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Dedication to
William Francis Ganong

illiam Francis (“Fran”) Ganong was an outstanding scientist, educator,

W and writer. He was completely dedicated to the field of physiology and

medical education in general. Chairman of the Department of
Physiology at the University of California, San Francisco, for many years, he
received numerous teaching awards and loved working with medical students.
Over the course of 40 years and some 22 editions, he was the sole author of
the best selling Review of Medical Physiology, and a co-author of 5 editions of
Pathophysiology of Disease: An Introduction to Clinical Medicine. He was one
of the “deans” of the Lange group of authors who produced concise medical text
and review books that to this day remain extraordinarily popular in print and
now in digital formats. Dr. Ganong made a gigantic impact on the education of
countless medical students and clinicians.
A general physiologist par excellence and a neuroendocrine physiologist by
subspecialty, Fran developed and maintained a rare understanding of the entire
field of physiology. This allowed him to write each new edition (every 2 years!)
of the Review of Medical Physiology as a sole author, a feat remarked on and
admired whenever the book came up for discussion among physiologists. He
was an excellent writer and far ahead of his time with his objective of distilling a
complex subject into a concise presentation. Like his good friend, Dr. Jack
Lange, founder of the Lange series of books, Fran took great pride in the many
different translations of the Review of Medical Physiology and was always
delighted to receive a copy of the new edition in any language.
He was a model author, organized, dedicated, and enthusiastic. His book was
his pride and joy and like other best-selling authors, he would work on the next
edition seemingly every day, updating references, rewriting as needed, and
always ready and on time when the next edition was due to the publisher. He did
the same with his other book, Pathophysiology of Disease: An Introduction to
Clinical Medicine, a book that he worked on meticulously in the years following
his formal retirement and appointment as an emeritus professor at UCSF.

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 Fran Ganong will always have a seat at the head table of the greats of the art
of medical science education and communication. He died on December 23,
2007. All of us who knew him and worked with him miss him greatly.

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About the Authors

KIM E. BARRETT

Kim Barrett received her PhD in biological chemistry from University College
London in 1982. Following postdoctoral training at the National Institutes of
Health, she joined the faculty at the University of California, San Diego, School
of Medicine in 1985, rising to the rank of Professor of Medicine in 1996, and
was named Distinguished Professor of Medicine in 2015. From 2006 to 2016,
she also served the University as Dean of the Graduate Division. Her research
interests focus on the physiology and pathophysiology of the intestinal
epithelium, and how its function is altered by commensal, probiotic, and
pathogenic bacteria as well as in specific disease states, such as inflammatory
bowel diseases. She has published more than 250 articles, chapters, and reviews,
and has received several honors for her research accomplishments including the
Bowditch and Davenport Lectureships from the American Physiological Society
(APS), the Bayliss-Starling Lectureship from The Physiological Society of the
UK and Ireland, and the degree of Doctor of Medical Sciences, honoris causa,
from Queens University, Belfast. She has been very active in scholarly editing,
serving currently as the Editor-in-Chief of the Journal of Physiology. She is also
a dedicated and award-winning instructor of medical, pharmacy, and graduate
students, and has taught various topics in medical and systems physiology to

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these groups for more than 30 years. Her efforts as a teacher and mentor were
recognized with the Bodil M. Schmidt- Nielson Distinguished Mentor and
Scientist Award from the APS in 2012, and she also served as the 86th APS
President from 2013 to 2014. Her teaching experiences led her to author a prior
volume (Gastrointestinal Physiology, McGraw-Hill, 2005; second edition
published in 2014) and she was honored to have been invited to take over the
helm of Ganong in 2007 for the 23rd and subsequent editions, including this one.

SUSAN M. BARMAN

Susan Barman received her PhD in physiology from Loyola University School
of Medicine in Maywood, Illinois. Afterward she went to Michigan State
University (MSU) where she is currently a Professor in the Department of
Pharmacology/Toxicology and the Neuroscience Program. She is also Chair of
the Institutional Animal Care and Use Committee and serves on the College of
Human Medicine (CHM) Curriculum Development Group for medical school
education. She has had a career-long interest in neural control of
cardiorespiratory function with an emphasis on the characterization and origin of
the naturally occurring discharges of sympathetic and phrenic nerves. She has
published about 150 research articles, invited review articles, and book chapters.
She was a recipient of a prestigious National Institutes of Health MERIT
(Method to Extend Research in Time) Award. She is also a recipient of an MSU
Outstanding University Woman Faculty Award, a CHM Distinguished Faculty
Award, a Distinguished Service Award from the Association of Chairs of
Departments of Physiology, and the Carl Ludwig Distinguished Lecture Award
from the Neural Control of Autonomic Regulation section of the American
Physiological Society (APS). She is also a Fellow of the APS and served as its
85th President. She has also served as a Councilor of APS and Chair of the
Women in Physiology and Section Advisory Committees of the APS. She is also
active in the Michigan Physiological Society, a chapter of the APS.

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HEDDWEN L. BROOKS

Heddwen Brooks received her PhD from Imperial College, University of

London and is a Professor in the Departments of Physiology and Pharmacology
at the University of Arizona (UA). Dr. Brooks is a renal physiologist and is best
known for her development of microarray technology to address in vivo
signaling pathways involved in the hormonal regulation of renal function. Dr.
Brooks’ many awards include the American Physiological Society (APS) Lazaro
J. Mandel Young Investigator Award, which is for an individual demonstrating
outstanding promise in epithelial or renal physiology. In 2009, Dr. Brooks
received the APS Renal Young Investigator Award at the annual meeting of the
Federation of American Societies for Experimental Biology. Dr. Brooks served
as Chair of the APS Renal Section (2011–2014) and currently serves as
Associate Editor for the American Journal of Physiology-Regulatory, Integrative
and Comparative Physiology, and on the Editorial Board for the American
Journal of Physiology-Renal Physiology (since 2001). Dr. Brooks has served on
study sections of the National Institutes of Health, the American Heart
Association, and served as a member of the Nephrology Merit Review Board for
the Department of Veterans’ Affairs.

JASON X.-J. YUAN

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Jason Yuan received his medical degree from Suzhou Medical College (Suzhou,
China) in 1983, his doctoral degree in cardiovascular physiology from Peking
Union Medical College (Beijing, China), and his postdoctoral training at the
University of Maryland at Baltimore. He joined the faculty at the University of
Maryland School of Medicine in 1993 and then moved to the University of
California, San Diego in 1999, rising to the rank of Professor in 2013. His
research interests center on pathogenic roles of membrane receptors and ion
channels in pulmonary vascular disease. He has published more than 300
articles, reviews, editorials and chapters, and has edited or co-edited nine books.
He has received several honors for his research accomplishments including the
Cournand and Comroe Young Investigator Award, the Established Investigator
Award and the Kenneth D. Bloch Memorial Lectureship from the American
Heart Association; the Guggenheim Fellowship Award from the John Simon
Guggenheim Memorial Foundation; the Estelle Grover Lectureship from the
American Thoracic Society; and the Robert M. Berne Memorial Lectureship
from The American Physiological Society. He is an elected Fellow of the
American Association for the Advancement of Science and an elected Member
of the American Society for Clinical Investigation and the Association of
American Physicians. He has served on many advisory committees including
Chair of the Respiratory Integrative Biology and Translational Research study
section of the National Institutes of Health and Chair of the Pulmonary
Circulation Assembly of the American Thoracic Society. He has also been very
active in scholarly editing serving currently as the Editor-in-Chief of the journal
Pulmonary Circulation and Associate Editor of the American Journal of
Physiology-Cell Physiology. He is a leading editor of the Textbook of Pulmonary
Vascular Disease (Springer, 2011).

