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69 Membrane Carbohydrates Attach to Both
Lipids and Proteins
69 Intracellular Compartments
69 Cells Are Divided into Compartments
69 The Cytoplasm Includes Cytosol, Inclusions, Fibers,
and Organelles
72 Inclusions Are in Direct Contact with the Cytosol
72 Cytoplasmic Protein Fibers Come in Three Sizes
73 Microtubules Form Centrioles, Cilia,
and Flagella
73 Emerging Concepts: Single Cilia
Are Sensors
74 The Cytoskeleton Is a Changeable Scaffold
74 Motor Proteins Create Movement
74 Organelles Create Compartments for
Specialized Functions
76 The Nucleus Is the Cell’s Control Center
76 Tissues of the Body
78 Extracellular Matrix Has Many Functions
78 Cell Junctions Hold Cells Together to Form Tissues
80 Epithelia Provide Protection and Regulate Exchange
85 Connective Tissues Provide Support
and Barriers
87 Biotechnology: Grow Your
Own Cartilage
89 Muscle and Neural Tissues Are Excitable
89 Tissue Remodeling
90 Apoptosis Is a Tidy Form of Cell Death
90 Stem Cells Can Create New
Specialized Cells
91 Focus On . . . the Skin
92 Organs
94 Chapter Summary
95 Questions
96 Answers
98 Energy and Cellular Chapter 4
Metabolism
99 Running Problem: Tay-Sachs Disease:
A Deadly Inheritance
99 Energy in Biological Systems
100 Energy Is Used to Perform Work
101 Energy Comes in Two Forms: Kinetic and Potential
101 Energy Can Be Converted from One Form to Another
102 Thermodynamics Is the Study of Energy Use
102 Chemical Reactions
102 Energy Is Transferred Between Molecules During
Reactions
103 Activation Energy Gets Reactions Started
103 Energy Is Trapped or Released during Reactions
104 Net Free Energy Change Determines Reaction
Reversibility
Contents vii
105 Enzymes
105 Enzymes Are Proteins
106 Biotechnology: Seeing Isozymes
106 Reaction Rates Are Variable
106 Enzymes May Be Activated, Inactivated,
or Modulated
107 Enzymes Lower Activation Energy
of Reactions
107 Enzymatic Reactions Can Be Categorized
109 Metabolism
110 Cells Regulate Their Metabolic Pathways
111 ATP Transfers Energy Between Reactions
111 Catabolic Pathways Produce ATP
116 One Glucose Molecule Can Yield 30–32 ATP
117 Anaerobic Metabolism Makes 2 ATP
117 Proteins Are the Key to Cell Function
118 DNA Guides the Synthesis of RNA
121 Alternative Splicing Creates Multiple
Proteins from One DNA Sequence
122 Emerging Concepts: Purple Petunias
and RNAi
122 mRNA Translation Links Amino Acids
122 Protein Sorting Directs Proteins to
Their Destination
123 Proteins Undergo Post-Translational
Modification
125 Chapter Summary
126 Questions
128 Answers
129 Membrane Dynamics Chapter 5
130 Homeostasis Does Not Mean Equilibrium
130 Running Problem: Cystic Fibrosis
132 Osmosis and Tonicity
132 The Body Is Mostly Water
132 The Body Is in Osmotic Equilibrium
133 Osmolarity Describes the Number of Particles in Solution
134 Tonicity Describes the Volume Change of a Cell
139 Transport Processes
140 Cell Membranes Are Selectively Permeable
141 Diffusion
142 Lipophilic Molecules Cross Membranes by
Simple Diffusion
144 Protein-Mediated Transport
145 Membrane Proteins Have Four Major
Functions
146 Emerging Concepts: Transporter
Gene Families
146 Channel Proteins Form Open, Water-Filled Passageways
viii Contents

148 Carrier Proteins Change Conformation to Move Molecules 181 Receptor-Enzymes Have Protein Kinase or Guanylyl
149 Facilitated Diffusion Uses Carrier Proteins Cyclase Activity

150 Active Transport Moves Substances against Their 183 Most Signal Transduction Uses G Proteins
Concentration Gradients 183 Many Lipophobic Hormones Use GPCR-cAMP
153 Carrier-Mediated Transport Exhibits Specificity, Pathways
Competition, and Saturation 183 G Protein–Coupled Receptors Also Use Lipid-Derived
Second Messengers
155 Vesicular Transport
185 Integrin Receptors Transfer Information from the
155 Phagocytosis Creates Vesicles Using the Cytoskeleton Extracellular Matrix
155 Endocytosis Creates Smaller Vesicles 185 The Most Rapid Signal Pathways Change Ion Flow
157 Clinical Focus LDL: The Lethal Through Channels
Lipoprotein
187 Novel Signal Molecules
157 Exocytosis Releases Molecules Too Large
for Transport Proteins 187 Calcium Is an Important Intracellular Signal
187 Gases Are Ephemeral Signal Molecules
157 Epithelial Transport
188 Biotechnology: Calcium Signals
158 Epithelial Transport May Be Paracellular Glow in the Dark
or Transcellular
188 Clinical Focus: From Dynamite
159 Transcellular Transport of Glucose Uses Membrane to Medicine
Proteins
188 Some Lipids Are Important Paracrine Signals
160 Transcytosis Uses Vesicles to Cross an Epithelium
189 Modulation of Signal Pathways
160 The Resting Membrane Potential
189 One Ligand May Have Multiple Receptors
161 Electricity Review
190 Receptors Exhibit Saturation, Specificity,
161 The Cell Membrane Enables Separation of and Competition
Electrical Charge in the Body
191 Up- and Down-Regulation Enable Cells to
163 The Resting Membrane Potential Is Due Modulate Responses
Mostly to Potassium
191 Cells Must Be Able to Terminate Signal Pathways
166 Changes in Ion Permeability Change
the Membrane Potential 191 Many Diseases and Drugs Target the Proteins of
Signal Transduction
166 Integrated Membrane Processes: Insulin Secretion
192 Homeostatic Reflex Pathways
169 Chapter Summary
170 Questions 192 Cannon’s Postulates Describe Regulated Variables
172 Answers and Control Systems
194 Long-Distance Pathways Maintain Homeostasis
197 Control Systems Vary in Their Speed
174 Communication, Integration, Chapter 6 and Specificity
and Homeostasis 198 Complex Reflex Control Pathways
Have Several Integrating Centers
175 Cell-to-Cell Communication
202 Chapter Summary
175 Running Problem: Diabetes Mellitus: 204 Questions
A Growing Epidemic 205 Answers
175 Gap Junctions Create Cytoplasmic Bridges
175 Contact-Dependent Signals Require
Cell-to-Cell Contact
177 Paracrine and Autocrine Signals Carry Out Unit 2
Local Communication
177 Long-Distance Communication May Be Electrical or Homeostasis and Control
Chemical
177 Cytokines May Act as Both Local and
Long-Distance Signals 206 Introduction to the Chapter 7
178 Signal Pathways Endocrine System
178 Receptor Proteins Are Located Inside the
Cell or on the Cell Membrane
207 Hormones
207 Running Problem: Graves’ Disease
180 Membrane Proteins Facilitate Signal Transduction
207 Hormones Have Been Known Since
Ancient Times

