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ISTUDY
 Human Physiology SIXTEENTH EDITION

STUART IRA FOX

Pierce College
Krista Rompolski
Moravian College

ISTUDY
 HUMAN PHYSIOLOGY
Published by McGraw Hill LLC, 1325 Avenue of the Americas, New York, NY 10121.
Copyright ©2022 by McGraw Hill LLC. All rights reserved. Printed in the United States
of America. No part of this publication may be reproduced or distributed in any form or by
any means, or stored in a database or retrieval system, without the prior written consent of
McGraw Hill LLC, including, but not limited to, in any network or other electronic storage or
transmission, or broadcast for distance learning.
Some ancillaries, including electronic and print components, may not be available to customers
outside the United States.
This book is printed on acid-free paper.
1 2 3 4 5 6 7 8 9 LWI 24 23 22 21 20
ISBN 978-1-260-59766-0
MHID 1-260-59766-0
Cover Image: ( Heart) ©Science Photo Library - PIXOLOGICSTUDIO/Getty Images
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All credits appearing on page or at the end of the book are considered to be an extension of the
copyright page.

The Internet addresses listed in the text were accurate at the time of publication. The inclusion of
a website does not indicate an endorsement by the authors or McGraw Hill LLC, and McGraw
Hill LLC does not guarantee the accuracy of the information presented at these sites.

mheducation.com/highered

ISTUDY
 Brief Contents

1 The Study of Body Function 1 13 Blood, Heart, and Circulation 403

2 Chemical Composition of the Body 24 14 Cardiac Output, Blood Flow, and Blood
Pressure 449
3 Cell Structure and Genetic Control 49
15 The Immune System 492
4 Enzymes and Energy 86
16 Respiratory Physiology 531
5 Cell Respiration and Metabolism 104
17 Physiology of the Kidneys 580
6 Interactions Between Cells and the Extracellular
Environment 128 18 The Digestive System 618

7 The Nervous System 160 19 Regulation of Metabolism 662

8 The Central Nervous System 204 20 Reproduction 702

9 The Autonomic Nervous System 242 Appendixes

Answers to Test Your Knowledge Questions A-1
10 Sensory Physiology 265
Medical and Pharmacological Abbreviations B-1

Glossary G-1
11 Endocrine Glands 315
Index I-1
12 Muscle 358

iii

ISTUDY
 About the Authors

Stuart Ira Fox earned a Ph.D. in human physiology

from the Department of Physiology, School of Medicine,
at the University of Southern California, after earning
degrees at the University of California at Los Angeles
(UCLA); California State University, Los Angeles; and UC
Santa Barbara. He has spent most of his professional life
teaching at Los Angeles City College; California State
University, Northridge; and Pierce College, where he
has won numerous teaching awards, including several
Golden Apples. Stuart has authored forty-two editions
of seven textbooks, which are used worldwide and have
been translated into several languages, and two novels.
When not engaged in professional activities, he likes to
hike, fly fish, and cross-country ski in the Eastern Sierra
Nevada Mountains.
To my wife, Ellen; and to Laura, Jacob, and Kayleigh. For
all the important reasons.

Krista Lee Rompolski earned her Ph.D. in exer-

cise physiology from the University of Pittsburgh, Depart-
ment of Health and Physical Activity, after earning her
bachelor’s and master’s degrees from Bloomsburg Uni-
versity, near her birthplace of Mount Carmel, PA. Krista
is currently an associate professor of Physical Therapy at © Ellen Fox

Moravian College in Bethlehem, PA, where she teaches

Gross Anatomy and Pathophysiology to the Physical
Therapy students, as well as Anatomy and Physiology to
undergraduate Health Sciences students. Prior to join-
ing Moravian College, Krista taught Anatomy, Physiology,
Pathophysiology, and clinical research courses at Drexel
University for seven years.
To the bravest person I know, my husband, Dan, for
always reminding me of what matters most.
© Katherine Coccagna

iv

ISTUDY
 Preface

The Story of the Sixteenth Edition This sixteenth edition marks a major addition to Human
Physiology: Krista Rompolski, Ph.D. (Moravian College) has
Stuart Fox, Ph.D., contributed significantly to the revision of chapters 8 and 18.
wrote the first edition As a very active physiology educator, Krista brings a new per-
(published 1983) to spective and her own expertise to make this edition an even
help students under- more exciting revision. This was achieved while maintaining
stand the concepts of the book’s tradition of remaining readable, accessible, and
human physiology, useful to students.
and this objective has To create this landmark sixteenth edition, Stuart had the
remained the guid- support of Krista Rompolski as coauthor and a superb team
ing principle through at McGraw-Hill. This team includes Matthew Garcia, Melisa
all of the subsequent Seegmiller, Sherry Kane, Brent Dela Cruz, Joan Weber, Angela
editions. All editions FitzPatrick, Valerie Kramer, Jim Connely, Kristine Rellihan,
have been lauded Beth Blech, and Lori Hancock. We are all incredibly grateful
for their readability, to the many reviewers who provided their time and expertise to
the currency of the critically examine individual chapters and be Board of Adviser
information, and the partners. These reviewers and advisers are listed on the pages
clarity of the presen- that follow.
tation. The sixteenth
edition continues this tradition by presenting human physi-
ology in the most current, readable, and student-oriented
way possible.

ISTUDY
 4
118 Chapter 5

Guided Tour
to 85% of the body’s energy is stored as fat, which amounts
Bile acids Steroids to about 140,000 kilocalories. Stored glycogen, by contrast,
accounts for less than 2,000 kilocalories, most of which
Cholesterol
(about 350 g) is stored in skeletal muscles and is available
for use only by the muscles. The liver contains between 80
REFRESH YOUR MEMORY and 90 C H A Pof
grams TEglycogen,
R O UTLI N E can be converted to glucose
which
and used by other organs. Protein accounts for 15% to 20% of
Ketone bodies Citric acid 4.1 Enzymes as Catalysts 87
cycle)to review 2these the stored calories in the body, but protein usually is not used
Acetyl CoA CO

WHAT MAKES THIS TEXT A MARKET LEADER?

Before you begin this chapter, you may
(Krebswant
extensively Mechanism
as an energyof source
Enzymebecause
Action 87
that would involve
concepts from previous chapters:
the loss of muscle
Naming mass.
of Enzymes 89
• Proteins 41
Clinical Applications—No
Fatty acids Other Human Physiology 4.2 TextControl
Has More!
of Enzyme Activity 90
• Lysosomes 58 White AdiposeEffects of Temperature and pH 90
Tissue
• Cell Nucleus
The framework of this
andtextbook is based on integrating clinically germane
Gene Expression
Triacylglycerol White adipose
Phospholipids
physiological processes. Examples of this abound throughout the book.
(triglyceride)
information
Cofactors
62 with 91
and Coenzymes knowledge of the body’s
tissue, or white fat, is where most of the tri-
Enzyme Activation 91
glycerides in the body are stored. When fat stored in adipose
Substrate Concentration and Reversible Reactions 92
tissue is going to be used as an energy source, lipase enzymes
hydrolyzeCLINICAL triglycerides intoINVESTIGATIONS glycerol and free fatty acids in IN
FIGURE 5.13 Divergent metabolic pathways for acetyl Metabolic Pathways 92
CLINICAL INVESTIGATION a
coenzyme A. Acetyl CoA is a common substrate that can be 4.3 Bioenergetics 95 molecules (primarily the free
process called lipolysis. These
used to produce a number of chemically related products.
Sheryl, an active 78-year-old, suddenly became greatly fatty acids) ALL serve CHAPTERS!
Endergonic as blood-borneandCell Exergonic energy
Respiration Reactions
carriers
and Metabolism 96that can be 109
fatigued and disoriented 114 Chapter 5
while skiing. When she was used by the liver, ⊳
Coupled skeletal
Chapter-Opening muscles,
Reactions:
respiration. Coupled Reactions: Oxidation-Reduction 98
ATP and 96 other organs
Clinical for aerobic Clues, and
Investigations,
brought to the hospital, blood tests revealed elevated levels however, the
In the formation INVESTIGATION
of fatty
TABLE acids,
5.1 | aATP number
Yield of acetic
per Glucose acid When adipocytes
in Aerobic Respiration
Summaries
energy(adipose
presence of100isanarereleased
enzyme
diagnostic
cells) in in
hydrolyze
case enzymatically
asmall,
studies found in each chapter.
sampletriglycerides,
can thus be detected by TABLE 4.1 | Example
CLINICAL CLUES Summary
of LDH, AST,
(two-carbon) ALT, and
subunits arethe MB together
joined isoform to of form
CK. the fatty acid controlled
the glycerol the Clues
oxidation
leaves are
through given
reactions, throughout
certain and a portion
protein and the
(38% caseto is
thefinallyby
40%) resolved at
job it does, and its concentration ATP channels
Made can by be in measured how Value of Some Enzym
chain.
Andrea Sixexperienced
Some acetyl
of theCoA
new molecules,
muscle
terms andfor
pain and example,
fatigueyou
concepts willwill
during produce a of the membrane
plasma energy thereleased
Review
rapidly endand of is
thecaptured
itActivities
enters
performs chapter.
theits 102
blood. in the
job. Thehigh-energy
Oxidative glycerol released
Phosphorylation
bonds
Phases of ATP Made Reduced
fatty acid
exercise, that
and is 12
later carbons
experienced long. When
chest pain three
that of these
required fatty of
into ATP.
the blood is mostly taken up by the liver, which Actual Yield**it
converts
encounter include: Respiration Directly Coenzymes Theoretical Yield*
acids condense with 1 glycerol (derived from phosphoglyceral- The aerobic respiration of glucose begins withglycerol glycoly-
medical attention. 2 ATP (net gain) into 2glucose through gluconeogenesis.
If from FADH2: By this means,
•••
Glucose to pyruvate NADH, but usually goes If from FADH2: Enzyme
dehyde),Enzymes,
a isoenzymes,
triglyceride (also (in coenzymes,
called triacylglycerol)
cytoplasm)
What caused muscle pain during exercise, and why did
and cofactorsmolecule is sis.
releasedGlycolysis
into from
mitochondria in both
adipocytes
as anaerobic
CLINICAL during
2 ATP (× metabolism
APPLICATION
exercise
2) = 4 ATP or and
fasting aerobic
1.5can
ATP be res-
(× 2)an = 3 ATP
produced.
LDH,
the TheAST,
trainer formation wasofCK
ALT,it and
say fat, or lipogenesis, occurs primar-
normal? piration
important
2 FADH results
source
2 in
of theliverproduction
or if stays NADH:
glucose. of 2 molecules orofif pyruvate, stays NADH: Alkaline phosphatase
When diseases 3 ATP (× 2) = 6 tissues,
damage ATP some 2.5 cells
ATP (×die 2) = and
5 ATPrelease
2 ATP, and 2the NADH + H+ per glucose molecule. In aerobic
ily in adipose
blood been •
glucose
tissue and in
What might her chestPyruvate the
is elevated following
liver
pain indicate, when
to acetyl CoA
a meal.
the
how concentration
(× might
2) it have
None
of However,
1 NADH (× 2) =
respiration,
lipolysis are the
their
however,
most
2 NADH enzymes
free fatty
significant
3 ATP
the acids. into
electrons
(× 2)the
energy
Most
= 6blood.
ATP
in fatty
NADH
carriers
Theare
acids 2.5
provided
2.5 ATP (×
activity
by
= 5 ATP
of2)these
not donated
consist of=a15 ATP
enzymes,
⊲ Fat
Clinical stored
produced, and why
attention??
in adiposeBoxes
Application
did cycle
Citric acid Andrea
cells (adipocytes)
(× 2) need medical
are in-depth of boxed
1 ATP (× 2) = 2 ATP
essays thatlong
white adipose to pyruvate
1 hydrocarbon
FADH2 (× 2) and
= 6 NADH their
3 NADH (× 2) reflecting
= 2lactate
chain
FADH
measured
is 2not
2 with ATP
in a test
(× 6) = 18 ATP
3 ATPconcentrations
a formed,
carboxyl
(× 2) = 4 ATP
tube
asgroup
happens
by adding
in the
(COOH) 1.5
blood
ATP (× 6)plasma,
inATPthe(×atlactate
their specific
2)one
= 3 ATP
can be
substrates.
Acid phosphatase

tissue
explore (or relevant serves as
white fat) topics of the
Total majorinterest
clinical
ATP form of energy and4are storage
ATP pathway.
placed atend. In a process Instead, the pyruvate
known as32β-oxidation will
(or 34) ATP move to a
(β is the 26different
Greek cellular
(or 28)letter
ATP
location and Becausea an
undergo increase
different in certain
reaction; the NADH enzymes in the blood can
produced Amylase
in the body. One gram of *fat Theprovides 9 iskilocalories of ATPof energy, oxidativebeta), enzymes remove two-carbon acetic
Detailed acid molecules from
key points in the chapter to support
theoretical yield the
the surrounding
number produced material.
by phosphorylation
by glycolysis
inside the
will
mitochondria
indicate damage
eventually
(explained in the
be to specific
oxidized,
Accounting
organs,
but that
section).
such
occurs testslater may aid the Aldolase
compared to 4 kilocalories for ayieldgram
**The actual ofaccount
takes into carbohydrates
the energy cost of or the
transporting ATPacid end
out of the of a fatty
mitochondria and intoacid chain. This results in the forma-
the cytoplasm.
Subjects
protein. Incovered
a nonobese include pathologies,
70-kilogram currentman,
(155-pound) research,
80% in the
tion of story.
diagnosis of diseases. For example, an increase in a man’s
acetyl CoA, as the third carbon from the end becomes Creatine kinase (or
pharmacology, and a variety of clinical diseases. In aerobic blood levels of acid
respiration, pyruvate phosphatase may result from disease of
oxidized to produce a new carboxyl group.leaves The fatty theacid cellchain cyto- creatine phosphokinas
the 36 to 38 ATP produced per glucose in the mitochondrion, plasm and the
enters from prostate
FADH
the 2 (table
interior result (the 4.1).
in thematrix)
pumping of of only 6 protons (4 by the
mitochondria. (CK or CPK)
C H E C K P O I NT S only 30 to 32 ATP actually enter the cytoplasm of the cell.
is thus decreased in length by 2 carbons. The
second pump and 2 by the third pump). Because 1 ATP is pro-
process of oxida-
Roughly 3 protons must pass through the respiratory
Once
tion pyruvateduced
continues
assem- until
is theinside
for entire
a protons
every 4 fatty
mitochondrion,
acid
pumped,molecule carbon
electrons is derived dioxide
converted from FADH2
Lactate dehydrogenase
1a. Define the termAPPLICATION
glycolysis in terms of its initial
LIFESTYLE
substrates and products. blies andExplain
activate whyATP synthase
there is to a produce
net
is
to enzymatically
acetyl CoA
1 ATP. However, the Lifestyle ⊳ (fig. removed
5.14).
result in the Application from
formation of 6 ÷Boxes each three-carbon-long
4 = 1.5 are ATP.readings
Each citric acid thatcycle
explore
(LDH)

