticothalamic fibers
 All these fibers between thalamus and cerebral cortex pass through internal capsule
 Fibers of thalamic radiation are divided into four groups which are called thalamic peduncle or stalks
1) Anterior (frontal) thalamic peduncle or radiation
2) Superior (centroparietal) thalamic peduncle or radiation
3) Posterior (occipital) thalamic peduncle or radiation
4) Inferior (temporal) thalamic peduncle or radiation
 1) Anterior thalamic peduncle
• Fibers connects the frontal lobe of cerebral cortex with medial & lateral thalamic nuclei
• Contains mostly motor nerve fibers
 2) Superior thalamic peduncle
• Fibers connects postcentral gyrus of parietal lobe & adjacent area in frontal cortex with lateral mass of
thalamic nuclei
• It contains mainly the sensory fibers
 3) Posterior thalamic peduncle
• Connects occipital lobe of cerebral cortex with pulvinar & lateral geniculate
body
• It contains the nerve fibers concerned with vision
 4) Inferior thalamic peduncle
• Connect temporal lobe & insula with pulvinar & medial geniculate body
• Contains the nerve fibers concerned with hearing
Functions
 Thalamus is primarily concerned with somatic functions and it plays little role in the visceral functions.
Following are the various functions of thalamus:
 1) RELAY CENTER
 Thalamus forms the relay center for the sensations.
 Impulses of almost all the sensations reach the thalamic nuclei, particularly in the ventral posterolateral
nucleus.
 After being processed in the thalamus, the impulses are carried to cerebral cortex through thalamocortical
fibers.
 2. CENTER FOR PROCESSING OF SENSORY INFORMATION
 Thalamus forms the major center for processing the sensory information.
 All the peripheral sensory impulses reaching thalamus are integrated and modified before being sent to specific
areas of cerebral cortex.
 This function of thalamus is usually called the processing of sensory information.
 Functional Gateway for Cerebral Cortex
 Almost all the sensations are processed in thalamus before reaching cerebral cortex.
 Very little information of somatosensory function is sent directly to cerebral cortex without being processed by
the thalamic nuclei.
 Because of this function, thalamus is usually called ‘functional gateway’ for cerebral cortex.
 3) CENTER FOR DETERMINING QUALITY OF SENSATIONS
 Thalamus is also the center for determining the quality of sensations, i.e. to determine the affective nature of
sensations. Usually the sensations have two qualities:
i. Discriminative nature
ii. Affective nature.
 i. Discriminative Nature
 Discriminative nature is the ability to recognize the type, location and other details of the sensations and it is
the function of cerebral cortex.
 ii. Affective Nature
 Affective nature is the capacity to determine whether a sensation is pleasant or unpleasant and agreeable or
disagreeable.
 Determining the affective nature of sensations is the function of thalamus.
 4) CENTER FOR SEXUAL SENSATIONS
 Thalamus forms the center for perception of sexual sensations.
 5) ROLE IN AROUSAL AND ALERTNESS REACTIONS
 Because of its connections with nuclei of reticular formation, thalamus plays an important role in arousal and
alertness reactions.
 6) CENTER FOR REFLEX ACTIVITY
 Since the sensory fibers relay here, thalamus forms the center for many
reflex activities.
 7) CENTER FOR INTEGRATION OF MOTOR ACTIVITY
 Through the connections with cerebellum and basal ganglia, thalamus serves
as a center for integration of motor functions.
Physiological Classification
Applied
 THALAMIC LESION
 Thalamic lesion occurs mainly because of blockage (due to thrombosis) in thalamogeniculate branch of posterior
cerebral artery.
 Mostly, posteroventral nuclei of thalamus are affected because the thalamogeniculate branch of posterior cerebral
artery supply this part of thalamus.
 Lesion of thalamus leads to a condition called thalamic syndrome.
 THALAMIC SYNDROME
 Thalamic syndrome is the neurological disease caused by infarction of posteroventral part of thalamus.
