ATOC10 3/17/06 10:06 AM Page 137 CHAPTER 10 Ascending Sensory Pathways CLINICAL CASE CLINICAL CASE SENSORY RECEPTORS ANTEROLATERAL SYSTEM A 60-year-old woman com- normal. plains of falls, imbalance, and On examination, the patient shows TACTILE SENSATION AND numbness and tingling in her hands and normal mental status. Strength seems PROPRIOCEPTION legs. There is also some incoordination of essentially normal throughout. Sensation, hand use and she has difficulty manipu- particularly to vibration and joint posi- SENSORY PATHWAYS TO THE lating small items such as buttons. She is tion, is severely diminished in the distal CEREBELLUM unable to play the piano now since she upper and lower limbs (arms, legs, hands, cannot position her fingers correctly on and feet). Tendon reflexes are normal in CLINICAL CONSIDERATIONS the piano keys. She thinks the strength in the arms, but somewhat brisk in the legs her arms and legs is adequate. Symptoms at the knees and ankles. Gait is moder- MODULATION OF NOCICEPTION started with very slight tingling sensa- ately ataxic and she has to reach out for tions, which she noticed about 5 years support by touching the walls of the NEUROPLASTICITY ago. The falls and difficulty walking have hallway at times. Fine movements of been present for about 2 months. Higher the fingers are performed poorly, even SYNONYMS AND EPONYMS order cognitive functions are intact though finger and wrist strength seems according to her husband. Her vision is normal. FOLLOW-UP TO CLINICAL CASE QUESTIONS TO PONDER A variety of sensory receptors scattered throughout the body progresses, via the ascending sensory systems (pathways), can become activated by exteroceptive, interoceptive, or to the cerebral cortex or to the cerebellum. Sensory informa- proprioceptive input. Exteroceptive input relays sensory tion is also relayed to other parts of the CNS where it may information about the body’s interaction with the external function to elicit a reflex response, or may be integrated into environment. Interoceptive input relays information about pattern-generating circuitry. the body’s internal state, whereas proprioceptive input con- The ascending sensory pathways are classified according veys information about position sense from the body and to the functional components (modalities) they carry as well its component parts. Each receptor is specialized to detect as by their anatomical localization. The two functional categor- mechanical, chemical, nociceptive (L. nocere, “to injure,” ies are the general somatic afferent (GSA) system, which “painful”), or thermal stimuli. Activation of a sensory recep- transmits sensory information such as touch, pressure, vibra- tor is converted into nerve impulses and this sensory input is tion, pain, temperature, stretch, and position sense from then conveyed via the fibers of the cranial or spinal nerves to somatic structures; and the general visceral afferent (GVA) their respective relay nuclei in the central nervous system system, which transmits sensory information such as pressure, (CNS). The sensory information is then further processed as it pain, and other visceral sensation from visceral structures. ·· ATOC10 3/17/06 10:06 AM Page 138 138 === CHAPTER 10 Anatomical system Anatomical tracts Functional component(s) Anterolateral (ALS) Spinothalamic* Pain, temperature, nondiscriminative Spinoreticular (crude) touch, pressure, and some Spinomesencephalic proprioceptive sensation Spinotectal Spinohypothalamic Dorsal column–medial Fasciculus gracilis Discriminative (fine) touch, vibratory lemniscal (DCML)* Fasciculus cuneatus sense, position sense Somatosensory to the Anterior spinocerebellar Primarily proprioceptive information cerebellum Posterior spinocerebellar (also some pain and pressure Rostral spinocerebellar information) Cuneocerebellar Table 10.1 = A general description of *Indicates conscious level. the anatomical and functional aspects of the ascending sensory pathways. Anatomically, the ascending sensory systems consist of ascending (somatosensory) pathways, the cerebellum, and three distinct pathways: the anterolateral system (ALS), the the somatosensory cortex, as well as the motor cortex and dorsal column–medial lemniscal (DCML) pathway, and the descending (motor) pathways. Furthermore, descending somatosensory pathways to the cerebellum. projections from the somatosensory cortex, as well as from The anterolateral system, which includes the spinothala- the raphe nucleus magnus and the dorsolateral pontine mic, spinoreticular, spinomesencephalic, spinotectal, and reticular formation to the somatosensory relay nuclei of the spinohypothalamic tracts, relays predominantly pain and brainstem and spinal cord, modulate the transmission of temperature sensation, as well as