abnormal matrixpyruvate andAmust to form

16-carbon-long a fatty
two-carbon-long
acid, get organic
forweexample, yields acid—acetic
8 acetyl
newly formed ATP is in the Transaminases (AST
Metabolic
gain ofsyndrome
2 moleculesisof aATP combination
in this process. of mitochondrial be produces
physiological
1 FADH
CLINICAL 2 and
principles INVESTIGATIONas
two citric
applied
acid
to
cycles from 1 glu-
well-being,
CLUES sports and ALT)
measurements—including moved into the cytoplasm; this transport also uses
central obesity (excess abdominal acid.
the The
proton enzyme cose,
CoA molecules. Each of these can enter that
so there catalyzes
are 2 FADH this reaction
that give 2 × combines
1.5
2 a citric acid cycle and ATP = 3 the
ATP.
1b. What are the initialgradient substrates and final
and costs 1 more products
proton. The of ATP and acetic H+ are trans-
produce acid
10 medicine,
with
ATP a
per The
coenzyme 23exercise
turn ATP
of subtotal
(derived
the physiology,
cycle, from
from oxidative
producing the and aging.
phosphorylation
vitamin
8 × 10 pan-
= They we are also
fat), hypertension (high ported
blood into pressure),
the cytoplasm insulin resistance
in exchange for ADP and Pi, which Sheryl’s
have atatblood
this point tests revealonly
includes elevated
the NADH levels ofFADHCK, LDH, AST,
anaerobic metabolism?
(prediabetes), type 2 diabetes mellitus, high plasma triglyc- tothenic
80 ATP. Inacid) placed called
addition,
and ALT. relevant
each coenzymetime points anA.acetyl Thein theCoA text
combination toand
molecule highlight
thus 2 pro-
is which concepts just
are transported
1c. Describe the physiological functions of lactate into the mitochondrion. Thus, it effectively takes duced in the mitochondrion. Remember that glycolysis,
produced
formed and isthe called
covered
end inthe
acetyl
carbon thecytoplasm,
chapter.
ofcoenzyme
the fattyalso A, abbreviated
acid chain isNADH. oxidized,acetyl TABLE 4.2 | Selecte
erides, and high LDL cholesterol—that
fermentation. In whichTo
4 protons to produce
tissue(s)
summarize:
greatly
1 ATPincrease
is anaerobic
that entersthe
The mellitus,
theoretical
the cytoplasm.
andATP yield 1CoA
NADHis (fig.
36 andto5.5). •
occurs
1 plasmic
FADH What in enzymes
are
2NADH produced. do theseproduces
cannot directly Oxidativeletters 2indicate,
enter phosphoryla-
Thesewhat
and
the mitochondrion, but
cyto-
the Reactions They C
⊳
risk of coronary heart disease, stroke, diabetes diseases do elevated blood levels of these enzymes
metabolism normal? 38 InATP which
per tissue isThe
glucose. it abnormal?
actual ATP Glycolysis
tion produces
yield, allowing for the 2.5 converts
there Systems
ATP is aper 1 NADH
processglucose
Interactions
by which molecule
and 1.5 pages
their ATP into atper
electrons 2themolecules
end
FADH
can be of 2. chapters have
“shuttled”
other conditions. The incidence of metabolic syndrome has Enzyme
costs of transport into the cytoplasm, is about For of to
30 pyruvate.
a32 ATP Because in. suggest?
The each pyruvate molecule is converted into
sixteen-carbon-long
been a net effect
traditionfatty of
ofthecytoplasm
acid, thismost
these common
textbook4 ATP shuttle
molecules
since is that
the a mol-
earliest editions. Catalase
increased alarmingly in recent
in obesity. Eating excessive
years because of the increase
per glucose. The details of how these numberswould
calories,next.
are described particularly in the form
are obtained
1 moleculebe formed •
ecule
of acetyl
Now, seven
of FADH
of
HowCoA
withtimes
symptoms?
NADH
mightand
2 this
in the
in
thesethe
1 CO
(producing
sixteenth
test2,results
mitochondrion. edition,
is translated
7 =relate
42×molecules 28 ATP).
Thea2new NADH
to into
ofSheryl’s
Not
Systems
a
acetyl molecule
produced Interactions
in
CoA andthe
counting 2 molecules
single ATP of CO
used toare derived
start from
β-oxidation each
(fig. glucose.
5.14),
5.2 AEROBIC RESPIRATION
of sugars (including high fructose corn syrup), stimulates
insulin secretion. Insulin then promotes the uptake of blood These
this fatty acetyl
acid icon
CoA
glycolysis,
could has been
molecules
yield
therefore,
a grand
2
addedserve
usually
for
total as the become
first
substrates
of 28
1.5 ATP = 3 ATP by oxidative phosphorylation. (An alterna- + 80,
2 FADH
timefor
or to and yield major
2relevant
mitochon-
108 ATP
2× sections
Carbonic anhydrase
Amylase
glucose into adipose cells, Detailed
whereAccounting
(through lipogenesis) it is molecules!
drial enzymes of inthethe
tive in-chapter
aerobic
pathway, where text.
pathway, theThese while
cytoplasmicalert thereaders
carbonisto
NADH diox-those sections that
transformed
In the aerobic respiration Eachof glucose,
NADH pyruvic
formed
converted into stored triglycerides (see figs. 5.12 and 5.13). in acid
the is formed donates
mitochondrion ide2 is carried
electrons Naming
specifically
by
into the blood
mitochondrial of
discuss toEnzymes
the
NADH howlungs the
and forchapter’s
produceselimination.
2 × body
2.5 ATP Itsystem
=is5 ATP,interacts with Lactate dehydrogenase
86by glycolysis and then to the electron
converted intotransport
acetyl system
coenzyme at the first
A. proton pump (see
important toother is less
notesystems common;
that theinoxygen however, this
in CO is the dominant
is derived pathway
from in the Ribonuclease
Conversely, the lowering of insulin secretion, by diets that the are service of2 total bodyactive.)function. The new
This begins
prevent the aplasma
fig. 5.9).pathway
cyclic metabolic
glucose from
The electrons
rising called arethe
sharply,
thencitric
passed
promotes acidto the secondBrown
pyruvic acid,In
and Adipose
third thefrom
liver
not
Systems
past,
Tissue
and heart, enzymes
oxygen
Interactions
which gas.were icon
given
metabolically
also
names
signals
highlythat
essay
were somewhat
questions in the
proton pumps, activating each of them in turn until the 2 elec- arbitrary. We The now have modern a totalsystem
of 26 ATP for(or,naming enzymes,
less commonly, established
28 ATP)
lipolysis (the breakdown
(Krebs) cycle. ofoffat)
As a resulttrons theseand weight loss.
pathways, a large amount Brown adipose tissue, or brown fat, has a different primary func-
are ultimately passed to oxygen. The first and second by
tion transports
than white adiposeend-chapter
produced
an internationalbyReview
tissue:
oxidative
it add Activities
phosphorylation
committee,
is the major siteis that
more ask
from
forbythermogenesis students
one
orderly molecule
and about
of specific
informative.
pumps transport
and 4FADH protons each, and the third pump glucose. We can thesome2 ATPolder made direct (substrate-level)
of reduced NAD and FAD (NADH 2) is generated. Citric Acid
2 protons, for a total of 10. Dividing 10 protons by the 4 it takes interactions
With Cycle
the exception
phosphorylation of the of
discussed
in glycolysis andbody
enzyme
the 2 ATP systems. names (such as pep-
made directly by
affect the active sites (ot
These reduced coenzymes
▲ provide electrons for a process sin, trypsin, and renin), all enzyme names end with the suffix the same reaction), but t
Learning Outcomes are numbered for easy referencing in Once acetyl-ase
to produce an ATP (in the cytoplasm) gives 2.5 ATP that are ▲ the
CoA(table two citric
has 4.2), acid
been and cycles
formed, to give a
theofacetic grand total
acidare of 30
subunit ATP (or,
that drives the formationproduced of ATP. for every pair of electrons donated by an NADH. Learning less commonly, Outcome 32 ATP) numbers
classes produced areby tied
enzymes directly
the aerobic named
respirationCheckpointat other locations so that
toaccording
digital material! (There is no such thing as half an ATP; the decimal fraction to
(2 carbons long) their combines
of glucose activity,
(see table
with
or “job oxaloacetic acid (4 carbons
5.1). category.” Hydrolases, for example, separated by standard bi
long) to formpromote numbers!
a molecule of citric acid (6 carbons long). Coen-
simply indicates an average.) hydrolysis reactions. Other enzyme categories include are useful in the diagnos
zymeeach A acts onlyC H Eas C KaP O transporter of aceticthe acidremovalfrom one
LE A R N I N G O UTCO M EThree S molecules of NADH are formed with citric phosphatases, I which
NT S catalyze of phosphate
acid cycle, and 1 NADH is also produced whenenzyme pyruvatetois another (similar to the transport of hydrogen by
After studying this section, you should
converted into acetylbe ableCoA to: (see fig. 5.5). StartingNAD). from 1 The
groups; 2a.synthases
Compare the and fate
glu- formation of citric acid begins a cyclic metabolic
of pyruvate inwhich
synthetases, aerobiccatalyze
and dehydration
CLINICAL AP
cose, two citric acid cycles (producing 6 NADH) and 2 pyru- synthesis reactions; dehydrogenases, which remove hydrogen
anaerobic cell respiration.
2. Describe the aerobic cell respiration of glucose through atoms 2b.from Draw their
a substrates;
simplified citric and
acid cycle and
kinases, which
indicate the add a phos-
vates converted to acetyl CoA (producing 2 NADH) yield Different diseased or
the citric acid cycle. 8 NADH. Multiplying by 2.5 ATP per NADH gives 20 ATP. phate group high-energy products.
to (phosphorylate) particular molecules. Enzymes
Electrons systemfrom FADH are donated later in the electron- called3a. Explain how NADH and FADH2 contribute to oxidative matic forms of an enz
3. Describe the electron transport and2 oxidative isomerases rearrange atoms within their substrate mol-
transport system than those donated by NADH; H
consequently, phosphorylation. H nosis of disease. Crea
phosphorylation, explaining the role of oxygen in this ecules to formNAD structural isomers, + H+ such as glucose and fructose
these electrons activate only the second and third proton pumps. 3b. Explain how NADH ATP is produced in oxidative example, is normally lo
process. H C H (chapter H 2; see fig. 2.13). H C H
Because the first proton pump is bypassed, the electrons passed phosphorylation. production of ATP (se
The aerobic respiration of glucose (C6H12O6) is given in the fol- C O + S
The names CoA
of many enzymes Cspecify O + CO
both the substrate isoenzymatic forms ar
vi of the enzyme and the job category of the enzyme. Lactic acid 2
lowing overall equation: ent chromosomes, and
C dehydrogenase, for example, removesS hydrogens CoA from lactic acid. bodies to distinguish b
ISTUDY
C H O + 6 O → 6 CO + 6 H O
HO O Enzymes that do exactly the same job (that catalyze the same form (or CK-1) is relea
6 12 6 2 2 2
 New to This Edition
CHAPTER CHANGES IN THE SIXTEENTH EDITION OF HUMAN PHYSIOLOGY
The following list includes relatively major changes in each
chapter and does not include changes involving a sentence or
two, word or phrase changes, or label changes in figures.
All Chapters
⦁. Boxes on Exercise Applications changed to Lifestyle
Applications throughout.
⦁. Chapters 8 and 18 modified extensively by Krista Rompolski,
Ph.D.
⦁. Section subheadings that specifically deal with interactions
between different body systems are called out and identified
with a distinctive Systems Interactions icon.
⦁. Review Activities questions that relate to the Systems
Interactions are identified with the Systems Interactions icons.
Chapter 1
⦁. Modifications in section on Scientific Method.
⦁. New citation in Table 1.1 for the 2019 Nobel Prize in
Physiology or Medicine.
⦁. Modifications in section on The Primary Tissues.
⦁. Modifications in the section on the skin.
Chapter 2
⦁. Additional information added to the section on buffers.
Chapter 3
⦁. New description of glycocalyx added.
⦁. Explanation of lysosomes and autophagy rewritten and
updated.
⦁. Information on mitochondria updated and expanded.
⦁. Expanded and updated information in the section describing the
genome, genes, and genetic expression.
⦁. Description of CRISPR-Cas9 updated.
⦁. Section on epigenetic inheritance expanded and updated.
Chapter 4
⦁. Updated and expanded information on gene therapy and genome
editing.
⦁. New information added to on specific autosomal recessive
diseases added to the Clinical Applications box on
phenylketonuria.
⦁. Chemical structure of NADH in figure 4.17 modified.
Chapter 5
⦁. Information about the uses of the lactic acid pathway expanded
and updated.
⦁. Information about the respiratory complexes expanded.
⦁. Respiratory complexes identified in figure 5.9.
⦁. Added information on essential amino acids.
ISTUDY
Chapter 6
⦁. Expanded explanation of carrier proteins and channel proteins,
with different listing order.
⦁. Addition of a description of the importance of Na+ in body
fluid osmolality.
⦁. Legend for figure 6.16 rewritten.
⦁. New discussion of sodium-coupled glucose transporters
(SGLT 1 and 2).
⦁. Added description of autocrine signaling.
Chapter 7
⦁. Updated and expanded description of microglia.
⦁. Updated discussion of oligodendrocytes.
⦁. Updated discussion of central nervous system (CNS) axon
regeneration.
⦁. Expanded and updated discussion of astrocytes.
⦁. Expanded and updated discussion of CNS capillaries with
addition of pericytes.
⦁. Updated and expanded discussion of the endogenous opioids.
Chapter 8
⦁. Clinical Investigation updated throughout the chapter.
⦁. Description of cerebrospinal fluid formation updated.
⦁. Figure 8.5 updated for most up-to-date terminology.
⦁. Updated description of autism spectrum disorder.
⦁. New research on the clinical applications of PET scans and fMRI.
⦁. Addition of glymphatic system and its association with sleep
and neurodegenerative disease.
⦁. Updated Clinical Application box on Huntington’s disease and
Parkinson’s disease.
⦁. Added information on the subthalamic nucleus.
⦁. Updated and expanded description on Wernicke’s aphasia and
conduction aphasia.
⦁. Arcuate fasciculus added as a label on figure 8.14.
⦁. New discussion on the amygdaloid body and the famous patient
“S.M.”
⦁. New evidence on metabolic activity of neurons as well as
activity in the hippocampus during memory consolidation.
⦁. Updated and additional research on the genetics, pathogenesis,
and prevention of Alzheimer’s disease.
⦁. Expanded discussion on the role of limbic system structures in
learning and memory formation.
⦁. Addition of the 2017 Nobel Prize in Physiology or Medicine,
which led to the discovery of circadian clock genes in humans.
⦁. Updated discussion in a Lifestyle Application box on
benzodiazepine prescription, use, and abuse.
⦁. Table 8.6 updated for accuracy on cranial nerve composition
and function.
vii
 New to This Edition

⦁. New figure of the cranial nerves added to Section 8.6. ⦁. Single-Unit and Multi-Unit Smooth Muscle heading given a
⦁. Additional information provided on the structure and function of Systems Interactions icon.
the anterior corticospinal tracts. ⦁. Autonomic Innervation of Smooth Muscles heading given a
Systems Interactions icon.
Chapter 9
⦁. New Systems Interactions question added to the Review
⦁. Figure 9.11 updated and modified for increased accuracy.
Activities.
⦁. Legend for figure 9.11 updated and expanded.
Chapter 13
Chapter 10
⦁. Updated information on aplastic anemia in the Clinical
⦁. Updated and expanded description of neural pathways for
Applications box on anemias.
somesthetic sensation.
⦁. Updated information on the origin of platelets.
⦁. New Clinical Application box on the gate control theory of pain.
⦁. Updated information on thrombopoietin.
⦁. Descriptions of the “labeled line” concept of taste transmission
⦁. Updated information on hemophilia A and B.
updated and expanded.
⦁. Figure 13.23 labels and legend modified to emphasize
⦁. Updated and expanded discussion of the physiology of sour
mechanical correlates of electrical activity.
taste.
⦁. Cardioprotective effects of exercise added to a Lifestyle
⦁. Descriptions of the structure and functions of the cupula in the
Application box.
semicircular canals updated and expanded.
⦁. Updated information on potential repairs of myocardial infarction.
⦁. Addition of motion sickness explanation.
⦁. Updated information on ECG changes in myocardial infarction.
⦁. Description of sensorineural deafness updated and expanded.
⦁. New information added on paroxysmal supraventricular
⦁. Figure 10.37 modified to show the direction of light.
tachycardia.
⦁. Description of intrinsically photosensitive retinal ganglion cells
⦁. New information on pericytes added.
added.
Chapter 14
Chapter 11
⦁. Expanded and updated discussion of the control of cardiac rate.
⦁. Updated discussion of the role of heat shock proteins in steroid
⦁. Heading Regulation of Blood Volume by the Kidneys given a
hormone action.
Systems Interactions icon.
⦁. Updated discussion in a Clinical Applications box of the Her2
⦁. Heading Extrinsic Regulation of Blood Flow given a Systems
receptor and the action of Herceptin in the treatment of breast
Interactions icon.
cancer.
⦁. Updated discussion of changes in coronary blood flow with
⦁. Figure 11.11 labels modified, and legend expanded and updated.
exercise.
⦁. Description of the osmoreceptor neurons and control of ADH
⦁. Updated and expanded explanation of how aerobic exercise
secretion updated and expanded.
improves cardiovascular health.
⦁. Heading Hypothalamic Control of the Anterior Pituitary given a
⦁. Heading Baroreceptor Reflex given a Systems Interactions icon.
Systems Interactions icon.
⦁. Updated discussion of hypertension.
⦁. Heading Functions of the Adrenal Cortex given a Systems
⦁. Table 14.8 updated to display latest classification system for
Interactions icon.
hypertension.
⦁. Stages of the General Adaptation Syndrome description
⦁. Updated discussion of septic shock.
expanded and updated.
⦁. Three new Systems Interactions questions added to the Review
⦁. Heading Pancreatic Islets given a Systems Interactions icon.
Activities.
⦁. Three new questions regarding Systems Interactions added to
the Test Your Understanding section of the Review Activities. Chapter 15
⦁. Updated information on danger-associated molecular patterns
Chapter 12
(DAMPs).
⦁. Clinical Application box on muscular dystrophy updated.
⦁. Updated information on macrophages.
⦁. Updated and expanded discussion of titin in muscle contraction.
⦁. Expanded and updated explanation of the causes and functions
⦁. Updated and expanded discussion of muscle fatigue.
of a fever.
⦁. New discussion of the myokine irisin added.
⦁. Expanded and updated description of the sources and functions
⦁. New Lifestyle Application box added regarding the healthful
of gamma interferon.
consequences of a change from a sedentary to a more active
⦁. Updated and expanded description of neutrophil actions in an
lifestyle.
infection.
⦁. Updated discussion of muscle satellite cells.
⦁. Expanded and updated information on regulatory
⦁. Updated information on muscle spindle.
T lymphocytes.
⦁. Heading Skeletal Muscle Reflexes given a Systems Interactions
⦁. Updated information on sepsis in a Clinical Applications box.
icon.

viii

ISTUDY
 ⦁. Updated and expanded explanation of the nature of B cell clones
in the development of secondary immune responses.
⦁. Updated information on immunological competence and
immunological tolerance.
⦁. Updated and expanded information on chimeric antigen
receptors and immune checkpoint blockade in the treatment of
cancer.
⦁. Updated information on the viral causes of cancer.
⦁. New information on anaphylaxis.
Chapter 16
⦁. Updated and expanded description of control of the vocal cords.
⦁. New information about lower respiratory tract infections added.
⦁. Updated and expanded information about how allergens
stimulate asthma.
⦁. Description of the role of the pons in the control of breathing
updated.
⦁. Updated description of the role of the central chemoreceptors in
the control of breathing.
⦁. New description of the hypoxic ventilatory response.
⦁. Updated and expanded description of obstructive sleep apnea.
⦁. New discussion of cardiovascular changes during breath-holding
and hyperventilation.
⦁. New information added about treatments for sickle cell anemia
and thalassemia.
⦁. Heading Principles of Acid–Base Balance gets Systems
Interactions icon.
⦁. Expanded and updated description of the role of the kidneys in
assisting the lungs in the control of acid–base balance.
⦁. Updated and expanded discussion of the mechanisms of
acclimatization to high altitude.
⦁. New Systems Interactions question added to the Review
Activities.
Chapter 17
⦁. Updated and expanded explanation of autoregulation of renal
blood flow and tubuloglomerular feedback.
⦁. Expanded and updated description of the secretion of ADH.
⦁. Figure 17.21 modified to show reabsorption from and secretion
into blood vessels.
⦁. Heading Role of Aldosterone in Na+/K+ Balance given a
Systems Interactions icon.
⦁. Figure 17.25 modified to show reabsorption from and secretion
into blood vessels.
⦁. Heading Renal Acid–Base Regulation given a Systems
Interaction icon.
⦁. Figure 17.29 modified to show Na+ being cotransported with
bicarbonate out of the proximal tubule.
⦁. Added explanation of sodium-bicarbonate cotransport from
proximal tubule.
⦁. Updated and expanded explanation of the renal generation of
bicarbonate and ammonia.
⦁. Updated description of acute mountain sickness.
ISTUDY
⦁. Two new Systems Interactions questions added to the Review
Activities.
Chapter 18
⦁. Clinical Investigation updated throughout the chapter.
⦁. Updated descriptions of the mucosal and serosal layers of the
alimentary tract.
⦁. New detail included on the nerve composition of the vagus nerve.
⦁. Clinical Application box on Barrett’s esophagus updated to
include the role of esophageal stem cells in the development of
esophageal cancer.
⦁. Clinical Application box on gastroesophageal reflux disease
(GERD) updated to include additional risk factors.
⦁. Discussion on celiac disease added to the updated Clinical
Application box on lactose intolerance.
⦁. New and expanded description of the function of the
interconnected cells of Cajal in the stomach and intestines.
⦁. Additional information on the structure of the large intestine.
⦁. Label for taenia coli added to figure 18.16.
⦁. Section on Intestinal Microbiota revitalized with new headings
and up-to-date research on its development, role in immune
function, metabolism and disease development.
⦁. Clinical Application box on inflammatory bowel disease
(IBD) and irritable bowel syndrome (IBS) updated to include
distinguishing features of ulcerative colitis and Chron’s disease.
⦁. Updated description of salt and water transport in the large intestine.
⦁. Description on the steps of defecation reflex updated.
⦁. Clinical Application box on cirrhosis, nonalcoholic fatty liver
disease, and liver diseases caused by chronic alcohol use updated.
⦁. Table 18.3 updated to include storage functions of the liver.
⦁. Legend for figure 18.22 on metabolism of heme and bilirubin
updated for more detail and accuracy.
⦁. Urobilinogen changed to sterocobilin in text and figure 18.23 for
accuracy in terminology.
⦁. Updated and expanded description of the metabolism and
circulation of bile acids and the role of bile acids in the
secretion of hormones by the small intestine.
⦁. Lifestyle Application on exercise and the timing of meals
updated to include the autonomic nervous system’s influence on
digestion during exercise.
⦁. New information added to the structure and function of the
enteric nervous system.
⦁. New information on the role of secretin in metabolism of brown
adipose tissue.
⦁. Updated explanation of the transport of chylomicrons by the
lacteals of the intestinal villi.
Chapter 19
⦁. Updated and expanded discussion of the development of adipose
tissue.
⦁. New discussion of subcutaneous adipose tissue, visceral fat, and
ectopic fat.
⦁. New heading, Hormonal Signals from the GI Tract.
ix
 New to This Edition

⦁. Updated and expanded discussion of enteroendocrine regulation Chapter 20

of hunger. ⦁. Updated information on anti-Müllerian hormone.
⦁. Information on leptin and obesity updated. ⦁. Updated information added on the mechanisms of the pubertal
⦁. New information on meal-induced thermogenesis added. growth spurt.
⦁. Description of thermoregulation updated and expanded. ⦁. Updated information added regarding the 5 α-reduced
⦁. Updated and expanded description regarding insulin and androgens.
glucagon during the postabsorptive state. ⦁. New and updated information regarding the role of estradiol in
⦁. Updated and expanded section on the role of autonomic nerves male physiology.
in pancreatic islet regulation. ⦁. New and expanded description of primary follicles and anti-
⦁. Updated and expanded description of insulin action during the Müllerian hormone in adult women.
absorptive state. ⦁. New and updated information regarding polycystic ovarian
⦁. Updated explanation of increased metabolism stimulated by syndrome.
thyroxine. ⦁. Updated information added regarding the sperm’s contribution
⦁. New information added regarding psychosocial effects on to the mitochondria of the zygote.
children’s levels of growth hormone and IGF-1. ⦁. New paragraph added distinguishing the embryonic and fetal
⦁. New and expanded description of the regulation of osteoclast stages of development.
development. ⦁. Current information added regarding the use of iPS cells in
⦁. Updated description of the effects of sex steroids on osteoblasts regenerative medicine.
and osteoclasts.
⦁. New and updated information added regarding the effects of
osteocalcin.