208 Clinical Focus: Diabetes:
The Discovery of Insulin
208 What Makes a Chemical a Hormone?
209 Hormones Act by Binding to Receptors
211 Hormone Action Must Be Terminated
211 The Classification of Hormones
212 Most Hormones Are Peptides or Proteins
214 Steroid Hormones Are Derived from Cholesterol
216 Some Hormones Are Derived from Single Amino Acids
216 Control of Hormone Release
216 Hormones Can Be Classified by Their Reflex Pathways
217 The Endocrine Cell Is the Sensor in the Simplest
Endocrine Reflexes
219 Many Endocrine Reflexes Involve the Nervous System
219 Neurohormones Are Secreted into the Blood by Neurons
219 The Pituitary Gland Is Actually Two Fused Glands
219 The Posterior Pituitary Stores and Releases Two
Neurohormones
221 The Anterior Pituitary Secretes Six Hormones
223 A Portal System Delivers Hormones from
Hypothalamus to Anterior Pituitary
223 Anterior Pituitary Hormones Control Growth,
Metabolism, and Reproduction
223 Feedback Loops Are Different in
the Hypothalamic-Pituitary Pathway
225 Hormone Interactions
225 In Synergism, the Effect of Interacting Hormones
Is More Than Additive
226 A Permissive Hormone Allows Another Hormone
to Exert Its Full Effect
227 Antagonistic Hormones Have Opposing Effects
227 Endocrine Pathologies
227 Hypersecretion Exaggerates a Hormone’s Effects
228 Hyposecretion Diminishes or Eliminates a Hormone’s
Effects
228 Receptor or Second Messenger Problems Cause
Abnormal Tissue Responsiveness
228 Diagnosis of Endocrine Pathologies Depends on the
Complexity of the Reflex
230 Hormone Evolution
231 Focus On . . . The Pineal Gland
233 Chapter Summary
234 Questions
236 Answers
237 Neurons: Cellular and Network Chapter 8
Properties
238 Running Problem: Mysterious Paralysis
239 Organization of the Nervous System
239 Cells of the Nervous System
Contents ix
239 Neurons Carry Electrical Signals
243 Establishing Synapses Depends on
Chemical Signals
244 Glial Cells Provide Support for Neurons
245 Can Stem Cells Repair Damaged Neurons?
248 Electrical Signals in Neurons
248 The Nernst Equation Predicts Membrane
Potential for a Single Ion
249 The GHK Equation Predicts Membrane Potential
Using Multiple Ions
249 Ion Movement Creates Electrical Signals
250 Gated Channels Control the Ion Permeability
of the Neuron
251 Clinical Focus: Mutant Channels
251 Current Flow Obeys Ohm’s Law
251 Graded Potentials Reflect Stimulus Strength
254 Action Potentials Travel Long Distances
255 Na+ and K+ Move Across the Membrane During Action
Potentials
257 One Action Potential Does Not Alter Ion Concentration
Gradients
257 Axonal Na+ Channels Have Two Gates
258 Action Potentials Will Not Fire During the Absolute
Refractory Period
259 Action Potentials Are Conducted
261 Larger Neurons Conduct Action Potentials Faster
261 Conduction Is Faster in Myelinated Axons
263 Biotechnology: The Body’s Wiring
265 Chemical Factors Alter Electrical Activity
266 Cell-to-Cell Communication in the
Nervous System
266 Neurons Communicate at Synapses
266 Neurons Secrete Chemical Signals
267 Neurotransmitters Are Highly Varied
269 Clinical Focus: Myasthenia Gravis
270 Biotechnology: Of Snakes, Snails,
Spiders, and Sushi
270 Neurotransmitters Are Released
from Vesicles
273 Stronger Stimuli Release More Neurotransmitter
274 Integration of Neural Information Transfer
274 Postsynaptic Responses May Be Slow or Fast
277 Neural Pathways May Involve Many Neurons
277 Synaptic Activity Can Be Modified
280 Long-Term Potentiation Alters Synapses
281 Disorders of Synaptic Transmission
Are Responsible for Many Diseases
283 Chapter Summary
284 Questions
286 Answers
x Contents
288 The Central Nervous System Chapter 9
289 Emergent Properties of Neural Networks
289 Running Problem: Infantile Spasms
289 Evolution of Nervous Systems
291 Biotechnology: Tracing Neurons
in a Network
291 Anatomy of the Central Nervous System
291 The CNS Develops from a Hollow Tube
293 The CNS Is Divided into Gray Matter and White Matter
293 Bone and Connective Tissue Support the CNS
293 The Brain Floats in Cerebrospinal Fluid
296 The Blood-Brain Barrier Protects the Brain
297 Neural Tissue Has Special Metabolic
Requirements
298 Clinical Focus: Diabetes:
Hypoglycemia and the Brain
298 The Spinal Cord
299 The Brain
301 The Brain Stem Is the Oldest Part of the Brain
303 The Cerebellum Coordinates Movement
303 The Diencephalon Contains the Centers for Homeostasis
304 The Cerebrum Is the Site of Higher Brain Functions
305 Brain Function
306 The Cerebral Cortex Is Organized into Functional Areas
307 The Spinal Cord and Brain Integrate Sensory Information
308 Sensory Information Is Processed into Perception
308 The Motor System Governs Output from the CNS
309 The Behavioral State System Modulates Motor Output
309 Why Do We Sleep?
312 Physiological Functions Exhibit Circadian
Rhythms
312 Emerging Concepts: Adenosine
and That “Java Jolt”
312 Emotion and Motivation Involve Complex
Neural Pathways
314 Moods Are Long-Lasting Emotional States
314 Learning and Memory Change Synaptic
Connections in the Brain
314 Learning Is the Acquisition of Knowledge
315 Memory Is the Ability to Retain
and Recall Information
317 Language Is the Most Elaborate Cognitive Behavior
318 Personality Is a Combination of Experience
and Inheritance
320 Chapter Summary
322 Questions
323 Answers
325 Sensory Physiology Chapter 10
326 Running Problem: Ménière’s Disease
326 General Properties of Sensory Systems
327 Receptors Are Sensitive to Particular Forms of Energy
328 Sensory Transduction Converts Stimuli into Graded
Potentials
328 A Sensory Neuron Has a Receptive Field
329 The CNS Integrates Sensory Information
331 Coding and Processing Distinguish Stimulus Properties
335 Somatic Senses
335 Pathways for Somatic Perception Project to the
Cortex and Cerebellum
335 Touch Receptors Respond to Many Different Stimuli
337 Temperature Receptors Are Free Nerve Endings
337 Nociceptors Initiate Protective Responses
338 Pain and Itching Are Mediated
by Nociceptors
340 Clinical Focus: Natural Painkillers
341 Chemoreception: Smell and Taste
341 Olfaction Is One of the Oldest Senses
344 Taste Is a Combination of Five Basic Sensations
344 Taste Transduction Uses Receptors and Channels
346 The Ear: Hearing
346 Hearing Is Our Perception of Sound
348 Sound Transduction Is a Multistep Process
349 The Cochlea Is Filled with Fluid
352 Sounds Are Processed First in the Cochlea
352 Auditory Pathways Project to the Auditory Cortex
353 Hearing Loss May Result from Mechanical or Neural
Damage
354 Biotechnology: Artificial Ears
354 The Ear: Equilibrium
354 The Vestibular Apparatus Provides Information about
Movement and Position
356 The Semicircular Canals Sense Rotational Acceleration
356 The Otolith Organs Sense Linear Acceleration and Head
Position
356 Equilibrium Pathways Project Primarily to the Cerebellum
357 The Eye and Vision
357 The Skull Protects the Eye
358 Clinical Focus: Glaucoma
358 Light Enters the Eye through the Pupil
359 The Lens Focuses Light on the Retina
361 Phototransduction Occurs at the Retina
365 Emerging Concepts: Melanopsin
365 Photoreceptors Transduce Light into
Electrical Signals

367 Signal Processing Begins in the Retina
372 Chapter Summary
374 Questions
375 Answers
377 Efferent Division: Autonomic
and Somatic Motor Control
378 Running Problem: A Powerful
Addiction
378 The Autonomic Division
379 Autonomic Reflexes Are Important for Homeostasis
380 Antagonistic Control Is a Hallmark of the
Autonomic Division
380 Autonomic Pathways Have Two Efferent
Neurons in Series
381 Sympathetic and Parasympathetic Branches
Originate in Different Regions
383 The Autonomic Nervous System Uses a Variety of
Chemical Signals
384 Autonomic Pathways Control Smooth and
Cardiac Muscle and Glands
385 Autonomic Neurotransmitters Are Synthesized
in the Axon
385 Autonomic Receptors Have Multiple Subtypes
386 The Adrenal Medulla Secretes Catecholamines
387 Autonomic Agonists and Antagonists Are
Important Tools in Research and Medicine
388 Clinical Focus: Diabetes: Autonomic
Neuropathy
388 Primary Disorders of the Autonomic
Nervous System Are Relatively Uncommon
389 Summary of Sympathetic and Parasympathetic
Branches
391 The Somatic Motor Division
391 A Somatic Motor Pathway Consists of One Neuron
391 The Neuromuscular Junction Contains Nicotinic
Receptors
394 Chapter Summary
395 Questions
396 Answers
398 Muscles
400 Running Problem: Periodic
Paralysis
400 Skeletal Muscle
400 Skeletal Muscles Are Composed of Muscle Fibers
401 Myofibrils Are Muscle Fiber Contractile Structures
Contents xi
404 Muscle Contraction Creates Force
406 Actin and Myosin Slide Past Each Other During
Contraction
407 Myosin Crossbridges Move Actin Filaments
407 Calcium Signals Initiate Contraction
Chapter 11 408 Myosin Heads Step Along Actin Filaments
410 Biotechnology: Watching Myosin
Work
410 Acetylcholine Initiates Excitation-
Contraction Coupling
413 Skeletal Muscle Contraction Requires
a Steady Supply of ATP
414 Fatigue Has Multiple Causes
415 Skeletal Muscle Is Classified by Speed and
Fatigue Resistance
417 Resting Fiber Length Affects Tension
417 Force of Contraction Increases with Summation
418 A Motor Unit Is One Motor Neuron
and Its Muscle Fibers
420 Contraction Force Depends on the Types and
Numbers of Motor Units
421 Mechanics of Body Movement
421 Isotonic Contractions Move Loads; Isometric
Contractions Create Force Without Movement
421 Bones and Muscles Around Joints Form Levers and
Fulcrums
423 Muscle Disorders Have Multiple Causes
426 Smooth Muscle
427 Smooth Muscle Is More Variable
Than Skeletal Muscle
429 Smooth Muscle Lacks Sarcomeres
429 Myosin Phosphorylation Controls Contraction
431 MLCP Controls Ca2+ Sensitivity
431 Calcium Initiates Smooth Muscle Contraction
433 Some Smooth Muscles Have Unstable Membrane
Potentials
433 Chemical Signals Influence Smooth
Muscle Activity
435 Cardiac Muscle
436 Chapter Summary
437 Questions
439 Answers
Chapter 12
Chapter 13
441 Integrative Physiology I:
Control of Body Movement
442 Neural Reflexes
442 Neural Reflex Pathways Can Be Classified
in Different Ways
442 Running Problem: Tetanus
xii Contents