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Contents

Preface

SECTION I Cellular & Molecular Basis for Medical Physiology

1 General Principles & Energy Production in Medical Physiology

2 Overview of Cellular Physiology

3 Immunity, Infection, & Inflammation

4 Excitable Tissue: Nerve

5 Excitable Tissue: Muscle

6 Synaptic & Junctional Transmission

7 Neurotransmitters & Neuromodulators

SECTION II Central & Peripheral Neurophysiology

8 Somatosensory Neurotransmission: Touch, Pain, & Temperature

9 Smell & Taste

10 Vision

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11 Hearing & Equilibrium

12 Reflex & Voluntary Control of Posture & Movement

13 Autonomic Nervous System

14 Electrical Activity of the Brain, Sleep–Wake States, & Circadian Rhythms

15 Learning, Memory, Language, & Speech

SECTION III Endocrine & Reproductive Physiology

16 Basic Concepts of Endocrine Regulation

17 Hypothalamic Regulation of Hormonal Functions

18 The Pituitary Gland

19 The Adrenal Medulla & Adrenal Cortex

20 The Thyroid Gland

21 Hormonal Control of Calcium & Phosphate Metabolism & the Physiology of

Bone

22 Reproductive Development & Function of the Female Reproductive System

23 Function of the Male Reproductive System

24 Endocrine Functions of the Pancreas & Regulation of Carbohydrate

Metabolism

SECTION IV Gastrointestinal Physiology

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25 Overview of Gastrointestinal Function & Regulation

26 Digestion & Absorption of Nutrients

27 Gastrointestinal Motility

28 Transport & Metabolic Functions of the Liver

SECTION V Cardiovascular Physiology

29 Origin of the Heartbeat & the Electrical Activity of the Heart

30 The Heart as a Pump

31 Blood as a Circulatory Fluid & the Dynamics of Blood & Lymph Flow

32 Cardiovascular Regulatory Mechanisms

33 Circulation Through Special Regions

SECTION VI Respiratory Physiology

34 Introduction to Pulmonary Structure & Mechanics

35 Gas Transport & pH

36 Regulation of Respiration

SECTION VII Renal Physiology

37 Renal Function & Micturition

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38 Regulation of Extracellular Fluid Composition & Volume

39 Acidification of the Urine & Bicarbonate Excretion

Answers to Multiple Choice Questions

Index

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Preface

It is difficult to believe that this preface signifies the fourth edition of Ganong’s
Review of Medical Physiology that our author group has overseen, and the 26th
edition overall of this important reference work aimed at medical and other
health professional students. As always, we have tried to maintain the highest
standards of excellence that were promulgated by the original author, Fran
Ganong, over the 46 years where he served, remarkably, as the sole author of the
textbook.
In this new edition, we have cast a fresh eye on the pedagogical approach
taken in each chapter and section, and have focused particularly on including
only material that is of the highest yield. We have thoroughly revised the
learning objectives for every chapter, reorganized and updated the text to ensure
that all objectives are clearly addressed in a logical order, aligned chapter
summaries so that the take-home messages quickly address each learning
objective in turn, and expanded the number of review questions so that readers
also have the ability to check their understanding and retention of every
objective covered. As a discipline evolves and new information emerges, there is
a tendency simply to concatenate these concepts such that chapter structure
degrades inevitably over time. With in-depth discussions amongst the author
team and significant “spring-cleaning,” we believe we have freshened and
simplified the volume while also making sure that important new developments
are incorporated. We are immensely thankful to Erica Wehrwein, PhD, Assistant
Professor of Physiology and an award-winning instructor at Michigan State
University, who took on the task of reviewing the book as a whole and providing
specific and detailed feedback to us on each chapter.
This new edition also welcomes a new member to the author team. We are
delighted to have been able to recruit Jason X.-J. Yuan, MD, PhD, Professor of
Medicine and Physiology as well as Chief of the Division of Translational and
Regenerative Medicine and Associate Vice President for Translational Health
Sciences at the University of Arizona, who has assumed responsibility for some

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cell physiology and cardiovascular topics, as well as the respiratory physiology
section. We are particularly excited to have a physician-scientist on the team,
who can guide us overall to focus on material that is of most benefit to those
preparing for a career incorporating patient care. We are most grateful for the
past contributions of Scott Boitano, PhD, whose other obligations meant that he
could no longer serve as an author.
We continue to be gratified by the many colleagues and students who contact
us from all over the world to request clarification of material covered in the text,
or to point out errors or omissions. We are especially grateful to Rajan Pandit,
Lecturer in Physiology at Nepal Medical College, who has painstakingly offered
dozens of suggestions for revision over the years. His efforts, and those of the
many others whom we have not named, allow us to engage in a process of
continual improvement. While, as always, any errors that remain in the book
(inevitable in a complex project such as this) are the sole responsibility of the
authorship team, we greatly value critical input, and urge readers once again to
contact us with any suggestions or critiques. We thank you in advance both for
such feedback, and also for your support of this new edition.

This edition is a revision of the original works of Dr. Francis Ganong.

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 SECTION I
Cellular & Molecular Basis for
Medical Physiology

Study of physiological system structure and function, as well as

pathophysiological alterations, has its foundations in physical and chemical laws
and the molecular and cellular makeup of each tissue and organ system.
Ganong’s Review of Medical Physiology is structured into seven sections. This
first section provides an overview of the basic building blocks or bases that
provide the important framework for human physiology. It is important to note
here that the seven chapters in this initial section are not meant to provide an
exhaustive understanding of biophysics, biochemistry, or cellular and molecular
physiology; rather, they are to serve as a reminder of how the basic principles
from these disciplines contribute to medical physiology discussed in later
sections associated with physiological functions of organs and systems.

In the first two chapters of this section, the following basic building blocks are
introduced and discussed: electrolytes; carbohydrates, lipids, and fatty acids;
amino acids and proteins; and nucleic acids. Students are reminded of some of
the basic principles and building blocks of biophysics and biochemistry and how
they fit into the physiologic environment. Examples of direct clinical
applications are provided in the clinical boxes to help bridge the gap between
basic principles and human cell, tissue, and organ functions. These basic
principles are followed up with a discussion of the generic cell and its
components. It is important to realize the cell is the basic functional unit within
the body, and it is the collection and fine-tuned interactions among and between
these fundamental units that allow for proper tissue, organ, and organism
function.

