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Human Physiology From Cells To Systems 9th Edition Sherwood Solutions Manual 1
Human Physiology From Cells To Systems 9th Edition Sherwood Solutions Manual 1
Chapter 6
The Peripheral Nervous System: Afferent Division; Special Senses
CHAPTER OUTLINE
Receptors have differential sensitivities to various stimuli. Each receptor will respond to one type of stimulus and can
be categorized according to this responsiveness.
Photoreceptors respond to visible wavelengths of light.
Mechanoreceptors are sensitive to mechanical energy such as stretch and pressure.
Thermoreceptors are sensitive to heat and/or cold.
Osmoreceptors detect changes in solute concentration in the ECF.
Chemoreceptors are sensitive to specific chemicals.
Nociceptors (pain receptors) are sensitive to tissue damage such as cutting, tearing, or burning.
The CNS can process inputs from several different types of simultaneously activated primary receptors to produce an
integrated sensation such as wetness (touch, pressure, and thermal receptor inputs).
Stimuli of the same intensity do not always produce the same magnitude of receptor potentials. Some receptors
diminish the extent of their depolarization despite sustained stimulus strength. This is known as receptor adaptation.
Acuity is influenced by receptive field size and lateral inhibition. Each somesthetic sensory neuron responds to stimuli
only in specific regions of skin surface surrounding the receptor, the receptive field. Smaller receptive fields with
higher receptor density lead to greater acuity or discriminative ability. Acuity is also influenced by lateral inhibition,
where information from the receptive fields surrounding the area of most intense stimulation is inhibited within the
CNS. This facilitates the ability to localize from where the stimulus is emanating.
6.2 PAIN
Pain is considered a protective mechanism meant to bring about awareness of tissue damage that is occurring or about
to occur. Pain receptors (nociceptors) do not adapt to sustained stimulus strength. Experience with pain helps us avoid
potentially harmful events in the future.
Stimulation of nociceptors elicits the perception of pain plus motivational and emotional responses. Pain perception
can be modified by past or present experiences. Fear of re-experiencing painful stimuli may heighten the perceived
pain, while other situations may reduce the perception of pain such as during a competitive event.
Nociceptors are categorized by the energy to which they react. Mechanical nociceptors respond to cutting, crushing, or
pinching. Thermal nociceptors respond to temperature extremes, especially heat. Polymodal nociceptors respond
equally to damaging inputs including irritating chemicals released from injured tissue.
Pain signals are transmitted from the receptors to the CNS via either fast pain pathways ( A- fibers; 30 m/sec.) or slow
pain pathways (C fibers; 12 m/sec. or slower). Afferent pain fibers utilize two neurotransmitters. Substance P is unique
to pain fibers and activates ascending pathways that transmit nociceptive signals to higher levels for further processing.
Glutamate, a major excitatory neurotransmitter, is released from primary afferent pain terminals.
The brain has a built-in analgesic system. This system suppresses transmission in the pain pathways as they enter the
spinal cord. Three brainstem regions are part of this system. The periaqueductal gray matter stimulates inhibitory
neurons to release enkephalin (an endogenous opiate peptide). There are specific analgesia-inducing nuclei in the
medulla and the reticular formation.
Protective mechanisms help prevent eye injuries. Specific protective structures include the eyelids, tear-producing
lacrimal glands, and eyelashes.
The eye refracts the entering light to focus the image on the retina. The cornea and flexible lens provide the refractive
mechanism. The shape of the lens is altered by the ciliary muscle. This change in shape alters the refractive qualities of
the lens (accommodation) and allows one to focus on objects at varying distances from the eye. The greater the
curvature is of the lens, the greater the strength of the lens to “bend” light rays and focus on closer objects.
Light must pass through several retinal layers before reaching the photoreceptors. The retina is an extension of the
CNS, not a separate peripheral organ. The photoreceptors, rods and cones, are located in the outermost layer of the
neural retina. The light-sensitive ends of these photoreceptors face away from the incoming light (face the pigmented
Chapter Six
layer of the retina). The middle layer of the neural retina contains bipolar cells and interneurons. The inner layer is
composed of ganglion cells whose axons join to form the optic nerve.
Light must pass through the layers of ganglion and bipolar cells to reach the photoreceptors in all areas of the retina
except the fovea, where the inner two layers are pulled aside to allow light to strike the photoreceptors (only cones are
found in the fovea), directly providing the most distinct vision.
Rods provide indistinct gray vision in low-light situations, whereas cones provide sharp color vision during periods of
high light intensity. There are 120 million rods and 6 million cones per retina. Rods are more numerous in the periphery
of the retina and are highly sensitive, but have low acuity. Cones are concentrated in or near the fovea. They have low
sensitivity to light, but have high acuity. There are four different photopigments, one specific to the rods and one in
each of the three different types of cones—red, green, and blue cones. The photopigments are composed of retinal and
four different opsins. As each opsin binds retinal differently, the four photopigments absorb different wavelengths of
light. Each photopigment absorbs maximally at a particular wavelength (strong stimulation), but also absorbs a range of
wavelengths, with those wavelengths farther from the peak wavelength producing weaker stimulation.