444 Autonomic Reflexes 479 Cardiac Muscle Contraction Can Be Graded

444 Skeletal Muscle Reflexes
446 Muscle Spindles Respond to Muscle Stretch
446 Clinical Focus: Reflexes
and Muscle Tone
448 Golgi Tendon Organs Respond
to Muscle Tension
449 Stretch Reflexes and Reciprocal Inhibition Control
Movement Around a Joint
451 Flexion Reflexes Pull Limbs away
from Painful Stimuli
452 The Integrated Control of Body Movement
452 Movement Can Be Classified as Reflex, Voluntary,
or Rhythmic
454 The CNS Integrates Movement
456 Emerging Concepts: Visualization
Techniques in Sports
457 Symptoms of Parkinson’s Disease
Reflect Basal Ganglia Function
457 Control of Movement in Visceral Muscles
459 Chapter Summary
460 Questions
461 Answers
Unit 3
Integration of Function
462 Cardiovascular Physiology Chapter 14
463 Running Problem: Myocardial
Infarction
463 Overview of the Cardiovascular System
464 The Cardiovascular System Transports
Materials throughout the Body
464 The Cardiovascular System Consists of the Heart, Blood
Vessels, and Blood
466 Pressure, Volume, Flow, and Resistance
467 The Pressure of Fluid in Motion Decreases over Distance
467 Pressure Changes in Liquids without a Change in Volume
467 Blood Flows from Higher Pressure to Lower Pressure
469 Resistance Opposes Flow
470 Velocity Depends on the Flow Rate and the
Cross-Sectional Area
471 Cardiac Muscle and the Heart
471 The Heart Has Four Chambers
474 Heart Valves Ensure One-Way Flow in the Heart
475 Cardiac Muscle Cells Contract without Innervation
477 Calcium Entry Is a Feature of Cardiac EC Coupling
479 Myocardial Action Potentials Vary
483 The Heart As a Pump
483 Electrical Signals Coordinate Contraction
485 Pacemakers Set the Heart Rate
485 Clinical Focus: Fibrillation
486 The Electrocardiogram Reflects
Electrical Activity
487 The Heart Contracts and Relaxes during a Cardiac Cycle
492 Clinical Focus: Gallops, Clicks,
and Murmurs
493 Pressure-Volume Curves Represent
One Cardiac Cycle
495 Stroke Volume Is the Volume of Blood Pumped per
Contraction
495 Cardiac Output Is a Measure of Cardiac Performance
495 The Autonomic Division Modulates Heart Rate
497 Multiple Factors Influence Stroke Volume
498 Contractility Is Controlled by the Nervous and
Endocrine Systems
500 Emerging Concepts: Stem Cells
for Heart Disease
500 EDV and Arterial Blood Pressure
Determine Afterload
503 Chapter Summary
504 Questions
506 Answers
Chapter 15
508 Blood Flow and the Control
of Blood Pressure
509 Running Problem: Essential
Hypertension
510 The Blood Vessels
510 Blood Vessels Contain Vascular Smooth Muscle
511 Arteries and Arterioles Carry Blood Away
from the Heart
511 Exchange Takes Place in the Capillaries
512 Blood Flow Converges in the Venules and Veins
512 Angiogenesis Creates New Blood Vessels
513 Blood Pressure
513 Blood Pressure Is Highest in Arteries and Lowest in Veins
515 Arterial Blood Pressure Reflects the Driving Pressure for
Blood Flow
515 Blood Pressure Is Estimated by Sphygmomanometry
516 Cardiac Output and Peripheral Resistance Determine
Mean Arterial Pressure
518 Changes in Blood Volume Affect Blood Pressure
519 Clinical Focus: Shock

519 Resistance in the Arterioles
519 Myogenic Autoregulation Automatically
Adjusts Blood Flow
520 Paracrines Alter Vascular Smooth Muscle
Contraction
522 The Sympathetic Branch Controls Most
Vascular Smooth Muscle
524 Distribution of Blood to the Tissues
524 Regulation of Cardiovascular Function
525 The Baroreceptor Reflex Controls Blood Pressure
527 Orthostatic Hypotension Triggers the
Baroreceptor Reflex
528 Other Systems Influence Cardiovascular Function
528 Exchange at the Capillaries
528 Velocity of Blood Flow Is Lowest in the Capillaries
529 Most Capillary Exchange Takes Place by
Diffusion and Transcytosis
529 Capillary Filtration and Absorption Take Place by
Bulk Flow
532 The Lymphatic System
533 Edema Results from Alterations in Capillary
Exchange
534 Cardiovascular Disease
534 Risk Factors Include Smoking and Obesity
534 Clinical Focus: Diabetes
and Cardiovascular Disease
535 Atherosclerosis Is an Inflammatory
Process
535 Emerging Concepts: Inflammatory
Markers for Cardiovascular Disease
536 Hypertension Represents a Failure of Homeostasis
538 Chapter Summary
540 Questions
542 Answers
544 Blood Chapter 16
545 Running Problem: Blood Doping
in Athletes
545 Plasma and the Cellular Elements of Blood
545 Plasma Is Extracellular Matrix
547 Cellular Elements Include RBCs, WBCs, and Platelets
547 Blood Cell Production
549 Blood Cells Are Produced in the Bone Marrow
549 Hematopoiesis Is Controlled by Cytokines
550 Colony-Stimulating Factors Regulate Leukopoiesis
551 Thrombopoietin Regulates Platelet Production
551 Erythropoietin Regulates RBC Production
551 Focus On . . . Bone Marrow
551 Red Blood Cells
551 Mature RBCs Lack a Nucleus
Contents xiii
554 Hemoglobin Synthesis Requires Iron
554 RBCs Live About Four Months
554 Clinical Focus: Diabetes: Hemoglobin
and Hyperglycemia
556 RBC Disorders Decrease Oxygen Transport
557 Platelets and Coagulation
557 Platelets Are Small Fragments of Cells
557 Hemostasis Prevents Blood Loss from Damaged Vessels
558 Emerging Concepts: Platelet-Rich
Plasma Therapy
560 Platelet Activation Begins the
Clotting Process
561 Coagulation Converts a Platelet Plug
into a Clot
562 Anticoagulants Prevent Coagulation
566 Chapter Summary
566 Questions
567 Answers
568 Mechanics of Breathing Chapter 17
569 Running Problem: Emphysema
569 The Respiratory System
570 Bones and Muscles of the Thorax Surround the Lungs
570 Pleural Sacs Enclose the Lungs
571 Airways Connect Lungs to the External Environment
571 The Airways Warm, Humidify, and Filter Inspired Air
574 Alveoli Are the Site of Gas Exchange
575 Pulmonary Circulation Is High-Flow,
Low-Pressure
576 Clinical Focus: Congestive
Heart Failure
576 Gas Laws
578 Air Is a Mixture of Gases
578 Gases Move Down Pressure Gradients
578 Boyle’s Law Describes Pressure-Volume Relationships
578 Ventilation
578 Lung Volumes Change During Ventilation
580 During Ventilation, Air Flows Because of Pressure
Gradients
580 Inspiration Occurs When Alveolar Pressure Decreases
583 Expiration Occurs When Alveolar Pressure Increases
583 Intrapleural Pressure Changes During Ventilation
585 Lung Compliance and Elastance May
Change in Disease States
586 Surfactant Decreases the Work of Breathing
587 Airway Diameter Determines Airway Resistance
588 Rate and Depth of Breathing Determine the
Efficiency of Breathing
590 Gas Composition in the Alveoli Varies Little During
Normal Breathing
590 Ventilation and Alveolar Blood Flow Are Matched
xiv Contents
592 Auscultation and Spirometry Assess
Pulmonary Function
595 Chapter Summary
596 Questions
598 Answers
Chapter 18
599 Gas Exchange and Transport
600 Running Problem: High Altitude
600 Gas Exchange in the Lungs and Tissues
602 Lower Alveolar PO2 Decreases Oxygen Uptake
604 Diffusion Problems Cause Hypoxia
604 Biotechnology: The Pulse Oximeter
605 Gas Solubility Affects Diffusion
606 Gas Transport in the Blood
608 Hemoglobin Binds to Oxygen
608 Oxygen Binding Obeys the Law of Mass Action
608 Hemoglobin Transports Most Oxygen
to the Tissues
609 Emerging Concepts: Blood
Substitutes
609 PO2 Determines Oxygen-Hb Binding
610 Oxygen Binding Is Expressed As a Percentage
612 Several Factors Affect Oxygen-Hb Binding
613 Carbon Dioxide Is Transported in Three Ways
615 Regulation of Ventilation
617 Neurons in the Medulla Control Breathing
618 Carbon Dioxide, Oxygen, and pH Influence Ventilation
620 Protective Reflexes Guard the Lungs
620 Higher Brain Centers Affect Patterns of Ventilation
622 Chapter Summary
623 Questions
625 Answers
626 The Kidneys Chapter 19
627 Running Problem: Gout
627 Functions of the Kidneys
628 Anatomy of the Urinary System
628 The Urinary System Consists of Kidneys,
Ureters, Bladder, and Urethra
628 Clinical Focus: Urinary Tract Infections
628 The Nephron Is the Functional Unit
of the Kidney
629 Overview of Kidney Function
629 Kidneys Filter, Reabsorb, and Secrete
633 The Nephron Modifies Fluid Volume and Osmolarity
634 Filtration
634 The Renal Corpuscle Contains Filtration Barriers
636 Capillary Pressure Causes Filtration
636 GFR Is Relatively Constant
636 GFR Is Subject to Autoregulation
638 Hormones and Autonomic Neurons
Also Influence GFR
638 Emerging Concepts: Diabetes:
Diabetic Nephropathy
640 Reabsorption
640 Reabsorption May Be Active or Passive
642 Biotechnology: Artificial Kidneys
642 Renal Transport Can Reach Saturation
644 Peritubular Capillary Pressures Favor Reabsorption
644 Secretion
646 Competition Decreases Penicillin Secretion
646 Excretion
647 Clearance Is a Noninvasive Way
to Measure GFR
649 Clearance Helps Us Determine Renal Handling
650 Micturition
652 Chapter Summary
654 Questions
655 Answers
657 Integrative Physiology II: Chapter 20
Fluid and Electrolyte Balance
658 Fluid and Electrolyte Homeostasis
658 Running Problem: Hyponatremia
658 ECF Osmolarity Affects Cell Volume
658 Multiple Systems Integrate Fluid
and Electrolyte Balance
659 Water Balance
660 Daily Water Intake and Excretion Are Balanced
660 The Kidneys Conserve Water
661 The Renal Medulla Creates Concentrated
Urine
662 Clinical Focus: Diabetes:
Osmotic Diuresis
663 Vasopressin Controls Water Reabsorption
663 Blood Volume and Osmolarity Activate Osmoreceptors
666 The Loop of Henle Is a Countercurrent Multiplier
668 Sodium Balance and ECF Volume
669 Aldosterone Controls Sodium Balance
671 Low Blood Pressure Stimulates Aldosterone Secretion
671 ANG II Has Many Effects
673 ANP Promotes Na+ and Water Excretion
675 Potassium Balance