In the third to seventh chapters of this introductory section, we take a cellular

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approach to lay a groundwork of understanding groups of cells that interact with
many of the systems discussed in future chapters. The first group of cells
presented contribute to inflammatory reactions in the body. These individual
players, their coordinated behavior, and the net effects of the “open system” of
inflammation in the body are discussed in detail. The second group of cells
discussed are responsible for the excitatory responses in human physiological
function and include both neuronal and muscle cells. A fundamental
understanding of the inner workings of these cells, and how they are controlled
by their neighboring cells, helps the student to understand their eventual
integration into individual systems discussed in later sections.

This first section serves as an introduction, refresher, and quick source of

material to best understand organ functions and systems physiology presented in
the later sections. For detailed understanding of any of the chapters within this
section, several excellent and current textbooks that provide more in-depth
reviews of principles of biochemistry, biophysics, cell physiology, and muscle
and neuronal physiology are provided as resources at the end of each individual
chapter. Students are encouraged to visit these texts for a more thorough
understanding of these basic principles.

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CHAPTER 1
General Principles & Energy Production in
Medical Physiology

OBJECTIVES

After studying this chapter, you should be able to:

Define functional units used in measuring physiological properties.

Define pH and buffering.
Understand electrolytes and define diffusion, osmosis, and tonicity.
Define and explain the significance of resting membrane potential.
Understand in general terms the basic building blocks of the cell (eg,
nucleotides, amino acids, carbohydrates, and fatty acids) to cell
metabolism, proliferation, and function.
Understand higher-order structures of the basic building blocks of the cell
(eg, DNA, RNA, proteins, and lipids) to cell replication, proliferation, and
signal transduction.
Understand the basic contributions of the basic building blocks of the cell to
its structure, function, and energy balance.

INTRODUCTION

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In unicellular organisms, all vital processes occur in a single cell. As the
evolution of multicellular organisms progressed, various cell groups organized
into tissues, and organs have taken over particular functions. In humans and
other vertebrate animals, there are a number of specialized collections of cells
that consist in organ systems serving for different functions. For example, a
gastrointestinal system to digest and absorb food; a respiratory system to take up
O2 and eliminate CO2; a urinary system to remove wastes; a cardiovascular
system to distribute nutrients, O2, and the products of metabolism; a
reproductive system to perpetuate the species; and nervous and endocrine
systems to coordinate and integrate the functions of the other systems. This book
is concerned with the way these systems function and the way each contributes
to the functions of the body as a whole. This first chapter lays a foundation for
the discussion of these organ systems with a review of basic biophysical and
biochemical principles at the cellular level and the introduction of the molecular
building blocks that contribute to cell physiological function within these organ
systems.

GENERAL PRINCIPLES
THE BODY AS ORGANIZED “SOLUTIONS”
In the average young adult male, 18% of the body weight is protein and related
substances, 7% is mineral, and 15% is fat. The remaining 60% is water. The
distribution of the body water is shown in Figure 1–1A.

FIGURE 1–1 Organization of body fluids and electrolytes into

compartments. A) Body fluids can be divided into intracellular and
extracellular fluid compartments (ICF and ECF, respectively). Their contribution

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to percentage body weight (based on a healthy young adult male; slight
variations exist with age and gender) emphasizes the dominance of fluid makeup
of the body. Transcellular fluids, which constitute a very small percentage of
total body fluids, are not shown. Arrows represent fluid movement between
compartments. B) Electrolytes and proteins are unequally distributed among the
body fluids. This uneven distribution is crucial to physiology. Prot−, protein,
which tends to have a negative charge at physiologic pH.

The cells that make up the bodies of all but the simplest multicellular animals,
both aquatic and terrestrial, exist in an “internal sea” of extracellular fluid
(ECF) enclosed within the integument of the animal. From this fluid, the cells
take up O2 and nutrients; into it, they discharge metabolic waste products. The
ECF is more dilute than present-day seawater, but its composition closely
resembles that of the primordial oceans in which, presumably, all life originated.
In animals with a closed vascular system, the ECF is divided into the
interstitial fluid, the circulating blood plasma, and the lymph fluid that
bridges these two domains. The interstitial fluid is the part of the ECF that is
outside the vascular and lymph systems, bathing the cells. The plasma and the
cellular elements of the blood, principally red blood cells, fill the vascular
system, and together they constitute the total blood volume. About one-third of
the total body water is extracellular; the remaining two-thirds is intracellular
(intracellular fluid). Inappropriate compartmentalization of the body fluids can
result in edema (Clinical Box 1–1).

CLINICAL BOX 1–1

Edema
Edema is the buildup of body fluids extracellularly or interstitially in tissues.
The increased fluid is related to an increased leak from the blood and/or
reduced removal by the lymph system. Edema is often observed in the feet,
ankles, and legs, but can happen in many areas of the body in response to
disease, including those of the heart, lung, liver, kidney, or thyroid.

THERAPEUTIC HIGHLIGHTS
The best treatment for edema includes reversing the underlying disorder. Thus,
proper diagnosis of the cause of edema is the primary first step in therapy. More

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general treatments include restricting dietary sodium to minimize fluid retention
and using appropriate diuretic therapy.

The intracellular component of the body water accounts for about 40% of
body weight and the extracellular component for about 20%. Approximately
25% of the extracellular component is in the vascular system (plasma = 5% of
body weight) and 75% outside the blood vessels (interstitial fluid = 15% of body
weight). The total blood volume is about 8% of body weight. Flow between
these compartments is tightly regulated.

UNITS FOR MEASURING CONCENTRATION OF

SOLUTES
In considering the effects of various physiologically important substances and
the interactions among them, the number of molecules, electrical charges, or
particles of a substance per unit volume of a particular body fluid are often more
meaningful than simply the weight of the substance per unit volume. For this
reason, physiological concentrations are frequently expressed in moles,
equivalents, or osmoles.

Moles
A mole is the gram-molecular weight of a substance, that is, the molecular
weight (MW) of the substance in grams. Each mole (mol) consists of 6 × 1023
molecules. The millimole (mmol) is 1/1000 of a mole, and the micromole (µmol)
is 1/1,000,000 of a mole. Thus, 1 mol of NaCl = 23 g + 35.5 g = 58.5 g and 1
mmol = 58.5 mg. The mole is the standard unit for expressing the amount of
substances in the SI unit system.
The molecular weight of a substance is the ratio of the mass of one molecule
of the substance to the mass of one-twelfth the mass of an atom of carbon-12.
Because molecular weight is a ratio, it is dimensionless. The dalton (Da) is a unit
of mass equal to one-twelfth the mass of an atom of carbon-12. The kilodalton (1
kDa = 1000 Da) is a useful unit for expressing the molecular mass of proteins.
Thus, for example, one can speak of a 64-kDa protein or state that the molecular
mass of the protein is 64,000 Da. However, because molecular weight is a
dimensionless ratio, it is incorrect to say that the molecular weight of the protein

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is 64 kDa.

