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How does different sensory stimuli differ in nerve signal?

How does different sensory stimuli differ in nerve signal?



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Every day, we experience different types of sensory stimulus, like heat, pain, cold, etc. However, in each case, the transmission of the stimulus to the brain through the neurones is carried on in the same way: through propagated depolarisation and repolarisation in axons, while through the release of acetylcholine at the synapses.

So, how does the different sensory stimulus differ if they are transmitted the same way? (I mean to say that how does the nerve signals differ for two different kinds of stimulus?)

Let's put the question simply:

How do two nerve signals originated from two different stimuli differ? If they do not differ, then how does the brain differentiate between different nerve signals if they are all alike? How does it understand what nerve signal carries what message?


I think the short answer to your question is that the contents of different types of sensory stimulation are communicated to different populations of cells in the brain (note this transmission is mostly glutamatergic, not cholinergic). From there, the remaining associations and experiences derive from connectivity with other networks of brain neurons that result in different types of experience, based on the networks that those networks form and learned associations over development and a lifetime.

Computer analogies aren't always appropriate to understanding things about the brain, but maybe it could work out a bit here. How does a computer distinguish between input on your keyboard vs your mouse? On many modern computers, it is common for both to be connected with USB. The computer has some software that distinguishes between the different inputs - the brain has hardware that does the same.

Later on in the brain, or at a different stage in a computer program, those inputs can be mapped to functions that underlie experiences or functional outputs. Movement of the mouse can be converted to the movement of a screen cursor ("motor output"). Proprioception (the sensation of the relative position of the body from receptors in the muscles, tendons, and joints) can be combined with motor plans to create an error signal to move a particular muscle to match the intended movement.

Explaining any further than this probably requires entire textbooks of material, so I will leave it there.


This is exactly the area of study known as neural coding. The representation of information is not universal and differs for different modality, different stages of information processing, and what not. It's an active area of research.

That said, for the peripheral nervous system, the simplest code is pretty much rate code with label line. In other words, knowing which nerve fiber is firing what rate carries much of the sensory information. There are adaptation and some timing information if you look closer.


18.3: Sensory Perception- Taste and Olfaction

A major role of sensory receptors is to help us learn about the environment around us, or about the state of our internal environment. Stimuli from varying sources, and of different types, are received and changed into the electrochemical signals of the nervous system. This occurs when a stimulus changes the cell membrane potential of a sensory neuron. The stimulus causes the sensory cell to produce an action potential that is relayed into the central nervous system (CNS), where it is integrated with other sensory information&mdashor sometimes higher cognitive functions&mdashto become a conscious perception of that stimulus. The central integration may then lead to a motor response.

Describing sensory function with the term sensation or perception is a deliberate distinction. Sensation is the activation of sensory receptor cells at the level of the stimulus. Perception is the central processing of sensory stimuli into a meaningful pattern. Perception is dependent on sensation, but not all sensations are perceived. Receptors are the cells or structures that detect sensations. A receptor cell is changed directly by a stimulus. A transmembrane protein receptor is a protein in the cell membrane that mediates a physiological change in a neuron, most often through the opening of ion channels or changes in the cell signaling processes. Transmembrane receptors are activated by chemicals called ligands.

For example, a molecule in food can serve as a ligand for taste receptors. Other transmembrane proteins, which are not accurately called receptors, are sensitive to mechanical or thermal changes. Physical changes in these proteins increase ion flow across the membrane, and can generate an action potential or a graded potential in the sensory neurons.


Reception

The first step in sensation is reception , which is the activation of sensory receptors by stimuli such as mechanical stimuli (being bent or squished, for example), chemicals, or temperature. The receptor can then respond to the stimuli. The region in space in which a given sensory receptor can respond to a stimulus, be it far away or in contact with the body, is that receptor&rsquos receptive field . Think for a moment about the differences in receptive fields for the different senses. For the sense of touch, a stimulus must come into contact with body. For the sense of hearing, a stimulus can be a moderate distance away (some baleen whale sounds can propagate for many kilometers). For vision, a stimulus can be very far away for example, the visual system perceives light from stars at enormous distances.


Transduction

The most fundamental function of a sensory system is the translation of a sensory signal to an electrical signal in the nervous system. This takes place at the sensory receptor, and the change in electrical potential that is produced is called the receptor potential . How is sensory input, such as pressure on the skin, changed to a receptor potential? In this example, a type of receptor called a mechanoreceptor (as shown in [link]) possesses specialized membranes that respond to pressure. Disturbance of these dendrites by compressing them or bending them opens gated ion channels in the plasma membrane of the sensory neuron, changing its electrical potential. Recall that in the nervous system, a positive change of a neuron’s electrical potential (also called the membrane potential), depolarizes the neuron. Receptor potentials are graded potentials: the magnitude of these graded (receptor) potentials varies with the strength of the stimulus. If the magnitude of depolarization is sufficient (that is, if membrane potential reaches a threshold), the neuron will fire an action potential. In most cases, the correct stimulus impinging on a sensory receptor will drive membrane potential in a positive direction, although for some receptors, such as those in the visual system, this is not always the case.


Sensory receptors for different senses are very different from each other, and they are specialized according to the type of stimulus they sense: they have receptor specificity. For example, touch receptors, light receptors, and sound receptors are each activated by different stimuli. Touch receptors are not sensitive to light or sound they are sensitive only to touch or pressure. However, stimuli may be combined at higher levels in the brain, as happens with olfaction, contributing to our sense of taste.


Transduction and Perception

Transduction is the process that converts a sensory signal to an electrical signal to be processed in a specialized area in the brain.

Learning Objectives

Explain how stimuli are converted to signals that are carried to the central nervous system

Key Takeaways

Key Points

  • Sensory signals are converted to electrical signals via depolarization of sensory neuron membranes upon stimulus of the receptor, which causes opening of gated ion channels that cause the membrane potential to reach its threshold.
  • The receptor potentials are classified as graded potentials the magnitude of these potentials is dependent on the strength of the stimulus.
  • The sensory system shows receptor specificity although stimuli can be combined in processing regions of the brain, a specific receptor will only be activated by its specific stimulus.
  • The brain contains specific processing regions (such as the somatosensory, visual, and auditory regions) that are dedicated to processing the information which has previously passed through the thalamus, the ‘clearinghouse and relay station’ for both sensory and motor signals.
  • The four major components of encoding and transmitting sensory information include: the type of stimulus, the stimulus location within the receptive field, the duration, and the intensity of the stimulus.

Key Terms

  • membrane potential: the difference in electrical potential across the enclosing membrane of a cell
  • action potential: a short term change in the electrical potential that travels along a cell
  • transduction: the translation of a sensory signal in the sensory system to an electrical signal in the nervous system

Transduction

The most fundamental function of a sensory system is the translation of a sensory signal to an electrical signal in the nervous system. This takes place at the sensory receptor. The change in electrical potential that is produced is called the receptor potential. How is sensory input, such as pressure on the skin, changed to a receptor potential? As an example, a type of receptor called a mechanoreceptor possesses specialized membranes that respond to pressure. Disturbance of these dendrites by compressing them or bending them opens gated ion channels in the plasma membrane of the sensory neuron, changing its electrical potential. In the nervous system, a positive change of a neuron’s electrical potential (also called the membrane potential), depolarizes the neuron. Receptor potentials are graded potentials: the magnitude of these graded (receptor) potentials varies with the strength of the stimulus. If the magnitude of depolarization is sufficient (that is, if membrane potential reaches a threshold), the neuron will fire an action potential. In most cases, the correct stimulus impinging on a sensory receptor will drive membrane potential in a positive direction, although for some receptors, such as those in the visual system, this is not always the case.

Mechanoreceptor activation: (a) Mechanosensitive ion channels are gated ion channels that respond to mechanical deformation of the plasma membrane. A mechanosensitive channel is connected to the plasma membrane and the cytoskeleton by hair-like tethers. When pressure causes the extracellular matrix to move, the channel opens, allowing ions to enter or exit the cell. (b) Stereocilia in the human ear are connected to mechanosensitive ion channels. When a sound causes the stereocilia to move, mechanosensitive ion channels transduce the signal to the cochlear nerve.