675 Behavioral Mechanisms in Salt and Water Balance
675 Drinking Replaces Fluid Loss
676 Low Na+ Stimulates Salt Appetite
676 Avoidance Behaviors Help Prevent Dehydration
676 Integrated Control of Volume and Osmolarity
676 Osmolarity and Volume Can Change Independently
677 Dehydration Triggers Homeostatic Responses
681 Acid-Base Balance
681 pH Changes Can Denature Proteins
682 Acids and Bases in the Body Come from Many Sources
682 pH Homeostasis Depends on Buffers, Lungs, and Kidneys
683 Buffer Systems Include Proteins, Phosphate Ions,
and HCO3-
684 Ventilation Can Compensate for pH Disturbances
684 Kidneys Use Ammonia and Phosphate Buffers
685 The Proximal Tubule Secretes H+ and Reabsorbs HCO3-
686 The Distal Nephron Controls Acid Excretion
688 Acid-Base Disturbances May Be
Respiratory or Metabolic
691 Chapter Summary
693 Questions
694 Answers
Unit 4
Metabolism, Growth, and Aging
696 The Digestive System Chapter 21
697 Running Problem: Cholera in Haiti
697 Digestive Function and Processes
699 Anatomy of the Digestive System
699 The Digestive System Is a Tube
702 The GI Tract Wall Has Four Layers
703 Motility
703 GI Smooth Muscle Exhibits Different
Patterns of Contraction
703 GI Smooth Muscle Contracts Spontaneously
703 Clinical Focus: Diabetes:
Delayed Gastric Emptying
705 Secretion
705 The Digestive System Secretes Ions and Water
707 Digestive Enzymes Are Secreted into the Lumen
708 Specialized Cells Secrete Mucus
708 Saliva Is an Exocrine Secretion
708 The Liver Secretes Bile
709 Focus On . . . The Liver
710 Digestion and Absorption
710 Carbohydrates Are Absorbed As Monosaccharides
Contents xv
712 Proteins Are Digested into Small Peptides and Amino Acids
712 Some Larger Peptides Can Be Absorbed Intact
712 Bile Salts Facilitate Fat Digestion
714 Nucleic Acids Are Digested into Bases and
Monosaccharides
714 The Intestine Absorbs Vitamins and Minerals
715 The Intestine Absorbs Ions and Water
715 Regulation of GI Function
717 Emerging Concepts: Taste
Receptors in the Gut
717 The Enteric Nervous System Can Act
Independently
718 GI Peptides Include Hormones,
Neuropeptides, and Cytokines
719 Integrated Function: The Cephalic Phase
719 Chemical and Mechanical Digestion Begins in the Mouth
720 Swallowing Moves Food from Mouth to Stomach
722 Integrated Function: The Gastric Phase
722 The Stomach Stores Food
722 The Stomach Secretes Acid and Enzymes
725 The Stomach Balances Digestion and Protection
725 Integrated Function: The Intestinal Phase
725 Bicarbonate Neutralizes Gastric Acid
727 Most Fluid Is Absorbed in the Small Intestine
727 Most Digestion Occurs in the Small
Intestine
728 Clinical Focus: Lactose Intolerance
728 The Large Intestine Concentrates Waste
729 Diarrhea Can Cause Dehydration
730 Immune Functions of the GI Tract
730 M Cells Sample Gut Contents
730 Vomiting Is a Protective Reflex
732 Chapter Summary
734 Questions
735 Answers
Chapter 22
736 Metabolism and Energy
Balance
737 Appetite and Satiety
737 Running Problem: Eating Disorders
738 Biotechnology: Discovering Peptides:
Research in Reverse
739 Energy Balance
739 Energy Input Equals Energy Output
740 Oxygen Consumption Reflects Energy Use
740 Clinical Focus: Estimating Fat—
The Body Mass Index
741 Many Factors Influence Metabolic Rate
742 Energy Is Stored in Fat and Glycogen
xvi Contents
742 Metabolism
743 Ingested Energy May Be Used or Stored
744 Enzymes Control the Direction of Metabolism
745 Fed-State Metabolism
745 Carbohydrates Make ATP
745 Amino Acids Make Proteins
745 Fats Store Energy
748 Plasma Cholesterol Predicts Heart Disease
749 Fasted-State Metabolism
749 Glycogen Converts to Glucose
749 Clinical Focus: Antioxidants Protect
the Body
750 Proteins Can Be Used to Make ATP
751 Lipids Store More Energy than Glucose or Protein
753 Homeostatic Control of Metabolism
753 The Pancreas Secretes Insulin and Glucagon
754 The Insulin-to-Glucagon Ratio Regulates Metabolism
754 Insulin Is the Dominant Hormone of the Fed State
755 Insulin Promotes Anabolism
759 Glucagon Is Dominant in the Fasted State
760 Diabetes Mellitus Is a Family of Diseases
761 Type 1 Diabetics Are Prone to Ketoacidosis
763 Type 2 Diabetics Often Have Elevated Insulin Levels
765 Metabolic Syndrome Links Diabetes and
Cardiovascular Disease
765 Regulation of Body Temperature
765 Body Temperature Balances Heat Production,
Gain, and Loss
767 Body Temperature Is Homeostatically Regulated
768 Movement and Metabolism Produce Heat
769 The Body’s Thermostat Can Be Reset
772 Chapter Summary
773 Questions
774 Answers
775 Endocrine Control of Growth
and Metabolism
776 Review of Endocrine Principles
776 Running Problem:
Hyperparathyroidism
776 Biotechnology: Mutant
Mouse Models
777 Adrenal Glucocorticoids
777 The Adrenal Cortex Secretes Steroid Hormones
777 Cortisol Secretion Is Controlled by ACTH
780 Cortisol Is Essential for Life
780 Cortisol Is a Useful Therapeutic Drug
781 Cortisol Pathologies Result from Too Much or
Too Little Hormone
781 CRH and ACTH Have Additional
Physiological Functions
782 Emerging Concepts: Melanocortins
and the Agouti Mouse
782 Thyroid Hormones
782 Thyroid Hormones Contain Iodine
784 TSH Controls the Thyroid Gland
784 Thyroid Pathologies Affect Quality of Life
787 Growth Hormone
787 Growth Hormone Is Anabolic
788 Clinical Focus: New Growth Charts
788 Growth Hormone Is Essential
for Normal Growth
788 Genetically Engineered hGH Raises Ethical Questions
790 Tissue and Bone Growth
790 Tissue Growth Requires Hormones and Paracrines
790 Bone Growth Requires Adequate Dietary Calcium
792 Calcium Balance
792 Plasma Calcium Is Closely Regulated
793 Three Hormones Control Calcium Balance
795 Calcium and Phosphate Homeostasis Are Linked
796 Osteoporosis Is a Disease of Bone Loss
798 Chapter Summary
798 Questions
800 Answers
Chapter 24
801 The Immune System
802 Overview
802 Running Problem: HPV:
To Vaccinate or Not?
802 Pathogens of the Human Body
803 Bacteria and Viruses Require Different Defense
Mechanisms
804 Viruses Can Replicate Only inside Host Cells
804 The Immune Response
Chapter 23
805 Anatomy of the Immune System
805 Lymphoid Tissues Are Everywhere
807 Focus On . . . The Spleen
808 Leukocytes Mediate Immunity
811 Innate Immunity: Nonspecific Responses
811 Barriers Are the Body’s First Line of Defense
811 Phagocytes Ingest Foreign Material
812 NK Cells Kill Infected and Tumor Cells
812 Cytokines Create the Inflammatory Response
814 Acquired Immunity: Antigen-Specific Responses
814 Lymphocytes Mediate the Acquired Immune Response
816 B Lymphocytes Become Plasma Cells and Memory Cells
816 Antibodies Are Proteins Secreted by Plasma Cells