Equivalents
The concept of electrical equivalence is important in physiology because many
of the solutes in the body are in the form of charged particles. One equivalent
(Eq) is 1 mol of an ionized substance divided by its valence. One mole of NaCl
dissociates into 1 Eq of Na+ and 1 Eq of Cl−. One equivalent of Na+ = 23 g, but
1 Eq of Ca2+ = 40 g ÷ 2 = 20 g. The milliequivalent (mEq) is 1/1000 of 1 Eq.
Electrical equivalence is not necessarily the same as chemical equivalence. A
gram equivalent is the weight of a substance that is chemically equivalent to 8.0
g of oxygen. The normality (N) of a solution is the number of gram equivalents
in 1 liter (L). A 1 N solution of hydrochloric acid (HCl) contains both H+ (1.0 g)
and Cl− (35.5 g) equivalents, = (1.0 g + 35.5 g)/L = 36.5 g/L.

WATER, ELECTROLYTES, & ACID/BASE

The water molecule (H2O) is an ideal solvent for physiological reactions. H2O
has a dipole moment where oxygen slightly pulls away electrons from the
hydrogen atoms and creates a charge separation that makes the molecule polar.
This allows water to dissolve a variety of charged atoms and molecules. It also
allows the H2O molecule to interact with other H2O molecules via hydrogen
bonding. The resulting hydrogen bond network in water allows for several key
properties relevant to physiology: (1) water has a high surface tension, (2) water
has a high heat of vaporization and heat capacity, and (3) water has a high
dielectric constant. In layman’s terms, H2O is an excellent biological fluid that
serves as a solute; it provides optimal heat transfer and conduction of current.
Electrolytes (eg, NaCl) are molecules that dissociate in water to their cation
(Na+) and anion (Cl−) equivalents. Because of the net charge on water
molecules, these electrolytes tend not to reassociate in water. There are many
important electrolytes in physiology, notably Na+, K+, Ca2+, Mg2+, Cl−, and
HCO3−. It is important to note that electrolytes and other charged compounds
(eg, proteins) are unevenly distributed in the body fluids (Figure 1–1B). These
separations play an important role in physiology, for example, in the
establishment of membrane potential and generation of action potential.

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pH & BUFFERING
The maintenance of a stable hydrogen ion concentration ([H+]) in body fluids is
essential to life. The pH of a solution is defined as the logarithm to the base 10
of the reciprocal of the H+, that is, the negative logarithm of the [H+]. The pH of
water at 25°C, in which H+ and OH− ions are present in equal numbers, is 7.0
(Figure 1–2). For each pH unit less than 7.0, the [H+] is increased 10-fold; for
each pH unit above 7.0, it is decreased 10-fold. In the plasma of healthy
individuals, pH is slightly alkaline, maintained in the narrow range of 7.35–7.45
(Clinical Box 1–2). Conversely, gastric fluid pH can be quite acidic (on the
order of 3.0) and pancreatic secretions can be quite alkaline (on the order of 8.0).
Enzymatic activity and protein structure are frequently sensitive to pH; in any
given body or cellular compartment, pH is maintained to allow for maximal
enzyme/protein efficiency.

FIGURE 1–2 Proton concentration and pH. Relative proton (H+)

concentrations for solutions on a pH scale are shown.

Molecules that act as H+ donors in solution are considered acids, while those
that tend to remove H+ from solutions are considered bases. Strong acids (eg,
HCl) or bases (eg, NaOH) dissociate completely in water and thus can most
change the [H+] in solution. In physiological compounds, most acids or bases are
considered “weak,” that is, they contribute or remove relatively few H+ from

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solution. Body pH is stabilized by the buffering capacity of the body fluids. A
buffer is a substance that has the ability to bind or release H+ in solution, thus
keeping the pH of the solution relatively constant despite the addition of
considerable quantities of acid or base. Of course there are a number of buffers
at work in biological fluids at any given time. All buffer pairs in a homogenous
solution are in equilibrium with the same [H+]; this is known as the isohydric
principle. One outcome of this principle is that by assaying a single buffer
system, we can understand a great deal about all of the biological buffers in that
system.
When acids are placed into solution, there is dissociation of some of the
component acid (HA) into its proton (H+) and free acid (A−). This is frequently
written as an equation:

According to the laws of mass action, a relationship for the dissociation can
be defined mathematically as:

CLINICAL BOX 1–2

Acid–Base Balance and Disorders

Excesses of acid (acidosis) or base (alkalosis) exist when the blood or blood
plasma is outside the normal pH range (7.35–7.45). Such changes impair the
delivery of O2 to and removal of CO2 from tissues. There are a variety of
conditions and diseases that can interfere with pH control in the body and
cause blood pH to fall outside of healthy limits. Acid–base disorders that
result from respiration to alter CO2 concentration are called respiratory
acidosis and respiratory alkalosis. Respiratory acidosis is often caused by
respiratory failure or ventilator failure, while respiratory alkalosis is caused
by alveolar hyperventilation and often found in patients with chronic liver
disease. Nonrespiratory disorders that affect HCO3− concentration are
referred to as metabolic acidosis and metabolic alkalosis. Metabolic acidosis
or alkalosis can be caused by electrolyte disturbances, severe vomiting or
diarrhea, ingestion of certain drugs and toxins, kidney disease, and diseases
that affect normal metabolism (eg, diabetes).

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 THERAPEUTIC HIGHLIGHTS
Proper treatments for acid–base disorders are dependent on correctly
identifying the underlying causal process(es). This is especially true when
mixed disorders are encountered. Treatment of respiratory acidosis should be
initially targeted at restoring ventilation, whereas treatment for respiratory
alkalosis is focused on the reversal of the primary causes (eg, alveolar
hyperventilation associated with head injury and anxiety, hypoxemia due to
peripheral chemoreceptor stimulation, pulmonary embolism, and edema).
Bicarbonate (via intravenous injection) is typically used as a treatment for acute
metabolic acidosis. An adequate amount of a chloride salt can restore acid–base
balance to normal over a matter of days for patients with a chloride-responsive
metabolic alkalosis whereas chloride-resistant metabolic alkalosis requires
treatment of the underlying disease.

where Ka is a constant, and the brackets represent concentrations of the

individual species (elements). In layman’s terms, the product of the proton
concentration ([H+]) and the free acid concentration ([A−]) divided by the bound
acid concentration ([HA]) is a defined constant (K). This can be rearranged to
read:

If the logarithm of each side is taken:

Both sides can be multiplied by –1 to yield:

This can be written in a more conventional form known as the Henderson-

Hasselbalch equation:

This relatively simple equation is quite powerful. One thing that can be
discerned right away is that the buffering capacity of a particular weak acid is

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trace of a digestive tract at any stage of the life-history of Cestodes. For
nourishment they absorb, through the skin, the previously-digested food (of the
host) that bathes them. In a few Cestodes the body is simple and not divided into
"proglottides" or generative segments, but in most cases it is jointed in such a way
that the last segment is the oldest, and each contains a set of reproductive organs.
The life-histories of Cestodes are most remarkable. The proglottides containing the
eggs pass out of the final host along with the faeces and enter the intermediate
host with the food. The larvae hatch, and boring their way into the blood-vessels,
are carried by the circulation to various internal organs. Here they usually become
"bladder-worms," and develop the "head" of the future sexual form. Then, if, as is
usually the case, the intermediate host is preyed upon by the final host, the larval
Cestodes enter the alimentary canal of the latter. The head of the larva alone
survives digestion, and from it the mature worm is formed.