Color vision depends on the ratios of stimulation of the three cone types.
As light is absorbed by the photoreceptor, retinal is changed from the 11-cis form to the all-trans form. This sets off a
cascade of events leading the breakdown of the photopigment. This ultimately hyperpolarizes the photoreceptor, closes
Ca++ channels, and reduces the amount of neurotransmitter released from the photoreceptor. This decrease in
neurotransmitter release depolarizes on-center bipolar cells and subsequently the on-center ganglion cells leading to
propagation of impulses along the optic nerve. However, off-center bipolar cells and the subsequent off-center ganglion
cells are hyperpolarized. This bipolar and ganglion cell processing allows for a distinct visual image to be formed.
The sensitivity of the eyes can vary markedly through dark and light adaptation. The sensitivity depends on the amount
of light-responsive photopigments present in the rods and cones. Exposure to strong light depletes the photopigments,
thus reduces the sensitivity. This is why when one comes into a dark room from a brightly lit area, there is a temporary
blindness. As the photopigments are regenerated, vision in the low-light areas increases. This is called dark adaptation.
Visual information is modified and separated before reaching the visual cortex. The optic nerves carry information
from lateral and medial areas of the retinas they serve. As the two optic nerves meet at the optic chiasm, fibers from the
lateral regions project to the ipsilateral side of the brain, while fibers from the medial regions project to the
contralateral side of the brain for processing.
Visual input goes to other areas of the brain not involved in vision perception. Example of non-sight related activities
include: (1) contribution to cortical alertness and attention e.g., getting drowsy in a dimly lit room; (2) control of pupil
size (constriction in bright light, dilation in dim light); and (3) control of eye movements. Furthermore, some ganglion
cells make melanopsin, a light-sensitive pigment involved in helping set the body’s biological clock.
Some sensory input may be detected by multiple sensory-processing areas in the brain.
The external ear plays a role in sound localization. The external ear consists of the pinna (ear), the external auditory
meatus (ear canal), and the tympanic membrane (eardrum). The pinna collects sound waves and directs them to the ear
canal where the sound waves are channeled to the eardrum. Humans cannot move their pinna, but because the pinna
partially shields sound arising from behind, a person can distinguish sounds coming from directly in front or behind.
The middle ear bones convert tympanic membrane vibrations into fluid movements in the inner ear.
The tympanic membrane vibrates in unison with sound waves in the external ear. The middle-ear bones act as a bridge
between the eardrum and the cochlea. The malleus is attached to the tympanic membrane. As the eardrum vibrates it
causes the malleus to vibrate. The vibrating malleus is attached to the incus, which functions to amplify the movements
of the malleus. The incus transmits the amplified vibrations to the stapes, which conveys the vibrations to the oval
window at the entrance to the fluid-filled cochlea.
The cochlea contains the organ of Corti, the sense organ for hearing. As the oval window vibrates the sound energy is
transferred to the fluid. Some of the movement of this fluid is to compensate for pressure changes in the cochlea, while
some of the energy is transferred to the hair cells in the organ of Corti for transduction to neural signals.
Hair cells in the organ of Corti transduce fluid movements into neural signals.
Pitch discrimination depends on the region of the basilar membrane that vibrates.
The vestibular apparatus is important for equilibrium by detecting head position and motion.
The inner ear has a non-hearing component called the vestibular apparatus, important for the sense of equilibrium and
for coordinating head movements with eye and postural movements. The semicircular canals detect rotational or
angular acceleration and deceleration of the head. The otolith organs provide information concerning the position of the
head relative to gravity.
The olfactory receptors in the nose are specialized endings of renewable afferent neurons.
Various parts of an odor are detected by different olfactory receptors and sorted into “smell files.” Humans have five
million olfactory receptors, of which there are 1000 different types. The odors are dissected into different components
based on the ratio of response of the various types of olfactory receptors. Afferent fibers arising from the olfactory
receptor endings synapse in the olfactory bulb for processing.
The olfactory system adapts quickly and odorants are rapidly cleared.
Chapter Six
LIST OF KEY TERMS
1. Obtain preserved specimens of bovine or sheep eyes and dissect them for the demonstration of their important
structures.
2. The shark’s vestibular apparatus and the middle ear bones of mammals are excellent demonstration materials.
3. Students can localize receptors in their skin for touch, pressure, pain, and temperature. See any general physiology
lab manual for techniques and supplies. For example, calipers can be used for two-point discrimination tests on
several regions of the skin.