818 T Lymphocytes Use Contact-Dependent Signaling
819 Focus on . . . The Thymus Gland
821 Immune Response Pathways
821 Bacterial Invasion Causes Inflammation
821 Viral Infections Require Intracellular Defense
824 Specific Antigens Trigger Allergic Responses
825 MHC Proteins Allow Recognition
of Foreign Tissue
827 The Immune System Must Recognize “Self”
827 Immune Surveillance Removes Abnormal Cells
828 Biotechnology: Engineered
Antibodies
828 Neuro-Endocrine-Immune Interactions
829 Stress Alters Immune System Function
830 Modern Medicine Includes Mind-Body Therapeutics
831 Chapter Summary
833 Questions
834 Answers
835 Integrative Physiology III:
Exercise
836 Metabolism and Exercise
836 Running Problem: Heat Stroke
838 Hormones Regulate Metabolism During Exercise
838 Oxygen Consumption Is Related to Exercise Intensity
839 Several Factors Limit Exercise
839 Ventilatory Responses to Exercise
840 Cardiovascular Responses to Exercise
840 Cardiac Output Increases During Exercise
841 Muscle Blood Flow Increases During Exercise
842 Blood Pressure Rises Slightly During Exercise
843 The Baroreceptor Reflex Adjusts to Exercise
843 Feedforward Responses to Exercise
843 Temperature Regulation During Exercise
844 Exercise and Health
844 Exercise Lowers the Risk of Cardiovascular Disease
845 Type 2 Diabetes Mellitus May Improve with Exercise
845 Stress and the Immune System May Be
Influenced by Exercise
847 Chapter Summary
848 Questions
849 Answers
850 Reproduction and
Development
851 Running Problem: Infertility
851 Sex Determination
851 Sex Chromosomes Determine Genetic Sex
Contents xvii
852 Sexual Differentiation Occurs Early in Development
853 Clinical Focus: X-Linked Inherited
Disorders
856 Basic Patterns of Reproduction
856 Clinical Focus: Determining Sex
857 Gametogenesis Begins in Utero
857 The Brain Directs Reproduction
860 Environmental Factors Influence Reproduction
860 Male Reproduction
861 Testes Produce Sperm and Hormones
865 Spermatogenesis Requires Gonadotropins and
Testosterone
865 Male Accessory Glands Contribute Secretions to Semen
866 Androgens Influence Secondary Sex Characteristics
866 Female Reproduction
866 Females Have an Internal Uterus
867 The Ovary Produces Eggs and Hormones
867 A Menstrual Cycle Lasts about One Month
867 Hormonal Control of the Menstrual Cycle Is Complex
Chapter 25
874 Hormones Influence Female Secondary Sex
Characteristics
874 Procreation
874 The Human Sexual Response Has Four Phases
874 The Male Sex Act Includes Erection and Ejaculation
876 Sexual Dysfunction Affects Males and Females
876 Contraceptives Are Designed to Prevent Pregnancy
877 Infertility Is the Inability to Conceive
878 Pregnancy and Parturition
878 Fertilization Requires Capacitation
878 The Developing Embryo Implants in the Endometrium
880 The Placenta Secretes Hormones During Pregnancy
881 Pregnancy Ends with Labor and Delivery
882 The Mammary Glands Secrete Milk During Lactation
885 Prolactin Has Other Physiological Roles
885 Growth and Aging
885 Puberty Marks the Beginning of the Reproductive Years
885 Menopause and Andropause Are a Consequence of Aging
887 Chapter Summary
888 Questions
890 Answers
Appendices
Appendix A: Answers to Review Questions A-1
Appendix B: Physics and Math A-18
Chapter 26
Appendix C: Genetics A-21
Glossary/Index GI-1
Photo Credits C-1
Owner’s Manual: How to Use This Book
Welcome to Human Physiology!
As you begin your study of the human body, you should be pre-
pared to make maximum use of the resources available to you,
including your instructor, the library, the Internet, and your
textbook. One of my goals in this book is to provide you not
only with information about how the human body functions
but also with tips for studying and problem solving. Many of
these study aids have been developed with the input of my stu-
dents, so I think you may find them particularly helpful. On the
following pages, I have put together a brief tour of the special
features of the book, especially those that you may not have en-
countered previously in textbooks. Please take a few minutes to
read about them so that you can make optimum use of the book
as you study.
One of your tasks as you study will be to construct for
yourself a global view of the body, its systems, and the many
processes that keep the systems working. This “big picture”
is what physiologists call the integration of systems, and it is
a key theme in the book. To integrate information, however,
you must do more than simply memorize it. You need to truly
understand it and be able to use it to solve problems that you
have never encountered before. If you are headed for a career in
the health professions, you will do this in the clinics. If you plan
a career in biology, you will solve problems in the laboratory,
field, or classroom. Analyzing, synthesizing, and evaluating
information are skills you need to develop while you are in
school, and I hope that the features of this book will help you
with this goal.
In this edition we have continued to update and focus on
basic themes and concepts of physiology. Chapter 1 introduces
you to the key concepts in physiology that you encounter re-
peatedly as you study different organ systems. It also includes
several special features: one on mapping, a useful study skill that
is also used for decision-making in the clinics, and one on con-
structing and interpreting graphs. The new Chapter 1 Running
xviii
Problem introduces you to effective ways to find information on
the Internet.
A new element in this edition’s art program is the Essentials
and Review figures. These figures distill the basics about a topic
onto one or two pages, much as the Anatomy Summaries do. My
students tell me they find them particularly useful for review
when there isn’t time to go back and read all the text.
We have also retained the four approaches to learning
physiology that proved so popular since this book was first
published in 1998.
1. Cellular and Molecular Physiology
Most physiological research today is being done at the cellular
and molecular level, and there have been many exciting devel-
opments in molecular medicine and physiology in the ten years
since the first edition. For example, now scientists are paying
more attention to primary cilia, the single cilium that occurs on
most cells of the body. Primary cilia are thought to play a role in
some kidney and other diseases. Look for similar links between
molecular and cellular biology, physiology, and medicine
throughout the book.
2. Physiology as a Dynamic Field
Physiology is a dynamic discipline, with numerous unanswered
questions that merit further investigation and research. Many of
the “facts” presented in this text are really only our current theo-
ries, so you should be prepared to change your mental models as
new information emerges from scientific research.
3. An Emphasis on Integration
The organ systems of the body do not work in isolation, although
we study them one at a time. To emphasize the integrative nature
of physiology, three chapters (Chapters 13, 20, and 25) focus on
how the physiological processes of multiple organ systems coor-
dinate with each other, especially when homeostasis is challenged.
4. A Focus on Problem Solving
One of the most valuable life skills students
should acquire is the ability to think critically and
use information to solve problems. As you study
physiology, you should be prepared to practice
these skills. You will find a number of features in this book, such
as the Concept Check questions and figure and graph questions,
that are designed to challenge your critical thinking and analysis
skills. In each chapter, read the Running Problem as you work It is my hope that by reading this book, you will develop
through the text and see if you can apply what you’re reading to an integrated view of physiology that allows you to enter your
the clinical scenario described in the problem. chosen profession with respect for the complexity of the human
Also, be sure to look at the back of the text, where we have body and a clear vision of the potential of physiological and bio-
combined the index and glossary to save time when you are medical research. May you find physiology as fun and exciting I
looking up unfamiliar words. The appendices have the answers do. Good luck with your studies!
to the end-of-chapter questions, as well as reviews of physics,
logarithms, and basic genetics. The back end papers include a
periodic table of the elements, diagrams of anatomical positions Warmest regards,
of the body, and tables with conversions and normal values
of blood components. Take a few minutes to look at all these Dr. Dee (as my students call me)
features so that you can make optimum use of them. silverthorn@mail.utexas.edu

xix
xx
Engaging Art Helps You Visualize Processes and Concepts—

NEW REVIEW ART FEATURES visually pull together foundational concepts so you can
quickly review key topics.

Fig. 2.6 R E V I E W

Molecular Bonds
When two or more atoms link by sharing electrons, they make units known as
molecules. The transfer of electrons from one atom to another or the sharing of
electrons by two atoms is a critical part of forming bonds, the links between atoms.

Covalent Bonds
Covalent bonds result when atoms share electrons. These
bonds require the most energy to make or break.

(a) Nonpolar Molecules

Nonpolar molecules have an even Hydrogen

Fatty acid
distribution of electrons. For example,
molecules composed mostly of carbon
and hydrogen tend to be nonpolar. Carbon

(b) Polar Molecules

Bonds
Polar molecules have regions of Negative pole - -
partial charge (δ+ or δ– ). The most Water molecule -
-

important example of a polar δ– δ– -

O
molecule is water. -
- -
- -
H H

δ+ δ+ O
H O H =
H H = H2O
Positive pole

Noncovalent Bonds

(c) Ionic Bonds

Ionic bonds are electrostatic attractions between ions. A common example is sodium chloride.

+ –

Na CI Na CI

Sodium ion (Na+ )

Sodium atom Chlorine atom Chloride ion (CI– )
Sodium gives up its one weakly held electron The sodium and chloride ions both have stable
to chlorine, creating sodium and chloride ions, outer shells that are filled with electrons. Because
Na+ and Cl-. of their opposite charges, they are attracted to
each other and, in the solid state, the ionic bonds
form a sodium chloride (NaCl) crystal.

(d) Hydrogen Bonds

Hydrogen bonds form between

a hydrogen atom and a nearby
oxygen, nitrogen, or fluorine
atom. So, for example, the
polar regions of adjacent
water molecules allow them to
form hydrogen bonds with Hydrogen
one another. bonding Hydrogen bonding
between water molecules
is responsible for the
surface tension of water.

(e) Van der Waals Forces

Van der Waals forces are weak, nonspecific attractions between atoms.

xx
and Focus on the Key Ideas

NEW ESSENTIALS ART FEATURES show each chapter’s core concepts, helping you
connect ideas visually and see the big picture of human physiology.

Fig. 7.3 E S S E N T I A L S

Peptide Hormone Synthesis and Processing

Peptide hormones are made as large, inactive preprohormones
that include a signal sequence, one or more copies of the
hormone, and additional peptide fragments.

(a) Preprohormones
1 mRNA 1 Messenger RNA on the
ribosomes binds amino
PreproTRH (thyrotropin-releasing acids into a peptide
hormone) has six copies of the chain called a
3-amino acid hormone TRH. Preprohormone Ribosome
preprohormone. The
Preprohormone chain is directed into
Endoplasmic the ER lumen by a
reticulum (ER) signal sequence of
PreproTRH (242 amino acids) 2 amino acids.

processes to
Signal Prohormone Transport
sequence vesicle
2 Enzymes in the ER
chop off the signal
6 TRH sequence, creating an
(3 amino acids each) inactive prohormone.
+

3
Other peptide fragments 3 The prohormone
passes from the ER
+ through the Golgi
complex.
Signal sequence Golgi
complex
4 Secretory vesicles
containing enzymes
and prohormone bud
(b) Prohormones off the Golgi. The
enzymes chop the
Prohormones, such as pro- prohormone into one
opiomelanocortin, the prohormone for or more active
ACTH, may contain several peptide peptides plus
sequences with biological activity. 4
Secretory additional peptide
vesicle Active hormone fragments.

Pro-opiomelanocortin Cytoplasm Peptide fragment

Release
processes to
5 The secretory vesicle
signal releases its contents
by exocytosis into the
ECF extracellular space.
ACTH 𝛄 lipotropin 𝛃 endorphin 5

+
Plasma Capillary 6 The hormone moves
Peptide fragment 6 To target into the circulation for
endothelium
transport to its target.

