Vanders Human Physiology The Mechanisms of Body Function 14Th Edition Widmaier Solutions Manual Full Chapter PDF

Vanders Human Physiology The
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6: NEURONAL SIGNALING AND THE STRUCTURE

OF THE NERVOUS SYSTEM
CHAPTER OVERVIEW
The chapters preceeding Chapter 6 provided an important overview of physical and chemical
properties that contribute to human physiological function. In Chapter 6, these principles are
applied to the first organ system to be studied, the Nervous System. The Nervous System is an
important means for communication across the body, and this communication occurs via
changes in the electrical activity of neurons. This chapter will first describe the cells of the
Nervous System. Next, the mechanisms that determine the difference in charge across a
neuron cell membrane and how it is used as a means for communication from one neuron to
another is described. This chapter also summarizes the different classes of neurotransmitters,
one of the intercellular signaling molecules, that are used for neuron-to-neuron (or effector)
communication. The last section of this chapter summarizes the structural and functional
organization of the Nervous System
CONTEXT FOR CHAPTER 6

One of the most fascinating stories in the history of physiology is the discovery of
neurotransmitters by the German scientist Otto Loewi, who was awarded the 1936 Nobel Prize
for Physiology or Medicine for his discovery. Part of the fascination lies in the report that the
idea for the critical experiment proving the existence of chemical messengers from nerves
came to Loewi in a dream. Loewi knew that stimulating the vagus nerve in frogs, as in
humans, causes the heart rate to slow. The question was: how? Was the cause an electrical
“spark” that inhibited the heart's pacemaker? Or was it a chemical messenger released by the
nerve? Loewi also knew that the frog heart can keep beating long after its removal from the
animal if placed in an isotonic salt solution approximating extracellular fluid. Loewi
performed a simple, yet brilliant, experiment: He placed one frog heart, with its vagus nerve
still attached, into a dish of isotonic saline (dish A) and another heart without the vagus into
another dish with the same saline (dish B). He then electrically stimulated the vagus nerve of
the heart in dish A and watched the heart rate slow. After some time, he removed the
stimulated heart and transferred the other heart from dish B to dish A—and observed that the
beating of the second heart slowed to match that of the first one! Something in the saline acted
like a stimulated vagus nerve in affecting heart rate. The signal from nerve to muscle thus had
to be a chemical.
The junction between a nerve and a muscle is a neuromuscular junction, not a synapse. But
scientists soon realized that most nerve cells “talk” to each other as well as to the heart and
other muscles. This communication takes place by means of chemical messengers called
neurotransmitters.
©2017 by McGraw-Hill Education. This is proprietary material solely for authorized instructor use. Not authorized for sale or distribution in any
manner. This document may not be copied, scanned, duplicated, forwarded, distributed, or posted on a website, in whole or part.
2
Chapter 6 is divided into four different sections to provide an overview of the structures and
functions of the Nervous System.
Section A: Cells of the Nervous System. Generalized cell structure was presented in Chapter
3. This chapter describes the specific cellular structures and functional classes of neurons, the
“nerve cells”. Glial cells, the second type of cells found in the Nervous System are also
described.
Section B: Membrane Potentials. This section brings together electrical principles from
physics and the movement of ions across the neurolemma to explain the how neurons changes
in membrane potential for communication. The differences between graded potentials and
action potentials are especially important to highlight and will be applied to many organ
systems: the Muscular System, the Cardiovascular System, and so on.
Section C: Synapses. Once a “message” has reached an axon terminal in one neuron, how can
it be relayed to a second neuron or other target tissue? The answer is via neurotransmitter
release into the synapse, or the junction between the presynaptic neuron and a postsynaptic
neuron/effector. If the neurotransmitter binds to receptors in the postsynaptic neuron, graded
potentials are generated and if they reach threshold an action potential will be created for the
continued propagation of the “message”. The different classes of neurotransmitters are also
described.
Section D: Structure of the Nervous System. This section begins with an overview of the
structural and functional organization of the Nervous System. It highlights information flow
between different divisions. Importantly, the autonomomic division of the Nervous System is
highlighted. The autonomic nervous system is a vital component of many homeostatic reflex
mechanisms as it innervates many different organs throughout the body. Upcoming
discussion of other organ systems throughout the body will hinge upon the understanding of
autonomic function.
CONCEPTS COMMONLY FOUND TO BE CHALLENGING – TEACHING HINTS

This is an expansive chapter covering many important aspects of Nervous System function.
Instructors may find areas to highlight or skim over depending upon the specific focus of the
course and time availability.
1. To give context, instructors may want to provide a very general overview of the Nervous
System as an introduction: explain that communication among the different divisions of the
Nervous System is vital for homeostatic regulation and human function. How is this
communication possible? To answer this question, students must understand neuron
function.
2. The factors contributing to resting membrane potential and how it is maintained are
important and tie together several concepts from chemistry, physics, and previous
chapters: the difference between the intracellular and extracellular fluid compartments,
membrane transport mechanisms and specifically voltage- gated ion channels, the Na+/K+-
ATPase pump, and as a positive feedback example during action potential generation.
There are numerous instructor resources available for this very important topic; pay
specific attention to figures and tables in the text and consider the inclusion of animations
during class sections or as recommendations for out-of-class activities.
3. It is common for students to have questions about the similarities/differences for graded
potentials and action potentials. Temporal and spatial summation are sometimes also not
immediately clear to students. One suggestion is to approach summation like it is a math
problem, and actually find the sum of the changes in membrane potential in response to
EPSPs and IPSPs to see if it reaches threshold.
4. Fig. 6.27 is a great one to carefully review. It summarizes the sequence of events at a
chemical synapse, which can lead into a discussion of mechanisms for pre/post synaptic
modulation of the “message” being sent, EPSP/ IPSP generation on the postsynaptic
membrane, etc. These factors can be added in to the model depicted in Fig. 6.27.
5. Fig. 6.46 is also an excellent review to compare and contrast the somatic and autonomic
divisions of the peripheral nervous system. It highlights: general structural organization,
neurotransmitters released, classes of neurotransmitter receptors, epinephrine as a
neurohormone, dual innervation and antagonistic control of autonomic effectors.
6. Suggested topic for discussion:
 Research efforts toward restoring function to damaged brain and spinal cord
 Drugs and the blood-brain barrier. Suggested examples: Treatment of Parkinson’s
disease includes administering L-dopa but not dopamine because dopamine cannot
cross the blood-brain barrier. Allergy sufferers benefit from antihistamines but many of
the classical drugs (e.g., Benadryl) cross the blood-brain barrier and make the person
drowsy. Newer drugs (e.g., Claritin, Allegra) are effective antihistamines for sinus and
nasal allergy symptoms, but they do not cross the blood-brain barrier and thus cannot
affect one’s state of alertness.
LECTURE OUTLINE
SECTION A: Cells of the Nervous System
I. The central and peripheral nervous systems

II. Neurons
A. Structures: cell body, dendrites, axons, initial segment, axon terminals (Fig. 6.1)
B. Sequence of information flow (electrical signals) across neurons
C. Myelination (Fig. 6.2)
D. Axon transport (Fig. 6.3)
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III. Functional classes of neurons (Fig. 6.4, Table 6.1)

IV. Neuronal communication: the synapse (Fig. 6.5)
V. Glial cells (Fig. 6.6)
VI. Neuronal growth
A. Undifferentiated precursor/stem cells
B. Migration
C. Growth cone
D. Neurotrophic factors
E. Plasticity
VII. Neuronal regeneration
LECTURE OUTLINE
SECTION B: Membrane Potentials
I. Basic principles of electricity