Sensory receptors for the various senses work differently from each other. They are specialized according to the type of stimulus they sense thus, they have receptor specificity. For example, touch receptors, light receptors, and sound receptors are each activated by different stimuli. Touch receptors are not sensitive to light or sound they are sensitive only to touch or pressure. However, stimuli may be combined at higher levels in the brain, as happens with olfaction, contributing to our sense of taste.

Encoding and Transmission of Sensory Information

Four aspects of sensory information are encoded by sensory systems: the type of stimulus, the location of the stimulus in the receptive field, the duration of the stimulus, and the relative intensity of the stimulus. Thus, action potentials transmitted over a sensory receptor’s afferent axons encode one type of stimulus. This segregation of the senses is preserved in other sensory circuits. For example, auditory receptors transmit signals over their own dedicated system. The electrical activity in the axons of the auditory receptors will be interpreted by the brain as an auditory stimulus: a sound.

The intensity of a stimulus is often encoded in the rate of action potentials produced by the sensory receptor. Thus, an intense stimulus will produce a more rapid train of action potentials. Reducing the stimulus will likewise slow the rate of production of action potentials. A second way in which intensity is encoded is by the number of receptors activated. An intense stimulus might initiate action potentials in a large number of adjacent receptors, while a less intense stimulus might stimulate fewer receptors. Integration of sensory information begins as soon as the information is received in the central nervous system.

Perception

Perception is an individual’s interpretation of a sensation. Although perception relies on the activation of sensory receptors, perception happens, not at the level of the sensory receptor, but at the brain level. The brain distinguishes sensory stimuli through a sensory pathway: action potentials from sensory receptors travel along neurons that are dedicated to a particular stimulus.

All sensory signals, except those from the olfactory system, are transmitted though the central nervous system: they are routed to the thalamus and to the appropriate region of the cortex. The thalamus is a structure in the forebrain that serves as a clearinghouse and relay station for sensory (as well as motor) signals. When the sensory signal exits the thalamus, it is conducted to the specific area of the cortex dedicated to processing that particular sense.

Sensation processing: The brain has dedicated areas to the processing of stimuli, including: (a) thalamus and (b) the auditory, visual and somatosensory processing regions.


Transduction

The most fundamental function of a sensory system is the translation of a sensory signal to an electrical signal in the nervous system. This takes place at the sensory receptor, and the change in electrical potential that is produced is called the receptor potential. How is sensory input, such as pressure on the skin, changed to a receptor potential? In this example, a type of receptor called a mechanoreceptor (as shown in Figure 1) possesses specialized membranes that respond to pressure. Disturbance of these dendrites by compressing them or bending them opens gated ion channels in the plasma membrane of the sensory neuron, changing its electrical potential. Recall that in the nervous system, a positive change of a neuron’s electrical potential (also called the membrane potential), depolarizes the neuron. Receptor potentials are graded potentials: the magnitude of these graded (receptor) potentials varies with the strength of the stimulus. If the magnitude of depolarization is sufficient (that is, if membrane potential reaches a threshold), the neuron will fire an action potential. In most cases, the correct stimulus impinging on a sensory receptor will drive membrane potential in a positive direction, although for some receptors, such as those in the visual system, this is not always the case.

Figure 1. (a) Mechanosensitive ion channels are gated ion channels that respond to mechanical deformation of the plasma membrane. A mechanosensitive channel is connected to the plasma membrane and the cytoskeleton by hair-like tethers. When pressure causes the extracellular matrix to move, the channel opens, allowing ions to enter or exit the cell. (b) Stereocilia in the human ear are connected to mechanosensitive ion channels. When a sound causes the stereocilia to move, mechanosensitive ion channels transduce the signal to the cochlear nerve.

Sensory receptors for different senses are very different from each other, and they are specialized according to the type of stimulus they sense: they have receptor specificity. For example, touch receptors, light receptors, and sound receptors are each activated by different stimuli. Touch receptors are not sensitive to light or sound they are sensitive only to touch or pressure. However, stimuli may be combined at higher levels in the brain, as happens with olfaction, contributing to our sense of taste.

Encoding and Transmission of Sensory Information

Four aspects of sensory information are encoded by sensory systems: the type of stimulus, the location of the stimulus in the receptive field, the duration of the stimulus, and the relative intensity of the stimulus. Thus, action potentials transmitted over a sensory receptor’s afferent axons encode one type of stimulus, and this segregation of the senses is preserved in other sensory circuits. For example, auditory receptors transmit signals over their own dedicated system, and electrical activity in the axons of the auditory receptors will be interpreted by the brain as an auditory stimulus—a sound.

The intensity of a stimulus is often encoded in the rate of action potentials produced by the sensory receptor. Thus, an intense stimulus will produce a more rapid train of action potentials, and reducing the stimulus will likewise slow the rate of production of action potentials. A second way in which intensity is encoded is by the number of receptors activated. An intense stimulus might initiate action potentials in a large number of adjacent receptors, while a less intense stimulus might stimulate fewer receptors. Integration of sensory information begins as soon as the information is received in the CNS, and the brain will further process incoming signals.


Nerve Cells and Synapses: Grade 9 Understanding for IGCSE Biology 2.88 2.89

There is very little in the iGCSE specification about nerve cells and synapses. This is a shame since neuroscience is going to be one of the massive growth areas in Biology in the 21st century. There is a syllabus point about reflex acs and I draw your attention to this blog post about that: https://pmgbiology.wordpress.com/2014/04/22/a-simple-reflex-arc/

But in this new post I am going to give you a tiny bit more detail about the types of nerve cells (neurones) that you might encounter, together with an explanation about the most important component of the nervous systems: the chemical synapse.

Neurones are the cells in the nervous system that are adapted to send nerve impulses. You won’t fully understand what the nerve impulse is until year 13 but it is correct so that it is a temporary electrical event that can be transmitted over large distances within a cell with no loss of signal strength. The upshot of this is that neurones can be very long indeed…..

There are three basic types of neurone that are grouped according to their function:

Motor neurones (efferent neurones) take nerve impulses from the CNS to skeletal muscle causing it to contract

Sensory neurones (afferent neurones) take nerve impulses from sensory receptors into the CNS

Relay (or sometimes Inter) neurones are found within the CNS and basically link sensory to motor neurones.

These three types of neurone also have different structures although many features are shared….

This is a diagram of a generalised motor neurone: I know it is a motor neurone since the cell body is at one end of the cell. The cell body contains the nucleus, most of the cytoplasm and many organelles. Structures that carry a nerve impulse towards the cell body are called dendrites (if there are lots of them) and a dendron if there is only one. The axon is the long thin projection of the cell that takes the nerve impulse away from the cell body. The axon will finish with a collection of nerve endings or synapses.

Neurones can only send nerve impulses in one direction. In the diagram above these two cells can only send impulses from left to right as shown. This is due to the nature of the junction between the cells, the synapse (see later on….)

The diagram above shows a sensory neurone. You can tell this because it has receptors at one end collecting sensory information to take to the CNS. The position of the cell body is also different in sensory neurones: in all sensory neurones the cell body is off at right angles to the axon/dendron.

You can see from the diagrams that motor and sensory neurones tend to be surrounded by a myelin sheath. Myelin is a type of lipid that acts as an insulator, speeding up the nerve impulse from around 0.5m/s in unmyelinated neurones to about 100 m/s in the fastest myelinated ones. The myelin sheath is made from a whole load of cells (glial cells) but there are gaps between glial cells called nodes of Ranvier. These will become important in Y12/13 when you study how the impulse manages to travel so fast in a myelinated neurone.

Relay neurones, also known as interneurones, have a much simpler structure. They are only found in the CNS, almost always unmyelinated and have their cell body in the centre of the cell.

The diagram above shows the three types of neurone and indeed how they are linked up in a simple reflex arc. The artist hasn’t really shown the interneurone structure very well, but it was the best I could find just now…..

Nerve cells are linked together (and indeed linked to muscle cells) by structures known as synapses. There are a lot of synapses in your nervous system. The human brain contains around 100 billion neurones and each neurone is linked by synapses to around 1000 other cells: a grand total of 100 trillion synapses. 100 000 000 000 000 is a big number.

The big idea with synapses is that the two neurones do not actually touch. There is a tiny gap called the synaptic cleft between the cells. The nerve impulse does not cross this tiny gap as an electrical event but instead there are chemicals called neurotransmitters that diffuse across the synaptic cleft.