nondiscriminative (crude incoming sensory impulses to higher brain centers. or poorly localized) touch, pressure, and some propriocep- This chapter includes a description of the sensory recep- tive sensation (Table 10.1). tors and the ascending sensory pathways from the body, The dorsal column–medial lemniscal pathway (which whereas the ascending sensory pathways from the head, includes the fasciculus gracilis, fasciculus cuneatus, and transmitted mostly by the trigeminal system, are described medial lemniscus) relays discriminative (fine) tactile sense, in Chapter 15. vibratory sense, and position sense (Table 10.1). The somatosensory pathways to the cerebellum, which include the anterior, posterior, and rostral spinocerebellar, SENSORY RECEPTORS as well as the cuneocerebellar tracts, relay primarily propri- Although sensory receptors vary, they Although sensory receptors oceptive (but also some pain and pressure) information generally all function in a similar vary according to their mor- (Table 10.1). fashion phology, the velocity of con- The ascending sensory pathways are the main avenues by duction, and the modality to which information concerning the body’s interaction with the which they respond, as well as to their location in the body, external environment, its internal condition, and the position they generally all function in a similar fashion. The stimulus and movement of its parts, reach the brain. One similarity to which a specific receptor responds causes an alteration in shared by all three ascending sensory pathways from the the ionic permeability of the nerve endings, generating a body (not including the head or face) is that the first order receptor potential that results in the formation of action neuron cell bodies reside in the dorsal root ganglia. It is potentials. This transformation of the stimulus into an elec- interesting to note that conscious perception of sensory trical signal is referred to as sensory transduction. information from external stimuli is mediated by the Some receptors that respond quickly and maximally at spinothalamic and DCML pathways to the ventral posterior the onset of the stimulus, but stop responding even if the lateral nucleus of the thalamus, whereas sensations that do stimulus continues, are known as rapidly adapting (phasic) not reach consciousness are mediated by the spinoreticular, receptors. These are essential in responding to changes but spinomesencephalic, spinotectal, spinohypothalamic, and they ignore ongoing processes, such as when one wears a the anterior, posterior, and rostral spinocerebellar, and wristwatch and ignores the continuous pressure on the skin cuneocerebellar tracts. These tracts terminate in the reticular of the wrist. However, there are other receptors, slowly formation, mesencephalon, hypothalamus and cerebellum, adapting (tonic) receptors, that continue to respond as long respectively. as the stimulus is present. Sensory input may ultimately elicit a reflex or other Sensory receptors are classified according to the source motor response because of the functional integration of the of the stimulus or according to the modality to which they ·· ATOC10 3/17/06 10:06 AM Page 139 ASCENDING SENSORY PATHWAYS === 139 respond. It is important to note that, in general, receptors do Nociceptors not transmit only one specific sensation. Nociceptors are rapidly adapting receptors that are sensitive to noxious or painful stimuli Classification according to stimulus source Nociceptors are rapidly adapting receptors that are sensitive Receptors that are classified according to the source of the to noxious or painful stimuli. They are located at the peri- stimulus are placed in one of the following three categories: pheral terminations of lightly myelinated free nerve endings exteroceptors, proprioceptors, or interoceptors. of type Aδ fibers, or unmyelinated type C fibers, transmitting pain. Nociceptors are further classified into three types. Exteroceptors are close to the body surface and are specialized to detect sensory information from the external environment 1 Mechanosensitive nociceptors (of Aδ fibers), which are sensitive to intense mechanical stimulation (such as 1 Exteroceptors are close to the body surface and are spe- pinching with pliers) or injury to tissues. cialized to detect sensory information from the external 2 Temperature-sensitive (thermosensitive) nociceptors environment (such as visual, olfactory, gustatory, audi- (of Aδ fibers), which are sensitive to intense heat and cold. tory, and tactile stimuli). Receptors in this class are sens- 3 Polymodal nociceptors (of C fibers), which are sensitive itive to touch (light stimulation of the skin surface), to noxious stimuli that are mechanical, thermal, or chem- pressure (stimulation of receptors in