(c) Prohormones Process to Active Hormone Plus Peptide Fragments

The peptide chain of

insulin's prohormone
folds back on itself with
the help of disulfide Proinsulin
(S—S) bonds. The
prohormone cleaves to
processes to
Insulin + C-peptide
S S S
insulin and C-peptide. S S
S
S S
S S
S
S

xxi
Everyday Applications Encourage Critical Thinking
RUNNING PROBLEM

RUNNING PROBLEMS begin each What to Believe?

chapter with a problem involving a disease Jimmy had just left his first physiology class when he got the
text from his mother: Please call. Need to ask you something.
or disorder that unfolds in segments on His mother seldom texted, so Jimmy figured it must be
important. “Hi, Mom! What’s going on?”
R U N N I N G P R O B L E M CO N C LU S I O N (continued)
subsequent pages. The questions in each “Oh, Jimmy, I don’t know what to do. I saw the doctor this
morning and he’s telling me that I need to take insulin. But
segment ask you to apply information you Question Answer and Commentary I don’t want to! My type of diabetes doesn’t need insulin.
I think he’s just trying to make me see him more by putting
have learned in the text. You can check your 2. What kinds of web sites
should Jimmy be looking for in
The best web sites for health information come from organizations that are part of the scientific
and health-care communities, such as the Nationalme on insulin.
Institutes Don’t (NIH),
of Health you think I’m right?”
nonprofi t groups dedi-
his results list, and how can he cated to supporting research on a particular disease (The paused
American forDiabetes Association, diabetes.
understanding by comparing your answers recognize them?
Jimmy a moment. “I’m not sure,
org), or clinics and universities where scientists and physicians are actively investigating causes and
probably
Mom. He’s
treatments for diseases. Treat commercial websites that endjust tryingwith
in *.com to do what’s
extra best for you. Didn’t you talk
caution.
with those in the Problem Conclusion at the 3. In The Doctors’ Medical Li-
to him about it?”
Dr. Kennedy claims that some patients with type 2 diabetes can be “successfully treated” by eat-
brary article called “Fiber,” what ing a high-fiber diet. (The classification of type 2“Well,
diabetesI tried but heonset”
as “adult didn’t have time to talk. You’re studying
is obsolete.)
end of each chapter. Three new Running does Dr. Kennedy say about
high-fiber diet and diabetes?
these things. Can’t you look it up and see if I really need
insulin?”
Problems appear in this edition: one about 4. How can Jimmy find out To learn more about who created a web site and“Iwhy,
guesslook
so. for
Letlinks at the
me see bottom
what I can fiof
ndthe page
out.” Jimmy hung up
more about who created the for HOME or ABOUT US. On the home page for The andDoctors’
thought.Medical Library you will learn that
finding reliable information on the internet, site and what their credentials
are?
“Now what?”
the site promotes “natural health care.” The link on Ron Kennedy, M.D., does not give you any
information about his medical training or other credentials.
(chapter 1), one on the human papillomavirus 5. Compare the number of The number of results will depend on when you do the2search 8because 10new articles
13 are
18 added 24 27
results from the PubMed constantly. But the number will probably be fewer than 50,000, far less than the millions of re-
and cervical cancer (chapter 24), and one on search to those for the Google
searches.
sults that came up following a Google search.

cholera in Haiti (chapter 21). 6. What did Jimmy tell his The articles published by these national organizations all say that people with type 2 diabetes
mother about her need to take may need to take insulin. Patients should always listen to their health-care providers and ask
insulin for her type 2 diabetes? questions if they are uncertain about what they should be doing.

7. Do the articles from NC-
CAM mention dietary fiber as
an alternative treatment for
diabetes?
The NCCAM article lists a number of alternative treatments that people have tried, but dietary
fiber is not among them. It also says that, so far, there is no scientific evidence supporting the
use of dietary supplements for treating diabetes. Patients should never stop their conven-
tional treatments when using complementary treatments, and they should always inform their
health-care providers about any vitamins or dietary supplements they are taking.

EMERGING CONCEPTS

Transporter Gene Families

One outcome of the Human Genome project has
been the recognition that many proteins are closely
related to each other, both within and across species. As
a result, scientists have discovered that most membrane
transporters for organic solutes belong to one of two
gene “superfamilies”: the ATP-binding cassette (ABC)
superfamily or the solute carrier (SLC) superfamily. The
ABC family transporters use ATP’s energy to transport
small organic molecules across membranes. Interestingly,
B I O T E C H N O LO G Y the CFTR chloride channel, which causes cystic fibrosis FOCUS BOXES highlight research
when absent or defective, is also a member of the
ABC family and is the only known ion channel in that in physiology and medicine.
Calcium Signals Glow in the Dark superfamily. The 43 families of the SLC superfamily family
include
oceanmost facilitated diffusion carriers, such as the
If you’ve ever run your hand through a tropical
GLUT
and seen the glow of bioluminescent jellyfish,
at night
sugar
you’ve transporters, as well as the secondary active
seen
Three kinds of focus boxes help you
transporters
a calcium signal. Aequorin, a protein complex isolated fromshown in Table 5.8.
understand the role of physiology
jellyfish, is one of the molecules that scientists use to monitor
the presence of calcium ions during a cellular response. When in science and medicine today:
aequorin combines with calcium, it releases light that can
be measured by electronic detection systems. Since the first Biotechnology boxes discuss
use of aequorin in 1967, researchers have been designing
increasingly sophisticated indicators that allow them to
applications and laboratory techniques
follow calcium signals in cells. With the help of molecules
C LI NI C A L F O C U S
called fura, Oregon green, BAPTA, and chameleons, we can
from the rapidly changing world of
now watch calcium ions diffuse through gap junctions and biotechnology; Clinical Focus boxes
flow out of intracellularLDL: The Lethal Lipoprotein
organelles. The sea nettle Chrysaora fuscescens.
“Limit the amount of cholesterol in your diet!” has
concentrate on clinical applications and
been the recommendation for many years. So why is
too much cholesterol bad for you? After all, cholesterol
pathologies; and Emerging Concepts
molecules are essential for membrane structure and for boxes describe upcoming advances in
making steroid hormones (such as the sex hormones). But
elevated cholesterol levels in the blood also lead to heart dis- physiological research.
ease. One reason some people have too much cholesterol in
their blood (hypercholesterolemia) is not diet but the failure
of cells to take up the cholesterol. In the blood, hydrophobic
cholesterol is bound to a lipoprotein carrier molecule to
make it water soluble. The most common form of carrier is
low-density lipoprotein (LDL). When the LDL-cholesterol com-
plex (LDL-C) binds to LDL receptors in caveolae, then it can
then enter the cell in a vesicle. When people do not have ad-
equate numbers of LDL receptors on their cell membranes,
LDL-C remains in the blood. Hypercholesterolemia due to
high levels of LDL-C predisposes these people to the devel-
opment of atherosclerosis, also known as hardening of the
arteries {atheroma, a tumor + skleros, hard + -sis, condition}.
In this condition, the accumulation of cholesterol in blood
vessels blocks blood flow and contributes to heart attacks.

xxii
Insightful Pedagogy Helps You Make Connections
THE REFLEX CONTROL OF VENTILATION

REFLEX PATHWAYS & CONCEPT MAPS Central and peripheral chemoreceptors monitor blood gases and pH.
Control networks in the brain stem regulate activity in somatic motor

organize physiological processes and details into a neurons leading to respiratory muscles. Emotions
and voluntary CO2 O2 and pH
control
logical, visual format. These figures use consistent 16

colors and shapes to represent processes and will 15

1
Higher
brain
Medullary
chemoreceptors
Carotid and aortic
chemoreceptors
2 centers
14
guide you to a better understanding of coordinated 13
3

physiological function. 4
Limbic
system
Afferent sensory
neurons
12
5

6 SPINAL REFLEXES

BACKGROUND BASICS , found on the chapter In a spinal refex, sensory information entering the spinal cord is acted
on without input from the brain. However, sensory information about

opening page, lists topics you will need to master 7

8
theoblongata
Medulla stimulus may be sent to the brain.
and pons

11
for understanding the material that follows. Page
references save study time, making the textbook an
Somatic Somatic
easy-to-use resource. 10
motor neurons
(inspiration)
motor neurons
(expiration)

9
Spinal Stimulus
cord
Scalene and External Internal Abdominal
Inspiration Expiration Diaphragm
sternocleidomastoid intercostals intercostals muscles
muscles
Sensory
information
KEY
FIGURE QUESTION
Integrating
Match the numbers on the Stimuli center Integrating centers
Interneuron
figure to the boxes of the map.
Sensors Efferent neurons

Afferent neurons Targets

Command to
A spinal reflex initiates muscles or
a response without input glands
from the brain.

Response

CONCEPT LINKS are blue page numbers [p. 321]

p p p
response by means of a signal transduction system ( Fig. 7.4). embedded in the text. They connect the concepts you
Many peptide hormones work through cAMP second messen- are reading about to topics discussed earlier in the book.
ger systems [p. 183]. A few peptide hormone receptors, such as Concept Links help you find material that you may have
that of insulin, have tyrosine kinase activity [p. 183] or work
forgotten or that may be helpful in understanding new
information.

(b) Filtration in systemic capillaries FIGURE AND GRAPH QUESTIONS promote

Net pressure = hydrostatic pressure (PH ) – colloid osmotic pressure (␲) KEY
analytical skills by encouraging you to interpret and apply
PH = 32 mm Hg PH = 25 mm Hg
PH = Hydrostatic pressure forces
fluid out of the capillary.
information presented in the art and graphs. Answers to
␲ = 25 mm Hg PH > ␲ PH < ␲ ␲ = 15 mm Hg
PH = ␲ ␲ = Colloid osmotic pressure of
proteins within the capillary
these questions appear at the end of each chapter.
pulls fluid into the capillary.
7200 L
day

FIGURE QUESTION Concept Check Answers: p. 000

Net filtration Net absorption
Suppose that the hydrostatic pressure PH at

Net flow out = 3 L/day

the arterial end of a capillary increases from
32 mm Hg to 35 mm Hg. If PH remains 15 mm
17. Why must insulin be administered as a shot and not as an oral pill?
Hg at the venous end, does net filtration in this
capillary decrease, increase, or stay the same? 18. Patients admitted to the hospital with acute diabetic ketoacidosis and
36(+=,36*0;@9,3(;065:/0705:2,3,;(34<:*3, dehydration are given insulin and fluids that contain K+ and other ions.
The acidosis is usually accompanied by hyperkalemia, so why is K+
A included in the rehydration fluids? (Hint: dehydrated patients may have
.9(7/8<,:;065: a high concentration of K+, but their total body fluid volume is low.)
1. At what point on the line is
contraction isometric?
19. In 2006 the FDA approved sitagliptin (Januvia®), a DPP4 inhibitor. This
Velocity of shortening

2. At what point is the muscle

contraction at maximum velocity? drug blocks action of the enzyme dipeptidyl peptidase-4, which
breaks down GLP-1 and GIP. Explain how sitagliptin is helpful in treating
diabetes.