A. Interaction of positive and negative charges (Fig. 6.7)
B. Physical principles: potential, current, resistance, Ohm’s Law (Fig. 6.8)
C. Applications to the fluid compartments of the human body (Fig. 6.8)
II. Resting membrane potential
A. Definition and measurement (Fig. 6.8)
B. Determinants
1. Ion concentration differences (Fig. 6.9, Fig. 6.10, Fig. 6.11, Fig. 6.12, Table 6.2)
2. Ion permeabilities and leak channels (Fig. 6.12)
3. Ion pumps: the Na+/K+-ATPase pump (Fig. 6.13)
III. Membrane potentials in excitable cells
A. Terminology (see Fig. 6.14 and Table 6.3)
1. Deplarization (Fig. 6.15, Fig. 6.18)
2. Repolarization (Fig. 6.18)
3. Overshoot and Repolarization
B. Properties of graded potentials (Figs. 6.16, Fig. 6.17, Table 6.4)
1. Graded response to increasing stimulus intensity
2. Decremental conduction of signal
3. Summation of responses to two or more stimuli
C. Action potentials (Fig. 6.19, Table 6.4)
1. Changes in membrane potential (Fig 6.21)
2. Changes in ion permeability and properties of involved ion channels (Fig. 6.18)
3. Properties of action potentials
a. Threshold potential
i. Ionic basis for reaching the threshold potential
ii. Voltage-gated channels and positive feedback (Fig. 6.20)
b. All-or-none response
c. Refractory periods: absolute and relative (Fig. 6.22)
4. Propagation of action potentials
a. Unidirectional, nondecremental propagation (Fig. 6.23)
b. Myelinated axons (Fig. 6.4)
LECTURE OUTLINE
SECTION C: Synapses
I. Transmission of neural signals from neuron to neuron: the synapse (Fig. 6.26)
A. Convergence and divergence in synaptic pathways (Fig. 6.25)
B. Mechanisms of neurotransmitter release (Fig. 6.27)
C. Excitatory and inhibitory postsynaptic potentials (Fig. 6.28, Fig. 6.29, Fig. 6.30, Fig. 6.32)
D. Synaptic integration (Fig. 6.31)
1. Temporal summation
2. Spatial summation
E. Regulation of synaptic strength (see Table 6.5)
1. Presynaptic mechanisms (Fig. 6.33)
2. Postsynaptic mechanisms
3. Modulation by drugs and disease (Fig. 6.34)
II. Classes of neurotransmitters or neuromodulators (see Table 6.6)
A. Acetylcholine (ACh) and cholinergic receptors
B. Biogenic amines
1. Catecholamines (Fig. 6.35)
a. Dopamine (DA)
b. Norepinephrine (NE)
c. Epinephrine (Epi)
2. Serotonin (5-hydroxytryptamine, 5-HT)
C. Amino acids
1. Excitatory: glutamate (Fig. 6.36)
2. Inhibitory
a. GABA (gamma-aminobutyric acid)
b. Glycine
D. Neuropeptides: endogenous opioids
E. Gases
1. Nitric oxide
2. Carbon monoxide
3. Hydrogen sulfide
F. Purines
1. ATP
2. Adenosine
G. Neuroeffector communication
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LECTURE OUTLINE
SECTION D: Structure of the Nervous System
I. Overview of the structures and general functions of the Nervous System (Fig. 6.37)
II. The Central Nervous System, CNS (see Fig. 6.37)
A. The Brain (Fig. 6.38, Table 6.7)
1. Cerebrum (see Fig. 6.38, Fig. 6.39)
a. Gray matter
i. Cerebral cortex
ii. Subcortical nuclei
b. White matter—fiber tracts and corpus callosum
2. Limbic system (Fig. 6.40)
3. Diencephalon—thalamus, hypothalamus, and epithalamus (Fig. 6.38)
4. Hindbrain: cerebellum (Fig. 6.38)
5. Brainstem—midbrain, pons, and medulla oblongata (Fig. 6.38)
B. Spinal cord (Fig. 6.41, 6-42)
1. Gray matter
a. Dorsal horns
b. Ventral horns
2. White matter—ascending and descending tracts
III. The Peripheral Nervous System, PNS: structure and general functions (see Fig. 6.7, Fig.
6.45, Fig. 6.46, Table 6.9)
A. Cranial and spinal nerves (Table 6.8, Fig. 6.42)
B. Somatic nervous system
B. Autonomic nervous system (Fig. 6.43, Fig. 6.44, Table 6.11)
1. Sympathetic division
2. Parasympathetic division
III. Protection of the brain
A. Cerebrospinal fluid (Fig. 6.47)
B. The blood-brain barrier
TEACHING/LEARNING OBJECTIVES BY SECTION

6.1 Structure and Maintenance of Neurons
Students should be able to:
explain that the nervous system is composed of Central Nervous System (CNS) and
Peripheral Nervous System (PNS) divisions: the brain and spinal cord comprise the
CNS and the cranial and spinal nerves comprise the PNS.
identify that the basic unit of the nervous system is a nerve cell or neuron, which is
capable of transmitting electrical signals that lead to the release of chemical
neurotransmitters.
identify the structures—cell body, dendrites, and axon—found on most neurons.
describe the general sequence of information flow via electrical signals across the
different regions of a neuron.
name the myelin-forming cells in the CNS and PNS. Explain how myelin is formed, and
what its general function is.
define axonal transport, describe its general function, and identify two motor proteins
associated with this process.
6.2 Functional Classes of Neurons

identify the three functional classes of neurons—afferent, efferent, and interneurons—
and identify the direction of information flow relative to the CNS and/or PNS.
know the ratio of 1 afferent neuron to 10 efferent neurons to 200,000 interneurons.
state where the cell bodies and axons for each class of neuron are located, and generally
what type of information each class of neurons relays.
distinguish between a nerve fiber and a nerve.
describe the relationship of one neuron to another at a synapse. Understand that all
interneurons are both postsynaptic cells and presynaptic cells.
6.3 Glial Cells

describe the general functions of glial cells with regard to Nervous System function.
list types of glial cells and generally describe their functions.
explain the functional role of the blood-brain barrier.
identify the structures associated with the blood-brain barrier.
recognize that some glial cells play a role in immune function in the CNS.
6.4 Neural Growth and Regeneration

understand that neurons develop from precursor cells, migrate to their final location,
and send out processes to their target cells.
describe the importance of plasticity during the first few years following birth.
recognize that recent research suggests that new neurons can be produced in some
brain regions throughout life.
explain that after damage to an axon, a peripheral neuron may regrow that axon to its
target organ, but that damaged neurons in the CNS do not regenerate (yet—but
scientists are working on how to promote axonal regeneration in the CNS).
8
6.5 Basic Principles of Electricity

identify the major positive and negative ions in the intracellular and extracellular fluids.
describe how positive and negative charges interact with each other.
recognize that a separation of charges results in an electrical potential, which has the
capacity to do biological work.
understand how current, voltage, and resistance are related (Ohm’s law).
explain why water is a good conductor of electricity.
6.6 The Resting Membrane Potential

recognize that in all body cells under resting conditions there is a potential difference
across the membrane such that the inside is negative with respect to the outside.
understand that the membrane potential is a result of two factors: (1) the uneven
distribution of (primarily) Na+ and K+ across the plasma membrane and (2) the unequal
permeabilities of membranes to those ions.
describe how a two-compartment model of solutions of NaCl and KCl separated by a
selectively permeable membrane demonstrates how a membrane potential can be
generated by having the membrane first permeable only to K +, and then permeable only
to Na+.
define equilibrium potential as the membrane potential needed to oppose the diffusion of
ions based upon a given concentration gradient, and at this point there is no net flux of
the ion. Recognize that each type of ion has its own equilibrium potential.
state the Nernst equation, and understand that it is used to determine the equilibrium
potential for a particular type of ion.
recognize that the Goldman-Hodgkin-Katz (GHK) equation can be used to calculate the
resting membrane potential of a cell by taking into consideration relative membrane
permeabilites to different ions.
understand why the resting membrane potential of cells is much closer to the
equilibrium potential for K+ than to the equilibrium potential for Na+.
define the term leak channel and describe how it influences membrane potential.
recognize that at the resting membrane potential, the driving force for Na + diffusion is
much greater than that for K+.
discuss the importance of the Na+/K+-ATPase in maintaining the concentration gradients
for Na+ and K+ and establishing the resting membrane potential.
describe how the movement of the Cl- is affected by the membrane potential.
6.7 Graded Potentials and Action Potentials

define the terms depolarize, repolarize, and hyperpolarize.
understand that depolarizing a membrane causes it to have a lesser potential difference
across (i.e. it becomes more positive).
define graded potential and describe how local current surrounding the depolarized
region produces depolarization of adjacent regions.
describe how the magnitude of the graded potential is affected by the magnitude of the
stimulus, the distance the potential has traveled, and summation.
describe an action potential and identify the types of cells with excitable membranes
capable of generating action potentials.
compare and contrast graded potentials and action potentials.
discuss the importance of ligand-gated channels and mechanically-gated channels to the
initiation of an action potential and the importance of voltage-gated channels to the
excitability of the membrane.
reproduce an action potential as membrane potential vs. time and label its various parts
in terms of being depolarized, repolarizing, and after-hyperpolarized. Superimpose the
changing patterns of Na+ and K+ permeabilities on the action potential.
describe the mechanism of ion channel changes that generate and terminate action
potentials. Appreciate that the action potential is a good example of a physiologically
important positive feedback mechanism.
explain why the threshold potential must be reached in order for a neuron to generate an
action potential.
explain why action potentials are said to be “all-or-none.”
understand the ionic basis of relative and absolute refractory periods.
differentiate between action potential propagation in myelinated axons and in
unmyelinated ones.
provide a physiological description of three different ways that action potentials can be
initiated.
SECTION C: Synapses
6.8 Functional Anatomy of Synapses
differentiate between excitatory and inhibitory synapses.
discuss how the functional anatomy of a synapse can account for the one-way
conduction of action potentials in a neural circuit.
draw a model illustrating convergence and divergens of neural inputs and discuss the
implications of these processes for synaptic transmission of information.
compare the structural and functional similarities and differences between electrical
and chemical synapses.
recognize that electrical synapses have recently been described in widespread locations
in the adult mammalian nervous system.
diagram a typical chemical synapse.
6.9 Mechanisms of Neurotransmitter Release