4. Use models of the eye and ear to illustrate their structures. During your presentation, describe the functions of
these structures as you locate them. Challenge the students to do the same with the models during their lab class.
The models can be obtained from Carolina Biological Supply Company, Burlington, NC.
5. To demonstrate the interaction of the senses of taste and smell, students can detect the difference between the
tastes of two foods (e.g., an apple and an onion) while pinching shut their nostrils.
6. Demonstrate the test for color blindness. See any general physiology lab manual for techniques and supplies.
Booklets with various charts are available to diagnose the different kinds of color blindness.
Chapter whereby one assumeth the form of the Chief god of the
Divine Cycle.
to be seen.
CHAPTER LXXX.
Chapter whereby one assumeth the form of the god who giveth
Light to the Darkness.
N .
1. Devouring monsters, .
I am the pure Lotus which cometh forth from the glory which is
at the nostril of Rā, and I make my journey and pursue it for
Horus, the great god beloved.
I am the pure Lotus which cometh forth in the field.
N .
This little chapter is not without its special difficulty. Are we to
read as a word implying motion, with as its
I fly like the Hawk, I cackle like the Smen-goose, I alight on the
right side of the Aat, on the feast of the Great One.
I execrate, I execrate: I eat it not. Dirt is what I execrate: I eat it
not. That which my Genius execrateth let it not enter into me.
Let me therefore live upon that which is put before them; the
gods and the glorified ones. Let me live and enjoy the bread and....
[90]
Let me then eat them in the presence of the gods and glorified
ones. Let me enjoy and eat them under the foliage of the date trees
of Hathor, my sovereign. Let the oblations be made, of bread and
beer in Tattu, and bendings of the head in Annu. Let my vesture
be girt upon me by Tait. Let me sit wherever it pleaseth me.
My head is that of Rā and I am summed up as Tmu: Four times
the arm’s length of Rā: four times the width of the world.(1)
I have come forth with the tongue of Ptah and the throat of
Hathor that I may record the words of my father Tmu with my
mouth, which draweth to itself the Spouse of Seb, and the
proclamation of whose lips inspireth fear.
I repeat the acclamations at my success on being declared the
heir of the Lord of Earth, Seb, from whom I issue.
Seb purifieth me, and giveth me his Theophanies.(2) The
dwellers in Annu bow their heads to me. I am their Master. I am
their Bull. More powerful am I than the Lord of Time; I am the
author and the master of endless years.
N .
90. The word seems to have been unintelligible to the copyists, who differ widely
from each other as to its orthography.
CHAPTER LXXXIII.
Let me wheel round in whirls, let me turn like the Turning One,
let me flourish like a flower and keep myself hidden like the Hider.
(1)
I am the Barley corn of every god.
I am the four Yesterdays of those seven Uræus deities who are
born in Amenta; Horus who giveth light by means of his own
body; the god who is against Sutu when Thoth is between them, as
in that dispute of the Prince of Sechem with the Spirits of Annu
where the river is between them.(2)
I come forth by day and disclose myself at the head of the gods.
I am the god who chaseth all boastfulness.(3)
N .
1. There is here a play on the words pa, ḫeper, ruṭ and šet. The
Turning One is the god Chepera. The Tortoise
derives its name (the hider), from the habit of drawing its
body within its shell. On the flight of the Bennu see the first note
of next chapter.
Thou who holdest the bound victims; ye knives over their heads
and locks and fleeces;(2) ye aged and bright ones who are armed
with the fated moment.
I come to heaven but I strike upon the earth; and conversely.
It is my power which produceth victory and raiseth the height of
heaven, and I make the lustrations which yield the extent of earth
to my feet against the sinful cities as I advance and cut in pieces(3)
those who are involved in rebellion.(4)
I leave the gods upon their paths but I strike the Wakers who
are in their coffins.
I know not Nu, I know not Tatunen, I know not the Red ones
when they bring opposition to me.
I know not a Word of Power to whose utterance I listen.
I am the Red Calf upon the tablets.
This is what the gods say when they raise their voice.
Let your countenances be without restraint towards him who
cometh to me.
The morning dawns are independent of you, ye have not the
charge of them; but my alternations are in my own hands. I say
not the wrong instead of the right.
Day after day unswervingly turneth back upon my eye-brow.
And Evening is the beginning of my voyage to celebrate the
solemnity of the Reclining and the Embrace of the Aged one who
hath charge of the Earth.
N .
1. Both the Bennu and the Shenshen (which I here translate
‘Hernshaw’) are Herons. They fly to a great height in spiral whirls.
N .
1. Soul. The Egyptian word which in our modern languages we
translate as Soul has already been explained as meaning Force. It
is so translated in this chapter in several passages where this sense
is emphatically required.