The nerve impulse arrives at the axon terminal of the presynaptic neurone. Inside this swelling are thousands of tiny membrane packets called vesicles, each one packed with a million or so molecules of neurotransmitter. When the impulse arrives at the terminal, a few hundred of these vesicles are stimulated to move towards and then fuse with the cell membrane, releasing the neurotransmitter into the synaptic cleft. The neurotransmitter will diffuse rapidly across the gap and when it reaches the post-synaptic membrane, it binds to specific receptor molecules embedded in the post-synaptic membrane. The binding of the neurotransmitter to the receptor often causes a new nerve impulse to form in the post-synaptic cell.

These chemical synapses are really beautiful things. They ensure the nerve impulse can only cross the synapse in one direction (can you see why?) and also they are infinitely flexible. They can be strengthened and weakened, their effects can be added together and when this is all put together, complex behaviour can emerge. I am going to exhibit some complex behaviour now by choosing to take my dogs for a walk… And it all happened due to synapses in my brain!


Biology A Level - Chapter 14 - Response to Stimuli

2 main functions:
-collect, process and respond to information in environment.
-co-ordinate the working of different organs and cells in the body.

-sends information TO the CNS from the outside world.
-transmits messages FROM CNS to muscles and glands in body.

2 parts:
-sensory nervous system (SNS)
-motor nervous system (MNS)

-carries nerve impulses away from the CNS, TO the effectors and receptors.

-governs vital functions in body such as breathing, digestion, heart rate, stress responses (AUTOMATIC processes).
-transmits nerve impulses to glands and muscle inside the body.
-is involuntary.

1. STIMULUS - an external or internal change that stimulates the receptors.
2. RECEPTOR - detects stimulus, and sends nerve impulse along sensory neurone.
3. SENSORY NEURONE- carries nerve impulse to the CNS.
4. COORDINATOR (RELAY NEURONE) - resides in the CNS and carries the nerve impulse from the sensory to the motor neurone.
5. MOTOR NEURONE - carries nerve impulses from CNS to an effector.
6. EFFECTOR (eg. muscle) - is stimulated by nerve impulse and carries out the response.
7. RESPONSE - (eg. muscle contraction).

2. The potential difference that exists because of this is called the RESTING POTENTIAL (is always quoted as inside vs outside).
The cell surface membrane is POLARISED.

3. In the presence of a stimulus, the K+ channel remains closed and the Na+ channel OPENS. Na+ ions DIFFUSE INTO THE RECEPTORS DOWN AN ELECTROCHEMICAL GRADIENT.

4. This causes the membrane to DEPOLARISE (the inside becomes more POSITIVE than the outside).

5. This depolarisation is called a GENERATOR POTENTIAL.

6. If the generator potential reaches a THRESHOLD value (due to a strong stimulus), then it would open VOLTAGE-GATED Na+ channels in the membrane of the SENSORY NEURONE.


Sensory Perception

A major role of sensory receptors is to help us learn about the environment around us, or about the state of our internal environment. Stimuli from varying sources, and of different types, are received and changed into the electrochemical signals of the nervous system. This occurs when a stimulus changes the cell membrane potential of a sensory neuron. The stimulus causes the sensory cell to produce an action potential that is relayed into the central nervous system (CNS), where it is integrated with other sensory information—or sometimes higher cognitive functions—to become a conscious perception of that stimulus. The central integration may then lead to a motor response.

Describing sensory function with the term sensation or perception is a deliberate distinction. Sensation is the activation of sensory receptor cells at the level of the stimulus. Perception is the central processing of sensory stimuli into a meaningful pattern. Perception is dependent on sensation, but not all sensations are perceived. Receptors are the cells or structures that detect sensations. A receptor cell is changed directly by a stimulus. A transmembrane protein receptor is a protein in the cell membrane that mediates a physiological change in a neuron, most often through the opening of ion channels or changes in the cell signaling processes. Transmembrane receptors are activated by chemicals called ligands. For example, a molecule in food can serve as a ligand for taste receptors. Other transmembrane proteins, which are not accurately called receptors, are sensitive to mechanical or thermal changes. Physical changes in these proteins increase ion flow across the membrane, and can generate an action potential or a graded potential in the sensory neurons.

Sensory Receptors

Stimuli in the environment activate specialized receptor cells in the peripheral nervous system. Different types of stimuli are sensed by different types of receptor cells. Receptor cells can be classified into types on the basis of three different criteria: cell type, position, and function. Receptors can be classified structurally on the basis of cell type and their position in relation to stimuli they sense. They can also be classified functionally on the basis of the transduction of stimuli, or how the mechanical stimulus, light, or chemical changed the cell membrane potential.

Structural Receptor Types

The cells that interpret information about the environment can be either (1) a neuron that has a free nerve ending , with dendrites embedded in tissue that would receive a sensation (2) a neuron that has an encapsulated ending in which the sensory nerve endings are encapsulated in connective tissue that enhances their sensitivity or (3) a specialized receptor cell , which has distinct structural components that interpret a specific type of stimulus ((Figure)). The pain and temperature receptors in the dermis of the skin are examples of neurons that have free nerve endings. Also located in the dermis of the skin are lamellated corpuscles, neurons with encapsulated nerve endings that respond to pressure and touch. The cells in the retina that respond to light stimuli are an example of a specialized receptor, a photoreceptor .

Another way that receptors can be classified is based on their location relative to the stimuli. An exteroceptor is a receptor that is located near a stimulus in the external environment, such as the somatosensory receptors that are located in the skin. An interoceptor is one that interprets stimuli from internal organs and tissues, such as the receptors that sense the increase in blood pressure in the aorta or carotid sinus. Finally, a proprioceptor is a receptor located near a moving part of the body, such as a muscle, that interprets the positions of the tissues as they move.

Functional Receptor Types

A third classification of receptors is by how the receptor transduces stimuli into membrane potential changes. Stimuli are of three general types. Some stimuli are ions and macromolecules that affect transmembrane receptor proteins when these chemicals diffuse across the cell membrane. Some stimuli are physical variations in the environment that affect receptor cell membrane potentials. Other stimuli include the electromagnetic radiation from visible light. For humans, the only electromagnetic energy that is perceived by our eyes is visible light. Some other organisms have receptors that humans lack, such as the heat sensors of snakes, the ultraviolet light sensors of bees, or magnetic receptors in migratory birds.

Receptor cells can be further categorized on the basis of the type of stimuli they transduce. Chemical stimuli can be interpreted by a chemoreceptor that interprets chemical stimuli, such as an object’s taste or smell. Osmoreceptors respond to solute concentrations of body fluids. Additionally, pain is primarily a chemical sense that interprets the presence of chemicals from tissue damage, or similar intense stimuli, through a nociceptor . Physical stimuli, such as pressure and vibration, as well as the sensation of sound and body position (balance), are interpreted through a mechanoreceptor . Another physical stimulus that has its own type of receptor is temperature, which is sensed through a thermoreceptor that is either sensitive to temperatures above (heat) or below (cold) normal body temperature.

Sensory Modalities

Ask anyone what the senses are, and they are likely to list the five major senses—taste, smell, touch, hearing, and sight. However, these are not all of the senses. The most obvious omission from this list is balance. Also, what is referred to simply as touch can be further subdivided into pressure, vibration, stretch, and hair-follicle position, on the basis of the type of mechanoreceptors that perceive these touch sensations. Other overlooked senses include temperature perception by thermoreceptors and pain perception by nociceptors.

Within the realm of physiology, senses can be classified as either general or specific. A general sense is one that is distributed throughout the body and has receptor cells within the structures of other organs. Mechanoreceptors in the skin, muscles, or the walls of blood vessels are examples of this type. General senses often contribute to the sense of touch, as described above, or to proprioception (body movement) and kinesthesia (body movement), or to a visceral sense , which is most important to autonomic functions. A special sense is one that has a specific organ devoted to it, namely the eye, inner ear, tongue, or nose.

Each of the senses is referred to as a sensory modality . Modality refers to the way that information is encoded, which is similar to the idea of transduction. The main sensory modalities can be described on the basis of how each is transduced. The chemical senses are taste and smell. The general sense that is usually referred to as touch includes chemical sensation in the form of nociception, or pain. Pressure, vibration, muscle stretch, and the movement of hair by an external stimulus, are all sensed by mechanoreceptors. Hearing and balance are also sensed by mechanoreceptors. Finally, vision involves the activation of photoreceptors.