Of these three branches of the phylum Platyhelminthes, the Turbellaria possess

features of special interest and importance. Not only do they furnish the
explanation of the structure of the two parasitic groups (which have probably arisen
from Turbellarian-like ancestors), but they occupy the lowest position in the whole
group of worms. There are reasons for thinking that this is the simplest group of
bilateral animals which adopt the habit of creeping. The Turbellaria are most
closely allied to that great extinct group from which they, the Nemertinea, Rotifera,
and even the Annelids, offer increasingly convincing evidence of having been
derived. Many questions relating to the affinities of, or the origin of organs in, the
Annelids, resolve themselves into similar questions about the Turbellaria. For these
reasons, this group is here dealt with at greater length than the others, the interest
of which is of a more special nature.

The history of our knowledge of the Cestodes dates back to ancient times, as the
presence and effects of tape-worms early attracted the attention of physicians.
Trematodes are first distinctly referred to in the sixteenth century, while Turbellaria
first figure in Trembley's memoir on Hydra (1744).[3] The whole subject of the
increase in our knowledge of parasitic Platyhelminthes is dealt with in the standard
work, The Parasites of Man, by Leuckart,[4] and a complete list of references in
zoological literature to Cestodes and Trematodes is to be found in Bronn's
Thierreich.[5] O. F. Müller[6] and Ehrenberg founded our knowledge of the
Turbellaria, but for a long time the group remained in a most neglected condition. In
this country Montagu, G. Johnston, and in Ireland, William Thompson, discovered
several marine species, one of which, Planocera folium (from Berwick), has not
again been met with on British shores. Dalyell[7] conducted classical researches on
the habits of Planarians, and Faraday[8] made interesting experiments on their
power of regenerating lost parts. The credit of assigning the correct interpretation
to most of the various organs of fresh-water Planarians belongs to von Baer[9] and
Dugès,[10] while Mertens[11] effected a similar service for the marine forms, or
Polyclads. The minute Rhabdocoels were first successfully investigated and
classified by Oscar Schmidt.[12] The great work on this group is, however, the
monograph by von Graff.[13] A similarly comprehensive and indispensable treatise
by Lang, on the Polycladida,[14] contains references to all previous publications on
the group, among which the papers by Quatrefages, Johannes Müller, Keferstein,
Minot, and Hallez stand out conspicuously. Moseley's work[15] on the Land
Planarians of Ceylon is undoubtedly the most revolutionary paper referring to this
group, and the best contribution towards elucidating the structure of the Tricladida
at a time when the subject was very obscure. A monograph on Land Planarians is
being prepared by von Graff.

The Turbellaria are divided into: (1) Polycladida, marine forms with multiple
intestinal branches; (2) Tricladida, marine, fresh-water, and terrestrial Planarians
with three main intestinal branches; (3) the Rhabdocoelida, as varied in habit as
the Triclads, but possessing a straight and simple or slightly lobed, intestine. A
detailed description of an example of the Polyclads, and then a comparative
account of each division, will now be given.

Fig. 1.—Leptoplana tremellaris O. F. M. Seen from the dorsal surface. The

alimentary canal runs down the middle line and sends branches to the margin
of the body. × 6.

Turbellaria. I. Polycladida.

Description of Leptoplana tremellaris.

Appearance and Habits.—An account of the Polyclad Turbellaria may be fitly

prefaced by a description of a very common representative, Leptoplana tremellaris,
so called on account of the thin, flat body which executes when disturbed,
quivering or tremulous swimming movements.

Like all Polyclads, Leptoplana is marine. It is probably found on all European

shores, northwards to Greenland and southwards to the Red Sea, while vertically it
ranges from the littoral zone down to fifty fathoms. There is, however, an
apparently well-marked difference between the littoral specimens, which vary from
three-quarters to one inch in length, are brownish in colour and firm in consistency,
and the more delicate examples half an inch long, white with a brown tinge, which
occur in deeper water.

Fig. 2.—Leptoplana tremellaris. Three-quarters view from the ventral surface. The
pharynx (ph) is widely protruded through the month (mo) as in the act of
attacking prey. br, Brain with nerves, close to which are the four groups of eyes;
mg, stomach; mgc, "marginal groove"; pe, penis; sc, sucker; ut, uterus; vd, vasa
deferentia; ♀ , female genital aperture surrounded by the shell-gland; ♂ , male
aperture. (Semi-diagrammatic, and × 6.)

At low water Leptoplana may be found buried in mud or on the under surface of
stones, in pools where darkness and dampness may be ensured till the return of
the tide. It is, however, by no means easy to detect and remove it from the
encrusting Polyzoa, Ascidians, or Sponges with which it is usually associated. The
flat, soft, unsegmented body is so closely appressed to the substratum that its
presence is usually only betrayed by its movement, an even gliding motion of the
mobile body, which suggested the apt name "la pellicule animée" to Dicquemare.
The creeping surface is called ventral, the upper one dorsal, and as the broader
end of the body always goes first, it is anterior as opposed to the more pointed
posterior extremity. With a lens the characters shown in Figs. 1 and 2 may be
observed. The eyes are seen as black dots near the anterior end, and are placed
at the sides of a clear oval space, the brain. Along the transparent margin of the
body, the ends of the intestinal branches may be seen. These ramify from a lobed
stomach or main-gut, and should the specimen be mature, the "uterus" loaded with
eggs forms a dark margin round the latter (Figs. 1 and 2, ut). The ventral surface is
whitish, and through it the "pharynx," a frilled protrusible structure, may be dimly
observed. The "mouth,"[16] through which the pharynx at the time of feeding is
thrust out (Fig. 2, mo), is almost in the centre of the ventral surface. Behind this, a
white, V-shaped mark (vd) indicates the ducts of the male reproductive organs, and
still further back is the irregular opaque mark of the "shell-gland," by which the egg-
shells are formed (Fig. 2, ♀).
 Fig. 3.—Leptoplana tremellaris in the act of swimming. A, Seen from the right side
during the downward stroke (the resemblance to a skate is striking); B, from
above, showing the upward stroke and longitudinal undulations of the swimming
lobes; C, side view during the upward stroke; D, transverse sections of the body
during the strokes. × 5.