10
understand that neurotransmitters are stored in vesicles within the axon terminal.
outline the sequence of events that links action potential propagation to
neurotransmitter release.
discuss the role of Ca2+ in stimulating neurotransmitter exocytosis from synaptic
vesicles.
6.10 Activation of the Postsynaptic Cell

describe the general ways in which neurotransmitters can directly or indirectly affect
ion channels on the postsynaptic membrane.
describe the three ways that unbound neurotransmitters can be removed from a
synaptic cleft.
distinguish between an excitatory postsynaptic potentials (EPSPs) and an inhibitory
postsynaptic potentials (IPSPs), discuss which ions are typically responsible for each, and
identify whether the membrane potential is increased or decreased.
6.11 Synaptic Integration

recognize that the membrane potential of a postsynaptic cell reflects the effects of
possibly hundreds of different, simultaneous synaptic inputs, both excitatory and
inhibitory.
understand how spatial and temporal summation interact to influence membrane
potential and may bring a postsynaptic cell to threshold.
discuss the significance of the initial segment of a postsynaptic cell’s axon and why
synapses near the initial segment have greater influence on the cell’s activation than do
synapses far removed from it.
6.12 Synaptic Strength

appreciate that the amount of neurotransmitter released following an action potential’s
arrival at the axon terminal is variable.
discuss the effects of greater or lesser amounts of neurotransmitter on the postsynaptic
cell.
appreciate the importance of presynaptic (axo-axonic) synapses and autoreceptors in
altering synaptic effectiveness.
differentiate presynaptic inhibition from presynaptic facilitation.
propose postsynaptic mechanisms that could varying synaptic effectiveness.
define receptor desensitization and explain how it influences the postsynaptic response.
identify eight mechanisms by which certain diseases and drugs can alter synaptic
effectiveness, and explain whether each mechanism affects pre- or postsynaptic neural
transmission.
6.13 Neurotransmitters and Neuromodulators
differentiate between neurotransmitters and neuromodulators.
list the six major classes of neurotransmitters and neuromodulators.
describe how and where acetylcholine is synthesized and metabolized.
discuss what is meant by the terms cholinergic neurons and cholinergic receptors; identify
nicotinic and muscarinic receptors as cholinergic receptors.
explain the link between Alzheimer’s Disease and cholinergic neuron function.
list five common biogenic amines.
describe the synthetic pathway for the catecholamines and how and where they are
metabolized. Understand why neurons that secrete norepinephrine or epinephrine are
called adrenergic neurons.
Identify the two major classes of adrenergic receptors.
describe the general functions/actions of the biogenic amine serotonin and the amino
acid neurotransmitters glutamate, GABA, and glycine.
discuss the role of glutamate in long-term potentiation and exitotoxicity.
discuss how the synthesis of neuropeptide neurotransmitters differs from that of other
types of neurotransmitters.
describe the general role of the opioid family of neurotransmitters. (The first one of
these to be discovered, beta-endorphin, was given its name because it resembled
“endogenous morphine” in relieving pain.)
discuss how gases such as nitric oxide, carbon monoxide, and hydrogen sulfide can act
as neurotransmitters.
recognize that the purines, ATP and adenosine, act principally as neuromodulators.
6.13 Neuroeffector Communication

understand what is meant by the term neuroeffector communication and neuroeffector
junctions.
understand the functional similarities for neuron-to-neuron communication and
neuron-effector communication.

Introduction
describe the organization of the central and peripheral nervous systems.
define the following terms: nerve, pathway/tract, commissure, ganglia, and (nervous system)
nuclei
6.15 Central Nervous System: Brain

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describe the general structural organization of the brain.

identify the structures and general functions associated with the cerebrum, limbic
system, diencephalon, cerebellum, and the brainstem.
differentiate between white matter and gray matter.
6.16 Central Nervous System: Spinal Cord

describe the basic anatomy of the spinal cord and its associated nerves. Differentiate
between the dorsal and ventral horns, and the dorsal and ventral roots.
explain the significance of the dorsal root ganglia.
6.17 Peripheral Nervous System

distinguish between the afferent and efferent divisions of the peripheral nervous system.
recognize that the efferent division is divided into the somatic and autonomic nervous
systems and that the autonomic division is divided into the parasympathetic, sympathetic,
and enteric divisions.
summarize the flow of information from the central nervous system to the somatic
nervous system. Identify the primary neurotransmitter associated with the somatic
motor system and its effector.
6.16 Autonomic Nervous System

describe the structural organization of the autonomic nervous system, and recognize
that autonomic efferents consist of two neurons: a preganglionic and postganglionic
neuron.
identify similarities and differences between sympathetic and parasympathetic
neurons, paying particular attention to neurotransmitters and postsynaptic receptors.
explain how dual innervation of autonomic effectors contributes to regulation of body
function. Appreciate the importance of dual innervation of organs to allow for fine-
tuned control of organ function (akin to having both an accelerator and a brake).
provide examples of autonomic effectors that exhibit dual innervation and effectors that
do not.
discuss the basic functions of the somatic and autonomic nervous systems (and
appreciate that they are not actually separate nervous systems but are members or
divisions of the peripheral nervous system). Understand how the two systems differ
anatomically and functionally.
6.19 Protective Elements Associated with the Brain

define and discuss the importance of cerebrospinal fluid.
recognize meningitis and hydrocephalus as examples of disease conditions related to
cerebrospinal fluid.
recognize that the brain is dependent upon a continuous supply of glucose and oxygen.
understand the importance of the blood-brain barrier in regulating the composition of the
brain’s extracellular fluid.
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CHAPTER 6 REVIEW QUESTIONS & ANSWERS
1. Describe the direction of information flow 3. Where are afferent neurons, efferent neurons,
through a neuron in response to input from and interneurons located in the nervous
another neuron. What is the relationship system? Are there places where all three could
between the presynaptic neuron and the be found?
postsynaptic neuron?
Afferent neurons have dendrites and axonal
Information flows into the dendrites and cell body projections that originate near the skin or organs.
regions of most neurons, and out from axons and Their cell bodies are located outside the CNS in the
axon terminals. The presynaptic neuron releases dorsal root ganglia. They also contain central
neurotransmitters into the synaptic cleft which then processes, which is the part of the axon that projects
bind to receptors on the postsynaptic neuron to cause into the CNS. Efferent neurons have cell bodies that
an effect. lie within the CNS and axons that project out to
muscles, glands, or other neurons. Interneurons lie
entirely within the CNS. The enteric nervous system
2. Contrast the two uses of the word receptor. contains all three types of neurons, and the CNS
“Receptor” refers both to structures at the peripheral contains parts of all three types of neurons - the
endings of afferent neurons that respond to physical central processes of afferent neurons, the
or chemical changes in their environment by causing interneurons, and the cell bodies and first part of the
electrical signals to be generated in the neuron, and axon of efferent neurons.
to the proteins within cells or on the plasma
membrane of cells that are activated by an
intercellular messenger.
1. Describe how negative and positive charges carry the current. Thus, the ion-containing
interact. intracellular and extracellular fluids are relatively
good conductors of electrical current, but membrane
Negative charges repel negative charges, positive
lipids contain very few charged groups and are very
charges repel positive charges, and negative and
poor conductors (they have high electrical
positive charges are mutually attractive. The
resistance).
electrical force of attraction between positive and
negative charges increases with the quantity of
charge and with a decrease in distance between them.
3. Draw a simple cell; indicate where the
concentrations of Na+, K+, and Cl- are high and
2. Contrast the abilities of intracellular and low and the electric potential difference across
extracellular fluids and membrane lipids to the membrane when the cell is at rest.
conduct electrical current.
Water that contains dissolved ions is a relatively
good conductor of electricity because the ions can
15
As stated above, (1) the relative ion concentrations
on either side of the membrane and (2) the relative
permeabilities of the membrane to the ions.
6. Explain why the resting membrane potential is