 It is a rare disease and it has many names
 Synonyms of thalamic syndrome
1. Dejerine-Roussy syndrome
2. Thalamic hyperesthetic anesthesia
3. Thalamic pain syndrome
4. Central pain syndrome
5. Central poststroke pain syndrome
6. Posterior thalamic syndrome
7. Retrolenticular syndrome
 In thalamic syndrome, whole body becomes hypersensitive to pain.
 Effects of thalamic lesion occur in the contralateral (opposite) side
 1. Loss of Sensations
 Loss of all sensations (anesthesia) occurs as the sensory relay system in thalamus is affected.
 2. Astereognosis
 Astereognosis is the loss of ability to recognize a known object by touch with closed eyes.
 It is due to the loss of tactile and kinesthetic sensations in thalamic syndrome.
 3. Ataxia
 Ataxia refers to incoordination of voluntary movements.
 It occurs due to loss of kinesthetic sensation.
 This type of ataxia due to loss of sensation is called sensory ataxia.
 It is very common in thalamic syndrome.
 4. Thalamic Phantom Limb
 The patient is unable to locate the position of a limb with closed eyes.
 The patient may search for the limb in air or may have the illusion that the limb is lost.
 This is called thalamic phantom limb.
 5. Anosognosia
 Anosognosia is the lack of awareness or denial of existance of a neurological defect or general illness or any
disability.
 6. Spontaneous Pain and Thalamic Over-reaction
 Spontaneous pain occurs often. Pain stimulus is felt more acutely than in normal conditions (hyperalgesia).
 Pain may be so intense, that it even resists the action of powerful sedatives like morphine.
 Threshold for pain is very much reduced.
 Even the light touch may be unpleasant.
 Sometimes, the patient feels pain even in the absence of pain stimulus.
 It becomes worst in conditions such as emotional disturbance and exposure to cold or heat.
 Pain is due to over activity of medial mass of nuclei of thalamus, which escape the lesion.
 Abnormal reaction to various stimuli is called thalamic over-reaction.
 7. Involuntary Movements
 Thalamic syndrome is always associated with some involuntary motor movements.
 Athetosis
 Athetosis means slow writhing and twisting movements.
 Chorea
 Chorea means quick, jerky, involuntary movements.
 Intention tremor
 Tremor is defined as rapid alternate rhythmic and involuntary movement of
flexion and extension in the joints of fingers and wrist or elbow.
 Intention tremor is the tremor that develops while attempting to do any
voluntary act.
 Intention tremor is the common feature of thalamic syndrome.
 8. Thalamic Hand or Athetoid Hand
 Athetoid hand is the abnormal attitude of hand in thalamic lesion.
 It is characterized by moderate flexion at wrist and hyperextension of all
fingers.