Listing all the different sensory modalities, which can number as many as 17, involves separating the five major senses into more specific categories, or submodalities , of the larger sense. An individual sensory modality represents the sensation of a specific type of stimulus. For example, the general sense of touch, which is known as somatosensation , can be separated into light pressure, deep pressure, vibration, itch, pain, temperature, or hair movement.

Gustation (Taste)

Only a few recognized submodalities exist within the sense of taste, or gustation . Until recently, only four tastes were recognized: sweet, salty, sour, and bitter. Research at the turn of the 20th century led to recognition of the fifth taste, umami, during the mid-1980s. Umami is a Japanese word that means “delicious taste,” and is often translated to mean savory. Very recent research has suggested that there may also be a sixth taste for fats, or lipids.

Gustation is the special sense associated with the tongue. The surface of the tongue, along with the rest of the oral cavity, is lined by a stratified squamous epithelium. Raised bumps called papillae (singular = papilla) contain the structures for gustatory transduction. There are four types of papillae, based on their appearance ((Figure)): circumvallate, foliate, filiform, and fungiform. Within the structure of the papillae are taste buds that contain specialized gustatory receptor cells for the transduction of taste stimuli. These receptor cells are sensitive to the chemicals contained within foods that are ingested, and they release neurotransmitters based on the amount of the chemical in the food. Neurotransmitters from the gustatory cells can activate sensory neurons in the facial, glossopharyngeal, and vagus cranial nerves.

Salty taste is simply the perception of sodium ions (Na + ) in the saliva. When you eat something salty, the salt crystals dissociate into the component ions Na + and Cl – , which dissolve into the saliva in your mouth. The Na + concentration becomes high outside the gustatory cells, creating a strong concentration gradient that drives the diffusion of the ion into the cells. The entry of Na + into these cells results in the depolarization of the cell membrane and the generation of a receptor potential.

Sour taste is the perception of H + concentration. Just as with sodium ions in salty flavors, these hydrogen ions enter the cell and trigger depolarization. Sour flavors are, essentially, the perception of acids in our food. Increasing hydrogen ion concentrations in the saliva (lowering saliva pH) triggers progressively stronger graded potentials in the gustatory cells. For example, orange juice—which contains citric acid—will taste sour because it has a pH value of approximately 3. Of course, it is often sweetened so that the sour taste is masked.

The first two tastes (salty and sour) are triggered by the cations Na + and H + . The other tastes result from food molecules binding to a G protein–coupled receptor. A G protein signal transduction system ultimately leads to depolarization of the gustatory cell. The sweet taste is the sensitivity of gustatory cells to the presence of glucose dissolved in the saliva. Other monosaccharides such as fructose, or artificial sweeteners such as aspartame (NutraSweet™), saccharine, or sucralose (Splenda™) also activate the sweet receptors. The affinity for each of these molecules varies, and some will taste sweeter than glucose because they bind to the G protein–coupled receptor differently.

Bitter taste is similar to sweet in that food molecules bind to G protein–coupled receptors. However, there are a number of different ways in which this can happen because there are a large diversity of bitter-tasting molecules. Some bitter molecules depolarize gustatory cells, whereas others hyperpolarize gustatory cells. Likewise, some bitter molecules increase G protein activation within the gustatory cells, whereas other bitter molecules decrease G protein activation. The specific response depends on which molecule is binding to the receptor.

One major group of bitter-tasting molecules are alkaloids. Alkaloids are nitrogen containing molecules that are commonly found in bitter-tasting plant products, such as coffee, hops (in beer), tannins (in wine), tea, and aspirin. By containing toxic alkaloids, the plant is less susceptible to microbe infection and less attractive to herbivores.

Therefore, the function of bitter taste may primarily be related to stimulating the gag reflex to avoid ingesting poisons. Because of this, many bitter foods that are normally ingested are often combined with a sweet component to make them more palatable (cream and sugar in coffee, for example). The highest concentration of bitter receptors appear to be in the posterior tongue, where a gag reflex could still spit out poisonous food.

The taste known as umami is often referred to as the savory taste. Like sweet and bitter, it is based on the activation of G protein–coupled receptors by a specific molecule. The molecule that activates this receptor is the amino acid L-glutamate. Therefore, the umami flavor is often perceived while eating protein-rich foods. Not surprisingly, dishes that contain meat are often described as savory.

Once the gustatory cells are activated by the taste molecules, they release neurotransmitters onto the dendrites of sensory neurons. These neurons are part of the facial and glossopharyngeal cranial nerves, as well as a component within the vagus nerve dedicated to the gag reflex. The facial nerve connects to taste buds in the anterior third of the tongue. The glossopharyngeal nerve connects to taste buds in the posterior two thirds of the tongue. The vagus nerve connects to taste buds in the extreme posterior of the tongue, verging on the pharynx, which are more sensitive to noxious stimuli such as bitterness.

Watch this video to learn about Dr. Danielle Reed of the Monell Chemical Senses Center in Philadelphia, Pennsylvania, who became interested in science at an early age because of her sensory experiences. She recognized that her sense of taste was unique compared with other people she knew. Now, she studies the genetic differences between people and their sensitivities to taste stimuli. In the video, there is a brief image of a person sticking out their tongue, which has been covered with a colored dye. This is how Dr. Reed is able to visualize and count papillae on the surface of the tongue. People fall into two groups known as “tasters” and “non-tasters” based on the density of papillae on their tongue, which also indicates the number of taste buds. Non-tasters can taste food, but they are not as sensitive to certain tastes, such as bitterness. Dr. Reed discovered that she is a non-taster, which explains why she perceived bitterness differently than other people she knew. Are you very sensitive to tastes? Can you see any similarities among the members of your family?

Olfaction (Smell)

Like taste, the sense of smell, or olfaction , is also responsive to chemical stimuli. The olfactory receptor neurons are located in a small region within the superior nasal cavity ((Figure)). This region is referred to as the olfactory epithelium and contains bipolar sensory neurons. Each olfactory sensory neuron has dendrites that extend from the apical surface of the epithelium into the mucus lining the cavity. As airborne molecules are inhaled through the nose, they pass over the olfactory epithelial region and dissolve into the mucus. These odorant molecules bind to proteins that keep them dissolved in the mucus and help transport them to the olfactory dendrites. The odorant–protein complex binds to a receptor protein within the cell membrane of an olfactory dendrite. These receptors are G protein–coupled, and will produce a graded membrane potential in the olfactory neurons.

The axon of an olfactory neuron extends from the basal surface of the epithelium, through an olfactory foramen in the cribriform plate of the ethmoid bone, and into the brain. The group of axons called the olfactory tract connect to the olfactory bulb on the ventral surface of the frontal lobe. From there, the axons split to travel to several brain regions. Some travel to the cerebrum, specifically to the primary olfactory cortex that is located in the inferior and medial areas of the temporal lobe. Others project to structures within the limbic system and hypothalamus, where smells become associated with long-term memory and emotional responses. This is how certain smells trigger emotional memories, such as the smell of food associated with one’s birthplace. Smell is the one sensory modality that does not synapse in the thalamus before connecting to the cerebral cortex. This intimate connection between the olfactory system and the cerebral cortex is one reason why smell can be a potent trigger of memories and emotion.

The nasal epithelium, including the olfactory cells, can be harmed by airborne toxic chemicals. Therefore, the olfactory neurons are regularly replaced within the nasal epithelium, after which the axons of the new neurons must find their appropriate connections in the olfactory bulb. These new axons grow along the axons that are already in place in the cranial nerve.

Olfactory System: Anosmia Blunt force trauma to the face, such as that common in many car accidents, can lead to the loss of the olfactory nerve, and subsequently, loss of the sense of smell. This condition is known as anosmia . When the frontal lobe of the brain moves relative to the ethmoid bone, the olfactory tract axons may be sheared apart. Professional fighters often experience anosmia because of repeated trauma to face and head. In addition, certain pharmaceuticals, such as antibiotics, can cause anosmia by killing all the olfactory neurons at once. If no axons are in place within the olfactory nerve, then the axons from newly formed olfactory neurons have no guide to lead them to their connections within the olfactory bulb. There are temporary causes of anosmia, as well, such as those caused by inflammatory responses related to respiratory infections or allergies.