ISTUDY
 Acknowledgments

REVIEWERS

Gwen Bachman Erin M. Itza

University of Nebraska-Lincoln University of South Alabama
Maegen A. Borzok Kimberly Jeckel
Ohio State University, Wexner Medical Center Colorado State University
Lindsay Calderon Kimberly D. Kyker
Eastern Kentucky University Oklahoma City Community College
Patricia Clark Greg M. Landry
Indiana University—Purdue University Indianapolis Massachusetts College of Pharmacy &
Health Sciences University
Barbara E. Davis
Eastern Kentucky University John McDaniel
San Jose State University
Stephen Dodd
University of Florida Lisa Middleton
Eastern Kentucky University
Maria Elena de Bellard
California State University, Northridge Mattia M. Migliore
MCPHS University- Boston
Michael L. Garcia
University of Missouri Sandra Lehmann Vierra
Merced College
Karri Haen Whitmer
Iowa State University Kate Ireton Walker
University of Arkansas
Tray W. Hamil
University of South Alabama Kennedy S. Wekesa
Alabama State University
Lisa M. Harrison-Bernard
Louisiana State University School of Medicine Sania Zaidi-Merchant
College of the Canyons
Carolyn Huffman
University of Kansas

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 Contents

Preface v 2.3 Proteins 40

Structure of Proteins 40

1 The Study of Body

Function 1
2.4
Functions of Proteins 43
Nucleic Acids 44
Deoxyribonucleic Acid 44

1.1 Introduction to Physiology 2

Ribonucleic Acid 45

Scientific Method 2 Summary 46

1.2 Homeostasis and Feedback Control 4
Review Activities 47

3 Cell
History of Physiology 4
Negative Feedback Loops 5 Structure and
Positive Feedback 8
Genetic Control 49
Neural and Endocrine Regulation 8
Feedback Control of Hormone Secretion 8 3.1 Plasma Membrane and Associated Structures 50

1.3 The Primary Tissues 10 Structure of the Plasma Membrane 51

Muscle Tissue 10 Phagocytosis 53
Nerve Tissue 11 Endocytosis 53
Epithelial Tissue 12 Exocytosis 54
Connective Tissue 15 Cilia and Flagella 54
1.4 Organs and Systems 17 Microvilli 55
An Example of an Organ: The Skin 18 3.2 Cytoplasm and its Organelles 55

Systems 19 Cytoplasm and Cytoskeleton 56

Body-Fluid Compartments 20 Lysosomes 57
Summary 21 Peroxisomes 57
Review Activities 22 Mitochondria 58
Ribosomes 59

2
Endoplasmic Reticulum 59
Chemical Composition Golgi Complex 60
of the Body 24 3.3 Cell Nucleus and Gene Expression 61
Genome and Proteome 62
2.1 Atoms, Ions, and Chemical Bonds 25
Chromatin 63
Atoms 25
RNA Synthesis 63
Chemical Bonds, Molecules, and Ionic
RNA Interference 66
Compounds 26
3.4 Protein Synthesis and Secretion 67
Acids, Bases, and the pH Scale 28
Transfer RNA 67
Organic Molecules 30
Formation of a Polypeptide 68
2.2 Carbohydrates and Lipids 32
Functions of the Endoplasmic Reticulum and Golgi
Carbohydrates 33
Complex 69
Lipids 36
Protein Degradation 69
xv

ISTUDY
 xvi Contents

3.5 Dna Synthesis and Cell Division 71 5.4 Metabolism of Lipids and Proteins 117
DNA Replication 71 Lipid Metabolism 117
The Cell Cycle 72 Amino Acid Metabolism 120
Mitosis 75 Uses of Different Energy Sources 121
Meiosis 77 Interactions 124
Epigenetic Inheritance 78 Summary 125
Interactions 82 Review Activities 126
Summary 83

6 Interactions
Review Activities 84
Between
Cells and the Extracellular
4 Enzymes and Energy
4.1 Enzymes as Catalysts 87
86

6.1
Environment
Extracellular Environment
128

129
Body Fluids 129
Mechanism of Enzyme Action 87
Extracellular Matrix 129
Naming of Enzymes 89
Categories of Transport Across the Plasma
4.2 Control of Enzyme Activity 90
Membrane 130
Effects of Temperature and pH 90
6.2 Diffusion and Osmosis 131
Cofactors and Coenzymes 91
Diffusion Through the Plasma Membrane 133
Enzyme Activation 91
Rate of Diffusion 134
Substrate Concentration and Reversible Reactions 92
Osmosis 134
Metabolic Pathways 92
Regulation of Blood Osmolality 139
4.3 Bioenergetics 95
6.3 Carrier-Mediated Transport 140
Endergonic and Exergonic Reactions 96
Facilitated Diffusion 141
Coupled Reactions: ATP 96
Active Transport 142
Coupled Reactions: Oxidation-Reduction 98
Bulk Transport 146
Summary 100
6.4 The Membrane Potential 147
Review Activities 102
Equilibrium Potentials 148

5 Metabolism
Resting Membrane Potential 150
Cell Respiration and 6.5 Cell Signaling 151
Second Messengers 152
104
G-Proteins 153
5.1 Glycolysis and the Lactic Acid Pathway 105 Interactions 155
Glycolysis 105 Summary 156
Lactic Acid Pathway 107 Review Activities 158

5.2 Aerobic Respiration

7
109
Citric Acid Cycle 109
Electron Transport and Oxidative Phosphorylation 110
The Nervous System 160

Coupling of Electron Transport to ATP Production 110 7.1 Neurons and Supporting Cells 161
ATP Balance Sheet 113 Neurons 161
5.3 Interconversion of Glucose, Lactic Acid, Classification of Neurons and Nerves 162
and Glycogen 115 Neuroglia 164
Glycogenesis and Glycogenolysis 115
Neurilemma and Myelin Sheath 165
Cori Cycle 115
Functions of Astrocytes 168

ISTUDY
 Contents xvii

7.2 Electrical Activity in Axons 170 Hindbrain 228

Ion Gating in Axons 171 Reticular Activating System in Sleep and Arousal 230
Action Potentials 172 8.5 Spinal Cord Tracts 231
Conduction of Nerve Impulses 175 Ascending Tracts 231
7.3 The Synapse 178 Descending Tracts 232
Electrical Synapses: Gap Junctions 179 8.6 Cranial and Spinal Nerves 234
Chemical Synapses 179 Cranial Nerves 234
7.4 Acetylcholine as a Neurotransmitter 182 Spinal Nerves 235
Chemically Regulated Channels 183 Summary 238
Acetylcholinesterase (AChE) 186 Review Activities 239

9 The
Acetylcholine in the PNS 187
Acetylcholine in the CNS 188 Autonomic Nervous
7.5 Monoamines as Neurotransmitters
System
188
242
Serotonin as a Neurotransmitter 190
Dopamine as a Neurotransmitter 191 9.1 Neural Control of Involuntary Effectors 243
Norepinephrine as a Neurotransmitter 191 Autonomic Neurons 243
7.6 Other Neurotransmitters 192 Visceral Effector Organs 244
Amino Acids as Neurotransmitters 192 9.2 Divisions of the Autonomic Nervous System 245
Polypeptides as Neurotransmitters 194 Sympathetic Division 245
Endocannabinoids as Neurotransmitters 195 Parasympathetic Division 246
Gases as Neurotransmitters 196 9.3 Functions of the Autonomic Nervous System 250
ATP and Adenosine as Neurotransmitters 196 Adrenergic and Cholinergic Synaptic
7.7 Synaptic Integration 197 Transmission 250
Synaptic Plasticity 197 Responses to Adrenergic Stimulation 251
Synaptic Inhibition 198 Responses to Cholinergic Stimulation 255
Summary 199 Other Autonomic Neurotransmitters 256
Review Activities 201 Organs With Dual Innervation 257
Organs Without Dual Innervation 258

8
Control of the Autonomic Nervous System by Higher
The Central Nervous Brain Centers 259

System 204 Interactions 261

Summary 262
8.1 Structural Organization of the Brain 205 Review Activities 263
8.2 Cerebrum 207

10 Sensory Physiology
Cerebral Cortex 207
Basal Nuclei 213 265

Cerebral Lateralization 214

10.1 Characteristics of Sensory Receptors 266
Language 216
Categories of Sensory Receptors 266
Limbic System and Emotion 217
Law of Specific Nerve Energies 267
Memory 218
Generator (Receptor) Potential 267
Emotion and Memory 223
10.2 Cutaneous Sensations 268
8.3 Diencephalon 224
Neural Pathways for Somatesthetic Sensations 270
Thalamus and Epithalamus 224
Receptive Fields and Sensory Acuity 271
Hypothalamus and Pituitary Gland 224
Lateral Inhibition 272
8.4 Midbrain and Hindbrain 227
Midbrain 227

ISTUDY
 xviii Contents

10.3 Taste and Smell 273 Hypothalamic Control of the Posterior Pituitary 332
Taste 273 Hypothalamic Control of the Anterior Pituitary 332
Smell 275 Feedback Control of the Anterior Pituitary 334
10.4 Vestibular Apparatus and Equilibrium 277 Higher Brain Function and Pituitary Secretion 335
Sensory Hair Cells of the Vestibular Apparatus 278 11.4 Adrenal Glands 336
Utricle and Saccule 279 Functions of the Adrenal Cortex 337
Semicircular Canals 279 Functions of the Adrenal Medulla 338
10.5 The Ears and Hearing 281 Stress and the Adrenal Gland 339
Outer Ear 282 11.5 Thyroid and Parathyroid Glands 340
Middle Ear 282 Production and Action of Thyroid Hormones 341
Cochlea 283 Parathyroid Glands 343
Spiral Organ (Organ of Corti) 285 11.6 Pancreas and Other Endocrine Glands 344
10.6 The Eyes and Vision 289 Pancreatic Islets 344
Refraction 293 Pineal Gland 346
Accommodation 294 Gastrointestinal Tract 348
Visual Acuity 295 Gonads and Placenta 348
10.7 Retina 296 11.7 Paracrine and Autocrine Regulation 348
Effect of Light on the Rods 298 Examples of Paracrine and Autocrine
Electrical Activity of Retinal Cells 299 Regulation 349
Cones and Color Vision 300 Prostaglandins 350
Visual Acuity and Sensitivity 302 Interactions 353
Neural Pathways from the Retina 303 Summary 354
10.8 Neural Processing of Visual Information 306 Review Activities 355

Ganglion Cell Receptive Fields 306

Lateral Geniculate Nuclei 307
Cerebral Cortex 307
Interactions 309
12 Muscle
12.1 Skeletal Muscles
358

359
Summary 310 Structure of Skeletal Muscles 359
Review Activities 313 Motor End Plates and Motor Units 360
12.2 Mechanisms of Contraction 363

11
Sliding Filament Theory of Contraction 366
Endocrine Glands 315 Regulation of Contraction 368
11.1 Endocrine Glands and Hormones 316 12.3 Contractions of Skeletal Muscles 373

Common Aspects of Neural and Endocrine Twitch, Summation, and Tetanus 373
Regulation 316 Types of Muscle Contractions 374
Chemical Classification of Hormones 318 Series-Elastic Component 375
Prohormones and Prehormones 319 Length-Tension Relationship 375
Hormone Interactions 320 12.4 Energy Requirements of Skeletal Muscles 376
Effects of Hormone Concentrations on Tissue Metabolism of Skeletal Muscles 377
Response 320 Slow- and Fast-Twitch Fibers 379
11.2 Mechanisms of Hormone Action 322 Muscle Fatigue 381
Hormones That Bind to Nuclear Receptor Adaptations of Muscles to Exercise Training 381
Proteins 322
Muscle Damage and Repair 383
Hormones That Use Second Messengers 325
12.5 Neural Control of Skeletal Muscles 383
11.3 Pituitary Gland 330
Muscle Spindle 384
Pituitary Hormones 330

ISTUDY
 Contents xix

14 Cardiac
Alpha and Gamma Motor Neurons 386
Coactivation of Alpha and Gamma Motor Neurons 386 Output, Blood Flow,
Skeletal Muscle Reflexes 386 and Blood Pressure 449
Upper Motor Neuron Control of Skeletal Muscles 389
14.1 Cardiac Output 450
12.6 Cardiac and Smooth Muscles 390
Regulation of Cardiac Rate 450
Cardiac Muscle 391
Regulation of Stroke Volume 451
Smooth Muscle 392
Venous Return 454
Interactions 397
14.2 Blood Volume 455
Summary 398
Exchange of Fluid Between Capillaries and Tissues 456
Review Activities 400
Regulation of Blood Volume by the Kidneys 458

13 Blood,
14.3 Vascular Resistance to Blood Flow 462

Heart, and Physical Laws Describing Blood Flow 463

Circulation 403
Extrinsic Regulation of Blood Flow 464
Paracrine Regulation of Blood Flow 465
13.1 Functions and Components of the Intrinsic Regulation of Blood Flow 466
Circulatory System 404 14.4 Blood Flow to the Heart and Skeletal Muscles 467
Functions of the Circulatory System 404 Aerobic Requirements of the Heart 467
Major Components of the Circulatory System 404 Regulation of Coronary Blood Flow 468
13.2 Composition of the Blood 405 Regulation of Blood Flow Through Skeletal Muscles 469
Blood Plasma 405 Circulatory Changes During Exercise 469
The Formed Elements of Blood 406 14.5 Blood Flow to the Brain and Skin 472
Hematopoiesis 408 Cerebral Circulation 472
Red Blood Cell Antigens and Blood Typing 411 Cutaneous Blood Flow 473
Blood Clotting 413 14.6 Blood Pressure 474
Dissolution of Clots 416 Baroreceptor Reflex 476
13.3 Structure of the Heart 417 Atrial Stretch Reflexes 478
Pulmonary and Systemic Circulations 417 Measurement of Blood Pressure 478
Atrioventricular and Semilunar Valves 418 Pulse Pressure and Mean Arterial Pressure 480
Heart Sounds 419 14.7 Hypertension, Shock, and Congestive
13.4 Cardiac Cycle 421 Heart Failure 481
Pressure Changes During the Cardiac Cycle 422 Hypertension 481
13.5 Electrical Activity of the Heart and Circulatory Shock 483
the Electrocardiogram 424 Congestive Heart Failure 485
Electrical Activity of the Heart 424 Interactions 487
The Electrocardiogram 427 Summary 488
13.6 Blood Vessels 430 Review Activities 489
Arteries 430

15 The Immune System

Capillaries 432
Veins 434 492
13.7 Atherosclerosis and Cardiac Arrhythmias 435
15.1 Defense Mechanisms 493
Atherosclerosis 435
Innate (Nonspecific) Immunity 493
Arrhythmias Detected by the Electrocardiograph 439
Adaptive (Specific) Immunity 496
13.8 Lymphatic System 441
Lymphocytes and Lymphoid Organs 498
Summary 444
Local Inflammation 499
Review Activities 446

ISTUDY
 xx Contents

15.2 Functions of B Lymphocytes 502 Effects of Blood PO2 on Ventilation 556

Antibodies 503 Effects of Pulmonary Receptors on Ventilation 557
The Complement System 505 16.6 Hemoglobin and Oxygen Transport 558
15.3 Functions of T Lymphocytes 506 Hemoglobin 558
Cytotoxic, Helper, and Regulatory T Lymphocytes 506 The Oxyhemoglobin Dissociation Curve 560
Interactions Between Antigen-Presenting Cells and T Effect of pH and Temperature on Oxygen Transport 561
Lymphocytes 510 Effect of 2,3-DPG on Oxygen Transport 562
15.4 Active and Passive Immunity 513 Inherited Defects in Hemoglobin Structure and
Active Immunity and the Clonal Selection Theory 514 Function 562
Immunological Tolerance 516 Muscle Myoglobin 563
Passive Immunity 517 16.7 Carbon Dioxide Transport 564

15.5 Tumor Immunology 518 The Chloride Shift 564

Innate Lymphoid Cells 519 The Reverse Chloride Shift 565
Effects of Aging and Stress 520 16.8 Acid-Base Balance of the Blood 566

15.6 Diseases Caused by the Immune System 520 Principles of Acid-Base Balance 567
Autoimmunity 520 Ventilation and Acid-Base Balance 568
Immune Complex Diseases 522 16.9 Effect of Exercise and High Altitude on
Respiratory Function 569
Allergy 522
Ventilation During Exercise 569
Interactions 526
Acclimatization to High Altitude 570
Summary 527
Interactions 574
Review Activities 528
Summary 575

16 Respiratory Physiology
Review Activities 577

17 Physiology of the Kidneys

531

16.1 The Respiratory System 532 580

Structure of the Respiratory System 532 17.1 Structure and Function of the Kidneys 581
Thoracic Cavity 535 Gross Structure of the Urinary System 581
16.2 Physical Aspects of Ventilation 535 Control of Micturition 583
Intrapulmonary and Intrapleural Pressures 536 Microscopic Structure of the Kidney 583
Physical Properties of the Lungs 537 17.2 Glomerular Filtration 586
Surfactant and Respiratory Distress Syndrome 539 Glomerular Ultrafiltrate 587
16.3 Mechanics of Breathing 539 Regulation of Glomerular Filtration Rate 588
Inspiration and Expiration 540 17.3 Reabsorption of Salt and Water 589
Pulmonary Function Tests 541 Reabsorption in the Proximal Tubule 590
Pulmonary Disorders 543 The Countercurrent Multiplier System 591
16.4 Gas Exchange in the Lungs 546 Collecting Duct: Effect of Antidiuretic Hormone (ADH) 594
Calculation of PO2 546 17.4 Renal Plasma Clearance 597
Partial Pressures of Gases in Blood 547 Transport Process Affecting Renal Clearance 598
Significance of Blood PO2 and PCO2 Measurements 549 Renal Clearance of Inulin: Measurement of GFR 599
Pulmonary Circulation and Ventilation/Perfusion Renal Clearance Measurements 600
Ratios 549
Reabsorption of Glucose 601
Disorders Caused by High Partial Pressures of Gases 551
17.5 Renal Control of Electrolyte and Acid–Base
16.5 Regulation of Breathing 552 Balance 603
Brain Stem Respiratory Centers 552 Role of Aldosterone in Na+/K+ Balance 603
Effects of Blood PCO2 and pH on Ventilation 554 Control of Aldosterone Secretion 605

ISTUDY
 Contents xxi

19 Regulation of Metabolism
Natriuretic Peptides 606
Relationship Between Na+, K+, and H+ 607 662

Renal Acid-Base Regulation 607 19.1 Nutritional Requirements 663

17.6 Diuretics and Renal Function Tests 610 Metabolic Rate and Caloric Requirements 663
Use of Diuretics 610 Anabolic Requirements 665
Renal Function Tests and Kidney Disease 612 Vitamins and Minerals 665
Interactions 613 Free Radicals and Antioxidants 668
Summary 614 19.2 Regulation of Energy Metabolism 670
Review Activities 615 Regulatory Functions of Adipose Tissue 671
Regulation of Hunger and Metabolic Rate 673

18
18.1
The Digestive System
Introduction to the Digestive System
618

619 19.3
Caloric Expenditures 675
Hormonal Regulation of Metabolism 676
Energy Regulation by the Pancreatic Islets 678
Layers of the Alimentary Tract 620 Regulation of Insulin and Glucagon Secretion 678
Regulation of the Alimentary Tract 622 Insulin and Glucagon: Absorptive State 680
18.2 From Mouth to Stomach 622 Insulin and Glucagon: Postabsorptive State 680
Esophagus 623 19.4 Diabetes Mellitus and Hypoglycemia 682
Stomach 624 Type 1 Diabetes Mellitus 683
Pepsin and Hydrochloric Acid Secretion 624 Type 2 Diabetes Mellitus 683
18.3 Small Intestine 628 Hypoglycemia 686
Villi and Microvilli 628 19.5 Metabolic Regulation by Adrenal Hormones,
Intestinal Enzymes 629 Thyroxine, and Growth Hormone 687
Intestinal Contractions and Motility 630 Adrenal Hormones 687
18.4 Large Intestine 632 Thyroxine 688
Intestinal Microbiota 633 Growth Hormone 689
Fluid and Electrolyte Absorption in the Intestine 635 19.6 Regulation of Calcium and Phosphate
Balance 691
Defecation 636
Bone Deposition and Resorption 691
18.5 Liver, Gallbladder, and Pancreas 636
Hormonal Regulation of Bone 693
Structure of the Liver 636
1,25-Dihydroxyvitamin D3 695
Functions of the Liver 639
Negative Feedback Control of Calcium and
Gallbladder 642
Phosphate Balance 696
Pancreas 643
Summary 698
18.6 Regulation of the Digestive System 645
Review Activities 699
Regulation of Gastric Function 646

20 Reproduction
Regulation of Intestinal Function 648
Regulation of Pancreatic Juice and Bile Secretion 650 702
Trophic Effects of Gastrointestinal Hormones 650
20.1 Sexual Reproduction 703
18.7 Digestion and Absorption of Food 651
Sex Determination 703
Digestion and Absorption of Carbohydrates 651
Development of Accessory Sex Organs and
Digestion and Absorption of Proteins 652
External Genitalia 706
Digestion and Absorption of Lipids 653
Disorders of Embryonic Sexual Development 707
Interactions 657
20.2 Endocrine Regulation of Reproduction 709
Summary 658
Interactions Among the Hypothalamus,
Review Activities 659 Pituitary Gland, and Gonads 710

ISTUDY
 xxii Contents

Onset of Puberty 711 20.6 Fertilization, Pregnancy, and Parturition 735

Pineal Gland 713 Fertilization 736
Human Sexual Response 713 Cleavage and Blastocyst Formation 738
20.3 Male Reproductive System 713 Implantation of the Blastocyst and Formation of the
Control of Gonadotropin Secretion 714 Placenta 741
Endocrine Functions of the Testes 715 Exchange of Molecules Across the Placenta 744
Spermatogenesis 716 Endocrine Functions of the Placenta 744
Male Accessory Sex Organs 719 Labor and Parturition 746
Erection, Emission, and Ejaculation 720 Lactation 747
Male Fertility 722 Concluding Remarks 750

20.4 Female Reproductive System 723 Interactions 751

Ovarian Cycle 725 Summary 752
Ovulation 727 Review Activities 753
Hypothalamic-Pituitary-Ovarian Axis 728
20.5 Menstrual Cycle 729 Appendixes
Phases of the Menstrual Cycle: Cyclic Changes in the Answers to Test Your Knowledge Questions A-1
Ovaries 729 Medical and Pharmacological Abbreviations B-1
Cyclic Changes in the Endometrium 732
Glossary G-1
Effects of Pheromones, Stress, and Body Fat 733
Contraceptive Methods 734 Index I-1

Menopause 735

ISTUDY
 1 The Study of Body
Function

C H A P TE R O UTLI N E
CLINICAL INVESTIGATION
1.1 Introduction to Physiology 2
As you study the sections of this chapter, you can see
Scientific Method 2
how your new knowledge can be applied to interesting
1.2 Homeostasis and Feedback Control 4
health issues that may be important to know in your future
History of Physiology 4
career as a health professional. This can add zest to your Negative Feedback Loops 5
studies and increase your motivation to truly understand Positive Feedback 8
physiological concepts, rather than to simply memorize Neural and Endocrine Regulation 8
facts for examinations. Each chapter begins with a medical Feedback Control of Hormone Secretion 8

mystery for you to solve, using information in the text of that 1.3 The Primary Tissues 10
Muscle Tissue 10
chapter and “Clinical Investigation Clues” within the chapter.
Nerve Tissue 11
For example, suppose Linda goes for a medical Epithelial Tissue 12
examination where her body temperature is measured, and Connective Tissue 15
she gives a fasting blood sample to test for glucose. Your 1.4 Organs and Systems 17
first Clinical Investigation challenge is to determine the An Example of an Organ: The Skin 18
medical significance of these physiological tests. Systems 19
Body-Fluid Compartments 20
Summary 21
Review Activities 22