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Thalamus

  • 2. Introduction  Situated at the rostral end of the brainstem  Function : Important relay & Integrative station for information passing to all areas of the cerebral cortex, the basal ganglia, the hypothalamus, and the brainstem
  • 3. General Appearances  Large, egg shaped mass of grey matter  Forms major part of the diencephalon  2 thalami situated on each side of the third ventricle  ANTERIOR END- is narrow and rounder - forms the posterior boundary of the interventricular foramen  POSTERIOR END- expanded to form Pulvinar, which overhangs the Superior colliculus  INFERIOR SURFACE- continuous with the tegmentum of the midbrain  MEDIAL SURFACE- forms part of lateral wall of the third ventricle & it usually connected to the opposite thalamus by a band of grey matter called the Interthalamic connection
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  • 5. Subdivision  Covered on its- superior surface by a thin layer of white matter, called the Stratum zonale & lateral surface by another layer, the External medullary lamina  Grey matter of thalamus is divided by a vertical sheet of white matter, the Internal medullary lamina, into medial & lateral halves  Internal medullary lamina consists of nerve fibers that pass from one thalamic nucleus to another  Anterosuperiorly, the internal medullary lamina splits, resembling a Y shape  Anterior part lies between the limbs of the Y  Medial & Lateral lie on the sides of the stem of the Y  Each of the three parts of the thalamus contains a group of thalamic nuclei  Smaller nuclear groups are situated within the internal medullary lamina & some are on medial & lateral side Thalamus Anterior part Medial Part Lateral Part
  • 6. Anterior Part  Contains the anterior thalamic nuclei  Receives mammillothalamic tract from the mammillary nuclei  Also receives reciprocal connections with the cingulate gyrus & hypothalamus  Function: it is closely associated with that of the limbic system & is concerned with emotional tone & the mechanism of recent memory
  • 7. Medial Part  Contains the large Dorsomedial nucleus & several smaller nuclei  It has two connections with the whole prefrontal cortex of frontal lobe of cerebral hemisphere  It also has connections with the hypothalamic nuclei  It is interconnected with all other groups of thalamic nuclei  Function: Integration of a large variety of sensory information, including somatic, visceral, and olfactory information, including somatic, visceral, and olfactory information, and the relation of this information to one’s emotional feelings & subjective states
  • 8. Lateral Part Lateral Part Dorsal tier of Nuclei Ventral Tier of Nuclei 1) Ventral anterior nucleus 2) Ventral lateral nucleus 3) Ventral posterior nucleus Ventral Posteromedial nucleus (VPM) Ventral Posterolateral nucleus (VPL)
  • 9. A) Dorsal Tier of the Nuclei  Includes the Lateral dorsal nucleus, the lateral posterior nucleus and the pulvinar  Nuclei detail is not clear  They are know, However, to have interconnections with other thalamus nuclei and with the parietal lobe, cingulate gyrus, and occipital & temporal lobes
  • 10. B) Ventral Tier of the Nuclei  In Craniocaudal sequence:  1) Ventral anterior nucleus  Connected to the reticular formation, the substantia nigra, the corpus striatum, and the premotor cortex and with many of the other nuclei  Since this nucleus lies on the pathway between the corpus striatum & the motor areas of the frontal cortex, it probably influences the activities of the motor cortex
  • 11.  2) Ventral lateral nucleus  Has connection with Reticular formation, substantia nigra, corpus striatum, premotor cortex, in addition, has a major input from the cerebellum & a minor input from the red nucleus  Main projections pass to the motor & premotor regions of cerebral cortex  This thalamic nucleus probably influences motor activity  3) Ventral posterior nucleus  Ventral Posteromedial nucleus (VPM)  Receives the ascending trigeminal & gustatory pathways  Ventral Posterolateral nucleus (VPL)  Receives the important ascending sensory tracts, the medial & lateral lemnisci  The thalamocortical projections from these important nuclei pass through the posterior limb of the internal capsule & corona radiate to the primary somatic sensory areas of the cerebral cortex in the postcentral gyrus (areas 3,1,2)
  • 12. Other Nuclei of the Thalamus  Intralaminar nuclei • Small collection of nerve cells within the internal medullary lamina • Receives afferent fibers from the reticular formation, spinothalamic & trigeminothalamic tract; they send efferent fibers to other thalamic nuclei, which in turn project to the cerebral cortex, & fibers to the corpus striatum • Nucleus influence the levels of consciousness and alertless  Midline nuclei • Groups of nerve cells adjacent to the third ventricle and in the interthalamic connection • Receives afferent fibers from the reticular formation • Precise functions are unknown  Reticular nucleus • It is a thin layer of nerve cells sandwiched between the external medullary lamina & posterior limb of the internal capsule • Afferent fibers converge on this nucleus from the cerebral cortex and the reticular formation, and its output is mainly to other thalamic nuclei • Function: it may be concerned with a mechanism by which the cerebral cortex regulates thalamic activity