not equal to the K+ equilibrium potential.
The resting membrane potential of neurons is close
to but not equal to the K+ equilibrium potential
because the membrane is not completely impermeable
to Na+. Therefore, at rest, Na+ diffuses into the cell
constantly along its electrochemical gradient as K+
diffuses out. (The diffusion force for Na+ is much
greater than that for K+.) Thus, the Na+/K+-ATPase
pumps must work constantly to maintain the
concentration gradients of the ions. (The fact that the
membrane at rest is much more permeable to K+ than
to Na+ is reflected in the fact that the resting
4. Explain the conditions that give rise to the membrane potential is much closer to the equilibrium
resting membrane potential. What effect does potential of K+ than to that of Na+.)
membrane permeability have on this potential?
What role do Na+/K+-ATPase membrane pumps
play in the membrane potential? Are these 7. Draw a graded potential and an action potential
functions direct or indirect? on a graph of membrane potential versus time.
Indicate zero membrane potential, resting
Membrane potentials are generated by the diffusion
membrane potential, and threshold potential;
of ions and are determined by the ionic concentration
indicate when the membrane is depolarized,
differences across the membrane and the membrane’s
repolarizing, and hyperpolarized.
relative permeabilities to different ions. In a neuron
at rest, the intracellular concentration of K+ is thirty
times greater than its extracellular concentration. Overshoot
Conversely, the extracellular concentrations of Na+ +30 Depolarized
and Cl- are about ten times greater than their
Repolarizing
intracellular concentrations. Moreover, the
permeability of the membrane to K+ at rest is much 0
greater (50-75 fold) than its permeability to Na+, and Threshold
potential 1
thus the resting membrane potential is much closer Resting
to the equilibrium potential for K+ than for Na+. The membrane
potential
Na+/K+-ATPase membrane pumps maintain the
membrane potential indirectly by pumping K+ in and
Na+ out, thereby maintaining the concentration
gradients across the cell membrane. Because the -70
2
pumps eject three Na+ for every two K+ they bring Graded
potential Hyperpolarized
into the cell, they also have a small direct
electrogenic effect on the membrane potential. TIME (milliseconds)
5. Which two factors involving ion diffusion 8. List the differences between graded potentials
determine the magnitude of the resting and action potentials.
membrane potential?
©2017 by McGraw-Hill Education. This is proprietary material solely for authorized instructor use. Not authorized for sale or distribution in
any manner. This document may not be copied, scanned, duplicated, forwarded, distributed, or posted on a website, in whole or part.
16
(1) Graded potentials (GPs) have variable favored by the positive membrane potential and the
amplitude; action potentials (APs) are all-or-none higher intracellular K+ concentration. K+ efflux
once threshold depolarization is reached. AP repolarizes the cell.
amplitude is independent of the initiating event.
(2) GPs can be summed; APs cannot be summed.
10. What determines the activity of the voltage-
(3) GPs have no threshold; APs have a threshold gated Na+ channel?
that is usually about 15 mV depolarized relative to
The Na+ channel is said to be voltage-gated because it
the resting potential.
possesses charged amino acid residues that make it
(4) GPs have no refractory period; APs have both an open and close in response to changes in the
absolute and a relative refractory period. membrane potential. Negative membrane potentials
tend to close the channel and depolarization opens
(5) GPs are conducted passively and decrementally;
the channel. Additionally, the channel can be closed
APs are conducted actively and without decrement;
by an inactivation gate that physically blocks the
the depolarization is amplified to a constant value at
channel shortly after it is opened by depolarization.
each point along the membrane.
(6) The duration of GPs varies with the initiating
conditions; the duration of APs is constant for a 11. Explain threshold and the relative and absolute
given cell type under constant conditions. refractory periods in terms of the ionic basis of
the action potential.
(7) A GP can be a depolarization or a
hyperpolarization; an AP is always a depolarization, The threshold is the level of depolarization at which
with an overshoot (the cell becomes positive, inside Na+ influx just exceeds K+ outflux, so that there is a
with respect to outside) and an after- net flux of positive charge into the cell. The driving
hyperpolarization phase in neurons. force for K+ out of the cell increases as the membrane
depolarizes, but at threshold the increased
(8) GPs are initiated by environmental stimuli
permeability of the membrane for Na+ assures that
(receptors), neurotransmitters (synapses), or
the positive-feedback cycle can be sustained.
spontaneously; APs are initiated by membrane
depolarization (i.e., GPs that depolarize to The absolute refractory period corresponds to the
threshold). period when voltage-gated Na+ channels are already
opened or are inactivated (roughly the first part of
(9) The mechanism for eliciting GPs depends on
the repolarizing phase of the action potential). The
ligand-gated channels or other chemical or physical
relative refractory period corresponds to the period of
changes; the mechanism for APs depends on voltage-
increased K+ permeability after the first repolarizing
gated channels.
phase, and this is also the timing when some of the
voltage-gated Na+ channels have returned to their
9. Describe how ion movement generates the closed state. The reason that a suprathreshold
action potential. stimulus (i.e., greater depolarization than at rest) is
necessary for an action potential to be initiated
Depolarization by a graded potential opens voltage- during the relative refractory period is because some
gated Na+ channels. The higher extracellular Na+ K+ channels are still open during this time, and net
concentration and the negative membrane potential K+ efflux favors membrane hyperpolarization. Thus,
favor the influx of Na+, which carries a positive Na+ influx resulting from a new stimulus must
charge into the cell. This depolarizes the cell further exceed K+ efflux, which is occurring simultaneously.
which opens more Na+ channels which allows more
Na+ to enter. This positive feedback mechanism
accounts for the "spike" or upstroke of the action 12. Describe the propagation of an action potential.
potential. Na+ channel inactivation prevents further Contrast this event in myelinated and
influx of Na+. As the slower, voltage-gated K+ unmyelinated axons.
channels open, the efflux of K+ begins. K+ efflux is
17
In unmyelinated axons, action potentials are saltatory conduction, and it is considerably faster
propagated by local current flow from the site of one than action potential propagation in unmyelinated
action potential to an adjacent part of the axon axons.
membrane. Positive charge flows from the
depolarized area in both directions along the
intracellular membrane (and negative charges do the 13. List three ways in which action potentials can
same along the outside of the membrane). In the be initiated in neurons.
“forward” direction, this current flow depolarizes the (1) receptor potentials
adjacent membrane to threshold, and an action
potential is generated. In the “backward” direction, (2) synaptic potentials
the membrane is still in the absolute refractory (3) pacemaker potentials
period, so it cannot be stimulated to generate another
action potential.
In myelinated axons the same process occurs, except
that current cannot flow across the membrane where
there is myelin because myelin has a very high
resistance to flow. Instead, the current flows along
the membrane from one interruption in the myelin (a
node of Ranvier) to the next. At the nodes, the
current flow causes depolarization to threshold and
an action potential is generated. This is called
SECTION C: Synapses
1. Describe the structure of presynaptic axon At inhibitory synapses, the binding of
terminals and the mechanism of neurotransmitter to its receptor results in a
neurotransmitter release. hyperpolarizing graded potential (an inhibitory
postsynaptic potential—IPSP) or a stabilization of
See Figure 6.27 and accompanying text for a
the membrane potential at (or below) its existing
description of the structure of presynaptic axon
value. The activated receptor at such synapses causes
terminals and the detailed mechanism of
ligand-sensitive channels for Cl- and/or K+ to open.
neurotransmitter release.
(In the cases where only Cl- channels open, a
hyperpolarizing potential is not recorded but the
2. Contrast the postsynaptic mechanisms of membrane is more stabilized at its resting level than
excitatory and inhibitory synapses. normal, because the increased membrane
permeability to Cl- makes it more difficult for other
At an excitatory synapse, the postsynaptic response ion types to change the membrane potential.)
to the released neurotransmitter is a depolarization Increasing the permeability to K+ causes the
(an excitatory postsynaptic potential—EPSP) postsynaptic membrane potential to become more
because binding of neurotransmitter to its receptor negative and closer to the K+ equilibrium potential.
causes ligand-sensitive channels for small, positively
charged ions to open. Even though the permeability
of the postsynaptic membrane is increased to both 3. Explain how synapses allow neurons to act as
Na+ and K+, both the electrical and concentration integrators; include the concepts of facilitation,
gradients favor Na+ influx, while the electrical temporal and spatial summation, and
gradient opposes the concentration gradient for K+. convergence in your explanation.
Therefore, there is net influx of Na+ and an increase
Neurons act as integrators because they receive
in membrane potential.
information in the form of synaptic activity from
more (often many, many more) than one presynaptic
18
cell, a property known as convergence. The amount and desensitization of receptors. (9) Certain drugs
of depolarization caused by the discharge of and diseases can affect both the presynaptic and
neurotransmitter from a single excitatory synapse is postsynaptic factors listed above. General factors
only about 0.5 mV, much less than the ~15 mV include: (10) the area of synaptic contact;
depolarization needed to reach threshold. Thus, (11) enzymatic destruction of the neurotransmitter;
activation of a postsynaptic neuron to the point that (12) the geometry of the diffusion path, and (13) the
it will generate action potentials requires summation rate of neurotransmitter reuptake.
of many excitatory synaptic events. This occurs in
two ways: The same synapse continues to be
repeatedly activated before the previous EPSPs or 5. Discuss differences between neurotransmitters
IPSPs have died away—temporal summation; and and neuromodulators.
more than one synapse may be activated Neurotransmitters cause the postsynaptic events
simultaneously or within a short time of the first described above, i.e., they elicit EPSPs or IPSPs
synapse—spatial summation. The postsynaptic cell when they bind to receptors in the postsynaptic
integrates IPSPs as well as EPSPs. In order for the membrane. Neuromodulators have more complex
postsynaptic cell to become activated to threshold, effects, including influencing the postsynaptic cell’s
excitatory synaptic activity must be considerably response to specific neurotransmitters, or changing
greater than inhibitory synaptic activity. the rate of synthesis, release, reuptake, or metabolism
Facilitation refers to one presynaptic cell’s of a transmitter by the presynaptic cell. In other
(cell A) positive influence on another presynaptic cell words, they alter the effectiveness of the synapse.
(cell B) through an axo-axonic synapse. If the The two kinds of messengers also have different time
neurotransmitter from cell A causes cell B to release courses: neurotransmitters initiate events that occur
more neurotransmitter to postsynaptic cell C when within milliseconds and are short-lived. The
an action potential reaches cell B’s axon terminal, activation of receptors for neuromodulators often
then cell A has increased cell B’s synaptic brings about changes in the metabolic processes in
effectiveness by presynaptic facilitation. (Another neurons via G proteins coupled to second-messenger
presynaptic cell with an axo-axonic synapse with systems. Such changes can occur over minutes,
cell B could decrease the amount of hours, or even days. Neuromodulators can be
neurotransmitter released by cell B. This is synthesized by the presynaptic cell and co-released
presynaptic inhibition.) with the neurotransmitter at synapses, or they can be
hormones, paracrine substances, or immune-system
messengers.
4. List at least eight ways in which the
effectiveness of synapses may be altered.
(1) The presynaptic facilitation and inhibition 6. List the major classes of neurotransmitters and
described above is one way. Other presynaptic give examples of each.
factors include: (2) availability of neurotransmitter, (1) Acetylcholine, (2) biogenic amines: the
which is in turn dependent upon the availability of catecholamines dopamine, norepinephrine, and
precursor molecules and the amount or activity of epinephrine; serotonin; 5-HT; and histamine, (3)
the rate-limiting enzyme in the pathway for amino acids: glutamate, GABA, and glycine, (4)
neurotransmitter synthesis; (3) the axon terminal neuropeptides: endogenous opioids and morphine, (5)
membrane potential—the more depolarized the gases: nitric oxide, carbon monoxide, and hydrogen
terminals are, the more voltage-gated Ca2+ channels sulfide, and (6) purines: adenosine and ATP.
open; (4) the residual Ca2+ in the terminal from
previous action potentials; and (5) the presence of
autoreceptors. Postsynaptic factors include: (6) the 7. Detail the mechanism of long-term potentiation,
immediate past history of the postsynaptic and explain what function it might have in in
membrane; (7) effects of other neurotransmitters or learning and memory. See Figure 6.36 and
neuromodulators; and (8) up- or down-regulation accompanying text description for a detailed
19
mechanism of long-term potentiation. This
mechanism might be central to learning and
memory, because it provides a way to strengthen
synapses that receive frequent, strong activation.

