第36章感覚系

Somatosensation is a mixed sensory category and includes all sensation received from the skin and mucous membranes, as well from as the limbs and joints. Somatosensation is also known as tactile sense, or more familiarly, as the sense of touch. Somatosensation occurs all over the exterior of the body and at some interior locations as well. A variety of receptor types—embedded in the skin, mucous membranes, muscles, joints, internal organs, and cardiovascular system—play a role.

Recall that the epidermis is the outermost layer of skin in mammals. It is relatively thin, is composed of keratin-filled cells, and has no blood supply. The epidermis serves as a barrier to water and to invasion by pathogens. Below this, the much thicker dermis contains blood vessels, sweat glands, hair follicles, lymph vessels, and lipid-secreting sebaceous glands (Figure 36.4). Below the epidermis and dermis is the subcutaneous tissue, or hypodermis, the fatty layer that contains blood vessels, connective tissue, and the axons of sensory neurons. The hypodermis, which holds about 50 percent of the body’s fat, attaches the dermis to the bone and muscle, and supplies nerves and blood vessels to the dermis.

Somatosensory receptors are classified into four categories: mechanoreceptors, thermoreceptors, pain receptors (nociceptors), and chemoreceptors. These categories are based on the nature of stimuli each receptor class transduces. What is commonly referred to as “touch” involves more than one kind of stimulus and more than one kind of receptor. Mechanoreceptors in the skin are described as encapsulated (that is, surrounded by a capsule) or unencapsulated (a group that includes free nerve endings). A free nerve ending, as its name implies, is an unencapsulated dendrite of a sensory neuron. Free nerve endings are the most common nerve endings in skin, and they extend into the middle of the epidermis. Free nerve endings are sensitive to painful stimuli, to hot and cold, and to light touch. They are slow to adjust to a stimulus and so are less sensitive to abrupt changes in stimulation.

There are three classes of mechanoreceptors: tactile, proprioceptors, and baroreceptors. Mechanoreceptors sense stimuli due to physical deformation of their plasma membranes. They contain mechanically gated ion channels whose gates open or close in response to pressure, touch, stretching, and sound. There are four primary tactile mechanoreceptors in human skin: Merkel’s disks, Meissner’s corpuscles, Ruffini endings, and Pacinian corpuscles; two are located toward the surface of the skin and two are located deeper. A fifth type of mechanoreceptor, Krause end bulbs, are found only in specialized regions. Merkel’s disks (shown in Figure 36.5) are found in the upper layers of skin near the base of the epidermis, both in skin that has hair and on glabrous skin, that is, the hairless skin found on the palms and fingers, the soles of the feet, and the lips of humans and other primates. Merkel’s disks are densely distributed in the fingertips and lips. They are slow-adapting, unencapsulated nerve endings, and they respond to light touch. Light touch, also known as discriminative touch, is a light pressure that allows the location of a stimulus to be pinpointed. The receptive fields of Merkel’s disks are small with well-defined borders. That makes them finely sensitive to edges and they come into use in tasks such as typing on a keyboard.

Meissner’s corpuscles, (shown in Figure 36.6) also known as tactile corpuscles, are found in the upper dermis, but they project into the epidermis. They, too, are found primarily in the glabrous skin on the fingertips and eyelids. They respond to fine touch and pressure, but they also respond to low-frequency vibration or flutter. They are rapidly adapting, fluid-filled, encapsulated neurons with small, well-defined borders and are responsive to fine details. Like Merkel’s disks, Meissner’s corpuscles are not as plentiful in the palms as they are in the fingertips.

Deeper in the epidermis, near the base, are Ruffini endings, which are also known as bulbous corpuscles. They are found in both glabrous and hairy skin. These are slow-adapting, encapsulated mechanoreceptors that detect skin stretch and deformations within joints, so they provide valuable feedback for gripping objects and controlling finger position and movement. Thus, they also contribute to proprioception and kinesthesia. Ruffini endings also detect warmth. Note that these warmth detectors are situated deeper in the skin than are the cold detectors. It is not surprising, then, that humans detect cold stimuli before they detect warm stimuli.

Pacinian corpuscles (seen in Figure 36.7) are located deep in the dermis of both glabrous and hairy skin and are structurally similar to Meissner’s corpuscles; they are found in the bone periosteum, joint capsules, pancreas and other viscera, breast, and genitals. They are rapidly adapting mechanoreceptors that sense deep transient (but not prolonged) pressure and high-frequency vibration. Pacinian receptors detect pressure and vibration by being compressed, stimulating their internal dendrites. There are fewer Pacinian corpuscles and Ruffini endings in skin than there are Merkel’s disks and Meissner’s corpuscles.

In proprioception, proprioceptive and kinesthetic signals travel through myelinated afferent neurons running from the spinal cord to the medulla. Neurons are not physically connected, but communicate via neurotransmitters secreted into synapses or “gaps” between communicating neurons. Once in the medulla, the neurons continue carrying the signals to the thalamus.

Muscle spindles are stretch receptors that detect the amount of stretch, or lengthening of muscles. Related to these are Golgi tendon organs, which are tension receptors that detect the force of muscle contraction. Proprioceptive and kinesthetic signals come from limbs. Unconscious proprioceptive signals run from the spinal cord to the cerebellum, the brain region that coordinates muscle contraction, rather than to the thalamus, like most other sensory information.

Baroreceptors detect pressure changes in an organ. They are found in the walls of the carotid artery and the aorta where they monitor blood pressure, and in the lungs where they detect the degree of lung expansion. Stretch receptors are found at various sites in the digestive and urinary systems.

In addition to these two types of deeper receptors, there are also rapidly adapting hair receptors, which are found on nerve endings that wrap around the base of hair follicles. There are a few types of hair receptors that detect slow and rapid hair movement, and they differ in their sensitivity to movement. Some hair receptors also detect skin deflection, and certain rapidly adapting hair receptors allow detection of stimuli that have not yet touched the skin.