  • 13.  Medial Geniculate body • Form part of the auditory pathway & is a swelling on the posterior surface of the thalamus beneath the pulvinar • Afferent fibers to the medial geniculate body form the inferior brachium & come from the inferior colliculus • Inferior colliculus receives the termination of the fibers of the lateral lemniscus • Medial geniculate body receives auditory information from both ears but predominantly from the opposite ear • Efferent fibers leave the medial geniculate body to form the auditory radiation, which passes to the auditory cortex of the superior temporal gyrus  Lateral Geniculate body • Forms part of the visual pathway & is a swelling on the undersurface of the pulvinar of the thalamus • Nucleus consists to six layers of nerve cells & is the terminus of all but a few fibers of the optic tract (except the fibers passing to the pretectal nucleus) • Fibers are the axons of the ganglion cell layer of the retina & come from the temporal half of the ipsilateral eye & from the nasal half of the contralateral eye, the latter fibers crossing the midline in the optic chiasma • Receives visual information from the opposite field of vision • Efferent fibers leave the lateral geniculate body to form the visual radiation, which passes to the visual cortex of the occipital lobe
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  • 17. Thalamic Radiations  Thalamic radiation is the collection of nerve fibers connecting thalamus & cerebral cortex  It contains both thalamocortical & corticothalamic fibers  All these fibers between thalamus and cerebral cortex pass through internal capsule  Fibers of thalamic radiation are divided into four groups which are called thalamic peduncle or stalks 1) Anterior (frontal) thalamic peduncle or radiation 2) Superior (centroparietal) thalamic peduncle or radiation 3) Posterior (occipital) thalamic peduncle or radiation 4) Inferior (temporal) thalamic peduncle or radiation  1) Anterior thalamic peduncle • Fibers connects the frontal lobe of cerebral cortex with medial & lateral thalamic nuclei • Contains mostly motor nerve fibers  2) Superior thalamic peduncle • Fibers connects postcentral gyrus of parietal lobe & adjacent area in frontal cortex with lateral mass of thalamic nuclei • It contains mainly the sensory fibers
  • 18.  3) Posterior thalamic peduncle • Connects occipital lobe of cerebral cortex with pulvinar & lateral geniculate body • It contains the nerve fibers concerned with vision  4) Inferior thalamic peduncle • Connect temporal lobe & insula with pulvinar & medial geniculate body • Contains the nerve fibers concerned with hearing
  • 19. Functions  Thalamus is primarily concerned with somatic functions and it plays little role in the visceral functions. Following are the various functions of thalamus:  1) RELAY CENTER  Thalamus forms the relay center for the sensations.  Impulses of almost all the sensations reach the thalamic nuclei, particularly in the ventral posterolateral nucleus.  After being processed in the thalamus, the impulses are carried to cerebral cortex through thalamocortical fibers.  2. CENTER FOR PROCESSING OF SENSORY INFORMATION  Thalamus forms the major center for processing the sensory information.  All the peripheral sensory impulses reaching thalamus are integrated and modified before being sent to specific areas of cerebral cortex.  This function of thalamus is usually called the processing of sensory information.  Functional Gateway for Cerebral Cortex  Almost all the sensations are processed in thalamus before reaching cerebral cortex.  Very little information of somatosensory function is sent directly to cerebral cortex without being processed by the thalamic nuclei.  Because of this function, thalamus is usually called ‘functional gateway’ for cerebral cortex.
  • 20.  3) CENTER FOR DETERMINING QUALITY OF SENSATIONS  Thalamus is also the center for determining the quality of sensations, i.e. to determine the affective nature of sensations. Usually the sensations have two qualities: i. Discriminative nature ii. Affective nature.  i. Discriminative Nature  Discriminative nature is the ability to recognize the type, location and other details of the sensations and it is the function of cerebral cortex.  ii. Affective Nature  Affective nature is the capacity to determine whether a sensation is pleasant or unpleasant and agreeable or disagreeable.  Determining the affective nature of sensations is the function of thalamus.  4) CENTER FOR SEXUAL SENSATIONS  Thalamus forms the center for perception of sexual sensations.  5) ROLE IN AROUSAL AND ALERTNESS REACTIONS  Because of its connections with nuclei of reticular formation, thalamus plays an important role in arousal and alertness reactions.