Leptoplana employs two kinds of movement, creeping and swimming. Creeping is

a uniform gliding movement, caused by the cilia of the ventral surface, aided
perhaps by the longitudinal muscular layers of this surface, and is effected on the
under side of the "surface-film" of water almost as well as on a solid substratum.
Swimming is a more rapid and elegant movement, employed when alarmed or in
pursuit of prey. The expanded fore-parts of the body act as lobes, which are
flapped rapidly up over the body and then down beneath it, undulations running
rapidly down them from before backwards. The action in fact is somewhat similar
to that by which a skate swims, a resemblance pointed out long ago by Dugès[17]
(Fig. 3).

We have few direct observations on the nature of the food of Leptoplana, or the
exact mode by which it is obtained. Dalyell,[18] who observed this species very
carefully, noticed that it was nocturnal and fed upon a Nereis, becoming greatly
distended and of a green colour after the meal, but pale after a long fast.
Keferstein[19] noticed a specimen in the act of devouring a Lumbriconereis longer
than itself, and also found the radulae of Chiton and Taenioglossate Molluscs in the
intestine. That such an apparently weak and defenceless animal does overpower
large and healthy Annelids and Mollusca, has not hitherto been definitely proved.
Weak or diseased examples may be chiefly selected. The flexible Leptoplana
adheres firmly to its prey, and the rapid action of the salivary glands of its mobile
pharynx quickly softens and disintegrates the internal parts of the victim. The food
passes into the stomach (Fig. 2, mg), and is there digested. It is then transferred to
the lateral branches of the intestine, and, after all the nutritious matters have been
absorbed, the faeces are ejected with a sudden contraction of the whole body
through the pharynx into the water.

Leptoplana probably does not live more than a year. In the spring or summer,
batches of eggs are laid and fixed to algae or stones by one individual, after having
been fertilised by another. Young Leptoplana hatch out in two to three weeks, and
lead a pelagic existence till they are three or four millimetres in length. In late
summer, numbers of such immature examples may be found among sea-weeds
and Corallina in tide pools. In the succeeding spring they develop first the male and
then the female reproductive organs.

Fig. 4.—Portion of a transverse section of Leptoplana tremellaris in the hinder part of

the body. × 100. bm, Basement (skeletal) membrane; cil, cilia; d.m, diagonal
muscles; d.v.m, dorso-ventral muscles; ep, epidermis; f.p, food particles; l.g,
lateral intestinal branches cut across; l.m ext, external, and l.m int, internal
longitudinal muscle layers; m.c, glandular (mucous) cells; md, their ducts; N,
longitudinal nerve; Nu, nuclei of the intestinal epithelium; ov, ovary; ovd,
oviduct; par, cells of the parenchyma; r.d, vasa deferentia, with spermatozoa;
rm, circular musculature; rh, rhabdites; sh, cells of the shell-gland; te, testes; ve,
vasa efferentia; y.c, "yellow cells." (After Lang.)

Anatomy of Leptoplana tremellaris.—Leptoplana may be divided into

corresponding halves only by a median vertical longitudinal plane. The body and all
the systems of organs are strictly bilaterally symmetrical. Excepting the cavities of
the organs themselves, the body is solid. A connective "parenchyma" (Fig. 4, par)
knits the various internal organs together, while it allows free play of one part on
another. These organs are enclosed in a muscular body-wall, clothed externally by
the ciliated epidermis, which is separated from the underlying musculature by a
strong membrane (Fig. 4, bm), the only skeletal element in the body.

Body-Wall.—The epidermis (Fig. 4, ep) is composed of a single layer of ciliated

cells, containing small, highly refractive, pointed rods or "rhabdites" (rh), and gives
rise to deeply-placed mucous cells (m.c), which are glandular and pour out on the
surface of the body a fluid in which the cilia vibrate. The tenacious hold on a stone
which Leptoplana exerts if suddenly disturbed, or when grasping its prey, is
probably due to the increased glutinous secretion of these glands, aided perhaps
by rhabdites, which on such occasions are shot out in great numbers. The
basement membrane is an elastic skeletal membrane composed of stellate cells
embedded in a firm matrix. It serves chiefly for the origin and insertion of the dorso-
ventral muscles (d.v.m). Under the basement membrane lies a very thin layer of
transverse muscular fibres (Fig. 4, rm), which are, however, apparently absent on
the ventral surface. Then follows a stout layer of longitudinal fibres (l.m ext), and
beneath this a diagonal layer (d.m), the fibres of which intersect along the median
line in such a way that the inner fibres of one side become the outer diagonal fibres
of the other. Lastly, within this again, on the ventral surface, is a second stout
longitudinal layer (l.m int). The sucker (sc, Figs. 2 and 5) is a modification of the
body-wall at that point. In addition to the dorso-ventral muscles, there exists a
complex visceral musculature regulating the movements of the pharynx, intestine,
and copulatory organs.

Parenchyma.—The spaces between the main organs of the body are filled by a
tissue containing various kinds of cells, salivary glands, shell-glands, and prostate
glands. Besides these, however, we find a vacuolated, nucleated, thick-walled
network, and to this the word parenchyma is properly applied. Besides its
connective function, the parenchyma confers that elasticity on the body which
Leptoplana possesses in such a high degree. Pigment cells are found in the
parenchyma in many Polyclads.

Digestive System.—The general arrangement of this system may be seen in Figs.

2, 5, and 7; and may be compared, especially when the pharynx is protruded, as in
Fig. 2, with the gastral system of a Medusa. The "mouth" (there is no anus) is
placed almost in the centre of the ventral surface. It leads (Fig. 7, B, phs) into a
chamber (the peripharyngeal space) divided into an upper and a lower division by
the insertion of a muscular collar-fold (the pharynx, ph), which may be protruded,
its free lips advancing, through the mouth (Fig. 2), and is then capable of enclosing
by its mobile frilled margin, prey as large as Leptoplana itself. The upper division of
the chamber communicates by a hole in its roof[20] (the true mouth, Figs. 5 and 7,
g.m) with the cavity of the main-gut or stomach (m.g), which runs almost the length
of the body in the middle line, forwards over the brain (Fig. 5, up). Seven pairs of
lateral gut-branches convey the digested food to the various organs, not directly
however, but only after the food mixed with sea-water has been repeatedly driven
by peristalsis first towards the blind end of the gut-branches and then back towards
the stomach. Respiration is probably largely effected by this means. The epithelium
of the intestine (Fig. 4, l.g) of a starving specimen is composed of separate
flagellated cells frequently containing "yellow cells."[21] After a meal, however, the
cell outlines are invisible. Gregarines, encysted Cercariae, and Orthonectida[22]
occur parasitically in the gut-branches.
An excretory system of "flame-cells" and fine vessels has hitherto been seen only
by Schultze[23] in this species, which will not, however, resist intact the
compression necessary to enable the details to be determined. They are probably
similar to those of Thysanozoon described on p. 25.