1. Make an organizational chart showing the CNS, 3. List two functions of the thalamus.
PNS, brain, spinal cord, spinal nerves, cranial
The thalamus is an important relay station for
nerves, forebrain, brainstem, cerebrum,
sensory pathways on their way to the cerebral cortex.
diencephalon, midbrain, pons, medulla
It also participates in control of body movement and
oblongata, and cerebellum.
skeletal muscle coordination, and it plays a key role
I. Central nervous system in awareness.
A. Brain
1. Forebrain
a. Cerebrum 4. List the functions of the hypothalamus, and
b. Diencephalon discuss how they relate to homeostatic control.
2. Brainstem (1) The hypothalamus is the “master gland” of the
a. Midbrain endocrine system and regulates the secretions of the
b. Pons pituitary gland. It also: (2) regulates water balance;
c. Medulla oblongata (3) participates in the regulation of the autonomic
3. Cerebellum nervous system; (4) regulates eating and drinking
B. Spinal cord behavior; (5) regulates the reproductive system and
II. Peripheral nervous system reproductive behavior; (6) reinforces certain
A. Cranial nerves behaviors; (7) generates and regulates circadian
B. Spinal nerves rhythms; (8) regulates body temperature; and
(9) participates in generation of emotional behavior.
2. Draw a cross section of the spinal cord showing The hypothalamus is the master command center for
the gray and white matter, dorsal and ventral neural and endocrine coordination. It is the single
roots, dorsal root ganglion, and spinal nerve. most important control area for homeostatic
Indicate the general location of pathways. regulation of the internal environment.
5. Make a PNS chart indicating the relationships