ISTUDY
 2 Chapter 1
1.1 INTRODUCTION TO
PHYSIOLOGY
Human physiology is the study of how the human body
functions, with emphasis on specific cause-and-effect
mechanisms. Knowledge of these mechanisms has been
obtained experimentally through applications of the scien-
tific method.
LE A R N I N G O UTCO M E S
After studying this section, you should be able to:
1. Describe the scientific study of human physiology.
2. Describe the characteristics of the scientific method.
Physiology (from the Greek physis = nature; logos = study)
is the study of biological function—of how the body works,
from molecular mechanisms within cells to the actions of tis-
sues, organs, and systems, and how the organism as a whole
accomplishes particular tasks essential for life. In the study of
physiology, the emphasis is on mechanisms—with questions
that begin with the word how and answers that involve cause-
and-effect sequences. These sequences can be woven into
larger and larger stories that include descriptions of the struc-
tures involved (anatomy) and that overlap with the sciences of
chemistry and physics.
The separate facts and relationships of these cause-and-
effect sequences are derived empirically from experimental
evidence. Explanations that seem logical are not necessar-
ily true; they are only as valid as the data on which they are
based, and they can change as new techniques are developed
and further experiments are performed. The ultimate objec-
tive of physiological research is to understand the normal
functioning of cells, organs, and systems. A related science—
pathophysiology—is concerned with how physiological pro-
cesses are altered in disease or injury.
Pathophysiology and the study of normal physiology
complement one another. For example, a standard technique
for investigating the functioning of an organ is to observe what
happens when the organ is surgically removed from an experi-
mental animal or when its function is altered in a specific
way. This study is often aided by “experiments of nature”—
diseases—that involve specific damage to the functioning of
an organ. The study of disease processes has thus aided our
understanding of normal functioning, and the study of nor-
mal physiology has provided much of the scientific basis of
modern medicine. This relationship is recognized by the Nobel
Prize committee, whose members award prizes in the category
“Physiology or Medicine.”
The physiology of invertebrates and of different vertebrate
groups is studied in the science of comparative physiology.
Much of the knowledge gained from comparative physiology has
benefited the study of human physiology. This is because ani-
mals, including humans, are more alike than they are different.
ISTUDY
This is especially true when comparing humans with other mam-
mals. The small differences in physiology between humans and
other mammals can be of crucial importance in the development
of pharmaceutical drugs (discussed later in this section), but these
differences are relatively slight in the overall study of physiology.
Scientific Method
All of the information in this text has been gained by people
applying the scientific method. Although many different tech-
niques are involved when people apply the scientific method,
all share three attributes: (1) confidence that the natural world,
including ourselves, is ultimately explainable in terms we can
understand; (2) descriptions and explanations of the natural
world that are honestly based on observations and that could
be modified or refuted by other observations; and (3) humility,
or the willingness to accept the fact that we could be wrong. If
further study should yield conclusions that refuted all or part
of an idea, the idea would have to be modified accordingly. In
order for the scientific enterprise to function, its practitioners
must honestly report their data and observations and be will-
ing to modify their ideas, sometimes long-held and cherished,
in response to new scientific information. Practicing scien-
tists may not always display these attributes, but the validity
of the large body of scientific knowledge that has been accu-
mulated—as shown by the technological applications and the
predictive value of scientific hypotheses—is ample testimony
to the fact that the scientific method works.
The scientific method involves specific steps. After cer-
tain observations regarding the natural world are made, a
hypothesis is formulated. In order for this hypothesis to be
scientific, it must be capable of being refuted by experiments
or other observations of the natural world. For example, one
might hypothesize that people who exercise regularly have a
lower resting pulse rate than other people. Experiments are
conducted, or other observations are made, and the results are
analyzed. Conclusions are then drawn as to whether the new
data either refute or support the hypothesis. If the hypoth-
esis survives such testing, it might be incorporated into a
more general theory. Scientific theories are thus not simply
conjectures; they are statements about the natural world that
incorporate a number of hypotheses that have been supported
by scientific evidence. They serve as a logical framework by
which these hypotheses can be interrelated and provide the
basis for predictions that may as yet be untested.
The hypothesis in the preceding example is scientific
because it is testable; the pulse rates of 100 athletes and 100
sedentary people could be measured, for example, to see if
there were statistically significant differences. If there were,
the statement that athletes, on the average, have lower resting
pulse rates than other people would be justified based on these
data. One must still be open to the fact that this conclusion
could be wrong. Before the discovery could become generally
accepted as fact, other scientists would have to consistently
replicate the results. Scientific theories are based on reproduc-
ible data.

It is quite possible that when others attempt to replicate the
experiment, their results will be slightly different. They may
then construct scientific hypotheses that the differences in rest-
ing pulse rate also depend on other factors, such as the nature
of the exercise performed. When scientists attempt to test these
hypotheses, they will likely encounter new problems requir-
ing new explanatory hypotheses, which then must be tested by
additional experiments.
In this way, a large body of highly specialized information
is gradually accumulated, and a more generalized explanation
(a scientific theory) can be formulated. This explanation will
almost always be different from preconceived notions. People
who follow the scientific method will then appropriately mod-
ify their concepts, realizing that their new ideas will probably
have to be changed again in the future as additional experi-
ments are performed.
Use of Measurements, Controls,
and Statistics
Suppose you wanted to test the hypothesis that a regular exer-
cise program causes people to have a lower resting heart rate.
First, you would have to decide on the nature of the exercise
program. Then, you would have to decide how the heart rate
(or pulse rate) would be measured. This is a typical problem in
physiology research because the testing of most physiological
hypotheses requires quantitative measurements.
The group that is subject to the testing condition—in this
case, exercise—is called the experimental group. A measure-
ment of the heart rate for this group would be meaningful only
if it is compared to that of another group, known as the control
group. How shall this control group be chosen? Perhaps the
subjects could serve as their own controls—that is, a person’s
resting heart rate could be measured before and after the exer-
cise regimen. If this isn’t possible, a control group could be
other people who do not follow the exercise program. The
choice of control groups is often a controversial aspect of phys-
iology studies. In this example, did the people in the control
group really refrain from any exercise? Were they comparable
to the people in the experimental group with regard to age, sex,
ethnicity, body weight, health status, and so on? You can see
how difficult it could be in practice to get a control group that
could satisfy any potential criticism.
Another possible criticism could be bias in the way that
the scientists perform the measurements. This bias could be
completely unintentional; scientists are human, after all, and
they may have invested months or years in this project. To pre-
vent such bias, the person doing the measurements often does
not know if a subject is part of the experimental or the control
group. This is known as a blind measurement.
Now suppose the data are in and it looks like the experi-
mental group indeed has a lower average resting heart rate than
the control group. But there is overlap—some people in the con-
trol group have measurements that are lower than some people
in the experimental group. Is the difference in the average mea-
surements of the groups due to a real physiological difference,
ISTUDY
The Study of Body Function 3
or is it due to chance variations in the measurements? Scien-
tists attempt to test the null hypothesis (the hypothesis that the
difference is due to chance) by employing the mathematical
tools of statistics. If the statistical results so warrant, the null
hypothesis can be rejected and the experimental hypothesis
can be deemed to be supported by this study.
The statistical test chosen will depend on the design of the
experiment, and it can also be a source of contention among
scientists in evaluating the validity of the results. Because of
the nature of the scientific method, “proof” in science is always
provisional. Some other researchers, employing the scientific
method in a different way (with different measuring tech-
niques, experimental procedures, choice of control groups, sta-
tistical tests, and so on), may later obtain different results. The
scientific method is thus an ongoing enterprise.
The results of the scientific enterprise are written up as
research articles, and these must be reviewed by other scien-
tists who work in the same field before they can be published
in peer-reviewed journals. More often than not, the reviewers
will suggest that certain changes be made in the articles before
they can be accepted for publication.
Examples of such peer-reviewed journals that pub-
lish articles in many scientific fields include Science
(www.sciencemag.org/), Nature (www.nature.com/nature/),
and Proceedings of the National Academy of Sciences
(www.pnas.org/). Review articles on physiology can be found
in Annual Review of Physiology (physiol.annualreviews.org/),
Physiological Reviews (journals.physiology.org), and Physi-
ology (physiologyonline.physiology.org). Medical research
journals, such as the New England Journal of Medicine
(content.nejm.org/) and Nature Medicine (www.nature.com/nm/),
also publish articles of physiological interest. There are also
many specialty journals in areas of physiology such as neuro-
physiology, endocrinology, and cardiovascular physiology.
Students who wish to look online for scientific articles
published in peer-reviewed journals that relate to a particular
subject can do so at the National Library of Medicine website,
PubMed (www.ncbi.nlm.nih.gov/entrez/query.fcgi).
Development of Pharmaceutical Drugs
The development of new pharmaceutical drugs can serve as
an example of how the scientific method is used in physiology
and its health applications. The process usually starts with
basic physiological research, often at cellular and molecular
levels. Perhaps a new family of drugs is developed using cells
in tissue culture (in vitro, or outside the body). For example,
cell physiologists studying membrane transport may discover
that a particular family of compounds blocks membrane chan-
nels for calcium ions (Ca2+). Because of their knowledge
of physiology, other scientists may predict that a drug of
this nature might be useful in the treatment of hypertension
(high blood pressure). This drug may then be tried in animal
experiments.
If a drug is effective at extremely low concentrations in
vitro (in cells cultured outside of the body), there is a chance
 4 Chapter 1
that it may work in vivo (in the body) at concentrations low
enough not to be toxic (poisonous). This possibility must be
thoroughly tested utilizing experimental animals, primarily
rats and mice. More than 90% of drugs tested in experimental
animals are too toxic for further development. Only in those
rare cases when the toxicity is low enough may development
progress to human/clinical trials.
Biomedical research is often aided by animal models of
particular diseases. These are strains of laboratory rats and
mice that are genetically susceptible to particular diseases that
resemble human diseases. Research utilizing laboratory ani-
mals typically takes several years and always precedes human
(clinical) trials of promising drugs. It should be noted that this
length of time does not include all of the years of “basic” phys-
iological research (involving laboratory animals) that provided
the scientific foundation for the specific medical application.
In phase I clinical trials, the drug is tested on healthy
human volunteers. This is done to test its toxicity in humans
and to study how the drug is “handled” by the body: how it is
metabolized, how rapidly it is removed from the blood by the
liver and kidneys, how it can be most effectively administered,
and so on. If significant toxic effects are not observed, the drug
can proceed to the next stage. In phase II clinical trials, the
drug is tested on the target human population (for example,
those with hypertension). Only in those exceptional cases
where the drug seems to be effective but has minimal toxicity
does testing move to the next phase. Phase III trials occur in
many research centers across the country to maximize the num-
ber of test participants. At this point, the test population must
include a sufficient number of subjects of both sexes, as well as
people of different ethnic groups. In addition, people are tested
who have other health problems besides the one that the drug
is intended to benefit. For example, those who have diabetes
in addition to hypertension would be included in this phase. If
the drug passes phase III trials, it goes to the Food and Drug
Administration (FDA) for approval. Phase IV trials test other
potential uses of the drug. These “post-marketing studies” often
reveal problems with the drug that were not previously evident.
Less than 10% of the tested drugs make it all the way
through clinical trials to eventually become approved and mar-
keted. This low success rate does not count those that fail after
approval because of unexpected toxicity, nor does it take into
account the great amount of drugs that fail earlier in research
before clinical trials begin. Notice the crucial role of basic
research, using experimental animals, in this process. Virtu-
ally every prescription drug on the market owes its existence
to such research.
C H E C K P O I NT S
1. How has the study of physiology aided, and been
aided by, the study of diseases?
2a. Describe the steps involved in the scientific method.
What would qualify a statement as unscientific?
2b. Describe the different types of trials a new drug must
undergo before it is “ready for market.”
ISTUDY
1.2 HOMEOSTASIS AND
FEEDBACK CONTROL
The regulatory mechanisms of the body can be understood
in terms of a single shared function: that of maintaining
constancy of the internal environment. A state of relative
constancy of the internal environment is known as homeo-
stasis, maintained by negative feedback loops.
LE A R N I N G O UTCO M E S
After studying this section, you should be able to:
3. Define homeostasis, and identify the components of
negative feedback loops.
4. Explain the role of antagonistic effectors in maintaining
homeostasis, and the nature of positive feedback loops.
5. Give examples of how negative feedback loops
involving the nervous and endocrine systems help to
maintain homeostasis.
History of Physiology
The Greek philosopher Aristotle (384–322 b.c.) speculated on
the function of the human body, but another ancient Greek,
Erasistratus (304–250 b.c.), is considered to be the first to
study physiology because he attempted to apply physical laws
to understand human function. Galen (a.d. 130–210) wrote
widely on the subject and was considered the supreme author-
ity until the Renaissance. Physiology became a fully experi-
mental science with the revolutionary work of the English
physician William Harvey (1578–1657), who demonstrated
that the heart pumps blood through a closed system of vessels.
However, the originator of modern physiology is the French
physiologist Claude Bernard (1813–1878), who observed that
the milieu intérieur (internal environment) remains remark-
ably constant despite changing conditions in the external envi-
ronment. In a book titled The Wisdom of the Body, published in
1932, the American physiologist Walter Cannon (1871–1945)
coined the term homeostasis to describe this internal con-
stancy. Cannon further suggested that the many mechanisms
of physiological regulation have but one purpose—the main-
tenance of internal constancy. In the early 1950s, James Har-
din extended Cannon’s concept by proposing that homeostatic
mechanisms maintain each physiological variable within a
normal range by comparing its value to a desired, or set point,
value (as will be described shortly).
Most of our present knowledge of human physiology has
been gained in the twentieth century. However, new knowl-
edge in the twenty-first century is being added at an ever more
rapid pace, fueled in more recent decades by the revolution-
ary growth of molecular genetics and its associated biotech-
nologies, and by the availability of more powerful computers
and other equipment. A very brief history of twentieth- and
 The Study of Body Function 5

twenty-first-century physiology, limited by space to only two

citations per decade, is provided in table 1.1.
Most of the citations in table 1.1 indicate the winners of
Nobel Prizes. The Nobel Prize in Physiology or Medicine
(a single prize category) was first awarded in 1901 to Emil
Adolf von Behring, a pioneer in immunology who coined the
term antibody and whose many other discoveries included the
use of serum (containing antibodies) to treat diphtheria. Many
scientists who might deserve a Nobel Prize never receive one,
and the prizes are given for particular achievements and not
others (Einstein didn’t win his Nobel Prize in Physics for rela-
tivity, for example) and are often awarded many years after
the discoveries were made. Nevertheless, the awarding of the
Nobel Prize in Physiology or Medicine each year is a cele-
brated event in the biomedical community, and the awards can
be a useful yardstick for tracking the course of physiological
research over time.
Negative Feedback Loops
The concept of homeostasis has been of immense value in the
study of physiology because it allows diverse regulatory mech-
anisms to be understood in terms of their “why” as well as
their “how.” The concept of homeostasis also provides a major
foundation for medical diagnostic procedures. When a par-
ticular measurement of the internal environment, such as a
blood measurement (table 1.2), deviates significantly from the
normal range of values, it can be concluded that homeostasis is
not being maintained and that the person is sick. A number of
such measurements, combined with clinical observations, may
allow the particular defective mechanism to be identified.
In order for internal constancy to be maintained, changes
in the body must stimulate receptors, which function as sen-
sors that can send information to an integrating center. This
allows the integrating center to detect changes from a set point.

TABLE 1.1 | History of Twentieth- and Twenty-First-Century Physiology (two citations per decade)
1900 Karl Landsteiner discovers the A, B, and O blood groups.
1904 Ivan Pavlov wins the Nobel Prize for his work on the physiology of digestion.
1910 Sir Henry Dale describes properties of histamine.
1918 Earnest Starling describes how the force of the heart’s contraction relates to the amount of blood in it.
1921 John Langley describes the functions of the autonomic nervous system.
1923 Sir Frederick Banting, Charles Best, and John Macleod win the Nobel Prize for the discovery of insulin.
1932 Sir Charles Sherrington and Lord Edgar Adrian win the Nobel Prize for their discoveries related to the functions of neurons.
1936 Sir Henry Dale and Otto Loewi win the Nobel Prize for the discovery of acetylcholine in synaptic transmission.
1939–47 Albert von Szent-Györgyi explains the role of ATP and contributes to the understanding of actin and myosin in muscle contraction.
1949 Hans Selye discovers the common physiological responses to stress.
1953 Sir Hans Krebs wins the Nobel Prize for his discovery of the citric acid cycle.
1954 Hugh Huxley, Jean Hanson, R. Niedergerde, and Andrew Huxley propose the sliding filament theory of muscle contraction.
1962 Francis Crick, James Watson, and Maurice Wilkins win the Nobel Prize for determining the structure of DNA.
1963 Sir John Eccles, Sir Alan Hodgkin, and Sir Andrew Huxley win the Nobel Prize for their discoveries relating to the nerve impulse.
1971 Earl Sutherland wins the Nobel Prize for his discovery of the mechanism of hormone action.
1977 Roger Guillemin and Andrew Schally win the Nobel Prize for their discoveries of the brain’s production of peptide hormone.
1981 Roger Sperry wins the Nobel Prize for his discoveries regarding the specializations of the right and left cerebral hemispheres.
1986 Stanley Cohen and Rita Levi-Montalcini win the Nobel Prize for their discoveries of growth factors regulating the nervous system.
1994 Alfred Gilman and Martin Rodbell win the Nobel Prize for their discovery of the functions of G-proteins in signal transduction in cells.
1998 Robert Furchgott, Louis Ignarro, and Ferid Murad win the Nobel Prize for discovering the role of nitric oxide as a signaling molecule
in the cardiovascular system.
2004 Linda B. Buck and Richard Axel win the Nobel Prize for their discoveries of odorant receptors and the organization of the olfactory
system.
2012 Sir John Gurdon and Shinya Yamanaka win the Nobel Prize for their discoveries that mature cells can be reprogrammed to become
pluripotent (like embryonic cells).

2019 William G. Kaelin, Gregg L. Semenza, and Peter J. Ratcliffe win the Nobel Prize for their discoveries of how cells sense the oxygen
levels in their environment and initiate physiological responses.

ISTUDY
 6 Chapter 1
TABLE 1.2 | Approximate Normal Ranges for
Measurements of Some Fasting Blood Values
Measurement Normal Range
Arterial pH 7.35–7.45
Bicarbonate 24–28 mEq/L
Sodium 135–145 mEq/L
Calcium 4.5–5.5 mEq/L
Oxygen content 17.2–22.0 ml/100 ml
Urea 12–35 mg/100 ml
Amino acids 3.3–5.1 mg/100 ml
Protein 6.5–8.0 g/100 ml
Total lipids 400–800 mg/100 ml
Glucose 70–99 mg/100 ml
The set point is analogous to the temperature set on a house
thermostat. In a similar manner, there is a set point for body
temperature, blood glucose concentration, the tension on a
tendon, and so on. The integrating center is often a particular
region of the brain or spinal cord, but it can also be a group
of cells in an endocrine gland. A number of different sensors
may send information to a particular integrating center, which
can then integrate this information and direct the responses of
effectors—generally muscles or glands. The integrating center
may cause increases or decreases in effector action to counter
the deviations from the set point and defend homeostasis.
The thermostat of a house can serve as a simple example.
Suppose you set the thermostat at a set point of 70° F. If the
temperature in the house rises sufficiently above the set point,
a sensor connected to an integrating center within the thermo-
stat will detect that deviation and turn on the air conditioner
(the effector in this example). The air conditioner will turn off
when the room temperature falls and the thermostat no longer
detects a deviation from the set-point temperature. However,
this simple example gives a wrong impression: the effectors in
the body are generally increased or decreased in activity, not
just turned on or off. Because of this, negative feedback con-
trol in the body works far more efficiently than does a house
thermostat.
If the body temperature exceeds the set point of 37° C,
sensors in a part of the brain detect this deviation and, act-
ing via an integrating center (also in the brain), stimulate
activities of effectors (including sweat glands) that lower the
temperature. For another example, if the blood glucose con-
centration falls below normal, the effectors act to increase
the blood glucose. One can think of the effectors as “defend-
ing” the set points against deviations. Because the activity of
the effectors is influenced by the effects they produce, and
because this regulation is in a negative, or reverse, direction,
this type of control system is known as a negative feedback
loop (fig. 1.1). (Notice that in figure 1.1 and in all subsequent
ISTUDY
1
Sensor Integrating center
X
–
X Effector
2
Sensor activated Effector activated
Normal 1 2
range
X Time
FIGURE 1.1 A rise in some factor of the internal
environment (↑X) is detected by a sensor. This information is
relayed to an integrating center, which causes an effector
to produce a change (1) in the opposite direction (↓X). The initial
deviation is thus reversed (2), completing a negative feedback
loop (shown by the dashed arrow and negative sign). The numbers
indicate the sequence of changes.
figures, negative feedback is indicated by a dashed line and a
negative sign.)
The nature of the negative feedback loop can be under-
stood by again referring to the analogy of the thermostat and
air conditioner. After the air conditioner has been on for some
time, the room temperature may fall significantly below the set
point of the thermostat. When this occurs, the air conditioner
will be turned off. The effector (air conditioner) is turned on
by a high temperature and, when activated, produces a nega-
tive change (lowering of the temperature) that ultimately
causes the effector to be turned off. In this way, constancy is
maintained.
It is important to realize that these negative feedback loops
are continuous, ongoing processes. Thus, a particular nerve
fiber that is part of an effector mechanism may always display
some activity, and a particular hormone that is part of another
effector mechanism may always be present in the blood. The
nerve activity and hormone concentration may decrease in
response to deviations of the internal environment in one direc-
tion (fig. 1.1), or they may increase in response to deviations
in the opposite direction (fig. 1.2). Changes from the normal
range in either direction are thus compensated for by reverse
changes in effector activity.
Because negative feedback loops respond after deviations
from the set point have stimulated sensors, the internal envi-
ronment is never absolutely constant. Homeostasis is best con-
ceived as a state of dynamic constancy in which conditions are
stabilized above and below the set point. These conditions can
be measured quantitatively, in degrees Celsius for body temper-
ature, for example, or in milligrams per deciliter (one-tenth of a
liter) for blood glucose. The set point can be taken as the aver-
age value within the normal range of measurements (fig. 1.3).