Nervous System.—The brain, which is enclosed in a tough capsule (Fig. 5, br), is

placed in front of the pharynx, but some distance behind the anterior margin of the
body. It is of an oval shape, subdivided superficially into right and left halves by a
shallow depression, and is provided in front with a pair of granular-looking
appendages, composed of ganglion-cells from which numerous sensory nerves
arise, supplying the eyes and anterior region. Posteriorly the brain gives rise to a
chiefly motor, nervous sheath (Fig. 5, nn), which invests the body just within the
musculature. This sheath is thickened along two ventral lines (Fig. 5, ln) and two
lateral lines (n.s), but is very slightly developed on the dorsal surface. Ganglion-
cells occur on the course of the nerves, and are particularly large at the point of
origin of the great motor nerves.
 Fig. 5.—Diagrammatic view of the structure of Leptoplana tremellaris
as a type of the Polycladida. The body is cut across the middle to
show the relative position of organs in transverse section. In the
posterior half the alimentary canal has been bisected and
removed from the left side, to exhibit the deeply placed nervous
sheath (nn) and the male reproductive organs. br, Brain; dp,
"diaphragm"; e, cerebral group of eyes; et, tentacular eye-group;
gr, marginal groove; gm, true mouth; lg, lateral gut-branch; ln,
longitudinal nerve stem; m, external mouth; mg, mg', main-gut,
whole, and bisected; n, sensory nerve supplying the eyes; nn,
nervous network lying on the ventral musculature; n.s, lateral
nerve; od, oviduct; ov, ovary; pe, penis (in section); ph, pharynx;
pr, prostate or "granule gland"; sc, sucker; sg, shell-gland; te,
testes; up, anterior unpaired gut-branch; ut, uterus; va, vagina (in
section); vd, vas deferens; ve, vasa efferentia; ♂ , male genital
pore; ♀, female pore.

Sense Organs.—Leptoplana possesses eyes, stiff tactile, marginal

cilia, and possibly a sense organ in the "marginal groove." The eyes,
which are easily seen as collections of black dots lying at the sides
of the brain, may be divided into two paired groups: (1) cerebral eyes
(Fig. 5, e), and (2) tentacle eyes (et), which indicate the position of a
pair of tentacles in allied forms (Fig. 8, A, t and B). Each ocellus
consists of a capsule placed at right angles to the surface of the
body in the parenchyma, below the dorsal muscles, and with its
convex face outwards. It is a single cell in which pigment granules
have accumulated. The light, however, can only reach the refractive
rods, which lie within it, obliquely at their outer ends. These rods are
in connexion with the retinal cells, and thus communicate by the
optic nerve with the brain. The cerebral eyes are really paired, and
are directed some upwards, some sideways, some downwards.

The "marginal groove" is a shallow depression of the epidermis (Fig.

5, gr) lined by cilia, and containing the ducts of very numerous gland-
cells. It runs almost parallel to the anterior margin of the body, a
short distance from it, but we have no observations on its functions.

Fig. 6.—Diagram of an eye of Leptoplana from the tentacle group. ×

600. (After Lang.)

Reproductive Organs.—Leptoplana is hermaphrodite, and, as in

most hermaphrodites, the reproductive organs are complicated. The
male organs are the first to ripen, but this does not appear to prevent
an overlapping of the periods of maturity of the male and female
products, so that when the eggs are being laid, the male organs are,
apparently, still in a functional state. The principal parts are seen in
Fig. 5. The very numerous testes (te) are placed ventrally, and are
connected with fine vasa efferentia (ve), which form a delicate
network opening at various points into the two vasa deferentia (vd).
These tubes, especially when distended with spermatozoa, may
easily be seen (Fig. 2, vd) converging at the base of the penis, and
connected posteriorly by a loop that runs behind the female genital
pore (Fig. 5). The penis (pe) is pyriform and muscular, and is divided
into two chambers, a large upper one for the spermatozoa, and a
smaller lower one for the secretion of a special "prostate" gland. The
apex of the penis is eversible and not merely protrusible, being
turned inside out when evaginated. The ovaries (Fig. 5, ov) are
numerous and somewhat spherical. They are dorsally placed, but
when fully developed extend deeply wherever they can find room to
do so, and they not only furnish the ova, but elaborate food-yolk in
the ova, as there are no special yolk-glands. The slender oviducts
(od) open at several points into the "uterus" (ut) (a misnomer, as no
development takes place within it), which encircles the pharynx, and
opens by a single duct into the vagina (va). Here the ova are
probably fertilised, and one by one invested by the shell-gland (sg)
with a secretion which hardens and forms a resistant shell. They are
then laid in plate-like masses which are attached to stones or shells.
The development is a direct one, and the young Leptoplana, which
hatches in about three weeks, has the outline of a spherical triangle,
and possesses most of the organs of the adult. After leading a
floating life for a few weeks it probably attains maturity in about nine
months.

Classification, Habits, and Structure of the Polycladida.

The Polyclads were so called by Lang on account of the numerous

primary branches of their intestine. They are free-living, purely
marine Platyhelminthes, possessing multiple ovaries, distinct male
and female genital pores (Digonopora), but no yolk-glands. The eggs
are small, and in many cases give rise to a distinct larval form,
known as "Müller's larva" (Fig. 12). The Polyclads, with one
exception,[24] fall into two sub-groups, Acotylea and Cotylea:—

Character. Acotylea. Cotylea.

Sucker A sucker absent.[25] A sucker always present
(Figs. 8, D, s; 7, A, sc).
 Mouth In the middle, or
behind the middle, of
the ventral surface.
Pharynx More or less intricately
folded.
Tentacles A pair of dorsal
tentacles usually
present.
Development Usually direct. Larva
when present, not a
typical Müller's larva.
In the middle, or in front
of the middle, of the
ventral surface.
Rarely folded. Usually
cylindrical or trumpet-
shaped.
A pair of marginal
tentacles (except in
Anonymus).
Müller's larva present.
Metamorphosis,
however, extremely
slight.

Fig. 8 shows that, starting with a member (A, D) of each division, in

which the mouth is almost in the middle of the ventral surface, and
the brain and sense organs somewhat remote from the anterior end,
we find in the Acotylea a series leading to an elongated form
(Cestoplanidae), in which the mouth, pharynx, and genital pores are
far back near the hinder end of the body; while in the Cotylea the
series leads similarly to the elongated Prosthiostomatidae, in which,
however, the pharynx and external apertures are in the front part of
the body. This view of the morphology of the Polyclads is due to
Lang, and is based on the assumption that the more radially-
constructed forms (Fig. 8, A, D) are the primitive ones.

Fig. 7.—Diagrammatic vertical longitudinal sections: A, Of

Prosthiostomum (type of Cotylea); B, of Leptoplana; C, of
Cestoplana (types of Acotylea). (After Lang.) These figures
illustrate the changes which follow the shifting of the mouth from a
 central position (B) to either end of the body. br, Brain; dphm,
"diaphragm"; gm, true mouth; lg, openings of lateral gut-branches;
m, mouth; mg, main-gut or stomach; mgbr, median gut-branch;
ph, pharynx; ph.m, aperture in pharyngeal fold; phs,
peripharyngeal sheath; sc, sucker; ♂, male, and ♀, female, genital
aperture.