among afferent and efferent divisions, somatic
and autonomic nervous systems, and
sympathetic and parasympathetic divisions.
The peripheral nervous system
I. Afferent division
II. Efferent division
A. Somatic nervous system
B. Autonomic nervous system
a. Sympathetic division
b. Parasympathetic division
Fig. 6.41
6. Contrast the somatic and autonomic divisions superior mesenteric, and inferior mesenteric
of the efferent nervous system; mention at least ganglia—are in the abdominal cavity, closer to the
three characteristics of each. innervated organ. The parasympathetic ganglia all
lie within the organs innervated by the
The neurons of the somatic nervous system innervate
postganglionic fibers or close to them, and so the
skeletal muscle, whereas autonomic neurons
postganglionic parasympathetic fibers are quite
innervate smooth and cardiac muscle, glands, and
short.
neurons in the gastrointestinal tract. The somatic
system consists of single neurons between the CNS The two divisions differ functionally as well. To
and skeletal muscle cells, whereas the autonomic some extent the sympathetic division acts as a single
system has two-neuron chains (connected by a unit, whereas the parasympathetic division is made
synapse in a ganglion) between the CNS and the up of relatively independent components. Perhaps
effector organ. The output of the somatic system is most importantly, the two divisions often innervate
always excitatory, whereas the autonomic output can the same organ and usually have opposite effects. In
be excitatory or inhibitory. general, the sympathetic division is activated in
situations requiring physical action—the fight-or-
flight response—and also when one is under
7. Name the neurotransmitter released at each psychological stress. The parasympathetic
synapse or neuroeffector junction in the somatic system is more dominant in times of rest-or-
and autonomic systems. digest.
Acetylcholine is the neurotransmitter released by: (1)
the somatic motor neurons at neuromuscular
9. Explain how the adrenal medulla can affect
junctions; (2) the preganglionic fibers of both the
receptors on various effector organs despite the
parasympathetic and sympathetic divisions; and (3)
fact that its cells have no axons.
postganglionic, parasympathetic fibers.
Norepinephrine is released by postganglionic The adrenal medulla is essentially a sympathetic
sympathetic fibers. The adrenal medulla releases ganglion. The postsynaptic cells within the adrenal
epinephrine into the blood; it is considered a medulla secrete their chemical messengers—
neurohormone. primarily epinephrine with some norepinephrine and
small amounts of dopamine—into the blood;
therefore, the substances released by the adrenal
8. Contrast the sympathetic and parasympathetic medulla are considered neurohormones. The
components of the autonomic nervous system; neurohormones are transported via the blood to
mention at least four characteristics of each. effector cells having receptors sensitive to them.
The first difference is mentioned above: the
postganglionic fibers of each division secrete different
10. The chemical composition of the CNS
neurotransmitters.
extracellular fluid is different from that of
The two divisions also differ anatomically: the nerve blood. Explain how this difference is achieved.
fibers of the two divisions leave the CNS from
A complex group of blood-brain barrier mechanisms
different levels—the sympathetic fibers from the
closely controls the kinds of substances that enter the
thoracic and lumbar areas of the spinal cord, and the
extracellular fluid of the brain and the rate at which
parasympathetic fibers from the brain and the sacral
they enter. The blood-brain barrier comprises the
portion of the spinal cord. The locations of ganglia
cells that line the smallest blood vessels in the brain
also differ for the two divisions: most of the
(the capillaries), and has both anatomical features
sympathetic ganglia lie close to the spinal cord and
such as tight junctions and physiological transport
form two chains of ganglia, the sympathetic trunks.
systems that handle different classes of substances in
The postsynaptic fibers from these ganglia can be
different ways.
quite long. Other sympathetic ganglia—the celiac,
21
SUMMARY OF INSTRUCTOR RESOURCES: CHAPTER 6
Figures
&
Chapter CONNECT
Section Tables Chapter Test CONNECT
Topics Review Question
(pages) (* denotes Questions Animations
Questions Bank
Physiological
Inquiry)
2. Neuron structure
3. Structure and
maintenance of
neurons
4. Functional classes of
neurons
Fig. 6.1, 6.2, 5. Anatomy of efferent,
Nervous Section A Recall and
6.1-6.5 6.3, 6.4, 6.5 afferent, and
System Comprehend:
(pp. 137-143) interneurons
Cells p. 143 #2
6. Glial cells
Table 6.1 7. The truth about glial
cells
8. Neural growth and
regeneration
9. The truth about neural
plasticity
1. General principles of
physiology
10. Basic principles of
electricity
11. Potential, current, and
resistance
12. Electrical properties of
membranes
13. Ion distribution
14. Nerst equation
15. Resting membrane
potential
16. Equilibrium potentials
Fig. 6.7, 6.8, Recall and
and ion flux
6.9, 6.10*, Comprehend:
17. Membrane potential
6.11*, 6.12*, #3, 4, 5, 6, 7,
terminology
6.13, 6.14,
18. Membrane potential
6.15*, 6.16, Apply,
glossary
6.17, 6.18, Section B Anayze, and
Membrane 6.5-6.7 19. Action potential
6.19*, 6.20, Evaluate:
Potentials (pp. 143-158) 20. Feedback control of
6.21, 6.22, pp. 157-158 # 1, 2, 5, 7, 8 Action potential
voltage-gated channels
6.23*, 6.24* propagation in an
21. Action potentials and
General unmyelinated axon
refractory periods
Principles
22. Graded vs. action
Table 6.2, Assessment:
potentials
6.3, 6.4 #2
23. Voltage clamp
experiment
24. Voltage clamp toxin
51. Charge movement
during an action
potential
52. Adjacent to an action
potential
53. One-way action
potential propagation
54. Directionality of action
potentials
55. Threshold definition
Chapter 6 22
physiology
25. Functional anatomy of
synapses
26. Label the synapse
Recall and 27. Neurotransmitter
Fig. 6.25, Comprehend: release
6.26, 6.27, #8, 9 28. Steps of
6.28, 6.29, neurotransmitter
6.30, 6.31*, Apply, signaling
6.32, 6.33, Section C Anayze, and 29. Excitatory chemical
6.8-6.14
Synapses 6.34, 6.35, Evaluate: synapses
(pp. 66-171)
6.36 p. 170 #6 30. Inhibitory chemical
synapses
General 31. Synaptic modification
Table 6.5, 6.6 Principles 32. EPSP vs. IPSP
Assessment: 33. Synaptic integration
#3 34. Synaptic facilitation
35. Long-term potentiation
36. Specific
neurotransmitters
37. Neuroeffector
communication
physiology
37. Neuroeffector
Recall and communication
Fig. 6.37*, Comprehend: 38. Brain anatomy
6.38, 6.39, #1, 10 39. Nervous system terms
6.40, 6.41, 40. Brain region functions
6.42, 6.43, Apply, 41. Spinal cord
Structure
6.44, 6.45, Section D Anayze, and 42. Spinal cord injuries
of the 6.15-6.19
6.46*, 6.47 Evaluate: 43. Cranial nerves
Nervous (pp. 171-185)
p. 182 #3, 4, 44. Spinal nerves
System
Table 6.7, 45. Somatic vs. autonomic
6.8, 6.9, 6.10, General nervous system
6.11 Principles 46. Divisions of the ANS
Assessment: 47. Autonomic drugs
#1, 3 48. Ventricles of the brain
49. The truth about the
blood-brain barrier
50. Clinical case study
Another random document with
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Tūmbaku (tumbākū).—Tobacco used for the kalian or water-
pipe.
Tūmbūn (tumban).—Petticoats (made as very loose drawers).
Umbar (ambār).—A cellar or store-room, a go-down.
Ūtū or Ootoo (atw).—An iron (flat or otherwise).
Utu-kesh (atw kash).—An ironer.
Vakeel-u-dowleh (wakīl ud Dawlah).—An agent of
Government; the empty title given to native newswriters,
who are supposed to act as English Consuls, and whose
offices are sinecures.
Yabū (yābū).—A pony, a common horse, a horse.
Yahū (?).—A kind of common house pigeon.
Yakhjal (Yakh-chāl).—An ice-store; a pond (and wall) for
making ice.
Yaourt (Turkish).—Curdled milk, (Persian) “mast.”
Yashmak (Turkish).—A kind of veil, or face covering.
Yawash (Turkish).—Gently, slowly.
Yessaoul (yasāwal).—A mounted mace-bearer.
Zalābi or Zalābieh (zalībiyā).—A sweet cake or fritter eaten
in Ramazan at night.
Zambūrek (zambūrak).—A tiny cannon carried on and fired
from a camel’s back (from Zambūr, a wasp).
Zangal (zangāl).—A legging.
Zenda-Rūd, Zendarūd (Zanda-Rud).—The river at Ispahan.
Zil-es-Sultan (Zill us Sultan) (title).—Shadow of the King.
Zoban-i-Gunghishk (zabāni gunjishk).—Lit. sparrow’s
tongue, a kind of willow.
Zūlf (zulf).—A long love-lock, a curl.
FOOTNOTES
[1] See Appendix D, page 417.
[2] Turcomanchai was the place where the treaty between
Persia and Russia was signed, February 22, 1828. Erivan and
Nakchewan were ceded to Russia, and two millions agreed to be
paid to her.
[3] The form of these was very various, though the principle of
action was always the same: the smoke was conducted to the
bottom of a pint or more of water and then sucked up in bubbles
through it, a gurgling noise being produced. Some used the long
“snake” or nehpeech, a spiral of copper wire covered with
coloured leather, and forming a flexible air-tight tube some four
yards long; this was the more old-fashioned way, and required
good lungs. A servant held the pipe itself at the side of the
master’s chair. Others affected the wooden stem with the pipe;
this as a rule is held by the smoker himself, and no great effort is
required in smoking, as the tube is only eighteen inches long and
air-tight, which the “nehpeech” or “snake” seldom is, save when
quite new.
The portion between the pipe-head and the water-holder is as a
rule always the same: a wooden tube some fourteen inches or
more long, with numerous indentations, turned in a lathe, and
coming to a point, so that any pipe-head will fit it; from the end of
this an inner tube goes to within an inch of the bottom of the
water. Sometimes this tube is made of ebony, at other times
covered with silver, and rarely with gold. In its side at the bottom
is the hole for the snake-like tube, or the stick.
The water-reservoir is usually of glass, either plain crystal, or
cut Bohemian; the shape of these glasses is that of a wide-
mouthed, long-necked decanter, and the neck serves as the place
by which the whole contrivance is held. In summer a porous clay
bottle is generally used as cooler by all classes, rich or poor.
Another kind of reservoir called a narghil (narghil, a cocoa-nut)
is made, having its shape like a cocoa-nut, with a spike or small
knob at the sharp end; this rests on the ground, and is meant for
travelling. It is made of brass, silver, or gold, and often in the two
latter cases enamelled; the “meāna,” or middle tube, to this kind
of pipe is often two and a half feet long, and the stem two.
Yet another form of kalian exists for travelling, and that is a
copy of the glass reservoir, of a rather squat shape, in buffalo or
rhinoceros hide; this is often, indeed usually, covered with
enamelled plates of gold and silver, often encrusted with gems,
and is only in use among the very rich.
As the great personages of Persia are constantly travelling,
these more elaborate forms of pipe are frequent; and, as a man’s
pipe often gives an idea of his social position, money is very
freely lavished on them. The mouth-piece is simply either
wooden, or else the end is shod with silver. The head consists of,
among the poor, a clay reservoir for the tobacco. These cost a
farthing. But most Persians, though only of the lower middle
class, manage to have a silver pipe-head; this consists of three
pieces, the handle or chōb (wood), a carved and turned piece of
wood pierced with a conical hole which fits the meāna (or stem)—
this may be represented by the lower two-thirds of an old-
fashioned wine-glass, with a small foot; the fire-holder, which is of
gold, silver, or stone, is fitted to this, and represents the upper
third of the wine-glass; and on this all the ingenuity of the
Persians is lavished in the matter of ornament. From its under
edge hang four or six little silver or gold chains four inches long,
terminated by flattened balls.
Lastly, the wind-guard, which prevents the fire from falling or
being blown up into an excessive state of incandescence, is
usually made of silver, and is an inverted cone of the same size
as the fire-holder, fitted to it with accuracy, and provided with two
holes to give the requisite amount of draught; at the side two pairs
of chains depend from the upper edge of this, and are made to
reach as far as do the lower set.
The fire-holder is lined with a mixture of clay and plaster of
Paris, on which is placed the tobacco, freshly moistened and
rubbed into coarse fragments (though connoisseurs prefer a more
elaborate preparation)—about three-quarters of an ounce is
required; it is flattened and smoothed, the surplus water being
squeezed away. Upon it are placed morsels of live charcoal,
which are blown into a fierce flame, and the excess of water in the
reservoir or bottle being driven out by blowing from the bottle,
which is always nearly filled. A few draws are taken by the pipe-
boy to see that all goes well, and to get rid of the taste of fresh
charcoal, and get the tobacco well alight, and it is then handed to
the smoker as under weigh.
On the fire-holder, however—perhaps because it is opposite
the eye and so most conspicuous—are seen the highest efforts of
Persian art. It is, whenever it can be afforded, of purest gold,
though often thin; some rare exceptions are unornamented; more
ordinarily it is chased or covered with high repoussé work, or
elaborately engraved. Or it may be so encrusted with turquoises
till little, if any, of the original metal shows; or it may be
ornamented with elaborate enamels of birds and flowers, or of
fruit; and a favourite pattern is vine-leaves of transparent enamel
let into the deeply-cut metal, and the bunches of grapes of varied
colours.
More often three or four ovals, some two inches long, are filled
by portraits of a girl or boy—of course fancy ones—and the
spaces between them filled with flowers and birds. These
enamels are very beautiful, very costly, and very brittle; ten
pounds being a common price paid to an enameller to decorate a
gold head, while as much as one hundred tomans, or forty
pounds, are given by great and rich amateurs.
Of the kalians, the heads and reservoirs of which are thickly
encrusted with gems, I do not speak at present; I had few
opportunities at that time of seeing such, and, as a rule, they are
only possessed by the Shah, his sons and uncles. I trust the
reader will bear with this long but needful detail as to pipes.
[4] As a rule, in Persia every one is up by six a.m.
[5] Those who feel curious on the subject of modern Persian
medicine, I must refer to my article on the subject in the British
Medical Journal.
[6] The English Legation or Embassy is always called “The
Mission” in Persia, by the members of it, and the English in the
country.
[7] Futteh-Ali Shah had over seventy sons and daughters, and
a prince’s son in Persia is a prince.
[8] As some confusion may be experienced in the matter of
money terms, I may append the following table of coins:—
s. d.
(Copper) 2 pūls = 1 shahi (or shaie) or 0 0½
English
banabat or half-keran
” 10 shahis = 1 ” 0 5
(silver)
20 shahis = 1 keran (silver) ” 0 10
10 kerans = 1 toman (tomaun), gold ” 7 6
Were the keran really tenpence, of course the tomaun would be