  • 21.  6) CENTER FOR REFLEX ACTIVITY  Since the sensory fibers relay here, thalamus forms the center for many reflex activities.  7) CENTER FOR INTEGRATION OF MOTOR ACTIVITY  Through the connections with cerebellum and basal ganglia, thalamus serves as a center for integration of motor functions.
  • 23. Applied  THALAMIC LESION  Thalamic lesion occurs mainly because of blockage (due to thrombosis) in thalamogeniculate branch of posterior cerebral artery.  Mostly, posteroventral nuclei of thalamus are affected because the thalamogeniculate branch of posterior cerebral artery supply this part of thalamus.  Lesion of thalamus leads to a condition called thalamic syndrome.  THALAMIC SYNDROME  Thalamic syndrome is the neurological disease caused by infarction of posteroventral part of thalamus.  It is a rare disease and it has many names  Synonyms of thalamic syndrome 1. Dejerine-Roussy syndrome 2. Thalamic hyperesthetic anesthesia 3. Thalamic pain syndrome 4. Central pain syndrome 5. Central poststroke pain syndrome 6. Posterior thalamic syndrome 7. Retrolenticular syndrome
  • 24.  In thalamic syndrome, whole body becomes hypersensitive to pain.  Effects of thalamic lesion occur in the contralateral (opposite) side  1. Loss of Sensations  Loss of all sensations (anesthesia) occurs as the sensory relay system in thalamus is affected.  2. Astereognosis  Astereognosis is the loss of ability to recognize a known object by touch with closed eyes.  It is due to the loss of tactile and kinesthetic sensations in thalamic syndrome.  3. Ataxia  Ataxia refers to incoordination of voluntary movements.  It occurs due to loss of kinesthetic sensation.  This type of ataxia due to loss of sensation is called sensory ataxia.  It is very common in thalamic syndrome.  4. Thalamic Phantom Limb  The patient is unable to locate the position of a limb with closed eyes.  The patient may search for the limb in air or may have the illusion that the limb is lost.  This is called thalamic phantom limb.
  • 25.  5. Anosognosia  Anosognosia is the lack of awareness or denial of existance of a neurological defect or general illness or any disability.  6. Spontaneous Pain and Thalamic Over-reaction  Spontaneous pain occurs often. Pain stimulus is felt more acutely than in normal conditions (hyperalgesia).  Pain may be so intense, that it even resists the action of powerful sedatives like morphine.  Threshold for pain is very much reduced.  Even the light touch may be unpleasant.  Sometimes, the patient feels pain even in the absence of pain stimulus.  It becomes worst in conditions such as emotional disturbance and exposure to cold or heat.  Pain is due to over activity of medial mass of nuclei of thalamus, which escape the lesion.  Abnormal reaction to various stimuli is called thalamic over-reaction.  7. Involuntary Movements  Thalamic syndrome is always associated with some involuntary motor movements.  Athetosis  Athetosis means slow writhing and twisting movements.  Chorea  Chorea means quick, jerky, involuntary movements.
  • 26.  Intention tremor  Tremor is defined as rapid alternate rhythmic and involuntary movement of flexion and extension in the joints of fingers and wrist or elbow.  Intention tremor is the tremor that develops while attempting to do any voluntary act.  Intention tremor is the common feature of thalamic syndrome.  8. Thalamic Hand or Athetoid Hand  Athetoid hand is the abnormal attitude of hand in thalamic lesion.  It is characterized by moderate flexion at wrist and hyperextension of all fingers.