0 B
Load on the muscle CONCEPT CHECK BOXES are placed at
intervals throughout the chapters, helping to test your
understanding before continuing to the next topic. You
can check your own answers using the key at the end of
every chapter.
xxiii
 FOCUS ON… figures highlight the anatomy
Fig. 7.16 F O C U S O N . . .
and physiology of important organs that are often
overlooked in physiology texts, including the skin, The Pineal Gland

the liver, the spleen, and the pineal gland. The pineal gland is a pea-sized structure
buried deep in the brain of humans.
Nearly 2000 years ago, this "seat of the
soul" was thought to act as a valve that
regulated the flow of vital spirits and
knowledge into the brain. By 1950,
Corpus however, scientists had decided that it
callosum was a vestigial structure with no
known function.
Thalamus

O
CH3O
CH2 CH2 NH C CH3

N
H

Melatonin is an amino acid–derived

hormone made from tryptophan.

ANATOMY SUMMARIES provide succinct 50

About 1957 one of the wonderful coincidences of scientific research
occurred. An investigator heard about a factor in beef pineal glands that

visual overviews of a physiological system from could lighten the skin of amphibians. Using the classical methodology of
endocrinology, he obtained pineal glands from a slaughterhouse and
started making extracts. His biological assay consisted of dropping
40

a macro to micro perspective. Whether you are pineal extracts into bowls of live tadpoles to see if their skin color

Melatonin (pg/mL plasma)

blanched. Several years and hundreds of thousands of pineal glands
later, he had isolated a small amount of melatonin.
30
learning the anatomy for the first time or refreshing Fifty years later, we are still learning about the functions of melatonin in
humans. In addition to its role in sleep-wake cycles and the body’s
internal clock, scientists have evidence that melatonin is a powerful

your memory, these summaries show you the 20

antioxidant. Some studies using mouse models of Alzheimer’s disease
suggest that melatonin may help slow the progression of the disease.
Melatonin has also been linked to sexual function, the onset of puberty,

essential features of each system in a single figure. 10 and depression in the darker winter months (seasonal affective disorder,
or SAD). In 2011 there were over 100 active clinical trials in the United
States testing the efficacy of melatonin in treating disorders associated
0 with sleep disturbances and depression.

6 P.M. 6 A.M. Noon 3 P.M. In 2009 European authorities approved the use of a melatonin receptor
Noon Midnight
agonist, agomelatine, for treating major depression. The U.S. Food and
Drug Administration has been slower to approve the drug, and it is
Melatonin is the "darkness hormone," secreted at night as we currently being tested in Phase II and Phase III clinical trials in the United
sleep. It is the chemical messenger that transmits information States. Phase II trials are usually placebo-controlled, double-blind
Fig. 14.5 A N A T O M Y S U M M A R Y about light-dark cycles to the brain center that governs the studies [p. 00]. Phase III trials include more patients and some
body's biological clock. uncontrolled studies. Some Phase III studies are “open-label,” meaning
that the patients and healthcare providers know what drug is being
(Adapted from J. Arendt, Clin. Endocrinol. 29:205-229, 1988.) administered.
The Heart
(a) The heart lies in the center of the thorax. Anatomy of the Thoracic Cavity Structure of the Heart
Position of (c) The heart is on the ventral side of the thoracic (e) The heart is encased within a membranous (f) The ventricles occupy the bulk of the heart. The arteries
Sternum semilunar valves cavity, sandwiched between the lungs. fluid-filled sac, the pericardium. and veins all attach to the base of the heart.

Trachea
Thyroid gland
Lung
First rib (cut) Aorta
Base of
heart

Apex of Pericardium Superior

heart vena cava Pulmonary artery

Position Auricle of left atrium

of AV
valves Coronary artery
Diaphragm
Diaphragm Right and vein
atrium

(b) Vessels that carry well- Diaphragm Apex of

oxygenated blood are heart Right
red; those with less ventricle
Left ventricle
well-oxygenated blood
are blue.
(g) One-way flow through the heart
is ensured by two sets of valves.

(d) Superior view of transverse plane in (b)

Superior Aorta (segment Pulmonary Pulmonary

Bronchus vena cava Esophagus removed) artery vein Pulmonary
Heart
Aorta semilunar valve
Aorta
Vena
Right
cava
pulmonary Left pulmonary
arteries arteries

Superior Left pulmonary (h) Myocardial muscle cells are branched, have
vena cava veins a single nucleus, and are attached to each
other by specialized junctions known as
intercalated disks.
Right atrium Left atrium
Intercalated Myocardial muscle
Cusp of left AV disks cell
(bicuspid) valve

Chordae tendineae
Cusp of right
AV (tricuspid)
Papillary muscles
valve

Right Right Pericardium Sternum Pericardial Left Left

atrium ventricle cavity ventricle atrium Right Left ventricle
ventricle

Inferior
vena cava

Descending aorta

xxiv
Another random document with
no related content on Scribd:
 CHARGE OF THE COSSACKS
Hai! roll up! Eagles brave,
To protect “the Tchighka” (Tchyka[24])
And gain glory newly.
Nobles all!
Or we fall.
Twice we die not, truly—
Hai! Take arms. On we go!

From our rifles we shall shout,

We shall roar from cannon,
With our sabres clashing—
Nobles all,
Or we fall!
’Gainst our foemen dashing.
Hai! Take arms. On we go!
 THE YOUNG RECRUITS
Along the hills lies the snow,
But the streams they melt and flow;
By the road the poppies blow—
Poppies? Nay, scarlet though they glow
These are no flowers—the young recruits!
They are the young recruits!

To Krym, to Krym they ride,

The soldiers, side by side—
And over the country wide
Sounds the beat of the horse’s stride.

One calls to her soldier son:

“Return, O careless one!
Of scrubbing wilt have none?
Let me wash thy head—then run!”

“Nay, mother, wash thine own,

Or make my sister groan.
Leave thou thy son alone!
Too swift the time has flown.

“My head the fine spring rain

Will soon wash clean again,
And stout thorns will be fain
To comb what rough has lain.

“The sun will make it dry,

Wind-parted it will lie—
So, mother mine, good-bye!”

· · · · ·

He could not hear her cry.

 MOTHER AND SON
(This song was composed before 1648)

All the oak forest is murmuring, murmuring:

Thick veils of fog o’er the fields and wide meadows cling.

“Go away, my son, from me—

May the raiding Turk take thee!”

“Mother, well the Sultan knows

Thy brave son. (This witness shows.)

“For he pays me from the mine

Tribute—gold and silver fine!”

“Go away, my son, from me—

May Litvà[25] soon capture thee!”

“Litvà knows me too—I feed

From her tribute, wine and mead.”

“Go away, my son, from me,

May the Tartars soon take thee!”

“Those wild Hordes take, in much fear,

Other roads when I draw near!”

“Go away, my son, from me—

Moscow! Let the Tzar take thee!”

“But the Tzar likes me so well,

With him I’ve been asked to dwell!”

“Ah, my son, come home instead.

Let me, dear one, wash thy head.”
 “Nay, my mother, nay. With rain
Washing it I’ll not complain.

“Winds will dry my dripping hair;

Teren-bush[26] will comb it fair.”

All the deebrova[27] is murmuring, murmuring—

Leaden clouds over heaven lowering masses fling.

“Farewell!” the sisters cry—for he must go with speed.

She who is eldest born leads out his splendid steed.

And then the second-born armour brings out to him:

Youngest of all entreats—asks with her eyes tear-dim:

“When, O my brother dear, comest thou back to us?”

“Ah, sister! Of the sand take thou a handful thus....

“Sow on a rock. Each dawn water it with thy tears.

That day the sand springs up—thy brother lost appears!”
 THE CAPTIVES
Cuckoo! calls the Cuckoo....

In the dawn, in the dawn the young Cossacks are crying,

Far away from their loves, in prison lying,
The dungeon’s dark, their hope is gone,
But the Cuckoo calls, in the dawn, in the dawn!

Blows the wind, blows the wind—From the sea were it blowing
’Twould bear us away beyond all knowing!
Our heavy chains we’d leave behind
If over the sea should come the wind.

O the sun! O the sun in Ukraine shining!

Take us to where our loves are pining....
The Cossacks have their dance begun,
The dance of joy, in the sun, in the sun.

Blue sea! On the sea with the wind they’re dancing—

Our brothers surely are advancing
From prison chains the sad to free.
O swiftly come, o’er the sea, o’er the sea!

Cuckoo, calls the Cuckoo....

In the dawn, in the dawn the Sultan sleeping

Is wakened by the sound of weeping—
“Bind stronger chains their limbs upon
That none may flee, in the dawn, in the dawn!”
 COSSACK MARCHING SONG
(Semi-historical)

The Harvesters are reaping on the hill-side,

And in the valley where the grass is green
The Cossacks leap astride their horses lean.

That gallant hetman, Doroshonko,

Is leading all his troop with right good-will—
Over at last the weary days of drill!

And see that captain stationed in the centre,

His steed is prancing, pawing up the ground ...
Brave Sahaidachni, at the rear, looks round.

In fair exchange for pipe and for tobacco

He’s said adieu to Priska, his good wife—
“Such a mistake! The greatest of my life!”

So is he thinking when he hears one calling:

“Come back, come back and take your wife once more;
My pipe and my tobacco please restore!”

“Ah, ha!” he shouts, “a wife I’ll not be needing—

But your carved pipe is handy on the road.
What a fine thing you have on me bestowed!