8s. 4d., but its value is really only ninepence at present exchange
(1883). Of these coins the pūls and shahis are copper, the kerans
and half-kerans or banabats silver, and the tomauns gold; though
for the past fifteen years, until just recently, the tomauns (in gold)
had nearly disappeared, and were merely nominal, or old coins
hoarded for the sake of the purity of their gold. Prices are given
indiscriminately in tomauns or kerans; the price in kerans as five
hundred kerans being mostly spoken of and always written as
kerans and not fifty tomauns. Till lately the tomaun has been only
a name. The merchant-class, too, use the dinar, an imaginary
coin (not now minted at least), as a convenient fraction for
calculation.
I on arrival took my servants’ accounts in tomauns and kerans,
afterwards in kerans and shaies, and at last in kerans and pūls;
while an English merchant friend actually wrote his house
accounts in dinars, and said it awed his servants! one thousand
dinars make a keran, so one dinar is the 1/1000 of 9d.
There are no bank-notes: and in The Times telegraphic news,
under the head of Persia, Friday, February 24th, 1883, is a
summary of a truly Persian edict. By it the Shah informs his
subjects that, “they are foolish to take dirty pieces of paper for
gold and silver, and that in future all Russian Rouble notes will be
confiscated!” Then follows a really useful prohibition forbidding
aniline dyes, and ordering such, when imported and discovered,
to be destroyed; these dyes, which are not fast, have been lately
much used by ignorant carpet-weavers in Persia.
[9] Hakim, a doctor or physician.
[10] This system accounts partly for the apparently very low
wages paid to the Persian servant, which are (I give those paid
latterly—1881—by myself; in the case of head-servants it is
sometimes, but very seldom, more, as the pay is of course
nothing to the modakel):—
A month. £ s. d.
A nazir or steward 50 kerans, or 2 0 0
A good cook 50 ” 2 0 0
A good peishkhidmut
(personal servant, waits at
40 to 50 kerans, or 30s. to 2 0 0
table, and valets one, and is
expected to dress well)
A farrash, i. e. sweeper or
25 kerans, or 1 0 0
message runner
A sherbet-dar, plate-
cleaner, maker of coffee, 25 ” 1 0 0
ices, etc.
A second farrash 20 ” 0 16 0
A third farrash 15 ” 0 12 0
A cook’s disciple, or
10 ” 0 7 6
scullery man
A washerman, or woman
who can wash and iron 35 ” 1 6 0
thoroughly
A woman-servant or nurse 25 ” 1 0 0
A head-groom 30 ” 1 5 0
An under-groom 20 ” 0 16 0
[11]? Mustela Sarmatica.

[12] More correctly munshi.
[13] Or nummud.
[14] Here are four tombs, cut in the face of the solid rock, those
of Darius, Xerxes I., Artaxerxes I., and Darius II. A detailed
description is to be found in Usher’s book. (See illustration.)
[15] Russian subjects are well protected in Persia, and no injury
or insult to them is allowed to pass by their embassy.
[16] The present comparative dearness of provisions, such as
bread, milk, eggs, etc., is compensated for by a corresponding
cheapness in the price of sugar, candles, etc., which formerly
were more expensive. I append a list of prices in Ispahan in 1882:
—
Kerans. s. d.
Rice (per maund, 14 lbs.) 2 1 6
Mutton ” ” 2 1 6
Beef ” ” 1½ 1 1½
Fowls (each) ¾ to 1 7d., 8d. and 0 9
Small chickens (each) ⅓ 0 3
Pigeons ” 0 2
Partridges ” ½ 0 4½
Eggs (40 to 60) 1 0 9
Butter (14 lbs.) 5 3 9
Clarified butter or ghee for
5 to 7 4s. to 5 0
cooking (14 lbs.)
Coffee, Mocha (per lb.) 1 9d. to 0 10
Tobacco (14 lbs.) 4 to 12 3s. to 10 0
Potatoes ” ½ to 2 4½d. to 1 6
Wood for firing (280 lbs.) 2½ 1 9
” broken, in small
5 3 9
quantities (280 lbs.)
Loaf-sugar, English (per lb.) ¾ to 1 6d. to 0 9
Charcoal, sifted (14 lbs.) ½ to 1 4½d. to 0 9
” unsifted ” ¼ to ½ 2d. to 0 4½
Grapes ” 7/20 to 15/20 3d. to 0 7
Dip candles ” 4 3 0
Commonest oil for servants
1½ 1 1½
(14 lbs.)
Bread (14 lbs.) 1 to 1½ 9d. to 1 1
The cost of horse-keep, including grooms’ wages, shoeing,

etc., is from 9d. to 1s. a day; this is supposing several are kept.
[17] I use this word for want of a better.
[18] On March 30th I left Erzeroum at nine a.m., reached
Hassan Kaleh, twenty-four miles, at three p.m.; started again at
four p.m. (all snow), reached Balakoohi, where a storm compelled
us to halt at seven p.m. Slept there.
March 31st.—Started at five a.m. for Kharassan, twenty-five
miles; arrived at half-past nine; made a detour of ten miles on
account of water. Started at half-past eleven on same horses;
stopped at a village twenty-four miles off, name unknown; horses
dead beat; road—water, mud, and thawing snow—twenty-four
miles.
April 1st.—Started at five a.m.; arrived at Moollah Suleiman,
eighteen miles (same sort of road), at ten a.m.; left at half-past
eleven for Kadikeesa, twenty miles; arrived at five p.m.; went on
through snow till nine p.m. to a village, twelve miles only; halted.
Slept in a sheep-shed full of tics.
April 2nd.—Started at six a.m.; arrived at three p.m. at
Desardūn, thirty-six miles. Here I saw Mount Ararat. Road very
bad, from melting snow. Arrived at Kizzil Deeza, twenty-four miles
(a wretched hole), at eight p.m.
April 3rd.—Five a.m. Road pretty fair over a long snow-pass,
twenty miles to Abajik, in Persia. Arrived at ten a.m. Quite a
pleasure to get among the Persians again, and to be able to
make myself understood. Then an easy twenty miles to Keranee
—half-past four p.m. Started at once; reached Zarabad, twenty-
two miles over a good road with capital horses, at half-past eight
p.m.
April 4th.—Left at half-past six a.m. for Khoi, a long twenty-four
miles; got in at eleven a.m. Khoi is a very large place, apparently
prosperous. Good dinner of bazaar kabobs. Arrived at Turseh,
twenty-four miles, ten p.m.; road good, but shocking horses, down
a tremendous pass, then along the shores of Lake Ooroomeyeh
—a kind of Dead Sea—it is very salt. Many bituminous fires
lighted it up at night, huge sheets of flame suddenly appearing.
April 5th.— Half-past two a.m., left Turseh for a place the name
of which has escaped me, arriving at half-past ten a.m. Arrived at
Sufian at half-past twelve noon. Left at once, reaching Tabriz,
twenty-four miles off, in four hours. Erzeroum to Tabriz, six days
and seven hours, three hundred and thirty-five miles.
I stopped with Colonel J⸺, V.C., our Consul-General, April
6th, 7th, and 8th. At three p.m. of April 9th I started for Teheran.
Stages between Tabriz and Teheran.
Miles.
Saoudabad 20
Hadji Aga 20
Darathiar 16
56
Slept.
April 10th.— Turcomanchai 24
Meana 16
Jemalabad 12
Tercham 16
Aga Mezar 12
Nikibeg 20
100
Slept four hours.

Zinjan (met one of our staff;
April 11th.— 26
breakfasted, and lost three hours)
Sultaneah 24
Khya 30
80
Slept four hours.

April 12th.— Khirve 18
Zeedaen 24
Kasvin 16
Abdulabad 18
76
Slept four hours.

April 13th.— Sufferkhoja 24
Shunkerabad 22
Meanjūb 20
Teheran 20
86
[19] This is the one standard weight of Persia, the other being
the miscal or sixth part of our ounce. This, for convenience, is
supposed to consist of twenty nokods—the nokod being a grain
similar to our pea in appearance. The nokod is subdivided into
three gundums or grains (of wheat); these again into four kērāts
(or carats)—these latter, however, are only used in weighing
gems. The Tabriz maund (or batman) and the miscal and its
subdivisions are in use throughout Persia in mercantile affairs.
Further north than Ispahan the sere and the gerewankeh—the
latter about a pound, and borrowed from the Russians—are in
use. Other local weights exist, only known in special places. As a
rule, each village has its special weight (literally stone, “sang”),
and their maunds get lighter and lighter as one gets away from
the large cities.
[20] The cost had been—
Kerans.
Thirty jars, at five kerans 150
Twenty loads of grapes 750
Carriage of same 60
Cost of labour, etc. 100
1060
Per contra.
Kerans.
Paid to me by Jews for refuse, for arrack-making 50
Resale of jars 140
190
Total cost, 870 kerans, or about 5½d. per bottle.