Loss of the sense of smell can result in food tasting bland. A person with an impaired sense of smell may require additional spice and seasoning levels for food to be tasted. Anosmia may also be related to some presentations of mild depression, because the loss of enjoyment of food may lead to a general sense of despair.

The ability of olfactory neurons to replace themselves decreases with age, leading to age-related anosmia. This explains why some elderly people salt their food more than younger people do. However, this increased sodium intake can increase blood volume and blood pressure, increasing the risk of cardiovascular diseases in the elderly.

Audition (Hearing)

Hearing, or audition , is the transduction of sound waves into a neural signal that is made possible by the structures of the ear ((Figure)). The large, fleshy structure on the lateral aspect of the head is known as the auricle . Some sources will also refer to this structure as the pinna, though that term is more appropriate for a structure that can be moved, such as the external ear of a cat. The C-shaped curves of the auricle direct sound waves toward the auditory canal. The canal enters the skull through the external auditory meatus of the temporal bone. At the end of the auditory canal is the tympanic membrane , or ear drum, which vibrates after it is struck by sound waves. The auricle, ear canal, and tympanic membrane are often referred to as the external ear . The middle ear consists of a space spanned by three small bones called the ossicles . The three ossicles are the malleus , incus , and stapes , which are Latin names that roughly translate to hammer, anvil, and stirrup. The malleus is attached to the tympanic membrane and articulates with the incus. The incus, in turn, articulates with the stapes. The stapes is then attached to the inner ear , where the sound waves will be transduced into a neural signal. The middle ear is connected to the pharynx through the Eustachian tube, which helps equilibrate air pressure across the tympanic membrane. The tube is normally closed but will pop open when the muscles of the pharynx contract during swallowing or yawning.

The inner ear is often described as a bony labyrinth, as it is composed of a series of canals embedded within the temporal bone. It has two separate regions, the cochlea and the vestibule , which are responsible for hearing and balance, respectively. The neural signals from these two regions are relayed to the brain stem through separate fiber bundles. However, these two distinct bundles travel together from the inner ear to the brain stem as the vestibulocochlear nerve. Sound is transduced into neural signals within the cochlear region of the inner ear, which contains the sensory neurons of the spiral ganglia . These ganglia are located within the spiral-shaped cochlea of the inner ear. The cochlea is attached to the stapes through the oval window .

The oval window is located at the beginning of a fluid-filled tube within the cochlea called the scala vestibuli . The scala vestibuli extends from the oval window, travelling above the cochlear duct , which is the central cavity of the cochlea that contains the sound-transducing neurons. At the uppermost tip of the cochlea, the scala vestibuli curves over the top of the cochlear duct. The fluid-filled tube, now called the scala tympani , returns to the base of the cochlea, this time travelling under the cochlear duct. The scala tympani ends at the round window , which is covered by a membrane that contains the fluid within the scala. As vibrations of the ossicles travel through the oval window, the fluid of the scala vestibuli and scala tympani moves in a wave-like motion. The frequency of the fluid waves match the frequencies of the sound waves ((Figure)). The membrane covering the round window will bulge out or pucker in with the movement of the fluid within the scala tympani.

A cross-sectional view of the cochlea shows that the scala vestibuli and scala tympani run along both sides of the cochlear duct ((Figure)). The cochlear duct contains several organs of Corti , which tranduce the wave motion of the two scala into neural signals. The organs of Corti lie on top of the basilar membrane , which is the side of the cochlear duct located between the organs of Corti and the scala tympani. As the fluid waves move through the scala vestibuli and scala tympani, the basilar membrane moves at a specific spot, depending on the frequency of the waves. Higher frequency waves move the region of the basilar membrane that is close to the base of the cochlea. Lower frequency waves move the region of the basilar membrane that is near the tip of the cochlea.

The organs of Corti contain hair cells , which are named for the hair-like stereocilia extending from the cell’s apical surfaces ((Figure)). The stereocilia are an array of microvilli-like structures arranged from tallest to shortest. Protein fibers tether adjacent hairs together within each array, such that the array will bend in response to movements of the basilar membrane. The stereocilia extend up from the hair cells to the overlying tectorial membrane , which is attached medially to the organ of Corti. When the pressure waves from the scala move the basilar membrane, the tectorial membrane slides across the stereocilia. This bends the stereocilia either toward or away from the tallest member of each array. When the stereocilia bend toward the tallest member of their array, tension in the protein tethers opens ion channels in the hair cell membrane. This will depolarize the hair cell membrane, triggering nerve impulses that travel down the afferent nerve fibers attached to the hair cells. When the stereocilia bend toward the shortest member of their array, the tension on the tethers slackens and the ion channels close. When no sound is present, and the stereocilia are standing straight, a small amount of tension still exists on the tethers, keeping the membrane potential of the hair cell slightly depolarized.

View the University of Michigan WebScope to explore the tissue sample in greater detail. The basilar membrane is the thin membrane that extends from the central core of the cochlea to the edge. What is anchored to this membrane so that they can be activated by movement of the fluids within the cochlea?

As stated above, a given region of the basilar membrane will only move if the incoming sound is at a specific frequency. Because the tectorial membrane only moves where the basilar membrane moves, the hair cells in this region will also only respond to sounds of this specific frequency. Therefore, as the frequency of a sound changes, different hair cells are activated all along the basilar membrane. The cochlea encodes auditory stimuli for frequencies between 20 and 20,000 Hz, which is the range of sound that human ears can detect. The unit of Hertz measures the frequency of sound waves in terms of cycles produced per second. Frequencies as low as 20 Hz are detected by hair cells at the apex, or tip, of the cochlea. Frequencies in the higher ranges of 20 KHz are encoded by hair cells at the base of the cochlea, close to the round and oval windows ((Figure)). Most auditory stimuli contain a mixture of sounds at a variety of frequencies and intensities (represented by the amplitude of the sound wave). The hair cells along the length of the cochlear duct, which are each sensitive to a particular frequency, allow the cochlea to separate auditory stimuli by frequency, just as a prism separates visible light into its component colors.

Watch this video to learn more about how the structures of the ear convert sound waves into a neural signal by moving the “hairs,” or stereocilia, of the cochlear duct. Specific locations along the length of the duct encode specific frequencies, or pitches. The brain interprets the meaning of the sounds we hear as music, speech, noise, etc. Which ear structures are responsible for the amplification and transfer of sound from the external ear to the inner ear?

Watch this animation to learn more about the inner ear and to see the cochlea unroll, with the base at the back of the image and the apex at the front. Specific wavelengths of sound cause specific regions of the basilar membrane to vibrate, much like the keys of a piano produce sound at different frequencies. Based on the animation, where do frequencies—from high to low pitches—cause activity in the hair cells within the cochlear duct?

Equilibrium (Balance)

Along with audition, the inner ear is responsible for encoding information about equilibrium , the sense of balance. A similar mechanoreceptor—a hair cell with stereocilia—senses head position, head movement, and whether our bodies are in motion. These cells are located within the vestibule of the inner ear. Head position is sensed by the utricle and saccule , whereas head movement is sensed by the semicircular canals . The neural signals generated in the vestibular ganglion are transmitted through the vestibulocochlear nerve to the brain stem and cerebellum.

The utricle and saccule are both largely composed of macula tissue (plural = maculae). The macula is composed of hair cells surrounded by support cells. The stereocilia of the hair cells extend into a viscous gel called the otolithic membrane ((Figure)). On top of the otolithic membrane is a layer of calcium carbonate crystals, called otoliths. The otoliths essentially make the otolithic membrane top-heavy. The otolithic membrane moves separately from the macula in response to head movements. Tilting the head causes the otolithic membrane to slide over the macula in the direction of gravity. The moving otolithic membrane, in turn, bends the sterocilia, causing some hair cells to depolarize as others hyperpolarize. The exact position of the head is interpreted by the brain based on the pattern of hair-cell depolarization.

