Chapter Summary
Study guidelines
- 1.
Recognise that the mature spinal cord is not segmented internally.
- 2.
Recall that the ventral horn cells take the form of columns rather than laminae.
- 3.
Recognise that ‘unconscious sensation’ simply means that the ascending afferent impulse activity concerned does not generate any kind of perception.
- 4.
Recognise that ‘conscious proprioception’ is more sensitive than either vision or the vestibular labyrinth in telling us when we are going off balance.
- 5.
Explain why muscles tell us more than joints do about the position of our limbs in space.
- 6.
Illustrate why it is clinically important to remember that one of the two ‘conscious’ pathways crosses the midline at all levels of the spinal cord, whereas the other crosses all at once, within the brainstem.
- 7.
Explain the meaning of the term ‘dissociated sensory loss’ and why it can occur.
General features
The arrangement of grey and white matter at different levels of the spinal cord is shown in Figure 15.1 . White matter consists mainly of axons and dendrites and is divided into ventral, lateral, and dorsal funiculi ( L. funiculus, ‘rope’), which are further divided into fasciculi ( L. fascis, ‘bundle’). The cervical (C5 to T1) and lumbosacral (L1 to S2) enlargements are produced by expansions of the grey matter required to innervate the corresponding limbs at those levels. White matter is most abundant in the upper reaches of the cord, which contain the sensory and motor pathways serving all four limbs. For example, within the dorsal funiculus, the gracile fasciculus carries information from the lower limb and is present at cervical as well as lumbosacral segmental levels, whereas the cuneate fasciculus carries information from the upper limb and is not seen at the lumbar level.
Although, as above, it is convenient to refer to different levels of the spinal cord in terms of numbered segments corresponding to the sites of attachment of the paired nerve roots, the cord shows no evidence of segmentation internally. The nuclear groups seen in transverse sections are in reality a series of discontinuous cell columns, most of them spanning several segments ( Figure 15.2 ).
Types of spinal neurons
The smallest neurons (soma diameters of 5 to 20 μm) are interneurons , and their cell bodies are contained within the cord. While the processes of some interneurons are confined within a single segment, others send their axons into the white matter surrounding the grey matter and ascend or descend two or more segments, interconnecting different spinal cord segments. These latter processes are termed propriospinal fibres and form the fasciculi proprii. Many of these smallest neurons participate in spinal reflexes. Others are intermediate cell stations interposed between fibre tracts descending from the brain and motor neurons projecting to cells controlling locomotion. Others again are so placed as to influence sensory transmission from lower to higher levels of the central nervous system (CNS).
Medium-sized neurons (soma diameters of 20 to 50 μm) are found in most parts of the grey matter. Most are relay (projection) cells receiving inputs from dorsal root afferents and projecting their axons to the brain. The projections are in the form of tracts , a tract being defined as a functionally homogeneous group of fibres. As will be seen, the term ‘tract’ is often used loosely because many projections originally thought to be ‘pure’ contain more than one functional class of fibre.
The largest neurons of all are the α motor neurons (soma diameters of 50 to 100 μm) for the supply of skeletal muscles. Scattered among them are smaller γ motor neurons supplying muscle spindles. In the medial part of the ventral horn are Renshaw cells , which exert tonic inhibition upon α motor neurons.
Spinal reflex arcs originating in muscle spindles and tendon organs have been described in Chapter 10 and the withdrawal reflex in Chapter 14 .
On the basis of cytoarchitectonic characteristics (e.g. neuronal size, staining characteristics, receptors, and connectivity), the spinal cord grey matter is divided into 10 layers, the laminae of Rexed that serve a descriptive but not necessarily functional purpose. Their configuration differs at various levels of the spinal cord; at some spinal cord levels, specific cell columns are recognised within the laminae, whereas in others, they are less clear ( Figure 15.3 ).
Spinal ganglia
The spinal or dorsal root ganglia are located on the dorsal root in the intervertebral foramina, where the ventral and dorsal roots come together to form the spinal nerves. Thoracic ganglia contain about 50,000 unipolar neurons, and spinocerebellar pathways serving the limbs contain about 100,000. These neurons are described as unipolar (or more correctly pseudounipolar). Their axons are morphologically indistinguishable from their dendrites because their somas are attached by a short stem axon . The individual ganglion cells are invested with modified Schwann cells called satellite cells ( Figure 15.4 ).
Central terminations of dorsal root afferents ( Figure 15.5 )
In the dorsal root entry zone close to the surface of the cord, the afferent fibres become segregated into medial and lateral divisions. The medial division comprises medium and large fibres that divide within the dorsal funiculus into ascending and descending branches. The branches swing into the dorsal grey horn and may synapse in the nucleus dorsalis (also known as the dorsal nucleus of Clarke). The largest ascending fibres run all the way to the dorsal column nuclei (gracilis/cuneatus) in the medulla oblongata, forming the bulk of the gracile and cuneate fasciculi.
The lateral division comprises small (Aδ and C) fibres, which upon entry divide into short ascending and descending branches within the Lissauer tract and synapse upon neurons of the substantia gelatinosa; some fibres synapse upon dendrites of cells belonging to the nucleus proprius. The nucleus proprius gives rise to the spinothalamic tract.
Ascending sensory pathways
Categories of sensation
In accordance with the flowchart in Figure 15.6 , neurologists speak of two kinds of sensation, conscious and unconscious. Conscious sensations are perceived at the level of the cerebral cortex. Unconscious sensations are not perceived; they are relayed to the cerebellum.
Conscious sensations
There are two kinds of conscious sensations: exteroceptive and proprioceptive . Exteroceptive sensations come from the external world; they impinge either on somatic receptors on the body surface or on telereceptors serving vision and hearing. Somatic sensations include touch, pressure, heat, cold, and pain.
Conscious proprioceptive sensations arise within the body. The receptors concerned are those of the locomotor system (muscles, joints, bones) and of the vestibular labyrinth. The pathways to the cerebral cortex form the substrate for position sense when the body is stationary, and for kinaesthetic sense during movement.
Unconscious sensations
There are also two kinds of unconscious sensations. Unconscious proprioception is the term used to describe afferent information reaching the cerebellum through the spinocerebellar pathways. This information is essential for smooth motor coordination. Second, interoception is a little-used term referring to unconscious afferent signals involved in visceral reflexes.
Sensory testing
Routine assessment of somatic exteroceptive sensation includes tests for the following:
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touch, by grazing the skin with a cotton swab
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pain, by applying the point of a pin
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thermal sense, by applying warm or cold test tubes to the skin
In alert and cooperative patients, active and passive tests of conscious proprioception can be performed. Active tests examine the patient’s ability to execute set-piece activities with the eyes closed:
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in the erect position, stand still, and with feet together ‘toe the line’ without swaying
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in the seated position, bring the index finger to the nose from the extended position of the arm (finger-to-nose test)
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in the recumbent position, place the heel of the foot on the opposite knee (heel-to-knee test)
Passive tests of conscious proprioception include the following:
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Joint sense . The clinician grasps the thumb or great toe by the sides and moves it while asking the patient to name the direction of movement (up or down). Joint sense is mediated in part by articular receptors but mainly by passive stretching of neuromuscular spindles. (If the nerves supplying a joint are anaesthetised or if the joint is completely replaced by a prosthesis, joint sense is only slightly impaired. Alternatively, activation of spindles by means of a vibrator creates the illusion of movement when the relevant joint is stationary.)
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Vibration sense . The clinician assesses the patient’s ability to detect the vibrations of a tuning fork applied proximal to the nail bed of the fingernail or toenail.