3. Here the chapter ends in Pc. The few words which follow in
other MSS. were unintelligible to the copyists and are written very
variously.
CHAPTER LXXXVI.
N .
1. The Swallow . The objection to this meaning is that
the bird in question was eaten; and that doves or pigeons would be
less meagre food than the Swallow, and therefore more probably
intended in the Egyptian texts. But Swallows are still eaten at
Rome, where like Clive Newcome we may be regaled not only with
“wild swans and ducks” but with “robins, owls, and οἰωνοῖσι τε
πᾶσι for dinner.” And Willughby, the naturalist, found a large
quantity of swallows being sold for food at Valencia in Spain.
The flat head, the short legs, and the tail of the bird are
characteristic not of the pigeon but of the swallow, and on many
pictures (e.g., pl. xxi, vignette from Leyden papyrus) we are
reminded of the song—
A N .
I am Seta—full of years.
I lay myself down [in death], and I am born daily.
I am Seta at the confines of the earth. I lay myself down [in
death], I restore myself and I renew myself daily.
N .
N .
1. Sebak is not always named in the papyri. The ideogram of the
crocodile was in some copies read emsuḥ and in others sebak.
Oh ye gods who draw along the Bark of the Eternal one: ye who
lift up above the Tuat, and who raise up the Sky: ye who enable the
Souls to enter into the mummied forms; ye whose hands grasp the
cordage, hold firm with your ropes and stop the adversaries that
the Bark may rejoice and the god proceed in peace.
And now grant that my Soul may come forth in your train from
the Eastern horizon of Heaven for ever and ever.
N .
The oldest papyri present a much shorter form than the later
ones. That portion which is here separated by a line from what
goes before it first appears on the sarcophagus of Seti I and in the
papyrus of Ani. The vignette is a very favourite decoration of
mummies.
1. Keep of which the regular variant in this chapter is
N .
Of this chapter we have unfortunately but one copy in Fa, of the
Musée Borély. This is defective both at the beginning and at the
end, and the text is inaccurate. The later copies are so inaccurate
that it is impossible to reconstitute the text. It is precisely on those
points where grammatical accuracy is required for fixing a definite
sense that the manuscripts are hopelessly defective. The preceding
translation is verbally correct, I trust, but I do not pretend that it
is intelligible. It stops where the papyrus Fa stops.
1. Restored. The reduplication in here gives the verb this
sense.
N .
1. There is no safe text here, ‘grandeur’ is only meant to indicate
the existence of in the original. But there certainly ought to
be something different from what any of the MSS. supply.
PLATE XXIII.
PLATE XXIV.
CHAPTER XCII.
Chapter whereby the Tomb is opened to the Soul and to the Shade of the person, that he
may come forth by day and may have mastery of his feet.
N .
1. I cannot agree with those who have hitherto translated this
chapter. The only grammatical interpretation which seems
possible for the first sentence depends upon the sense given to the
suffix tȧ. I take this as representing the second person
2. The words which follow are evidently the words of Osiris and
those attached to him, which are addressed to the deceased and
are repeated by him. The text here, as indeed everywhere, is very
corrupt.
N .
This chapter contains one of those threats (of which there are
other instances) made to the gods. The speaker is in fact so
identified with divinity that any evil which happens to him must
be conceived as involving the same calamity to the gods and to the
universe.
There is a very considerable difference between the earlier and
the later texts. There is very great confusion in the text of the
Turin Todtenbuch as compared with that of the Cadet papyrus.
later texts. The latter form, which has for determinative pearls or
globules of some kind, reminds one of the disease formerly called
gutta serena.
2. Destruction, . But this word is
Oh mighty one, who seest thy father, and who hast charge of the
Book of Thoth.
Here am I, I come and am glorified and filled with Soul and
Power and provided with the writings of Thoth, which I bring in
order to purify the tunnel which is in Sutu.(1) I bring the Palette
and I bring the Inkstand as the instruments of Thoth, the secrets
of which are divine.
Here am I, as the Scribe; I bring the remains of Osiris;(2) and
the writing which I have made upon them is decreed by the great
god to be good, daily, among the good. Thou hast decreed, Horus
of the Two Horizons, that I shall be the author of Maāt and
tend(3) it daily to Rā.
N .
1. In Sutu; that is, in Darkness. See chapter 96.
N .
The papyrus Ad gives this chapter the title of “assuming the
form of the Smen-goose,” and Dr. Birch published the text of this
papyrus in the Zeitschrift of 1869 (p. 25) as one of those additional
chapters which “do not occur in the Ritual of Turin.” This is of
course an error of oversight. This chapter is in the Turin
Todtenbuch, and the papyrus Ad merely gives it under an
erroneous title, which was evidently meant for another text.
1. The Dread one, . Instead of this Ad has