The semicircular canals are three ring-like extensions of the vestibule. One is oriented in the horizontal plane, whereas the other two are oriented in the vertical plane. The anterior and posterior vertical canals are oriented at approximately 45 degrees relative to the sagittal plane ((Figure)). The base of each semicircular canal, where it meets with the vestibule, connects to an enlarged region known as the ampulla . The ampulla contains the hair cells that respond to rotational movement, such as turning the head while saying “no.” The stereocilia of these hair cells extend into the cupula , a membrane that attaches to the top of the ampulla. As the head rotates in a plane parallel to the semicircular canal, the fluid lags, deflecting the cupula in the direction opposite to the head movement. The semicircular canals contain several ampullae, with some oriented horizontally and others oriented vertically. By comparing the relative movements of both the horizontal and vertical ampullae, the vestibular system can detect the direction of most head movements within three-dimensional (3-D) space.

Somatosensation (Touch)

Somatosensation is considered a general sense, as opposed to the special senses discussed in this section. Somatosensation is the group of sensory modalities that are associated with touch, proprioception, and interoception. These modalities include pressure, vibration, light touch, tickle, itch, temperature, pain, proprioception, and kinesthesia. This means that its receptors are not associated with a specialized organ, but are instead spread throughout the body in a variety of organs. Many of the somatosensory receptors are located in the skin, but receptors are also found in muscles, tendons, joint capsules, ligaments, and in the walls of visceral organs.

Two types of somatosensory signals that are transduced by free nerve endings are pain and temperature. These two modalities use thermoreceptors and nociceptors to transduce temperature and pain stimuli, respectively. Temperature receptors are stimulated when local temperatures differ from body temperature. Some thermoreceptors are sensitive to just cold and others to just heat. Nociception is the sensation of potentially damaging stimuli. Mechanical, chemical, or thermal stimuli beyond a set threshold will elicit painful sensations. Stressed or damaged tissues release chemicals that activate receptor proteins in the nociceptors. For example, the sensation of heat associated with spicy foods involves capsaicin , the active molecule in hot peppers. Capsaicin molecules bind to a transmembrane ion channel in nociceptors that is sensitive to temperatures above 37°C. The dynamics of capsaicin binding with this transmembrane ion channel is unusual in that the molecule remains bound for a long time. Because of this, it will decrease the ability of other stimuli to elicit pain sensations through the activated nociceptor. For this reason, capsaicin can be used as a topical analgesic, such as in products such as Icy Hot™.

If you drag your finger across a textured surface, the skin of your finger will vibrate. Such low frequency vibrations are sensed by mechanoreceptors called Merkel cells, also known as type I cutaneous mechanoreceptors. Merkel cells are located in the stratum basale of the epidermis. Deep pressure and vibration is transduced by lamellated (Pacinian) corpuscles, which are receptors with encapsulated endings found deep in the dermis, or subcutaneous tissue. Light touch is transduced by the encapsulated endings known as tactile (Meissner) corpuscles. Follicles are also wrapped in a plexus of nerve endings known as the hair follicle plexus. These nerve endings detect the movement of hair at the surface of the skin, such as when an insect may be walking along the skin. Stretching of the skin is transduced by stretch receptors known as bulbous corpuscles. Bulbous corpuscles are also known as Ruffini corpuscles, or type II cutaneous mechanoreceptors.

Other somatosensory receptors are found in the joints and muscles. Stretch receptors monitor the stretching of tendons, muscles, and the components of joints. For example, have you ever stretched your muscles before or after exercise and noticed that you can only stretch so far before your muscles spasm back to a less stretched state? This spasm is a reflex that is initiated by stretch receptors to avoid muscle tearing. Such stretch receptors can also prevent over-contraction of a muscle. In skeletal muscle tissue, these stretch receptors are called muscle spindles. Golgi tendon organs similarly transduce the stretch levels of tendons. Bulbous corpuscles are also present in joint capsules, where they measure stretch in the components of the skeletal system within the joint. The types of nerve endings, their locations, and the stimuli they transduce are presented in (Figure).

*No corresponding eponymous name.
Mechanoreceptors of Somatosensation
Name Historical (eponymous) name Location(s) Stimuli
Free nerve endings * Dermis, cornea, tongue, joint capsules, visceral organs Pain, temperature, mechanical deformation
Mechanoreceptors Merkel’s discs Epidermal–dermal junction, mucosal membranes Low frequency vibration (5–15 Hz)
Bulbous corpuscle Ruffini’s corpuscle Dermis, joint capsules Stretch
Tactile corpuscle Meissner’s corpuscle Papillary dermis, especially in the fingertips and lips Light touch, vibrations below 50 Hz
Lamellated corpuscle Pacinian corpuscle Deep dermis, subcutaneous tissue Deep pressure, high-frequency vibration (around 250 Hz)
Hair follicle plexus * Wrapped around hair follicles in the dermis Movement of hair
Muscle spindle * In line with skeletal muscle fibers Muscle contraction and stretch
Tendon stretch organ Golgi tendon organ In line with tendons Stretch of tendons

Vision

Vision is the special sense of sight that is based on the transduction of light stimuli received through the eyes. The eyes are located within either orbit in the skull. The bony orbits surround the eyeballs, protecting them and anchoring the soft tissues of the eye ((Figure)). The eyelids, with lashes at their leading edges, help to protect the eye from abrasions by blocking particles that may land on the surface of the eye. The inner surface of each lid is a thin membrane known as the palpebral conjunctiva . The conjunctiva extends over the white areas of the eye (the sclera), connecting the eyelids to the eyeball. Tears are produced by the lacrimal gland , located beneath the lateral edges of the nose. Tears produced by this gland flow through the lacrimal duct to the medial corner of the eye, where the tears flow over the conjunctiva, washing away foreign particles.

Movement of the eye within the orbit is accomplished by the contraction of six extraocular muscles that originate from the bones of the orbit and insert into the surface of the eyeball ((Figure)). Four of the muscles are arranged at the cardinal points around the eye and are named for those locations. They are the superior rectus , medial rectus , inferior rectus , and lateral rectus . When each of these muscles contract, the eye to moves toward the contracting muscle. For example, when the superior rectus contracts, the eye rotates to look up. The superior oblique originates at the posterior orbit, near the origin of the four rectus muscles. However, the tendon of the oblique muscles threads through a pulley-like piece of cartilage known as the trochlea . The tendon inserts obliquely into the superior surface of the eye. The angle of the tendon through the trochlea means that contraction of the superior oblique rotates the eye medially. The inferior oblique muscle originates from the floor of the orbit and inserts into the inferolateral surface of the eye. When it contracts, it laterally rotates the eye, in opposition to the superior oblique. Rotation of the eye by the two oblique muscles is necessary because the eye is not perfectly aligned on the sagittal plane. When the eye looks up or down, the eye must also rotate slightly to compensate for the superior rectus pulling at approximately a 20-degree angle, rather than straight up. The same is true for the inferior rectus, which is compensated by contraction of the inferior oblique. A seventh muscle in the orbit is the levator palpebrae superioris , which is responsible for elevating and retracting the upper eyelid, a movement that usually occurs in concert with elevation of the eye by the superior rectus (see (Figure)).

The extraocular muscles are innervated by three cranial nerves. The lateral rectus, which causes abduction of the eye, is innervated by the abducens nerve. The superior oblique is innervated by the trochlear nerve. All of the other muscles are innervated by the oculomotor nerve, as is the levator palpebrae superioris. The motor nuclei of these cranial nerves connect to the brain stem, which coordinates eye movements.

The eye itself is a hollow sphere composed of three layers of tissue. The outermost layer is the fibrous tunic , which includes the white sclera and clear cornea . The sclera accounts for five sixths of the surface of the eye, most of which is not visible, though humans are unique compared with many other species in having so much of the “white of the eye” visible ((Figure)). The transparent cornea covers the anterior tip of the eye and allows light to enter the eye. The middle layer of the eye is the vascular tunic , which is mostly composed of the choroid, ciliary body, and iris. The choroid is a layer of highly vascularized connective tissue that provides a blood supply to the eyeball. The choroid is posterior to the ciliary body , a muscular structure that is attached to the lens by suspensory ligaments, or zonule fibers . These two structures bend the lens, allowing it to focus light on the back of the eye. Overlaying the ciliary body, and visible in the anterior eye, is the iris —the colored part of the eye. The iris is a smooth muscle that opens or closes the pupil , which is the hole at the center of the eye that allows light to enter. The iris constricts the pupil in response to bright light and dilates the pupil in response to dim light. The innermost layer of the eye is the neural tunic , or retina , which contains the nervous tissue responsible for photoreception.