1
X Sensor Integrating center
– range
X Effector
2
X Time
Normal
range 1 2
Sensor activated Effector activated
FIGURE 1.2 A fall in some factor of the internal
environment (↓X) is detected by a sensor. (Compare this
negative feedback loop with that shown in fig. 1.1.)
– – – To study physiological mechanisms, scientists must measure
Set point
(average)
– –
FIGURE 1.3 Negative feedback loops maintain a state of
dynamic constancy within the internal environment. The
completion of the negative feedback loop is indicated by negative
signs.
Antagonistic Effectors
Most factors in the internal environment are controlled by sev-
eral effectors, which often have antagonistic actions. Control
by antagonistic effectors is sometimes described as “push-
pull,” where the increasing activity of one effector is accom-
panied by decreasing activity of an antagonistic effector. This
affords a finer degree of control than could be achieved by sim-
ply switching one effector on and off.
Room temperature can be maintained, for example, by
simply turning an air conditioner on and off, or by just turning
a heater on and off. A much more stable temperature, how-
ever, can be achieved if the air conditioner and heater are both
controlled by a thermostat. Then the heater is turned on when
the air conditioner is turned off, and vice versa. Normal body
temperature is maintained about a set point of 37° C by the
antagonistic effects of sweating, shivering, and other mecha-
nisms (fig. 1.4).
The blood concentrations of glucose, calcium, and other
substances are regulated by negative feedback loops involving
hormones that promote opposite effects. Insulin, for example,
lowers blood glucose, and other hormones raise the blood
glucose concentration. The heart rate, similarly, is controlled
by nerve fibers that produce opposite effects: stimulation of
one group of nerve fibers increases heart rate; stimulation of
another group slows the heart rate.
ISTUDY
The Study of Body Function 7
Sweat Sweat
37° C Normal
Shiver Shiver
FIGURE 1.4 How body temperature is maintained within
the normal range. The body temperature normally has a set
point of 37° C. This is maintained, in part, by two antagonistic
mechanisms—shivering and sweating. Shivering is induced when
the body temperature falls too low, and it gradually subsides
as the temperature rises. Sweating occurs when the body
temperature is too high, and it diminishes as the body temperature
falls. Most aspects of the internal environment are regulated by the
antagonistic actions of different effector mechanisms.
See the Test Your Quantitative Ability section of the Review Activities
at the end of this chapter.
Quantitative Measurements
Normal
–
range specific values and mathematically determine such statistics as
their normal range, their averages, and their deviations from
the average (which can represent the set point). For these and
other reasons, quantitative measurements are basic to the sci-
ence of physiology. One example of this, and of the actions of
antagonistic mechanisms in maintaining homeostasis, is shown
in figure 1.5. Blood glucose concentrations were measured in
five healthy people before and after an injection of insulin, a
hormone that acts to lower the blood glucose concentration. A
graph of the data reveals that the blood glucose concentration
decreased rapidly but was brought back up to normal levels
within 80 minutes after the injection. This demonstrates that
Insulin injected
100
concentration
Glucose
(mg/dl)
50
0
–80 –40 0 40 80 120
Time (min)
FIGURE 1.5 Homeostasis of the blood glucose
concentration. Average blood glucose concentrations of
five healthy individuals are graphed before and after a rapid
intravenous injection of insulin. The “0” indicates the time of the
injection. The blood glucose concentration is first lowered by
the insulin injection, but is then raised back to the normal range
(by hormones antagonistic to insulin that stimulate the liver to
secrete glucose into the blood). Homeostasis of blood glucose is
maintained by the antagonistic actions of insulin and several other
hormones.
 8 Chapter 1
negative feedback mechanisms acted to restore homeostasis in
this experiment. These mechanisms involve the action of hor-
mones whose effects are antagonistic to that of insulin—that
is, they promote the secretion of glucose from the liver (see
chapter 19).
Positive Feedback
Constancy of the internal environment is maintained by effec-
tors that act to compensate for the change that served as the
stimulus for their activation; in short, by negative feedback
loops. A thermostat, for example, maintains a constant tem-
perature by increasing heat production when it is cold and
decreasing heat production when it is warm. The opposite
occurs during positive feedback—in this case, the action of
effectors amplifies those changes that stimulated the effec-
tors. A thermostat that works by positive feedback, for exam-
ple, would increase heat production in response to a rise in
temperature.
It is clear that homeostasis must ultimately be maintained
by negative rather than by positive feedback mechanisms. The
effectiveness of some negative feedback loops, however, is
increased by positive feedback mechanisms that amplify the
actions of a negative feedback response. Blood clotting, for
example, occurs as a result of a sequential activation of clotting
factors; the activation of one clotting factor results in activation
of many in a positive feedback cascade. In this way, a single
change is amplified to produce a blood clot. Formation of the
clot, however, can prevent further loss of blood, and thus rep-
resents the completion of a negative feedback loop that restores
homeostasis.
Two other examples of positive feedback in the body are
both related to the female reproductive system. One of these
examples occurs when estrogen, secreted by the ovaries, stim-
ulates the woman’s pituitary gland to secrete LH (luteinizing
hormone). This stimulatory, positive feedback effect creates an
“LH surge” (very rapid rise in blood LH concentrations) that
triggers ovulation. Interestingly, estrogen secretion after ovula-
tion has an inhibitory, negative feedback, effect on LH secre-
tion (this is the physiological basis for the birth control pill,
discussed in chapter 20). Another example of positive feedback
is contraction of the uterus during childbirth (parturition). Con-
traction of the uterus is stimulated by the pituitary hormone
oxytocin, and the secretion of oxytocin is increased by sensory
feedback from contractions of the uterus during labor. The
strength of uterine contractions during labor is thus increased
through positive feedback. The mechanisms involved in labor
are discussed in more detail in chapter 20 (see fig. 20.50).
Neural and Endocrine Regulation
Homeostasis is maintained by two general categories of regu-
latory mechanisms: (1) those that are intrinsic, or “built into”
the organs being regulated (such as molecules produced in the
walls of blood vessels that cause vessel dilation or constric-
tion); and (2) those that are extrinsic, as in regulation of an
ISTUDY
organ by the nervous and endocrine systems. The endocrine
system functions closely with the nervous system in regulating
and integrating body processes and maintaining homeostasis.
The nervous system controls the secretion of many endocrine
glands, and some hormones in turn affect the function of the
nervous system. Together, the nervous and endocrine systems
regulate the activities of most of the other systems of the body.
Regulation by the endocrine system is achieved by the
secretion of chemical regulators called hormones into the
blood, which carries the hormones to all organs in the body.
Only specific organs can respond to a particular hormone, how-
ever; these are known as the target organs of that hormone.
Nerve fibers are said to innervate the organs that they
regulate. When stimulated, these fibers produce electrochemi-
cal nerve impulses that are conducted from the origin of the
fiber to its terminals in the target organ innervated by the fiber.
These target organs can be muscles or glands that may function
as effectors in the maintenance of homeostasis.
For example, we have negative feedback loops that help
maintain homeostasis of arterial blood pressure, in part by
adjusting the heart rate. If everything else is equal, blood
pressure is lowered by a decreased heart rate and raised by an
increased heart rate. This is accomplished by regulating the
activity of the autonomic nervous system, as will be discussed
in later chapters. Thus, a fall in blood pressure—produced
daily as we go from a lying to a standing position—is compen-
sated by a faster heart rate (fig. 1.6). As a consequence of this
negative feedback loop, our heart rate varies as we go through
our day, speeding up and slowing down, so that we can main-
tain homeostasis of blood pressure and keep it within limits.
Feedback Control of
Hormone Secretion
The nature of the endocrine glands, the interaction of the ner-
vous and endocrine systems, and the actions of hormones will
be discussed in detail in later chapters. For now, it is sufficient
to describe the regulation of hormone secretion very broadly,
because it so superbly illustrates the principles of homeostasis
and negative feedback regulation.
Hormones are secreted in response to specific chemi-
cal stimuli. A rise in the plasma glucose concentration, for
example, stimulates insulin secretion from structures in the
pancreas known as the pancreatic islets. Hormones are also
secreted in response to nerve stimulation and stimulation by
other hormones.
The secretion of a hormone can be inhibited by its own
effects in a negative feedback manner. Insulin, as previously
described, produces a lowering of blood glucose. Because a
rise in blood glucose stimulates insulin secretion, a lowering of
blood glucose caused by insulin’s action inhibits further insu-
lin secretion. This closed-loop control system is called nega-
tive feedback inhibition (fig. 1.7a).
Homeostasis of blood glucose is too important—the brain
uses blood glucose as its primary source of energy—to entrust
to the regulation of only one hormone, insulin. So, when blood
 The Study of Body Function 9

Sensor
Integrating center
Effector Lying down
Standing up

Negative –
feedback 4. Rise in blood pressure 1. Blood pressure falls Stimulus
response

3. Heart rate
increases
2. Blood pressure
receptors respond Sensor
Effector

Medulla Sensory
Motor oblongata nerve fibers
nerve fibers of brain
Integrating center

FIGURE 1.6 Negative feedback control of blood pressure. Blood pressure influences the activity of sensory neurons from the
blood pressure receptors (sensors); a rise in pressure increases the firing rate, and a fall in pressure decreases the firing rate of nerve
impulses. When a person stands up from a lying-down position, the blood pressure momentarily falls. The resulting decreased firing rate
of nerve impulses in sensory neurons affects the medulla oblongata of the brain (the integrating center). This causes the motor nerves to
the heart (effector) to increase the heart rate, helping to raise the blood pressure.

Sensor
Integrating center Fasting
Effector

Eating Blood glucose

Pancreatic islets
Blood glucose (of Langerhans)

Pancreatic islets
Insulin
(of Langerhans)
Glucagon
–

– Insulin
Cellular uptake of glucose
Glucose secretion into
Cellular uptake of glucose blood by liver

Blood glucose Blood glucose

(a) (b)

FIGURE 1.7 Negative feedback control of blood glucose. (a) The rise in blood glucose that occurs after eating carbohydrates is
corrected by the action of insulin, which is secreted in increasing amounts at that time. (b) During fasting, when blood glucose falls, insulin
secretion is inhibited and the secretion of an antagonistic hormone, glucagon, is increased. This stimulates the liver to secrete glucose
into the blood, helping to prevent blood glucose from continuing to fall. In this way, blood glucose concentrations are maintained within a
homeostatic range following eating and during fasting.

ISTUDY
 10 Chapter 1
glucose falls during fasting, several mechanisms prevent it from
falling too far (fig. 1.7b). First, insulin secretion decreases, pre-
venting muscle, liver, and adipose cells from taking too much
glucose from the blood. Second, the secretion of a hormone
antagonistic to insulin, called glucagon, increases. Glucagon
stimulates processes in the liver (breakdown of a stored, starch-
like molecule called glycogen; chapter 2, section 2.2) that cause
it to secrete glucose into the blood. Through these and other
antagonistic negative feedback mechanisms, the blood glucose
is maintained within a homeostatic range.
CLINICAL INVESTIGATION CLUES
Clinical Investigation Clues are placed immediately
following the text information that pertains to the Clinical
Investigation for the chapter. Use these to solve the medical
mystery—if you need to, reread the information preceding
the “Clues.” You can check your answers against the Clinical
Investigation Summaries at the end of the chapters. In this
case, Linda had a normal resting body temperature and
a normal fasting glucose concentration, suggesting that
homeostasis of these values was being maintained.
C H E C K P O I NT S
3a. Define homeostasis and describe how this concept can
be used to explain physiological control mechanisms.
3b. Define negative feedback and explain how it
contributes to homeostasis. Illustrate this concept by
drawing and labeling a negative feedback loop.
4. Describe positive feedback and explain how this
process functions in the body.
5. Explain how the secretion of a hormone is controlled
by negative feedback inhibition. Use the control of
insulin secretion as an example.
1.3 THE PRIMARY TISSUES
The organs of the body are composed of four different pri-
mary tissues, each of which has its own characteristic struc-
ture and function. The activities and interactions of these
tissues determine the physiology of the organs.
LE A R N I N G O UTCO M E S
After studying this section, you should be able to:
6. Distinguish the primary tissues and their subtypes.
7. Relate the structure of the primary tissues to their
functions.
Although physiology is the study of function, it is difficult to
properly understand the function of the body without some
knowledge of its anatomy, particularly at a microscopic level. The
ISTUDY
microscopic anatomy of tissues constitutes a field of study known
as histology. The anatomy and histology of specific organs will
be discussed together with their functions in later chapters. In this
section, the common “fabric” of all organs is described.
Cells are the basic units of structure and function in the
body. Cells that have similar functions are grouped into catego-
ries called tissues. The entire body is composed of only four
major types of tissues. These primary tissues are (1) muscle,
(2) nerve, (3) epithelial, and (4) connective tissues. Groupings of
these four primary tissues into anatomical and functional units
are called organs. Organs, in turn, may be grouped together by
common functions into systems. The systems of the body act in
a coordinated fashion to maintain the entire organism.
Muscle Tissue
Muscle tissue is specialized for contraction. There are three types
of muscle tissue: skeletal, cardiac, and smooth. Skeletal muscle
is often called voluntary muscle because its contraction is con-
sciously controlled. Both skeletal and cardiac muscle tissues are
striated muscle tissue; they have striations, or stripes, that extend
across the width of the muscle cell (figs. 1.8 and 1.9). These stria-
tions are produced by a characteristic arrangement of contractile
Muscle
fibers
Nucleus
FIGURE 1.8 Skeletal muscle fibers showing the
characteristic light and dark cross striations. Because of this
feature, skeletal muscle is also called striated muscle tissue.
Al Telser/McGraw-Hill Education
Nucleus
Intercalated
discs
FIGURE 1.9 Human cardiac muscle. Notice the striated
appearance and dark-staining intercalated discs.
Al Telser/McGraw-Hill Education

Nuclei
FIGURE 1.10 A photomicrograph of smooth muscle
cells. Notice that these cells contain single, centrally located
nuclei and lack striations. McGraw-Hill Education/Dennis Strete,
photographer
proteins, and for this reason skeletal and cardiac muscle have sim-
ilar mechanisms of contraction. Smooth muscle (fig. 1.10) lacks
these striations and has a different mechanism of contraction.
Skeletal Muscle
Skeletal muscles are generally attached to bones at both ends
by means of tendons; hence, contraction produces movements
of the skeleton. There are exceptions to this pattern, however.
The tongue, superior portion of the esophagus, anal sphincter,
and diaphragm are also composed of skeletal muscle, but they
do not cause movements of the skeleton.
Beginning at about the fourth week of embryonic devel-
opment, separate cells called myoblasts fuse together to form
skeletal muscle fibers, or myofiber (from the Greek myos =
muscle). Although myofibers are often referred to as skeletal
muscle cells, each is actually a syncytium, or multinucleate mass
formed from the union of separate cells. Despite their unique
origin and structure, each myofiber contains mitochondria and
other organelles (described in chapter 3) common to all cells.
The muscle fibers within a skeletal muscle are arranged
in bundles, and within these bundles the fibers extend in par-
allel from one end of the bundle to the other. The parallel
arrangement of muscle fibers (fig. 1.8) allows each fiber to be
controlled individually: one can thus contract fewer or more
muscle fibers and, in this way, vary the strength of contrac-
tion of the whole muscle. The ability to vary, or “grade,” the
strength of skeletal muscle contraction is needed for precise
control of skeletal movements.
Cardiac Muscle
Although cardiac muscle is striated muscle tissue, it differs
markedly from skeletal muscle in appearance. Cardiac muscle is
found only in the heart, where the heart muscle cells, or cardiac
muscle (myocardial) cells, are short, branched, and intimately
interconnected to form a continuous fabric. Special areas of
contact between adjacent cells stain darkly to show intercalated
discs (fig. 1.9), which are characteristic of heart muscle.
The intercalated discs couple cardiac muscle cells to­­
gether mechanically and electrically. Unlike skeletal muscles,
ISTUDY
The Study of Body Function 11
therefore, the heart cannot produce a graded contraction by
varying the number of cells stimulated to contract. Because of
the way the heart is constructed, the stimulation of one cardiac
muscle cell results in the stimulation of all other cells in the
mass and a “wholehearted” contraction.
Smooth Muscle
As implied by the name, smooth muscle cells (fig. 1.10) do not
have the striations characteristic of skeletal and cardiac muscle.
Smooth muscle is found in the digestive tract, blood vessels,
bronchioles (small air passages in the lungs), and the ducts of
the urinary and reproductive systems. Circular arrangements
of smooth muscle in these organs produce constriction of the
lumen (cavity) when the muscle cells contract. The digestive
tract also contains longitudinally arranged layers of smooth
muscle. Peristalsis is the coordinated wavelike contractions of
the circular and longitudinal smooth muscle layers that push
food from the oral to the anal end of the digestive tract.
The three types of muscle tissue are discussed further in
chapter 12.
Nerve Tissue
Nerve tissue consists of nerve cells, or neurons, which are
specialized for the generation and conduction of electrical
events, and neuroglial (or glial) cells, which compose the neu-
roglia. Neuroglia provide the neurons with structural support
and perform a variety of functions that are needed for the nor-
mal physiology of the nervous system.
Each neuron consists of three parts: (1) a cell body,
(2) dendrites, and (3) an axon (fig. 1.11). The cell body contains
Dendrites
Cell body
Neuroglial cells
Axon
FIGURE 1.11 A photomicrograph of nerve tissue. A single
neuron and numerous smaller neuroglial cells can be seen.
©Ed Reschke
 12 Chapter 1

the nucleus and serves as the metabolic center of the cell. The
dendrites (literally, “branches”) are highly branched cytoplasmic
extensions of the cell body that receive input from other neurons
or from receptor cells. The axon is a single cytoplasmic extension
of the cell body that can be quite long (up to a few feet in length).
It is specialized for conducting nerve impulses from the cell body
to another neuron or to an effector (muscle or gland) cell.
Neuroglia do not conduct impulses but instead serve to
bind neurons together, modify the extracellular environment of
the nervous system, and influence the nourishment and electri-
cal activity of neurons. In recent years, neuroglia have been
shown to cooperate with neurons in chemical neurotransmis-
sion (chapter 7), and to have many other roles in the normal
physiology (as well as disease processes) of the brain and spi-
nal cord. Neuroglia are now known to be about as numerous
as neurons and can divide by mitosis throughout life, whereas
mature neurons do not divide.
Neurons and neuroglia are discussed in detail in chapter 7.
Epithelial Tissue
Epithelial tissue consists of cells that form membranes,
which cover and line the body surfaces, and of glands, which
are derived from these membranes. There are two categories
of glands. Exocrine glands (from the Greek exo = outside)
secrete chemicals through a duct that leads to the outside of
a membrane, and thus to the outside of a body surface. Endo-
crine glands (from the Greek endon = within) secrete chemi-
cals called hormones into the blood. Endocrine glands are
discussed in chapter 11.
Epithelial Membranes
The general term membrane refers to a thin sheet of some kind
that acts as a barrier and separation. When we talk about a
plasma membrane around cells, we are referring to a molecular
structure so thin that it cannot be seen with a microscope. Epi-
thelial membranes, by contrast, are composed of cells that are in
close contact with each other in a sheet-like arrangement visible
in a microscope. Epithelial membranes are classified accord-
ing to the number of their layers and the shape of the cells in
the upper layer (table 1.3). Epithelial cells that are flattened in
shape are squamous; those that are as wide as they are tall are
cuboidal; and those that are taller than they are wide are colum-
nar (fig. 1.12a–c). Those epithelial membranes that are only one
cell layer thick are known as simple epithelia; those that are
composed of a number of layers are stratified epithelia.
Epithelial membranes cover all body surfaces and line the
cavity (lumen) of every hollow organ. Thus, epithelial mem-
branes provide a barrier between the external environment

TABLE 1.3 | Summary of Epithelial Membranes

Type
Simple Epithelia
Simple squamous epithelium
Simple cuboidal epithelium
Simple columnar epithelium
Simple ciliated columnar
epithelium
Pseudostratified ciliated
columnar epithelium
Stratified Epithelia
Stratified squamous epithelium
(keratinized)
Stratified squamous epithelium
(nonkeratinized)
Stratified cuboidal epithelium
Transitional epithelium
Structure and Function
Single layer of cells; function varies with type
Single layer of flattened, tightly bound cells; diffusion
and filtration
Single layer of cube-shaped cells; excretion, secretion,
or absorption
Single layer of nonciliated, tall, column-shaped cells;
protection, secretion, and absorption
Single layer of ciliated, column-shaped cells;
transportive role through ciliary motion
Single layer of ciliated, irregularly shaped cells; many
goblet cells; protection, secretion, ciliary movement
Two or more layers of cells; function varies with type
Numerous layers containing keratin, with outer layers
flattened and dead; protection
Numerous layers lacking keratin, with outer layers
moistened and alive; protection and pliability
Usually two layers of cube-shaped cells; strengthening
of luminal walls
Numerous layers of rounded, nonkeratinized cells;
distension
Location
Covering visceral organs; linings of body
cavities, tubes, and ducts
Capillary walls; pulmonary alveoli of lungs;
covering visceral organs; linings of body
cavities
Surface of ovaries; linings of kidney tubules,
salivary ducts, and pancreatic ducts
Lining of most of digestive tract
Lining of uterine tubes
Lining of respiratory passageways
Epidermis of skin; linings of body openings,
ducts, and urinary bladder
Epidermis of skin
Linings of oral and nasal cavities, vagina, and
anal canal
Large ducts of sweat glands, salivary glands,
and pancreas
Walls of ureters, part of urethra, and urinary
bladder

ISTUDY
 The Study of Body Function 13

environments. In order for a substance to get into the body,

it must pass through an epithelial membrane, and simple epi-
thelia are specialized for this function. For example, a simple
squamous epithelium in the lungs allows the rapid passage of
oxygen and carbon dioxide between the air (external environ-
ment) and blood (internal environment). A simple columnar
epithelium in the small intestine, as another example, allows
digestion products to pass from the intestinal lumen (external
environment) to the blood (internal environment).
Dispersed among the columnar epithelial cells are special-
ized unicellular glands called goblet cells that secrete mucus.
The columnar epithelial cells in the uterine tubes of females
Nucleus Connective
tissue and in the respiratory passages contain numerous cilia (hair-
Basement like structures, described in chapter 3) that can move in a coor-
(a) membrane
dinated fashion and aid the functions of these organs.
The epithelial lining of the esophagus and vagina that pro-
vides protection for these organs is a stratified squamous epi-
thelium (fig. 1.13). This is a nonkeratinized membrane, and all
Basement
layers consist of living cells. The epidermis of the skin, by con-
Nucleus membrane trast, is keratinized, or cornified (fig. 1.14). Because the epi-
dermis is dry and exposed to the potentially desiccating effects
of the air, the surface is covered with dead cells that are filled

(b)

Basement
membrane
Goblet Connective
cell Nucleus tissue

(a)

Cytoplasm
Nucleus

Squamous
(c) surface cells

FIGURE 1.12 Different types of simple epithelia.