“Hai! Who goes there? Pass, friend—and on we’re faring;

With flint and steel I’ll get a puff or two,
So then—don’t worry—and good-bye to you.”
 SONG OF VICTORY—1648
When the Cossacks under Khmelnitzky expelled the Poles from the
Ukraine

Hai, all ye good people! list what I tell ye,

What’s done in Ukraina’s plain—
There under Dashiev, across the Soroka,
What numbers of Poles now lie slain.

Hai, Perebiynees! But seven hundred

Cossacks he asked for that day.
Then he with sabres smote the Poles’ heads off—
The rest swept the river away.

Drink ye swamp water, Oi! all ye Poles now—

Quench thirst at each rain-pond ye see....
And once ye were drinking, in our Ukraina,
Wine and sweet mead flowing free!

Each Polish “Pan” is lost now in wonder;

“What do these brave Cossacks eat?
Verily, look ye, they live just on pike-flesh,
Solamàkha[28] with water their meat.”

Look now, ye Poles, whom our Hetman Khmelnitzky

Fought on the grim “Yellow Sands,”
Of all your army fighting young Cossacks
Not one has escaped from our hands.

See, Pole! A Cossack is dancing, is dancing

Upon a grey horse after thee!
When he stands with his musket thy heart sinks in anguish
In great fear of death thou dost flee.

We own the whole land e’en as far as Sluch river,

 Kostiana! As far as thy Hill—
O rude and uncourteous! Poles caused our revolting
So mourn they their lost Ukraine still.

As a thunder-cloud brooding on Vistula’s river

The Poles lie, expelled from Ukraine.
As long as we live they shall no more leave Poland,
They shall not come nigh us again.

Hai! All ye young Cossacks! Leap up now with shouting—

Akimbo our arms let us place.
We threw all the Poles across Vistula’s waters—
And here they won’t dare show their face!
 IN TURKISH CAPTIVITY
On the blue sea waves are roaring,
Mountain high they tower.
Crying in their Turkish dungeon
Wretched Cossacks cower.

“Why, O gracious God, this torture?

Two years now we lie here;
With the chains our hands are heavy—
Wilt Thou let us die here?

“Wings of Ukraina’s Eaglets,

Yanichars[29] cut, throwing
In the grave the living victims,
All their sorrow knowing.

“Hai! Ye youthful Zaporogians,[30]

Have ye not arisen?
Sons of Freedom, ever glorious,
Rescue us from prison!”
 LAMENT FOR MOROZENKO
(This Cossack song of the seventeenth century is sung to a
mournful air which makes a splendid funeral march.
Morozenko was an Ukrainian Governor of a province killed in
war with the Tartars.)

Trenches along the foot of the mountain—

They took Morozenko on Sunday morning.
The Tartars nor slashed him, nor pierced him with spears;
They tore out the heart from the white, white breast,
And they led him to Savour-Mohyla’s height:
“Look thou, O son of the foe, down there!
Look on thine Ukraine stretching far!”
They set him down on the yellow sands,
And they took off from him a red, red shirt.[31]

Oi, Moroze, Morozenko!

Thou glorious Cossack.
All Ukraine laments thee,
O brave Morozenko!
Much more thy bold army,
O glorious Cossack!

· · · · ·

On the way to the town Morozikha wept—

Sore wept Morozikha for her son.

“Don’t cry, Morozikha, don’t be sad.

Come with us Cossacks to drink wine-mead.”

“Drink your good health, if drink you would,

But around my head misfortune flies.
Drink your good health, if drink you may....
Oi, where does he fight, my son, my son!
Does he fight with the Tartars, one by one?”

“Don’t cry, Morozikha, don’t be sad;

Come with us Cossacks to look on....
For see! A horse walks behind a wagon,
A bloody wagon it walks behind.
It carries your glorious Moroze,
The white flesh cut, the brave head broken,
The face is covered with red kitayka.[32]...”

Oi, Moroze, Morozenko!

Thou glorious Cossack—
All Ukraine shall weep
And mourn for thee.
ROBBER SONGS
 [33]
THE DEATH OF DOBUSH
1745

Along the edges of the wooded height

Walks young Dobush;
Lame in one leg, he on his topir[34] leans
And calls his lads:

“O, ye Legini,[35] O, my boys!

We’ll council hold
Whom next are we to rob?
Kooty we must not miss,
Nor overlook Kossiev.[36]
Now sleep, my boys,
Because we rise at dawn;
Dress in a trice, skin postoli[37] put on,
Povoloki[38] of silk....

“Now run, boys—quick!

Snow covers all the paths;
To Dzveenka’s house go first,
Where we’ll see Stefan’s wife.”

“Oi, Dobush! Nay, my lord,

Sure mischief will befall.”

“Don’t trouble about me;

Load your good musket with
A double charge—stand by the gate—
I’ll to the window go
To see if she still sleeps.”

“My heart, dost thou sleep,

Dost thou hear?
Dost thou wish to receive Dobush?”
“I am not asleep. No. I hear
Each word that you say to me.
I’m working that I may sup—
Stefan is not at home....
The supper’s not ready yet,
But ’twill be a splendid one,
And a wonder for all the world.”

“Dost thou sleep, my heart,

Dost thou hear?
Wilt receive Dobush for the night?”

“I sleep not—I hear every word—

I will not let the robber in.”

“Wilt thou open the door, I say?

Dost tell me to storm it then?”

“I give no command to storm.

But—open it? No, not I.”

“Let me into the hut—thou fool!

Ere I break open the door.”

“My door is too strong for you—

My locks are of trusty steel.”

“Thy locks will not help thee much

When to them my shoulder I set.”

“The strength of full seven more years

You’ll need ere you burst my door.”

Dobush, Dobush pressed hard—

The locks fell in a heap,
And Dobush opened the door,
Just a little opened the door....

And then Dzveenka fired

From the attic where he hid,
He aimed at the heart of Dobush.

Not in the heart fell the blow;

Through shoulders the bright blood burst.

“Dog-catcher! You! Dzveenchuk!

You have eaten me up for her.”

“Why did you woo her? Why

Did you say you were Dobush?
Why tell her all the truth?
Knew you not woman’s truth
Is fast-running water’s foam?”

The Oprishki came to the hut

But they found Dzveenka was flown.

“Oi! Dobush, our good lord,

Why killed you not the wife?”

“How could I kill her, say,

If I loved her so much?”

“Oi, Dobush, our great lord!

Misfortune’s surely here.
Treachery ne’er before
To your Legini came,
But now there’s treachery.”

“Legini, Oi! my boys,

Lay me on your topirs,
Carry me down in the Chorna-Hora,
Where the Black Mountains be,
Then cut my body up as fine as poppy seed.
Let not the Germans mock,
Or quarter my body.

“Divide among yourselves the treasure that was ours—

Then singly go away.
But not to rob—
Not to shed human blood;
Blood is not water, mind,
Not meant to be poured down!”

But then the Germans came,

And Dzveenka led them on.

“Oi, Oi, Dobush, our lord,

What woeful fate is ours!
Where shall we winter spend,
Where all the summer days?”

“In Stanislav, my boys,

Yea, at the market-place!
Tortured, while, bound in irons,
Germans shall tear your flesh,
And there you’ll sleep for aye.”
 [39]
SONG OF THE OPRISHKI —(OUTLAWS)
Hai, Brethren, Oprishki—give me more horeevka![40]
On the camp-fire now heap on more wood.
If you tuned then my throat to the sound of Sopeevka,[41]
I’d sing for as long as I could.

We are safe just as long as the green grass is growing—

If the forest of leaves be not bare,
If behind the thick bush and green pine we are going,
Even Chorts[42] could not find us hid there.

As the heaven for birds, so for us are the hollows,

The caves in Carpathian crests.
We sleep till the stars, till our own shadow follows,
And then we creep out of our nests.

Tobacco we bring from far Hungary’s borders

(Fleet horsemen their chase may give o’er),
The Jew merchant clothing shall give at our orders,
Or else he’ll be nailed to his door.

Be joyful, my brothers, each day that is ours,

No life such as this can last long.
When snow falls our heads will hang down like the flowers;
No more shall be heard our glad song.

For Austrian soldiers, when first snow is falling,

In uniforms white will appear....
Kolomea![43] Thy bells as of old may be calling—
Their chiming we never shall hear.
 THE HAIDAMAKY—“KNIGHTS OF
VENGEANCE”
“Haidamaky” they call us, unrelenting and stern,
With the wrongs of our nation for vengeance we burn.

Our forebears were tortured; our grandsons shall be

Unless we will show them how men may be free.

Haidamaky they call us, forever the same,

And we lay down our lives, caring nothing for fame.

For the time long has passed when the yoke pressed us sore:
If a hundred shall fall there are yet thousands more.

Out of misery’s chains the trampled slaves rise,

And to Freedom’s bright flag they will lift dazzled eyes.

Truth and courage for oath, and our Vengeance for breath—
Haidamaky they call us, men who fear not their death.
 SONG OF KARMELŪK
(Written by himself)

From Siberia I return—

With no fortune I am come.
Not in chains, but yet not free.
Wife and children may be mine,
But their faces I can’t see.
When I think upon their fate
Then I weep most bitterly.

Good lads have I gathered round

(What concerns it any one?),
By the road lie on the ground!
Riders, when will ye pass by?
Tedious it is to wait—
No abode, no hut have I.
The police won’t make me wince
Though Assessors scan each nook,
Hunt in every likely den.
They themselves have killed more men
Than your Karmelūk has sins!

“Robber!” so good folk may cry—

“Murderer!” But I’ve killed none.
For, look ye, I have a soul.
I may take from rich my toll
(And I’ll do it too, be sure!),
Free from sin is Karmelūk
For he gives it to the poor.

Rising o’er Siberia

Shines the sun. Keep watch you must.
Yet in me put all your trust.
Rest your hopes on Karmelūk!
TCHUMAK SONGS