The eye is also divided into two cavities: the anterior cavity and the posterior cavity. The anterior cavity is the space between the cornea and lens, including the iris and ciliary body. It is filled with a watery fluid called the aqueous humor . The posterior cavity is the space behind the lens that extends to the posterior side of the interior eyeball, where the retina is located. The posterior cavity is filled with a more viscous fluid called the vitreous humor .

The retina is composed of several layers and contains specialized cells for the initial processing of visual stimuli. The photoreceptors (rods and cones) change their membrane potential when stimulated by light energy. The change in membrane potential alters the amount of neurotransmitter that the photoreceptor cells release onto bipolar cells in the outer synaptic layer . It is the bipolar cell in the retina that connects a photoreceptor to a retinal ganglion cell (RGC) in the inner synaptic layer . There, amacrine cells additionally contribute to retinal processing before an action potential is produced by the RGC. The axons of RGCs, which lie at the innermost layer of the retina, collect at the optic disc and leave the eye as the optic nerve (see (Figure)). Because these axons pass through the retina, there are no photoreceptors at the very back of the eye, where the optic nerve begins. This creates a “blind spot” in the retina, and a corresponding blind spot in our visual field.

Note that the photoreceptors in the retina (rods and cones) are located behind the axons, RGCs, bipolar cells, and retinal blood vessels. A significant amount of light is absorbed by these structures before the light reaches the photoreceptor cells. However, at the exact center of the retina is a small area known as the fovea . At the fovea, the retina lacks the supporting cells and blood vessels, and only contains photoreceptors. Therefore, visual acuity , or the sharpness of vision, is greatest at the fovea. This is because the fovea is where the least amount of incoming light is absorbed by other retinal structures (see (Figure)). As one moves in either direction from this central point of the retina, visual acuity drops significantly. In addition, each photoreceptor cell of the fovea is connected to a single RGC. Therefore, this RGC does not have to integrate inputs from multiple photoreceptors, which reduces the accuracy of visual transduction. Toward the edges of the retina, several photoreceptors converge on RGCs (through the bipolar cells) up to a ratio of 50 to 1. The difference in visual acuity between the fovea and peripheral retina is easily evidenced by looking directly at a word in the middle of this paragraph. The visual stimulus in the middle of the field of view falls on the fovea and is in the sharpest focus. Without moving your eyes off that word, notice that words at the beginning or end of the paragraph are not in focus. The images in your peripheral vision are focused by the peripheral retina, and have vague, blurry edges and words that are not as clearly identified. As a result, a large part of the neural function of the eyes is concerned with moving the eyes and head so that important visual stimuli are centered on the fovea.

Light falling on the retina causes chemical changes to pigment molecules in the photoreceptors, ultimately leading to a change in the activity of the RGCs. Photoreceptor cells have two parts, the inner segment and the outer segment ((Figure)). The inner segment contains the nucleus and other common organelles of a cell, whereas the outer segment is a specialized region in which photoreception takes place. There are two types of photoreceptors—rods and cones—which differ in the shape of their outer segment. The rod-shaped outer segments of the rod photoreceptor contain a stack of membrane-bound discs that contain the photosensitive pigment rhodopsin . The cone-shaped outer segments of the cone photoreceptor contain their photosensitive pigments in infoldings of the cell membrane. There are three cone photopigments, called opsins , which are each sensitive to a particular wavelength of light. The wavelength of visible light determines its color. The pigments in human eyes are specialized in perceiving three different primary colors: red, green, and blue.

At the molecular level, visual stimuli cause changes in the photopigment molecule that lead to changes in membrane potential of the photoreceptor cell. A single unit of light is called a photon , which is described in physics as a packet of energy with properties of both a particle and a wave. The energy of a photon is represented by its wavelength, with each wavelength of visible light corresponding to a particular color. Visible light is electromagnetic radiation with a wavelength between 380 and 720 nm. Wavelengths of electromagnetic radiation longer than 720 nm fall into the infrared range, whereas wavelengths shorter than 380 nm fall into the ultraviolet range. Light with a wavelength of 380 nm is blue whereas light with a wavelength of 720 nm is dark red. All other colors fall between red and blue at various points along the wavelength scale.

Opsin pigments are actually transmembrane proteins that contain a cofactor known as retinal . Retinal is a hydrocarbon molecule related to vitamin A. When a photon hits retinal, the long hydrocarbon chain of the molecule is biochemically altered. Specifically, photons cause some of the double-bonded carbons within the chain to switch from a cis to a trans conformation. This process is called photoisomerization . Before interacting with a photon, retinal’s flexible double-bonded carbons are in the cis conformation. This molecule is referred to as 11-cis-retinal. A photon interacting with the molecule causes the flexible double-bonded carbons to change to the trans– conformation, forming all-trans-retinal, which has a straight hydrocarbon chain ((Figure)).

The shape change of retinal in the photoreceptors initiates visual transduction in the retina. Activation of retinal and the opsin proteins result in activation of a G protein. The G protein changes the membrane potential of the photoreceptor cell, which then releases less neurotransmitter into the outer synaptic layer of the retina. Until the retinal molecule is changed back to the 11-cis-retinal shape, the opsin cannot respond to light energy, which is called bleaching. When a large group of photopigments is bleached, the retina will send information as if opposing visual information is being perceived. After a bright flash of light, afterimages are usually seen in negative. The photoisomerization is reversed by a series of enzymatic changes so that the retinal responds to more light energy.

The opsins are sensitive to limited wavelengths of light. Rhodopsin, the photopigment in rods, is most sensitive to light at a wavelength of 498 nm. The three color opsins have peak sensitivities of 564 nm, 534 nm, and 420 nm corresponding roughly to the primary colors of red, green, and blue ((Figure)). The absorbance of rhodopsin in the rods is much more sensitive than in the cone opsins specifically, rods are sensitive to vision in low light conditions, and cones are sensitive to brighter conditions. In normal sunlight, rhodopsin will be constantly bleached while the cones are active. In a darkened room, there is not enough light to activate cone opsins, and vision is entirely dependent on rods. Rods are so sensitive to light that a single photon can result in an action potential from a rod’s corresponding RGC.

The three types of cone opsins, being sensitive to different wavelengths of light, provide us with color vision. By comparing the activity of the three different cones, the brain can extract color information from visual stimuli. For example, a bright blue light that has a wavelength of approximately 450 nm would activate the “red” cones minimally, the “green” cones marginally, and the “blue” cones predominantly. The relative activation of the three different cones is calculated by the brain, which perceives the color as blue. However, cones cannot react to low-intensity light, and rods do not sense the color of light. Therefore, our low-light vision is—in essence—in grayscale. In other words, in a dark room, everything appears as a shade of gray. If you think that you can see colors in the dark, it is most likely because your brain knows what color something is and is relying on that memory.

Watch this video to learn more about a transverse section through the brain that depicts the visual pathway from the eye to the occipital cortex. The first half of the pathway is the projection from the RGCs through the optic nerve to the lateral geniculate nucleus in the thalamus on either side. This first fiber in the pathway synapses on a thalamic cell that then projects to the visual cortex in the occipital lobe where “seeing,” or visual perception, takes place. This video gives an abbreviated overview of the visual system by concentrating on the pathway from the eyes to the occipital lobe. The video makes the statement (at 0:45) that “specialized cells in the retina called ganglion cells convert the light rays into electrical signals.” What aspect of retinal processing is simplified by that statement? Explain your answer.

Sensory Nerves

Once any sensory cell transduces a stimulus into a nerve impulse, that impulse has to travel along axons to reach the CNS. In many of the special senses, the axons leaving the sensory receptors have a topographical arrangement, meaning that the location of the sensory receptor relates to the location of the axon in the nerve. For example, in the retina, axons from RGCs in the fovea are located at the center of the optic nerve, where they are surrounded by axons from the more peripheral RGCs.