(a) Simple squamous, (b) simple cuboidal, and (c) simple columnar
Mitotically active
epithelia. The basement membrane, composed of proteins and germinal area
cabohydrates, binds the epithelial membrane to the underlying
connective tissue. (a) Ray Simons/Science Source; (b) Ray Simons/Science Basement membrane
Source; (c) ©Ed Reschke
Connective tissue

and the internal environment of the body. Stratified epithe-
lia are specialized to provide protection. Simple epithelia, in
contrast, provide little protection; instead, they are specialized
for transport of substances between the internal and external
(b)
FIGURE 1.13 A stratified squamous nonkeratinized
epithelial membrane. This is a photomicrograph (a) and
illustration (b) of the epithelial lining of the vagina.
(a) ©Victor P. Eroschenko

ISTUDY
 14 Chapter 1
Keratinized
layer
Epidermis
Extracellular
material:
collagen fibers,
Dermis scattered cells,
tissue fluid
A blood
capillary
A lymph capillary,
which helps drain
off tissue fluid
FIGURE 1.14 The epidermis is a stratified, squamous,
keratinized epithelium. The upper cell layers are dead and
impregnated with the protein keratin, producing a cornified
epithelial membrane, which is supported by layers of living cells.
The epidermis is nourished by blood vessels located in the loose
connective tissue of the dermis.
with a water-resistant protein known as keratin. This protective
layer is constantly flaked off from the surface of the skin and
therefore must be constantly replaced by the division of cells in
the deeper layers of the epidermis.
The constant loss and renewal of cells is characteristic
of epithelial membranes. The entire epidermis is completely
replaced every two weeks; the stomach lining is renewed
every two to three days. Examination of the cells that are lost,
or “exfoliated,” from the outer layer of epithelium lining the
female reproductive tract is a common procedure in gynecol-
ogy (as in the Pap smear).
To form a strong membrane that is effective as a barrier
at the body surfaces, epithelial cells are very closely packed
and are joined together by intercellular junctions collectively
called junctional complexes (chapter 6; see fig. 6.22). There
is no room for blood vessels between adjacent epithelial cells.
The epithelium must therefore receive nourishment from the
tissue beneath, which has large intercellular spaces that can
accommodate blood vessels and nerves. This underlying tissue
is called connective tissue. Epithelial membranes are attached
to the underlying connective tissue by a layer of proteins and
polysaccharides known as the basement membrane. This
layer can be observed only under the microscope using special-
ized staining techniques.
Basement membranes are believed to induce a polarity
to the cells of epithelial membranes; that is, the top (apical)
ISTUDY
portion of epithelial cells has different structural and functional
components than the bottom (basal) portion. This is important
in many physiological processes. For example, substances are
transported in specific directions across simple epithelia (dis-
cussed in chapter 6; see fig. 6.21). In stratified epithelia, only
the basal (bottom) layer of cells is on the basement membrane,
and it is these cells that undergo mitosis to form new epithelial
cells to replace those lost from the top. Scientists recently dem-
onstrated that when these basal cells divide, one of the daugh-
ter cells is attached to the basement membrane (renewing the
basal cell population), while the other is not. The daughter cell
that is “unstuck” from the basement membrane differentiates
(becomes specialized) and migrates upward in the stratified
epithelium.
CLINICAL APPLICATION
Basement membranes consist primarily of the structural pro-
tein known as collagen (see fig. 1.17). The type of collagen in
basement membranes is a large protein assembled from six
different subunits. Alport syndrome is a genetic disorder of
the collagen subunits that, among other problems, results in
damage to the glomeruli (the filtering units) of the kidneys.
This is one of the most common causes of kidney failure.
In Goodpasture’s syndrome, the collagen in the basement
membranes of the glomeruli and the lungs is attacked by the
person’s own antibodies, leading to both kidney and lung
disease.
Exfoliative cytology is the collection and examination
of epithelial cells that are shed and collected by mechani-
cal scraping of the membranes, washing of the membranes,
or aspiration of body fluids containing the shed cells. Micro-
scopic examination of these desquamated (shed) cells, for
example in a Pap smear, may reveal a malignancy.
Exocrine Glands
Exocrine glands are derived from cells of epithelial mem-
branes. The secretions of these cells are passed to the outside
of the epithelial membranes (and hence to the surface of the
body) through ducts. This is in contrast to endocrine glands,
which lack ducts and which therefore secrete into capillaries
within the body (fig. 1.15). The structure of endocrine glands
will be described in chapter 11.
The secretory units of exocrine glands may be simple
tubes, or they may be modified to form clusters of units around
branched ducts (fig. 1.16). These clusters, or acini, are often
surrounded by tentacle-like extensions of myoepithelial cells
that contract and squeeze the secretions through the ducts. The
rate of secretion and the action of myoepithelial cells are sub-
ject to neural and endocrine regulation.
Examples of exocrine glands in the skin include the lacri-
mal (tear) glands, sebaceous glands (which secrete oily sebum
into hair follicles), and sweat glands. There are two types of
sweat glands. The more numerous, the eccrine (or merocrine)
 The Study of Body Function 15

Epithelium

Connective
tissue

Cells from
surface
Connecting epithelium Connecting
cells grow down cells
Epithelial into
persist to cord or disappear
form duct underlying
tubule tissue
Deepest If exocrine If endocrine
cells become gland forms gland forms
secretory
Capillary Deepest cells
remain to secrete
into capillaries
FIGURE 1.15 The formation of exocrine and endocrine glands from epithelial membranes. Note that exocrine glands retain
a duct that can carry their secretion to the surface of the epithelial membrane, whereas endocrine glands are ductless.

Exocrine glands are also prominent in the reproductive sys-

Duct
Secretory
portion
Simple Simple
tubular acinar Simple branched acinar
FIGURE 1.16 The structure of exocrine glands. Exocrine
glands may be simple invaginations of epithelial membranes, or
they may be more complex derivatives.
sweat glands, secrete a dilute salt solution that serves in ther-
moregulation (evaporation cools the skin). The apocrine sweat
glands, located in the axillae (underarms) and pubic region,
secrete a protein-rich fluid and become more active during
puberty. This provides nourishment for bacteria that produce
the characteristic odor of this type of sweat.
All of the glands that secrete into the digestive tract are also
exocrine. This is because the lumen of the digestive tract is a
part of the external environment, and secretions of these glands
go to the outside of the membrane that lines this tract. Mucous
glands are located throughout the length of the digestive tract.
Other relatively simple glands of the tract include salivary
glands, gastric glands, and simple tubular glands in the intestine.
The liver and pancreas are exocrine (as well as endocrine)
glands, derived embryologically from the digestive tract. The
exocrine secretion of the pancreas—pancreatic juice—contains
digestive enzymes and bicarbonate and is secreted into the
small intestine via the pancreatic duct. The liver produces and
secretes bile (an emulsifier of fat) into the small intestine via
the gallbladder and bile duct.
tem. The female reproductive tract contains numerous mucus-
secreting exocrine glands. The male accessory sex organs—the
prostate and seminal vesicles—are exocrine glands that con-
tribute to semen. The testes and ovaries (the gonads) are both
endocrine and exocrine glands. They are endocrine because
they secrete sex steroid hormones into the blood; they are exo-
crine because they release gametes (ova and sperm) into the
reproductive tracts.
Connective Tissue
Connective tissue is characterized by large amounts of extra-
cellular material between the different types of connective
tissue cells. The extracellular material, called the extracel-
lular matrix, varies in the four primary types of connective
tissues: (1) connective tissue proper; (2) cartilage; (3) bone;
and (4) blood. Blood is classified as a type of connective tis-
sue because about half its volume is an extracellular fluid, the
blood plasma (chapter 13, section 13.1).
Connective tissue proper, in which the extracellular
matrix consists of protein fibers and a proteinaceous, gel-like
ground substance, is divided into subtypes. In loose connec-
tive tissue, protein fibers composed of collagen are scattered
loosely in the ground substance (fig. 1.17), which provides
space for the presence of blood vessels, nerve fibers, and other
structures (see the dermis of the skin, shown in fig. 1.14, as
an example). Dense regular connective tissues are those in
which collagen fibers are oriented parallel to each other and
densely packed in the extracellular matrix, leaving little room
for cells and ground substance (fig. 1.18). Examples of dense
regular connective tissues include tendons (connecting bone
to bone) and ligaments (connecting bones together at joints).
Dense irregular connective tissues, forming tough capsules

ISTUDY
 16 Chapter 1

Extracellular matrix

Protein Ground
fibers (collagen) substance

Mesenchymal
cell

Elastic fibers (a)

Fibroblast Nucleus of
adipocyte

Collagen Fat
fibers globule

Reticular Cytoplasm
fibers
Cell
membrane
Macrophage (b)

FIGURE 1.19 Adipose tissue. Each adipocyte contains a

large, central globule of fat surrounded by the cytoplasm of the
adipocyte. (a) Photomicrograph and (b) illustration of adipose
tissue. (a) Al Telser/McGraw-Hill Education

Adipocyte (fat cell) Blood vessel

FIGURE 1.17 Loose connective tissue. This illustration
shows the cells and protein fibers characteristic of connective
tissue proper. The ground substance is the extracellular
background material, against which the different protein fibers
can be seen. The macrophage is a phagocytic connective tissue
cell, which can be derived from monocytes (a type of white
blood cell).
Collagen fibers
Fibroblast nucleus
FIGURE 1.18 Dense regular connective tissue. In this
photomicrograph, the collagen fibers in a tendon are packaged
densely into parallel groups. The ground substance is in the tiny
spaces between the collagen fibers. McGraw-Hill Education/Al
Telser, photographer
and sheaths around organs, contain densely packed collagen
bundles arranged in various orientations that resist forces
applied from different directions.
Adipose tissue is a specialized type of loose connective
tissue. In each adipose cell, or adipocyte, the cytoplasm is
stretched around a central globule of fat (fig. 1.19). The syn-
thesis and breakdown of fat are accomplished by enzymes
within the cytoplasm of the adipocytes.
Cartilage consists of cells, called chondrocytes, surrounded
by a semisolid ground substance that imparts elastic properties
to the tissue. Cartilage is a type of supportive and protective tis-
sue commonly called “gristle.” It forms the precursor to many
bones that develop in the fetus and persists at the articular (joint)
surfaces on the bones at all movable joints in adults.
Bone is produced as concentric layers, or lamellae, of cal-
cified material laid around blood vessels. The bone-forming
cells, or osteoblasts, surrounded by their calcified products,
become trapped within cavities called lacunae. The trapped
cells, which are now called osteocytes, remain alive because
they are nourished by “lifelines” of cytoplasm that extend from
the cells to the blood vessels in canaliculi (little canals). The
blood vessels lie within central canals, surrounded by concen-
tric rings of bone lamellae with their trapped osteocytes. These
units of bone structure are called osteons (fig. 1.20).
The dentin of a tooth (fig. 1.21) is similar in composition
to bone, but the cells that form this calcified tissue are located
in the pulp (composed of loose connective tissue). These cells
send cytoplasmic extensions, called dentinal tubules, into the
dentin. Dentin, like bone, is thus a living tissue that can be
remodeled in response to stresses. The cells that form the outer

ISTUDY
 The Study of Body Function 17

Enamel

Dentin

Pulp

Cementum

FIGURE 1.21 A cross section of a tooth showing pulp,

dentin, and enamel. The root of the tooth is covered by
cementum, a calcified connective tissue that helps anchor the
tooth in its bony socket. ©Southern Illinois University/Science Source
(a)

enamel of a tooth, by contrast, are lost as the tooth erupts.

Enamel is a highly calcified material, harder than bone or den-
tin, that cannot be regenerated; artificial “fillings” are therefore
required to patch holes in the enamel.

C H E C K P O I NT S
6a. List the four primary tissues and describe the
distinguishing features of each type.
6b. Compare and contrast the three types of muscle tissue.
6c. Describe the different types of epithelial membranes
and state their locations in the body.
7a. Explain why exocrine and endocrine glands are
considered epithelial tissues and distinguish between
(b)
these two types of glands.
7b. Describe the different types of connective tissues
and explain how they differ from one another in their
content of extracellular material.

Lamellae

1.4 ORGANS AND SYSTEMS

Central canal
Organs are composed of two or more primary tissues that serve
Osteocyte
within a the different functions of the organ. The skin is an organ that
lacuna has numerous functions provided by its constituent tissues.
Canaliculi
LE A R N I N G O UTCO M E S
(c)
After studying this section, you should be able to:
FIGURE 1.20 The structure of bone. (a) A diagram of a long 8. Use the skin as an example to describe how the
bone, (b) a photomicrograph showing osteons, and (c) a diagram
of osteons. Within each central canal, an artery (red), a vein (blue), different primary tissues compose organs.
and a nerve (yellow) is illustrated. (b) ©Ed Reschke 9. Identify the body fluid compartments.

ISTUDY
 18 Chapter 1
Hair Sebaceous
gland
Epidermis
(Epithelial
tissue)
Dermis
(Connective
tissue)
Hypodermis
Adipose tissue
Hair bulb
Motor nerve
(Nerve tissue)
FIGURE 1.22 A diagram of the skin. The skin is an organ that contains all four types of primary tissues.
An organ is a structure composed of at least two, and usually
all four, primary tissues. The largest organ in the body, in terms
of surface area, is the skin (fig. 1.22). In this section, the numer-
ous functions of the skin serve to illustrate how primary tissues
cooperate in the service of organ physiology.
An Example of an Organ: The Skin
The cornified epidermis protects the skin against water loss
and against invasion by disease-causing organisms. These
functions, as well as the function of body temperature regula-
tion (thermoregulation) through sweating and flushing, are
ways that the skin participates in maintaining homeostasis.
Invaginations of the epithelium into the underlying connec-
tive tissue dermis create the exocrine glands of the skin.
These include hair follicles (which produce the hair), sweat
glands, and sebaceous glands. The secretion of sweat glands
cools the body by evaporation and produces odors that, at
least in lower animals, serve as sexual attractants. Sebaceous
glands secrete oily sebum into hair follicles, which transport
the sebum to the surface of the skin. Sebum lubricates the
ISTUDY
Sweat
pore
Stratum corneum
Stratum granulosum
Stratum spinosum
Stratum basale
Arrector
pili muscle
(Muscle tissue)
Sweat gland
Arteriole
Venule
Sensory
nerve
cornified surface of the skin, helping to prevent it from dry-
ing and cracking.
The skin is nourished by blood vessels within the dermis.
In addition to blood vessels, the dermis contains wandering
white blood cells and other types of cells that protect against
invading disease-causing organisms. It also contains nerve
fibers and adipose (fat) cells; however, most of the adipose
cells are grouped together to form the hypodermis (a layer
beneath the dermis). Although adipose cells are a type of con-
nective tissue, masses of fat deposits throughout the body are
referred to as adipose tissue.
Sensory nerve endings within the dermis mediate the cuta-
neous sensations of touch, pressure, heat, cold, and pain. Motor
nerve fibers in the skin stimulate effector organs, resulting in, for
example, the secretions of exocrine glands and contractions of the
arrector pili muscles, which attach to hair follicles and surround-
ing connective tissue (producing goose bumps). The degree of
constriction or dilation of cutaneous blood vessels—and therefore
the rate of blood flow—is also regulated by motor nerve fibers.
The epidermis itself is a dynamic structure that can respond
to environmental stimuli. The rate of its cell division—and

consequently the thickness of the cornified layer—increases
under the stimulus of constant abrasion. This produces cal-
luses. The skin also protects itself against the dangers of ultra-
violet light by increasing its production of melanin pigment,
which absorbs ultraviolet light while producing a tan. In addi-
tion, the skin is an endocrine gland; it synthesizes and secretes
vitamin D (derived from cholesterol under the influence of
ultraviolet light), which functions as a hormone.
The architecture of most organs is similar to that of the
skin. Most are covered by an epithelium that lies immediately
over a connective tissue layer. The connective tissue con-
tains blood vessels, nerve endings, scattered cells for fighting
infection, and possibly glandular tissue as well. If the organ
is hollow—as with the digestive tract or blood vessels—the
lumen is also lined with an epithelium overlying a connective
tissue layer. The presence, type, and distribution of muscle tis-
sue and nerve tissue vary in different organs.
Stem Cells
The different tissues of an organ are composed of cells that
are highly specialized, or differentiated. The process of dif-
ferentiation begins during embryonic development, when the
fertilized ovum, or zygote, divides to produce three embry-
onic tissue layers, or germ layers: ectoderm, mesoderm, and
endoderm (chapter 20; see fig. 20.45a). During the course of
embryonic and fetal development, the three germ layers give
rise to the four primary tissues and their subtypes.
The zygote is totipotent—it can produce all of the differ-
ent specialized cells of the body. After a number of cell divi-
sions, when the embryo is at the stage where it implants into
the mother’s uterus, its cells are pluripotent—they can pro-
duce all of the body cells except those that contribute to the
placenta, and so have been called embryonic stem cells. As
development proceeds through successive cell divisions, the
cells become increasingly differentiated and lose the ability to
form unrelated cell types. However, genes are not lost during
differentiation. This was first demonstrated in the 1960s by
a British scientist, who found that he could produce a fully
formed frog by transplanting the nucleus from a differenti-
ated frog intestinal cell into the cytoplasm of an enucleated
ovum. In 2006 and later, Japanese scientists found that they
could transform differentiated fibroblasts into a pluripotent
state, similar to embryonic stem cells, by treating the fibro-
blasts with a few specific regulatory molecules. The 2012
Nobel Prize in Physiology or Medicine was awarded for these
discoveries.
Because the specialized cells have a limited life span,
many organs retain small populations of cells that are less dif-
ferentiated and more able to divide to become the specialized
cell types within the organ. These less-differentiated cells are
known as adult stem cells. In the red bone marrow, for exam-
ple, the stem cell population gives rise to all of the different
blood cells—red blood cells, white blood cells, and platelets
(chapter 13). Similarly, there are stem cells in the brain (chap-
ter 8), skeletal muscles (chapter 12), and intestine (chapter 18).
ISTUDY
The Study of Body Function 19
Hair
Epidermis
Basal layer
External root sheath
Sebaceous gland
Inner root sheath
Bulge region
with stem cells
Lower germinal matrix
Dermal papilla
FIGURE 1.23 The bulge region of the hair follicle
with stem cells. Stem cells in this region migrate to form the
differentiated cells of the hair follicle, sebaceous gland, and
epidermis.
Scientists have recently discovered that there are also stem
cells in the bulge region of the hair follicle (fig. 1.23). These
stem cells form keratinocytes, which migrate down to the lower
germinal matrix of the hair follicle and divide to form the hair
shaft and root sheath. Other stem cells in the region of the hair
follicle just above the bulge form new sebaceous gland cells,
which have a high turnover. Skin wounds stimulate the migra-
tion of stem cells from the hair follicles into the skin between
follicles to promote healing of the wounded skin.
The bulge region also contains melanocyte stem cells,
which migrate to the lower germinal matrix of the follicle and
give the hair its color. Scientists have now shown that gray-
ing of the hair with age is caused by loss of the melanocyte
stem cells in the bulge of the hair follicles. The melanocyte
stem cells appeared to be present in most of the hair follicles
of people aged 20 to 30 and absent from most hair follicles of
people aged 70 and older.
As demonstrated by the stem cells in the bulge of the hair fol-
licle, adult stem cells can form a variety of related cell types; the
adult stem cells are therefore described as multipotent. The topics
of embryonic and adult stem cells are discussed in more detail in
the context of embryonic development (chapter 20, section 20.6).
Systems
Organs that are located in different regions of the body and
that perform related functions are grouped into systems. These
include the integumentary system, nervous system, endocrine
system, skeletal system, muscular system, circulatory system,
 20 Chapter 1