[21] Must is a Persian word signifying “drunk;” it also means the
state of excitement of male camels at certain times.
[22] Strangled, after he had refused a cup of poisoned coffee, in
1882, by order of the Zil-es-Sultan, while an honoured guest in his
(the Zil-es-Sultan’s) house.
[23] Topi, a sun-helmet.
[24] The full title is Kawam-ul-Molk.
[25] The full title is Muschir-ul-Molk.
[26] Would it have been necessary to have explained to Bishop
Bonner the use of the thumbscrews after his cruelty to the
Reformers?
[27] Aug. 2nd, 1887. Dr. Odling writes me that this well is 849
feet deep. I fancy that there is some error in this, as I put down
the six hundred yards. Possibly there may be a second shaft.
[28] The murdered sons of Ali, considered by the Persians, and
all Shiah Mahommedans, as the rightful successors of the
prophet, consequently sainted martyrs.
[29] Bhang, an intoxicating drug used by dervishes.
[30] Azraël, the angel of death.
[31] Not of coral, as in the Levant.
[32] Origin of our word “hummums.”
[33] Now Sir Oliver St. John.
[34] See Ussher’s ‘London to Persepolis,’ p. 564.
[35] Muleteer.
[36] See Appendix D, page 417.
[37] Shah Abbas the Great built caravanserais of great size and
solidity all over Persia, hence a good and large caravanserai,
even though not built by him, is called a “Shah Abbas
caravanserai.”
[38] Mushir al Mulk, counsellor of the province.
INDEX.
Abadeh, 261, 356

carvings, 332
Abbah, the, 319
Abbas Kūli Khan, 215
the Great, 161
Abdul, 285
Abdul Hamid, 276, 353
Abdullah’s types, 9
Abdul Mahomed, 64
Ab-i-Rūkhni, 218
Ab-i-Zungi, 218
Ableh, 62
Abū Seif Mirza, 59, 84
Senna, 82
Accident to Mr. H⸺, 128
Accidents in driving, 374
Actors, Persian, 282
Aden, 343
Administration of justice, 146
Adulteration of opium, 180
Afghan poosseens, 319
(?) tiles, 198
Aflatoon, 82
Agha Hassan, 109
Ahs an Ahs, 96
Ahū, 167
Aid-i-No Ruz, 48, 51
Akbar Khan, 402
Alangū, 323
Alarm of robbers, 130
Alexander the Great, 378
coins of, 76
Algiers, 342
Ali Akbar, 282
death of, 283
Ali Oh! 43
Alison, His Excellency Mr., 48, 201
Alla Sung, 392
Alligators, 344
American missionaries, 144
Aminabad, 262
Ancient Armenian language, 140
buildings, disappearance of, 364
engraved ruby, 37
Julfa, 161
mud-houses, 137
Anderūn, 92
Anecdote of a dervish, 47
re smoking, 32
Aniline dyes, 149
dyes, prohibition of, 63
Animals, treatment of, 316
Antelope hunt, 86
Antelopes, 56
Apostate monk, 139
Appetites, large, 336
Apricots, 168
April, the 1st of, 330
Arab dress, 110
horse, my, 61
horses, 106
pipe, 33
Arachnoort, the, 138, 141, 159
Ararat, hailstones at, 391
Araxes river, 19, 313
‘Arcot,’ voyage in the, 341
“Armchair,” 136
Armenian Alsatia, 142
artificers, 162
baptism, 141
church, 160
converts, 164
cook, 363
fasts, 144
grateful, 93
graves, 162
jewellers, 162
Kaweh Khana, 163
loafers, 143
marriage of, 141
priest, 132
Protestant teacher, 140
schools, 144
scriptures, 140
theatre, 9
tribute, 376
village, 131
wine-sellers, 142
women, dress of, 132
women, industry of the, 360
Armenians, 72, 110
anecdote of, 73
apostatising, 111
bread, 336
carpenters, good, 123
character of, 316
disguised as Europeans, 72
education of, 144
former oppression of, 144
idleness of the, 359
improved position of, 144
of Hamadan, 72, 74
position of in Persia, 74
sanctity of, 73
successful, 143
taken to Julfa, 161
uncleanliness of, 316
Arms, 322
Arnold, Mr. Arthur, 273
Arrack, 141, 159, 360
Art of avoiding falls, 54
Arts, lost, 162
Asparagus, wild, 168
Ass, wild, the, 308
Assadabad Pass, 101
Astrachan, 405
As we turn in another turns out, 110
Attempts to proselytise among the Persians, 144
Audience at Tazzia, 281
Austrian officers, 371
Avadavats, 347
Avicenna, 82
Ayrton, Mr., 5
B⸺, Mr., 27, 213

B⸺, Rev. R., 340
Baab, cursing, 155
Baabi artificers, 164
conspiracy, 154
death of a, 154
revolt, 272
Baabiism, tenets of, 154, 339
Baabis, 144, 339
charges against, 154
visit to, 201
Baab, 153
Bad drainage, 153
Badraghah, 56
Bad water, 153, 241
Baghalli, 236
Bagh-i-No, 218
Bagh-i-Takht, 220, 292
Baker, an ungrateful, 183
Bakhtiaris, 262
Bakū, 403
Bamiah, 170
Bankers, 192
Bank-notes, edict as to, 63
Baptism, Armenian, 141
Barber’s Bridge, the, 389
Bargain for mules, a, 381
Bargains, 187
Barley, 102
Bastinado, the, 146
at Kūmishah, 254
degrees of, 148
Bath carpets, 152
at Constantinople, 212
the, 334
Bazaar, at Teheran, 372
breakfasts, 200
practice, 182
Bazaarcha Baland, 200
Bazū-band, 290, 323
Bear and dog fight, 227
Beards, 321
Beaters, 177
Bēbē Sakineh Sultan Khanūm, 215
Bedding for travelling, 55
Bedding horses, 101
Beef, 142, 299
Beetles, road, 215
Bell tower, Julfa, 139
Bells, substitute for, 139
Belly-dance, the, 115
Belt, the, 320
Belūchistan, 345
Berlin, 407
Besitūn, 109
Bewitched, 65
Bishop Moses, 138
of Julfa, the, 159
Thaddeus’s tomb, 158
Bishop’s, the, pictures, 159
Bits, native, 329
Black-wood furniture, 345
Black flags, 283
Sea, 9
Blandford, Mr. W., 321
Blowing from a mortar, 203
from guns, 202
Boat-building, 247
Boat journey, 210
Boiled to death, 272
Bombassi, 326
Bonaat, 77
Bookbinding, 288
Boorio, 197
Boots, 321
Borasjūn, 348
Boulevard at Teheran, 371
Bowin, 392
Boy dancers, 246
singers, 281
Boys, a mob of, 393
Bread, varieties of, 335
Breasts as ornaments, 132
Bribery, 189
Bribing postmaster, 13
Bricking up alive, 269
Brickwork, fine, 222
Bridge of tombstones, 163
Brigands, 263
Broom plant, 309
Bruce, Mrs., 164
Rev. Dr., 164
Bulbul, 114
Bull-terriers of Zil-es-Sultan, 366
Bunder Abbas, 345
Burial of a Christian child, 140
Burke, Captain, 344
Burmese Embassy, 376
Burning alive, 204
Bushire, 345

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Vanders Human Physiology The

Mechanisms of Body Function 14th

6: NEURONAL SIGNALING AND THE STRUCTURE

CONTEXT FOR CHAPTER 6

CONCEPTS COMMONLY FOUND TO BE CHALLENGING – TEACHING HINTS

I. The central and peripheral nervous systems

III. Functional classes of neurons (Fig. 6.4, Table 6.1)

I. Basic principles of electricity

TEACHING/LEARNING OBJECTIVES BY SECTION

SECTION A: Cells of the Nervous System

6.2 Functional Classes of Neurons

6.3 Glial Cells

6.4 Neural Growth and Regeneration

SECTION B: Membrane Potentials

6.5 Basic Principles of Electricity

6.6 The Resting Membrane Potential

6.7 Graded Potentials and Action Potentials

6.9 Mechanisms of Neurotransmitter Release

6.10 Activation of the Postsynaptic Cell

6.11 Synaptic Integration

6.12 Synaptic Strength

6.13 Neuroeffector Communication

SECTION D: Structure of the Nervous System

6.15 Central Nervous System: Brain

describe the general structural organization of the brain.

6.16 Central Nervous System: Spinal Cord

6.17 Peripheral Nervous System

6.16 Autonomic Nervous System

6.19 Protective Elements Associated with the Brain

CHAPTER 6 REVIEW QUESTIONS & ANSWERS

SECTION A: Cells of the Nervous System

SECTION B: Membrane Potentials

6. Explain why the resting membrane potential is

SECTION D: Structure of the Nervous System

5. Make a PNS chart indicating the relationships

SUMMARY OF INSTRUCTOR RESOURCES: CHAPTER 6

Were the keran really tenpence, of course the tomaun would be

[11]? Mustela Sarmatica.

The cost of horse-keep, including grooms’ wages, shoeing,

Slept four hours.

Slept four hours.

Slept four hours.

Total cost, 870 kerans, or about 5½d. per bottle.

Abadeh, 261, 356

B⸺, Mr., 27, 213

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