Spinal Nerves

Generally, spinal nerves contain afferent axons from sensory receptors in the periphery, such as from the skin, mixed with efferent axons travelling to the muscles or other effector organs. As the spinal nerve nears the spinal cord, it splits into dorsal and ventral roots. The dorsal root contains only the axons of sensory neurons, whereas the ventral roots contain only the axons of the motor neurons. Some of the branches will synapse with local neurons in the dorsal root ganglion, posterior (dorsal) horn, or even the anterior (ventral) horn, at the level of the spinal cord where they enter. Other branches will travel a short distance up or down the spine to interact with neurons at other levels of the spinal cord. A branch may also turn into the posterior (dorsal) column of the white matter to connect with the brain. For the sake of convenience, we will use the terms ventral and dorsal in reference to structures within the spinal cord that are part of these pathways. This will help to underscore the relationships between the different components. Typically, spinal nerve systems that connect to the brain are contralateral , in that the right side of the body is connected to the left side of the brain and the left side of the body to the right side of the brain.

Cranial Nerves

Cranial nerves convey specific sensory information from the head and neck directly to the brain. For sensations below the neck, the right side of the body is connected to the left side of the brain and the left side of the body to the right side of the brain. Whereas spinal information is contralateral, cranial nerve systems are mostly ipsilateral , meaning that a cranial nerve on the right side of the head is connected to the right side of the brain. Some cranial nerves contain only sensory axons, such as the olfactory, optic, and vestibulocochlear nerves. Other cranial nerves contain both sensory and motor axons, including the trigeminal, facial, glossopharyngeal, and vagus nerves (however, the vagus nerve is not associated with the somatic nervous system). The general senses of somatosensation for the face travel through the trigeminal system.

Chapter Review

The senses are olfaction (smell), gustation (taste), somatosensation (sensations associated with the skin and body), audition (hearing), equilibrium (balance), and vision. With the exception of somatosensation, this list represents the special senses, or those systems of the body that are associated with specific organs such as the tongue or eye. Somatosensation belongs to the general senses, which are those sensory structures that are distributed throughout the body and in the walls of various organs. The special senses are all primarily part of the somatic nervous system in that they are consciously perceived through cerebral processes, though some special senses contribute to autonomic function. The general senses can be divided into somatosensation, which is commonly considered touch, but includes tactile, pressure, vibration, temperature, and pain perception. The general senses also include the visceral senses, which are separate from the somatic nervous system function in that they do not normally rise to the level of conscious perception.

The cells that transduce sensory stimuli into the electrochemical signals of the nervous system are classified on the basis of structural or functional aspects of the cells. The structural classifications are either based on the anatomy of the cell that is interacting with the stimulus (free nerve endings, encapsulated endings, or specialized receptor cell), or where the cell is located relative to the stimulus (interoceptor, exteroceptor, proprioceptor). Thirdly, the functional classification is based on how the cell transduces the stimulus into a neural signal. Chemoreceptors respond to chemical stimuli and are the basis for olfaction and gustation. Related to chemoreceptors are osmoreceptors and nociceptors for fluid balance and pain reception, respectively. Mechanoreceptors respond to mechanical stimuli and are the basis for most aspects of somatosensation, as well as being the basis of audition and equilibrium in the inner ear. Thermoreceptors are sensitive to temperature changes, and photoreceptors are sensitive to light energy.

The nerves that convey sensory information from the periphery to the CNS are either spinal nerves, connected to the spinal cord, or cranial nerves, connected to the brain. Spinal nerves have mixed populations of fibers some are motor fibers and some are sensory. The sensory fibers connect to the spinal cord through the dorsal root, which is attached to the dorsal root ganglion. Sensory information from the body that is conveyed through spinal nerves will project to the opposite side of the brain to be processed by the cerebral cortex. The cranial nerves can be strictly sensory fibers, such as the olfactory, optic, and vestibulocochlear nerves, or mixed sensory and motor nerves, such as the trigeminal, facial, glossopharyngeal, and vagus nerves. The cranial nerves are connected to the same side of the brain from which the sensory information originates.

Interactive Link Questions

Watch this video to learn about Dr. Danielle Reed of the Monell Chemical Senses Center in Philadelphia, PA, who became interested in science at an early age because of her sensory experiences. She recognized that her sense of taste was unique compared with other people she knew. Now, she studies the genetic differences between people and their sensitivities to taste stimuli. In the video, there is a brief image of a person sticking out their tongue, which has been covered with a colored dye. This is how Dr. Reed is able to visualize and count papillae on the surface of the tongue. People fall into two large groups known as “tasters” and “non-tasters” on the basis of the density of papillae on their tongue, which also indicates the number of taste buds. Non-tasters can taste food, but they are not as sensitive to certain tastes, such as bitterness. Dr. Reed discovered that she is a non-taster, which explains why she perceived bitterness differently than other people she knew. Are you very sensitive to tastes? Can you see any similarities among the members of your family?

Answers will vary, but a typical answer might be: I can eat most anything (except mushrooms!), so I don’t think that I’m that sensitive to tastes. My whole family likes eating a variety of foods, so it seems that we all have the same level of sensitivity.

(Figure) The basilar membrane is the thin membrane that extends from the central core of the cochlea to the edge. What is anchored to this membrane so that they can be activated by movement of the fluids within the cochlea?

(Figure) The hair cells are located in the organ of Corti, which is located on the basilar membrane. The stereocilia of those cells would normally be attached to the tectorial membrane (though they are detached in the micrograph because of processing of the tissue).

Watch this video to learn more about how the structures of the ear convert sound waves into a neural signal by moving the “hairs,” or stereocilia, of the cochlear duct. Specific locations along the length of the duct encode specific frequencies, or pitches. The brain interprets the meaning of the sounds we hear as music, speech, noise, etc. Which ear structures are responsible for the amplification and transfer of sound from the external ear to the inner ear?

The small bones in the middle ear, the ossicles, amplify and transfer sound between the tympanic membrane of the external ear and the oval window of the inner ear.

Watch this animation to learn more about the inner ear and to see the cochlea unroll, with the base at the back of the image and the apex at the front. Specific wavelengths of sound cause specific regions of the basilar membrane to vibrate, much like the keys of a piano produce sound at different frequencies. Based on the animation, where do frequencies—from high to low pitches—cause activity in the hair cells within the cochlear duct?

High frequencies activate hair cells toward the base of the cochlea, and low frequencies activate hair cells toward the apex of the cochlea.

Watch this video to learn more about a transverse section through the brain that depicts the visual pathway from the eye to the occipital cortex. The first half of the pathway is the projection from the RGCs through the optic nerve to the lateral geniculate nucleus in the thalamus on either side. This first fiber in the pathway synapses on a thalamic cell that then projects to the visual cortex in the occipital lobe where “seeing,” or visual perception, takes place. This video gives an abbreviated overview of the visual system by concentrating on the pathway from the eyes to the occipital lobe. The video makes the statement (at 0:45) that “specialized cells in the retina called ganglion cells convert the light rays into electrical signals.” What aspect of retinal processing is simplified by that statement? Explain your answer.

Photoreceptors convert light energy, or photons, into an electrochemical signal. The retina contains bipolar cells and the RGCs that finally convert it into action potentials that are sent from the retina to the CNS. It is important to recognize when popular media and online sources oversimplify complex physiological processes so that misunderstandings are not generated. This video was created by a medical device manufacturer who might be trying to highlight other aspects of the visual system than retinal processing. The statement they make is not incorrect, it just bundles together several steps, which makes it sound like RGCs are the transducers, rather than photoreceptors.


Section Summary

A sensory activation occurs when a physical or chemical stimulus is processed into a neural signal (sensory transduction) by a sensory receptor. Perception is an individual interpretation of a sensation and is a brain function. Humans have special senses: olfaction, gustation, equilibrium, and hearing, plus the general senses of somatosensation.

Sensory receptors are either specialized cells associated with sensory neurons or the specialized ends of sensory neurons that are a part of the peripheral nervous system, and they are used to receive information about the environment (internal or external). Each sensory receptor is modified for the type of stimulus it detects. For example, neither gustatory receptors nor auditory receptors are sensitive to light. Each sensory receptor is responsive to stimuli within a specific region in space, which is known as that receptor’s receptive field. The most fundamental function of a sensory system is the translation of a sensory signal to an electrical signal in the nervous system.

All sensory signals, except those from the olfactory system, enter the central nervous system and are routed to the thalamus. When the sensory signal exits the thalamus, it is conducted to the specific area of the cortex dedicated to processing that particular sense.


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