TABLE 1.4 | Organ Systems of the Body The extracellular compartment is subdivided into two
parts. One part is the blood plasma, the fluid portion of the
System Major Organs Primary Functions
blood. The other is the fluid that bathes the cells within the
Integumentary Skin, hair, nails Protection, organs of the body. This is called interstitial fluid. In most parts
thermoregulation of the body, blood plasma and interstitial fluid communicate
Nervous Brain, spinal cord, Regulation of other freely through blood capillaries. The kidneys regulate the vol-
nerves body systems ume and composition of the blood plasma, and thus, indirectly,
Endocrine Hormone-secreting Secretion of the fluid volume and composition of the entire extracellular
glands, such as the regulatory compartment. The interactions between these fluid compart-
pituitary, thyroid, and molecules called ments and total body fluid regulation is explained in chapter 14
adrenal glands hormones and illustrated in figure 14.8.
Skeletal Bones, cartilages Movement and There is also selective communication between the intra-
support cellular and extracellular compartments through the movement
Muscular Skeletal muscles Movements of the of molecules and ions through the cell membrane, as described
skeleton in chapter 6. This is how cells obtain the molecules they need
for life and how they eliminate waste products.
Circulatory Heart, blood vessels, Movement of blood
lymphatic vessels and lymph
Immune Red bone marrow, Defense of the body C H E C K P O I NT S
lymphoid organs against invading 8a. State the location of each type of primary tissue in
pathogens
the skin.
Respiratory Lungs, airways Gas exchange 8b. Describe the functions of nervous, muscle, and
Urinary Kidneys, ureters, Regulation of blood connective tissue in the skin.
urethra volume and 8c. Describe the functions of the epidermis and explain
composition
why this tissue is called “dynamic.”
Alimentary Mouth, stomach, Breakdown of food 9. Distinguish between the intracellular and extracellular
intestine, liver, into molecules that
gallbladder, enter the body
compartments and explain their significance.
pancreas
Reproductive Gonads, external Continuation of the
genitalia, associated human species
glands and ducts CLINICAL INVESTIGATION SUMMARY

Homeostasis of body temperature and blood glucose is

immune system, respiratory system, urinary system, alimen-
tary (digestive) system, and reproductive system (table 1.4).
By means of numerous regulatory mechanisms, these systems
work together to maintain the life and health of the entire
organism.
Body-Fluid Compartments
Tissues, organs, and systems can all be divided into two major
parts, or compartments. The intracellular compartment is
that part inside the cells; the extracellular compartment is
that part outside the cells. Both compartments consist primar-
ily of water—they are said to be aqueous. About 65% of the
total body water is in the intracellular compartment, while
about 35% is in the extracellular compartment. The two com-
partments are separated by the plasma membrane surrounding
each cell (chapter 3, section 3.1).
maintained by physiological mechanisms that operate in
negative feedback loops. Health requires that the body
maintain homeostasis, and this can be assessed clinically
by specific measurements. If Linda’s fasting blood glucose
were above the normal range, it could indicate a disease
called diabetes mellitus. In this disease, the hormone insulin
does not lower the blood glucose concentration adequately
in response to a rise in blood glucose.
You can solve another medical mystery now by going
to the Connect site for this text. Look for Feedback Control
of Blood Glucose, which is based on this chapter’s material.
The more you play with physiological concepts, the better
you will understand them. The Clinical Investigation boxes
at the end of subsequent chapters will provide additional
Clinical Investigations on the Connect site that relate to
those chapters.

ISTUDY

SUMMARY
1.1 Introduction to Physiology 2 1.3 The Primary Tissues
A. Physiology is the study of how cells, tissues, and organs
function.
1. In the study of physiology, cause-and-effect sequences
are emphasized.
2. Knowledge of physiological mechanisms is deduced from
data obtained experimentally.
B. The science of physiology shares knowledge with the related
sciences of pathophysiology and comparative physiology.
1. Pathophysiology is concerned with the functions of
diseased or injured body systems and is based on
knowledge of how normal systems function, which is the
focus of physiology.
2. Comparative physiology is concerned with the
physiology of animals other than humans and shares
much information with human physiology.
C. All of the information in this book has been gained by
applications of the scientific method. This method has three
essential characteristics:
1. It is assumed that the subject under study can ultimately
be explained in terms we can understand.
2. Descriptions and explanations are honestly based on
observations of the natural world and can be changed as
warranted by new observations.
3. Humility is an important characteristic of the scientific
method; the scientist must be willing to change his or her
theories when warranted by the weight of the evidence.
1.2 Homeostasis and Feedback Control 4
A. Homeostasis refers to the dynamic constancy of the internal
environment.
1.4 Organs and Systems
1. Homeostasis is maintained by mechanisms that act
through negative feedback loops.
a. A negative feedback loop requires (1) a sensor
that can detect a change in the internal environment
and (2) an effector that can be activated by the sensor.
b. In a negative feedback loop, the effector acts to cause
changes in the internal environment that compensate
for the initial deviations that were detected by the
sensor.
2. Positive feedback loops serve to amplify changes and
may be part of the action of an overall negative feedback
mechanism.
3. The nervous and endocrine systems provide extrinsic
regulation of other body systems and act to maintain
homeostasis.
4. The secretion of hormones is stimulated by specific
chemicals and is inhibited by negative feedback
mechanisms.
B. Effectors act antagonistically to defend the set point against
deviations in any direction.
ISTUDY
The Study of Body Function 21
10
A. The body is composed of four types of primary tissues: mus-
cle, nerve, epithelial, and connective tissues.
1. There are three types of muscle tissue: skeletal, cardiac,
and smooth muscle.
a. Skeletal and cardiac muscle are striated muscle tissue.
b. Smooth muscle is found in the walls of the internal
organs.
2. Nerve tissue is composed of neurons and neuroglia.
a. Neurons are specialized for the generation and
conduction of electrical impulses.
b. Neuroglia provide the neurons with anatomical and
functional support.
3. Epithelial tissue includes membranes and glands.
a. Epithelial membranes cover and line the body
surfaces, and their cells are tightly joined by
intercellular junctions.
b. Epithelial membranes may be simple or stratified, and
their cells may be squamous, cuboidal, or columnar.
c. Exocrine glands, which secrete into ducts, and
endocrine glands, which lack ducts and secrete
hormones into the blood, are derived from epithelial
membranes.
4. Connective tissue is characterized by large intercellular
spaces that contain extracellular material.
a. Connective tissue proper is categorized into subtypes,
including loose, dense fibrous, adipose, and others.
b. Cartilage, bone, and blood are classified as connective
tissues because their cells are widely spaced with
abundant extracellular material between them.
17
A. Organs are units of structure and function that are composed
of at least two, and usually all four, of the primary types of
tissues.
1. The skin is a good example of an organ.
a. The epidermis is a stratified squamous keratinized
epithelium that protects underlying structures and
produces vitamin D.
b. The dermis is an example of loose connective tissue.
c. Hair follicles, sweat glands, and sebaceous glands
are exocrine glands located within the dermis.
d. Sensory and motor nerve fibers enter the spaces
within the dermis to innervate sensory organs and
smooth muscles.
e. The arrector pili muscles that attach to the hair
follicles are composed of smooth muscle.
2. Organs that are located in different regions of the body
and that perform related functions are grouped into
systems. These include, among others, the circulatory
system, alimentary system, and endocrine system.
 22 Chapter 1

3. Many organs contain adult stem cells, which are able to
differentiate into a number of related cell types.
a. Because of their limited flexibility, adult stem cells
are described as multipotent, rather than as totipotent
or pluripotent.
b. For example, the bulge region of a hair follicle
contains stem cells that can become keratinocytes,
epithelial cells, and melanocytes; the loss of the
melanocyte stem cells causes graying of the hair.
B. The fluids of the body are divided into two major
compartments.
1. The intracellular compartment refers to the fluid within
cells.
2. The extracellular compartment refers to the fluid outside
of cells; extracellular fluid is subdivided into plasma (the
fluid portion of the blood) and interstitial fluid.

REVIEW ACTIVITIES
Test your knowledge 9. A hormone called parathyroid hormone acts to help raise the
1. Glands are derived from blood calcium concentration. According to the principles
of negative feedback, an effective stimulus for parathyroid
a. nerve tissue. c. muscle tissue. hormone secretion would be
b. connective tissue. d. epithelial tissue. a. a fall in blood calcium.
2. Cells joined tightly together are characteristic of b. a rise in blood calcium.
a. nerve tissue. c. muscle tissue. c. either a fall or a rise in blood Ca2+ at different times.
b. connective tissue. d. epithelial tissue. d. unrelated to the blood Ca2+ levels.
3. Cells are separated by large extracellular spaces in 10. Which of these consists of dense parallel arrangements of
a. nerve tissue. c. muscle tissue. collagen fibers?
b. connective tissue. d. epithelial tissue. a. skeletal muscle tissue c. tendons
4. Blood vessels and nerves are usually located within b. nerve tissue d. dermis of the skin
a. nerve tissue. c. connective tissue. 11. The act of breathing raises the blood oxygen level,
b. muscle tissue. d. epithelial tissue. lowers the blood carbon dioxide concentration, and
5. Most organs are composed of raises the blood pH. According to the principles of
negative feedback, sensors that regulate breathing should
a. epithelial tissue. c. connective tissue. respond to
b. muscle tissue. d. all of these. a. a rise in blood oxygen.
6. Sweat is secreted by exocrine glands. This means that b. a rise in blood pH.
a. it is produced by endocrine cells. c. a rise in blood carbon dioxide concentration.
b. it is a hormone. d. all of these.
c. it is secreted into a duct. 12. Adult stem cells, such as those in the bone marrow, brain,
d. it is produced outside the body. or hair follicles, can best be described as       ,
7. Which of these statements about homeostasis is true? whereas embryonic stem cells are described as
      .
a. The internal environment is maintained absolutely constant.
a. totipotent; pluripotent
b. Negative feedback mechanisms act to correct deviations
from a normal range within the internal environment. b. pluripotent; multipotent
c. Homeostasis is maintained by turning effectors on c. multipotent; pluripotent
and off. d. totipotent; multipotent
d. All of these are true.
8. In a negative feedback loop, the effector produces changes Test Your Understanding
that are
13. Describe the structure of the various epithelial
a. in the same direction as the change produced by the initial membranes and explain how their structures relate to
stimulus. their functions.
b. opposite in direction to the change produced by the initial 14. Compare bone, blood, and the dermis of the skin in terms of
stimulus. their similarities. What are the major structural differences
c. in the same and opposite direction as the initial stimulus between these tissues?
at different times. 15. Describe the role of antagonistic negative feedback
d. unrelated to the initial stimulus. processes in the maintenance of homeostasis.

ISTUDY
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as it passes over the lower seventeen pipes through which it
originally entered as cold weak brine.
Direct Expansion.—Many plants are equipped completely with
direct expansion systems. This is quite satisfactory provided means
are arranged to dispose of the accumulating ice and snow on the
coils. This is particularly applicable to coils in pipe lofts where
moisture rapidly accumulates, and is accomplished by some
arrangement of defrosting such as circulating a brine over coils by
permitting it to drip over the coils, pumping over and over again.
Two-Stage Compressors.—The revising editor brought into
prominence the two-stage compression system by building a
successfully operating plant which functioned with quite astounding
results. The system consists of a large low temperature gas
compressor and a smaller second stage compressor, arranged the
reverse to a compound steam engine.
Ammonia gas which at the low temperatures resulting when low
back-pressures are required, becomes highly attenuated (light), it is
necessary to handle a very large volume per ton of refrigeration
developed. To provide for this, the low pressure cylinder is made
about double the volume of the high pressure cylinder.
In the low pressure cylinder the gas is compressed to a pre-
determined pressure and passed at a relatively high pressure,
making for a very high efficiency. The gas is chilled by the
introduction of expanded ammonia to take up the superheat.
The use of this compressor arrangement with its refinement
makes an economical cold producing unit. A record of one year’s run
follows:

Two-Stage Compression System.

Per Per
year. ton.
Tons received 60,000
Cu. ft. space cooled 2,930,000
Coal cost $17,309.11 .288
Labor 15,156.73 .252
Oil and waste 1,030.82 .017
Water and chemicals 2,103.21 .035
 Repairs and changes 1,698.38 .028
Total cost $37,298.25 .62
Expense per cu. ft. space per year .013

Ample Capacity.—All of the above factors must be considered in

the calculations for refrigerating requirements,
and it is best to make allowances for a considerable factor of safety
over and above the actual maximum, as well as for the economical
operation of the plant. It never pays to crowd a plant to its limit or
capacity. Guard against emergencies and possible abnormal
demand for refrigeration by providing ample equipment. The plant
should be constructed as far as possible in duplicate, not only as
regards the machinery but also in the apparatus, as a safeguard
against accidents and total cessation of refrigeration. With two
machines and duplicate apparatus one-half the maximum
refrigeration is always available, and the preservation of the product
in storage is assured, even if it be found necessary to stop killing for
a sufficient period to make the repairs on the broken machinery or
apparatus, while with one machine only available, a breakdown
might result in a very serious loss and considerable damage to the
stored products.
With regard to insulation, it may be said that the best is the most
economical in the long run. There is no such thing as absolute
insulation, some heat leakage must occur through the very best
insulation, and the reduction of this heat leakage to the minimum
should be the chief object or factor for consideration. First-class
insulation costs more in original investment, but it creates a
continuous saving and economy in refrigeration, resulting oftentimes
in a less investment also in refrigerating, pumping and steam
equipment.
 CHAPTER V.
REFRIGERATION REQUIREMENTS.
Refrigeration — Necessities — Ice Plants — Natural Ice — Chilling —
Quantity Refrigeration to Provide — Space Per Ton Machine —
Machinery Per Head — Ice Computation — Compressor Capacity —
Summary Computations — Low Temperature Brine System.

Refrigeration.—This word covers the “bugbear” and the “solace”

of the packers. No other one thing is of so much importance. Since
the abolishment of the use of chemicals, such as borax, boracic acid,
etc., as a preventive or retardant in the propagation of bacilli during
the curing process, the packer’s sole dependence is refrigeration.
Consequently there has been considerable development and
importance in its use.
From the moment the animal is killed until the parts are disposed
of, refrigeration is a necessity. The proportion of freezing space to be
provided or hired at a commercial storage house is growing daily. In
many plants the total volume of freezer space approaches or
exceeds, that given over for moderate temperature storage
purposes.
Necessities.—In the order of progress the first necessity for
refrigeration is the use of ice for chilling water for fats and meats.
Practically every packing house is now equipped with a small ice
making plant both for its use, and with commercial possibilities of
selling ice.
Ice Plants.—Ice plants are of various kinds:
(1) Common ice made from filtered water without processing.
(2) Raw water ice made from filtered or treated water, aerated or agitated,
while freezing.
 (3) Distilled water ice made from condensed steam, reboiled.
(4) Plate ice, made from raw water in large plates and cut up into blocks of
convenient size.

In the production of distilled water there are various methods

which will not be treated upon here.
For the icing of cars and for ordinary cooling purposes ice made
by process No. 1 is amply good except, perhaps, if the water from
which the ice is made should not be clear, it may be necessary to
use a portion of the bottom of the cake for car icing purposes only.
Natural Ice.—The situation and climate has much to do with the
determination as to whether the packer can afford to harvest natural
ice for general use. For example, if the plant be situated in a
northern climate alongside a suitable stream adjacent to the plant,
the harvesting of ice and its storage can be afforded. However, if the
ice is stored at a remote point and teaming or car shipment of ice is
necessary, then the manufacturing plant will probably be the more
economical.
Chilling.—The chilling of the meat immediately after killing is
perhaps the most important function in the whole refrigerating
process and one that cannot have too much attention. In the matter
of beef it is the practice of the best operators to see to it that
immediately after the hide is removed the beef is placed in the
coolers. At most it should not be allowed to stand out in the open air
to exceed half an hour. Sheep and veal should be treated in the
same manner. As for hogs in some climates, such as latitude equal
to Chicago or north thereof, if space is available an open air hanging
room is a valuable adjunct where the animal can be air dried,
depending upon seasons, from one hour to over night. This feature
also permits of increased slaughtering during the winter season. It is
a custom that can be abused, however, by careless handling—
resulting in sour meats.
Quantity of Refrigeration to Provide.—There are so many
factors which enter into the question of packing house refrigeration,
and so many details of construction and application of refrigerating
apparatus, that it is almost impossible to cover this subject practically
and intelligibly without devoting a volume to refrigeration alone.
 To give a fixed rule for packing house refrigeration, applicable to
all conditions, would be impossible, as much depends upon
conditions and surroundings, and the requirements would vary
accordingly. For instance, atmospheric conditions, quantity of space
to be cooled, temperature to be maintained, time for chilling, etc.,
these and many other factors affecting the general results must be
taken into consideration and provided for, and as these vary, so will
the refrigeration requirements vary. A few ideas will be set out to
assist in the computation of the quantity of equipment to provide.
The capacity or size of the refrigerating equipment required for any
given plant is variable but can be reasonably closely computed by
assuming certain conditions.
Space Per Ton Machine.—Practice has demonstrated that one
ton of mechanical refrigerating effect duty, as rated by manufacturer
used continuously during a period of twenty-four hours will maintain
at a temperature of 33° to 38° F. from 7,000 to 12,000 cubic feet of
storage space. Conditions of exposure, insulation and outside
temperature affect this.
The minimum for beef storage coolers, the maximum for curing
cellars. The area of room or quantity of space enters into each of
these assumptions. If small rooms are used the unit to be assumed
is smaller. Concrete buildings are easier to control due to the stored
cold in the building mass of floors, columns and girders.
Machinery Per Head.—It is customary to figure that in addition to
that needed for cooling the space, one ton of refrigeration for twenty-
four hours would be required for either one of the following items:
From fifteen to twenty-two hogs, average weight, 225 pounds.
From five to six head of cattle, average weight, 700 pounds.
About forty-five calves, average weight, 80 pounds.
From fifty to sixty-five sheep, average weight, 60 pounds.

For Cooled Meats.—For cold storage rooms, where meats which

have already been chilled or cooled are stored and held for a greater
or lesser period, the refrigeration requirements are not so great as in
chill and cooling rooms. The animal heat has been removed and the
meats cooled down to a low temperature, consequently but little
more refrigeration is required than that necessary to take care of the
heat leakage through the insulation, and possibly the recooling of the
meat through a range of a few degrees, when the meats may have
gained a little in temperature by exposure. Therefore it is estimated
that one ton of refrigerating duty will handle 12,000 cubic feet of
curing or storage space for temperature of 33° to 35° F.
Freezer Space.—As will be explained in the chapter relating to
refrigerating machine capacity, the capacity of a compressor very
rapidly decreases when operating under conditions necessary to
produce low freezer temperatures, and owing to the severe
conditions imposed it seems necessary to compute that one ton of
refrigerating duty will handle about 3,000 feet of freezer space.
Ice Computations.—Before the application of mechanical
refrigeration to packing house purposes, all artificial refrigeration was
accomplished by means of ice melting alone, and at that time the
packers computed ice melting requirements on a basis of cooling
three pounds of meat from 80° F. to as low as it could be cooled by
ice melting, for each pound of ice melted. While this rule undoubtedly
was the result of practical experience with well constructed coolers,
and was in no wise based upon theoretical or heat unit formulas, yet
it is interesting to note how close this old rule compares with modern
formulas of computing refrigeration. For example, the cooling of 100
head of hogs, averaging 250 pounds dressed, by the packer’s rule,
would require—
100 × 250
= 8,333 pounds, or 4.16 tons ice melting.
3
And on a heat unit basis, cooling the same number and weight of
hogs from 80° to 32° F. would require—
100 × 250 × (80 - 32)
= 4.166 tons refrigeration.
288,000
The above comparison shows that theory and practice approach
very closely to a common line, although in the calculation the factor
of specific heat of the meat is ignored, and in practice this may well
be left out, as specific heat of meats at various temperatures has by
no means been accurately established.
Chilling Lard.—The chilling of lard is a severe task on
refrigerating equipment owing to the very heavy duty imposed in a
short time. For example, with an eight-foot lard roll turning off 4,000
pounds per hour and changing the temperatures of the lard from 90°
to 50° F. requires a very large compressor capacity per hour.
Summary—Computations.—Assuming the killing of 100 cattle
and 250 hogs per day; making 10,000 pounds of lard, fifteen tons
ice; maintaining a freezer of 100,000 cubic feet and the small work
connected with the establishment:
For 100 cattle killed daily, and storage of 400 hanging, about 6,500 square feet
area of cooler would be required; the height of this with the lofts would be about
twenty-two feet or a capacity of 143,000 cubic feet.
For 250 hogs daily or 750 total hanging, there would be required 3,000 square
feet of floor area and a height of 18 feet or 54,000 cu. ft.
The above number of hogs daily would also require a storage or cellar capacity
based upon one hundred pounds meat sent to cellar for a turn over in sixty days
equal to storing 1,500,000 pounds of product, or 15,000 to 20,000 square feet of
floor area, requiring 200,000 cubic feet of building space.
Ten thousand pounds lard daily in four hours running would require the
extraction of about two tons of duty, but by reason of the heat transmission and the
low temperature brine necessary would require twelve tons duty while applied.
Fifteen tons daily ice making capacity requires about 30 tons refrigerating duty,
allowing for radiation and other losses.
A freezer capacity of 100,000 cubic feet which would store 1,500,000 pounds
would not appear excessive. From known results this would require about one ton
refrigerating duty for 3,000 cubic feet of space or thirty-five tons machine duty. We
have then:

SUMMARY OF REFRIGERATION REQUIRED

Tons
143,000 cubic feet beef cooler space ÷ 10,000 14.3
100 cattle daily 500 pounds = 50,000 pounds
50,000 pounds meat chilled 70° or 70 B.t.u. per pound
50,000 × 70 = 3,500,000 ÷ 288,000 B.t.u. 12.2
54,000 cubic feet hog cooler space ÷ 10,000 5.4
250 hogs 180 pounds each = 45,000 pounds
 45,000 × 70 ÷ 288,000 = 10.9
200,000 feet cellar space ÷ 12,000 16.6
15 tons ice × 2 tons refrigerating duty 30.0
10,000 pounds lard cooler 12.0
100,000 cubic feet freezer space 35.0
Total tonnage refrigerating effect 136.4

Low Temperature Brine System.—The use of low temperature

brine, cooled by two stage compressors, a new development, is in
the writer’s opinion preferable to direct expansion. (See description
of two stage compression in Chapter IV.) A lineal foot of pipe filled
with a liquid like chilled brine seems to have greater heat absorbing
power than the same pipe filled with a light gas and consequently
less piping can be used.
Preference Reasons.—Occasion arose to compare for a new large
installation the relative merits of direct expansion versus brine and
the following items suggested themselves:
(1) The direct expansion plant would require 150,000 lbs.
ammonia in excess of brine plant.
(2) There would be approximately 8,000 more joints to prevent
leaking.
(3) Unavoidable ammonia leakage through rods, glands, valve
stems, compressor rods and purging would amount to at least
$20,000 per year based on experience with good practice.
(4) The installation by the same manufacturer estimated to cost
15% more than a brine system.
(5) There would be no stored energy in the ammonia pipes as in
brine pipes should the refrigerating plant suspend operation
temporarily.
(6) The compressor hazard from returning liquid and possible
machine wrecking with line breaks, ammonia losses, and danger to
life.
(7) In case of extension or modification of piping, the difficulty with
ammonia pipes over brine is marked.
(8) In direct expansion plants the cooling effect is stopped
immediately when the compressor is stopped. It is required to
operate incessantly, or if speed be reduced the current is chiefly
wasted in resistance grids if electrically driven. Whereas in a brine
plant the compressors can be stopped several hours, the brine being
circulated rises in temperature but not sufficiently to vary room
temperatures. This is a great advantage, permitting keeping “off
peaks” on commercial electric lines and consequently earning lower
rates.
(9) With an electrically driven compressor, speed control is fixed
within quite limited bounds and if direct expansion system is used
more power is purchased than needed for reasons already
mentioned.
(10) The total cost for operating pumps and calcium taken
collectively will be far less than the uncontrollable ammonia leakage.
The coil attendants can be less skilled.
(11) Brine temperature 25 degrees below zero, Fahr., can be
carried, sufficiently low for all purposes. Rooms can not be controlled
at as exact temperatures with direct expansion as with brine, owing
to its sensitiveness.
(12) Lastly and very important, the complicated piping system to
carry several back pressures to produce varying temperatures and
the necessity of operating several compressor “ends” makes a
complicated installation of small units as against a simple system
carrying one temperature brine.
Plants of the character described usually operate on direct
expansion, require three sets of temperatures, three sets of suction
pressures and three sets of suction mains with various cross
connections and intricate pump-out arrangements. This necessitates
multiplicity of machines and connections to various cylinder ends.
Progressive authorities recognize the necessity of freezer storage
in meat plants. In combination pork and beef or pork plants, newer
installations will be made with far less curing space for sweet pickled
meats and far greater space for storing meats frozen either before or
after curing to the end of producing palatable meats, not salt soaked
and of inferior quality. There are slaughtering plants for export where
beef is shipped chilled or frozen, and mutton frozen. Some have
exclusive freezer buildings, others parts set aside for freezers.
 One temperature brine is circulated, the temperature in the rooms
being governed by the quantity of pipe in service. In all instances,
sharp freezers, storage rooms and curing rooms are equipped with
pipes therein—and the chilling rooms for beef and pork closed coils
or spray, optional.
Shell and tube coolers have proven so fitting that no hesitancy
need be made about using them, and the possibility of freezing is
negligible with reasonable prudence. Tanks with submerged coils
can be used, but it prevents the saving made by using balanced
brine pumping, on high buildings, a quite material one.
 CHAPTER VI
POWER PLANT REQUIREMENT.
Steam Making Equipment — Uses of Pumps — Types of Pumps — Pumps
for Brine — Water Supply — Pumps for Wells — Water Distribution
Systems — Pumps for Boiler Feeding — Heating Boiler Feed Water —
Wells — Cooling Towers — Economizers — Superheaters — Steam
Engines — Exhaust Steam — Advice as to Exhaust Steam — Boilers —
Conclusion.

Steam Making Equipment.—The quantity and type of boilers

needed for a plant is entirely dependent upon the quantity of steam
required and its economical or uneconomical use. The refrigerating
and electric producing apparatus in the average packing house are
usually looked upon as the chief using elements, but they each
frequently require less steam than is used for pumping purposes,
and less than is used for cooking and kindred operations. It would be
well to discuss the uses of steam and return to the boilers.
Uses of Pumps.—Pumps are used for water circulating, brine
circulating, boiler feeding and pumping products. These are named
in the order of their steam consuming demand as it usually occurs.
They consume from 40 to 120 pounds of steam per horse power
hour based on the kind of pump used and the skill in its operation.
The most economical are electrically operated centrifugal pumps
provided the current is economically generated, steam turbine
centrifugal pumps, next if in good condition, and reciprocating
simplex or duplex pumps third and fourth.
Types of Pumps.—The types chiefly used in packing house work
are simplex and duplex reciprocating, centrifugal single and multi-
stage operated by motor or steam turbine directly attached to the
motive power, also power pumps of triplex type and rotary or impeller
pumps for fats and oils. There are endless varieties of each type.
 Pumps for Brine.—For brine pumping chiefly reciprocating and
direct connected horizontally set motor driven
centrifugal pumps of one or more stages are used. At times power
operated triplex pumps are used but the most satisfactory are the
second named.
Pumping brine for refrigerating purposes is a never ending
process since it must be constantly performed. The common practice
of an open brine tank with submerged coils is being fast superseded
by the use of shell and tube type closed coolers which superseded
double pipe coolers. The use of either the latter two types enables
the operator to take advantage of hydraulic head by installing a
balanced system described in this work which very greatly reduces
the power necessary to exert to pump brine. The friction in an eight
to twelve pass brine cooler is from ten to fifteen pounds, and this
with the pipe friction on a balanced system will total to perhaps thirty
pounds on large systems, but less than the ordinary open system—
depending upon the building height.
Water Supply.—The water demand for packing house operation is
quite an important element in relation to steam consumption. The
quantity required for ammonia condensing, steam condensing (if
done), and for plant purposes, washing, etc., is, in the aggregate,
quite a large amount. For example: For ammonia condensing, from
1¹⁄₄ to 1¹⁄₂ gallons per minute is required per ton of refrigeration
developed; for steam condensing for steam engine from 32 to 40
pounds per pound of steam condensed; for boiler operation about
four gallons per horse power per hour; for plant purposes from 250
to 3,000 gallons or more per hour, depending upon the size of plant
and its intelligent use. It will be readily seen that those quantities will
mount in power consumption if the head or pressure pumped against
is high.
Pumps for Wells.—Water pumped from wells is usually pumped
by one of four methods:
(1) Suction type reciprocating pumps.
(2) Deep well heads with low set pumping barrel.
(3) Submerged centrifugal pumps.
(4) Compressed air.
 For small plants where one or two wells will supply the water, the
first named pump is usually used owing to cheapness of cost for
installing and particularly if the water rises in the well to a height
reachable by a suction type pump, viz: 23 to 25 feet. Power operated
suction pumps are also available for this service. If the water be too
low for the above named type to handle, deep well pumps with
steam head serve pretty well and usually cost less than air pumped
wells.
For larger plants, larger wells are installed and submerged
centrifugal pumps or compressed air is used, the latter always as a
last resort from a cost of pumping standpoint, although its simplicity
and reliability appeals to many.
Submerged centrifugal pumps can be operated with steam turbine
or motor set on top of shaft above ground. The motor operated set is
the more economical, particularly if the current be generated at low
cost.
Water Distribution Systems.—The intelligent laying out of water
systems and the pressure pumped against is a factor worth great
attention. It is far better to have pumps of proper proportions for the
service intended, even though it involves the use of a separated
piping system.
Taking advantage of water once raised to a high level in its return
to service is a power saving. For example: It is usual to locate
ammonia condensers at a point where there is a good circulation of
air. This is done to permit the winds to carry away the moist air from
about the condensers, together with the heat delivered up through it
and to the air by the discharged gas from the ammonia compressors.
From the catch pan beneath the ammonia condenser the water can
be diverted through the steam condenser and if the shell be bought
sufficiently strong in the case of a closed condenser, the water can
be delivered to the suction side of a pump which in turn will deliver it
for house use for cleansing purposes. If a barometric condenser is
used, the water is delivered to a hot well from whence it can be
pumped or flow to sewer.
Pumps For Boiler Feeding.—Dependent upon the size of plant
and the pressure carried pumps are usually provided as follows:
 Small boiler plant up to 500 H. P. low pressures. Steam 80 to 125 lbs.
simplex or duplex piston packed reciprocating pumps.
Medium Boiler Plant up to 1,000 to 1,500 H. P. steam 125 to 150 pounds
simplex or duplex outside piston packed reciprocating pumps.
Over 1,500 H. P. the same or centrifugal type multi-stage pump motor or
steam driven.

Heating Boiler Feed Water.—Every plant, large or small, should

be equipped with boiler water heating devices. These are so
common and the practice so usual that it is scarcely necessary to
mention it but the writer has seen plants where the exhaust steam
from pumps and engines was being thrown to the air and feed water
pumped into boilers at low temperatures. There are two types:
closed and open. In the first named the water is circulated through
tubes enclosed in a vessel, the tubes surrounded by steam. In the
open type the steam is brought in direct contact with the water and
performs the heating in this manner. The closed types are used on
the principle that no oil carried in the steam is passed to the boilers,
but the latter type are usually equipped with an oil separator that
eliminates the oil and prevents danger from this source.
Wells.—In most instances wells of various depths from 80 to
2,000 feet will find water. A few localities, however, do not yield water
via wells and this is a poor situation from a packing house
standpoint. The water from surface or shallow wells is, at times,
improper for use on meats owing to contamination, in which case,
resort must be made to other sources. In some localities the
neighboring streams yield suitable waters but in this case sewage
contamination must be guarded against. The quality of boiler supply
water is an important factor in the boiler operation in its bearing to
shutdowns for cleaning, blowing down, repairs, etc., all of which
contribute to or impair economy.
Cooling Towers.—Where it is necessary to husband water for
condensing ammonia or steam the use of cooling towers comes into
play. These function by the cooling of water by evaporation, the
evaporation abstracting the heat. There are various kinds—sprays
over ponds, water dripping over spatter boards made in various
ways, such as brush piled loosely; a checker work of wood slats;
screen wire, etc. These usually depend upon the circulation of air in
a natural way. There are various patented types, depending upon
fans, to pass air through falling water. The efficiency is dependent
upon the fineness of the water and the quantity of air passing
through it, the relative humidity of the air controlling greatly the
cooling effect.
Economizers.—Where the utmost economy of fuel is striven for
and on larger plants, these are installed. An economizer is a bank of
cast iron flues placed in a position between the boilers and the
smoke stack. Water is circulated through the tubes and the furnace
gases pass about them. They are usually calculated upon to
decrease the stack temperatures from 100 to 150 degrees—
dependent upon its initial temperature and to heat the feed water
one hundred degrees. This is a saving from a wasted source, since it
is impossible to take up this heat in any boiler yet designed.
Superheaters.—In an earlier chapter reference was made to
superheated steam. The office of this apparatus is to heat the steam
to a higher point than it rises to, under ordinary boiler conditions. For
instance, steam as delivered from the nozzle of a boiler is spoken of
as “saturated” steam, but, if before use this is subjected to a further
heating before it is used, it is said to be “superheated.”
Superheaters are of two types. They can be described as follows:
A bank of coils located in the flame pass in the boilers, the steam
within the coils and the flame on the outside; or a bank of coils in a
separate furnace, the flame or heat generated in the furnace passing
over the coils.
Steam superheated beyond 450° F. total sensible temperature
requires especial pipe fittings, type of valves, and engines for
reasonable safe operation, and renders an economy that a small
plant cannot afford, due to the expenditure, to take advantage of.
Steam Engines.—Owing to the cheapness of fuel in the United
States, users have been satisfied with equipment at a low initial cost
and in consequence pay bills continually by using uneconomical
prime movers, viz.: ordinary steam engines of the simple Corliss
type, using from 26 lbs. to 35 lbs. of steam per horsepower hour, or
compound condensing at 15 lbs. to 18 lbs. of steam per horsepower
hour or high speed direct connected slide valve engine generator
sets using from 35 to 50 lbs. steam per horsepower, depending upon
load.
European engineers, where fuel costs are double and triple
American fuel costs, have of necessity paid more attention to the
subject of steam economy and have outstripped this country in
design and use of economical steam engines, notably of the so-
called “uniflow,” “drop valve,” “poppet valve,” or “Lentz” type and with
the use of superheated steam, have reached much higher
efficiencies. The fast increasing costs of fuel in this country have
necessitated American builders resorting to high steam pressures
with independent or integral superheaters in conjunction with steam
boilers, and as a result American manufacturers are offering simple
uniflow engines for high speed (say 200 r. p. m.) direct connection to
electric generators. Also compound drop valve or poppet valve or
Lentz types for slowly moving engines, viz.: 60 r. p. m. for connecting
to ammonia compressors and in either case with steam at an initial
pressure of 160 lbs. and ordinary superheat of 100° operating
condensing, engines of the above types under the conditions named
can be readily bought under guarantees to produce an indicated
horsepower with the use of 12 lbs. of steam per horsepower hour.
The uniflow engine is now offered by several builders and is
adaptable to most any service, atmospheric exhaust, condensing or
against a back pressure using the exhaust for cooking or heating.
There is but little advantage, however, in a uniflow engine under this
circumstance unless it be the flexibility as to conditions of operation.
Exhaust Steam.—The practice of using exhaust steam, that is
steam which has passed through engines, for heating and cooking is
considered economical. It is based upon the theory that the quantity
of heat contained in a pound of steam that has been reduced in
pressure from its initial or high pressure, say 100, to a low one—
example 5 lbs., and then used for heating, etc., contains almost as
many heat units as it did originally.
There are many places where this low pressure or exhaust steam
can be used advantageously, but if extensively used, the likelihood
will be to have long mains into which certain equipment will pass its
exhaust and from which the demand will be spasmodic for cooking
purposes and intermittent for heating, except during winter seasons,
and as a result there is a considerable quantity of steam passed to
the air at most times.
A careful analysis is necessary for intelligent use of this idea, since
the maintaining of a balance owing to business fluctuations and
changing seasons makes an almost unmatchable combination.
During one season live steam will be used for make up, and at other
times there is a surplus.
Advice As To Exhaust Steam.—Aside from heating feed water
and drying where the requirement is constant, a better practice is
probably to limit the extent of exhaust steam systems and use live
steam for most purposes.
The sources of exhaust steam, engines or pumps, are operating
usually twenty-four hours daily and constantly pouring steam into the
exhaust mains, which is constantly radiating, leaking and the
accumulated moisture is being trapped away to a sewer or feed
water heater. The traps are usually just a little out of order, and the
little leaks in multiple mount up fast.
Again the apparatus in almost any department used for cooking
and manufacturing is irregular in its use and an actual record will
show hardly to exceed six hours’ demand or use. Why operate
engines and pumps at a disadvantage all the time for the sake of a
supply six or eight hours daily? The engine and pump requirement
increases rapidly with the exhaust steam pressure and requires
larger production from the boiler.
Any power like that needed for pumping can be transmitted on a
wire with far less loss than as steam through a pipe.
The recommendations then are: Produce electricity as cheaply as
possible; if electricity is made, use high class equipment operating
condensing and take advantage of superheat if possible; produce
refrigeration or compress air with the same intelligence; perform all
pumping with electrically operated pumps; distribute steam for all
uses at high pressure, at high velocity through small pipes, well
insulated and have the heat at the process when needed; fit boilers
so they can arise to a demand when needed with ample stack and
draft; install stokers, if the plant will justify it; insulate the boiler walls
and generate steam as cheaply as possible—and don’t waste it.
 Boilers.—The common types of boilers in the United States are
the fire tube boiler and the water tube boiler. The first
named kind is that which suggests itself to the reader when a boiler
is mentioned and consists of a shell cylindrical in form with a head in
each end into which tubes are expanded. In operation the water
surrounds the tubes and the fire passes through them. The water
tube boiler is the opposite of this. There are variations of these and
boilers that are built using both principles in part. There are also
Cornish, Lancashire and Yorkshire types used in England which are
fire tubes using only one or two tubes. Where a plant requirement is
up to 500 horsepower, the fire tube is the usual installation owing to
its cheaper cost, and more units which enables more continuous
operation, especially if a unit is out of service for cleaning, 100, 125
or 150 horse-power being the standard size units. If a plant requires
750 horse-power or more, installation of the water tube is preferable
owing to their requiring less floor space, higher pressures are usually
carried and they are regarded as more safe under these conditions.
They can be forced to a higher output if demanded, which is an
advantage in times of rush business.
Conclusion.—The mounting costs of fuel prompts the writer to
say that the management of a power plant of any packing house
demands intelligence in the selection of equipment and in its
operation. The losses that occur are silently passing on perpetually
and are immeasurable to the proprietor since they are intangible in
dollars and cents. In these days of educated engineers, a mixture of
education and common sense is obtainable and it takes this training
or the practical man who has augmented his practice with ability to
use a pencil to figure out his conditions, and suggest remedies that
will stop a leak of no inconsiderable consequence. It is a usually
neglected department by the proprietary interests.
 CHAPTER VII
COOLERS.
Meat Chilling — Dividing Beef Coolers — Regular Temperatures — Main
Cooler — Handling Beef — Fore-Cooler — Heavy Cattle — Domestic
Beef — Trimming of Beef — Skirt Trimming — Ribbing Beef — Freezing
Beef — Loading Beef — Weighing and Tagging — Care of Coolers —
Mutton and Veal — Chilling Hogs — Cellars — Freezing Meat.

Meat Chilling.—Meat chilling must be done intelligently with

opportunity afforded for gradual chilling without too rapid hardening.
The meat should not be subjected to sudden contracting low
temperature effects, but to a gradual tempering that will carry the
meat from blood heat down to the desired temperature. This effect is
readily attained by properly constructed chill rooms. Air circulation is
the great requisite.
Sufficient attention has not been paid in the past to provide ample
room above the hanging carcass, that is space between the carcass
and the underside of pan loft. Note slope on spray loft in Fig. 17
(Chap. IV), showing design of Balanced Brine System. Coil lofts
should follow the same principle.
Dividing Beef Coolers.—Regarding the subject of dividing beef
coolers to avoid freshly killed animals from being brought into
contact with chilled ones. This is a disputed point, but it would seem
that if space were available it would be a benefit to provide for it. Any
moisture that might be given off from the freshly killed animal would,
upon coming in contact with the chilled one naturally condense, and
moisture is a detriment to the preservation of meats. The nature of
the business being conducted and the length of time it might be
desired to store the meat should be considered.
Regular Temperature.—Temperature is the all important feature
in meat preservation. The best practice appears to be, for beef in
domestic meat trade, to reduce the temperature to 34° F. as quickly
as possible and endeavor to maintain it at a uniform temperature.
Main Cooler.—When these rooms are being filled from the fore
cooler they should be held to a maximum temperature of 38° F. As
soon as the filling of the cooler is finished the refrigeration should be
turned on full force with a view to bringing the temperature down to
34° F. in twelve to fifteen hours after the cooler is closed. While it
might be possible to bring it down in much less time it is not
advisable; neither should it be longer than this. At the end of twenty-
four hours for domestic shipment or city sales, the cooler should be
held at a temperature of 34° F.
Handling Beef.—When the beef is ready to go into coolers it is
supposedly well dressed and absolutely clean. Such being the case,
care should be taken to see that the carcasses, as they are moved
along the rails, are kept bone to bone, or back to back, for if they are
thrown in promiscuously (as more or less blood will run out of the
kidneys if run together, one front against another back of beef) they
will certainly become discolored. The beef should be handled
speedily and with as little jerking and throwing around as possible,
for when handled roughly veins will purge more or less and the beef,
even though clean, will be highly colored.
The sides of beef should be placed at least a foot apart in the
preliminary or fore-cooler. They should be placed so that under no
condition does one side of meat touch the other, for if it does touch it
will show an unchilled spot on the beef, which disfigures it.
Beef should always, if possible, be hung a foot apart the first
twenty-four hours; later it can be hung closer without detriment,
although it should not be allowed to touch.
Fore-Cooler.—When a fore-cooler is used beef should be held in
the fore-cooler, which consists of one-third of the length of the cooler
cut off by a partition or a separate room, a large amount of the
moisture and animal heat that is left in the carcass is here
abstracted. When killing choice cattle it is advisable to have the
temperature of the fore-cooler as near 38° F. as possible, 43° F. is
not detrimental. Sides of heavy cattle should hang at least eighteen