Fig. 8.—Chief forms of Polycladida: A-C, Acotylea; D-F, Cotylea. A,

Planocera graffii Lang, nat. size; B, Stylochoplana maculata
Stimps, × 7; C, Cestoplana rubrocincta Lang, × 4⁄3; D, Anonymus
virilis Lang, × 3, ventral surface; E, Thysanozoon brocchii Grube,
nat. size; the head is thrown back and the pharynx (ph) is
protruded. F, Prosthiostomum siphunculus Lang, × 3. Br, Brain;
CG, cerebral eye group; DM, true mouth; Ey, marginal eyes; m,
mouth; MG, main-gut or stomach; P, dorsal papillae; Ph, pharynx;
s, sucker (ventral); T, tentacles; UP, dorsal median gut-branch. ♂,
male, and ♀, female, genital aperture, except in D, where ♂ refers
to the multiple penes. (After Lang and Schmidt.)

Classification of Polycladida.

ACOTYLEA.
 Family. Genus. British
Representatives.
Planocera (Fig. 8,
Planocera folium
A).
Grube. Berwick-
Imogine.
on-Tweed.
Planoceridae. Conoceros.
Stylochoplana
With dorsal tentacles. Stylochus.
maculata Quatref.
Mouth sub-central. Stylochoplana
Among brown
(Fig. 8, B).
weeds in
Diplonchus.
Laminarian zone.
Planctoplana.
Leptoplana
tremellaris O. F.
Müll.
Discocelis.
L. fallax Quatref.
Leptoplanidae. Cryptocelis.
Plymouth.
Without dorsal Leptoplana.
L. droebachensis
tentacles. Penis Trigonoporus.
Oe. Plymouth
directed backwards. ?Polypostia (see
Sound.
p. 27).
L. atomata O. F.
Müll. Doubtful
species.
Cestoplana (Fig. 8,
Cestoplanidae. C).
No tentacles. Body In Mediterranean
elongated. Penis and on French
directed forwards. side of the
Channel.
Enantiidae.
No sucker. No
tentacles. Main-gut Enantia.
very short. External Adriatic Sea.
apertures as in
Euryleptidae.

COTYLEA.
Anonymidae. Anonymus (Fig. 8,
Mouth central. No D).
tentacles. With two Naples (two
rows of penes. specimens).
Pseudoceridae. Thysanozoon (Fig.
Marginal tentacles 8, E).
folded. Mouth in Pseudoceros.
anterior half. Yungia.
Prostheceraeus
vittatus Mont. On
west coast.
P. argus Quatref.
Guernsey.
Cycloporus
Euryleptidae. papillosus Lang.
Tentacles usually Prostheceraeus. On Ascidians in
present and pointed, Cycloporus. 2-30 fms.
or represented by Eurylepta. Eurylepta cornuta
two groups of eyes. Oligocladus. O.F. Müll. On
Mouth close to Stylostomum. sponges and
anterior end. Aceros. shells, 2-10 fms.
Pharynx cylindrical. Oligocladus
sanguinolentus
Quatref.
O. auritus Clap.
Doubtful.
Stylostomum
variabile Lang.
Prosthiostomatidae.
Tentacles absent.
Body elongated.
Prosthiostomum
Pharynx long,
(Fig. 8, F).
cylindrical. Penis
with accessory
muscular vesicles.
Appearance and Size of Polyclad Turbellaria.—Polyclads are
almost unique amongst animals in possessing a broad and thin,
delicate body that glides like a living pellicle over stones and weeds,
moulding itself on to any inequalities of the surface over which it is
travelling, yet so fragile that a touch of the finger will rend its tissues
and often cause its speedy dissolution. The dorsal surface in a few
forms is raised into fine processes (Planocera villosa), or into hollow
papillae (Thysanozoon brocchii), and in very rare cases may be
armed with spines (Acanthozoon armatum,[26] Enantia spinifera); in
others, again, nettle-cells (nematocysts) are found (Stylochoplana
tarda, Anonymus virilis). Some Polyclads, especially the pelagic
forms, are almost transparent; in others, the colour may be an
intense orange or velvety black, and is then due to peculiar deposits
in the epidermal cells. Between these two extremes the colour is
dependent upon the blending of two sources, the pigment of the
body itself and the tint of the food. Thus a starved Leptoplana is
almost or quite white, a specimen fed on vascular tissue reddish.
Many forms are coloured in such a way as to make their detection
exceedingly difficult, but this is probably not merely due, as Dalyell
supposed, to the substratum furnishing them with food and thus
colouring them sympathetically, but is probably a result of natural
selection.

The largest Polyclad, the bulkiest Turbellarian, is Leptoplana gigas

(6 inches long and 4 in breadth), taken by Schmarda, free-
swimming, off the coast of Ceylon. The largest European form is
Pseudoceros maximus, 3½ inches in length and stoutly built. A
British species, Prostheceraeus vittatus, attains a length of from 2 to
3 inches. These large forms, especially the Pseudoceridae (pre-
eminently the family of big Polyclads), are brightly coloured, and
usually possess good swimming powers, since, being broad and flat,
they are certainly not well adapted for creeping rapidly, and this is
well shown by the way these Polyclads take to swimming when in
pursuit of prey at night. The size of any individual is determined,
amongst other factors, by the period at which maturity sets in, after
which probably no increase takes place. Polyclads apparently live
about twelve months, and mature specimens of the same species
vary from ½ inch to 2½ inches in length (Thysanozoon brocchii),
showing that growth is, under favourable conditions, very rapid.

Habits of Polyclad Turbellaria.—Polyclads are exclusively marine,

and for the most part littoral, animals. Moreover, there is no evidence
of their occurrence in those inland seas where certain marine
animals (including one or two species of otherwise characteristically
marine Rhabdocoelida, p. 46) have persisted under changed
conditions. From half-tide mark down to 50 fathoms, some Polyclads
probably occur on all coasts, but as to their relative abundance in
different seas we have very little accurate information. The southern
seas of Europe possess more individuals and species than the
northern, and probably the maximum development of the group
takes place on the coasts and coral islands of the tropics.[27] No
Polyclads have been taken below 60 fathoms; but their delicacy and
inconspicuousness render this negative evidence of little value. Six
truly pelagic forms, however, are known,[28] and these are interesting
on account of their wide distribution (three occurring in the Atlantic,
Pacific, and Indian oceans), and also from the distinct modifications
they have undergone in relation to their pelagic existence.

Whatever may be the interpretations of the fact, Polyclads are

notoriously difficult to detect, and this fact doubtless explains the
scanty references to them by the older naturalists who collected
even in tropical seas. Lang, who worked seven years at Naples,
added to the Mediterranean fauna as many Polyclads as were
previously known for all Europe, in spite of the assiduous labours of
his predecessors, Delle Chiaje and Quatrefages. Again Hallez,
collecting at Wimereux at low-water, obtained some twenty
specimens of Leptoplana tremellaris in an hour, while some other
collectors working by his side could only find two or three. Yet, even
making allowance for the difficulty of finding Polyclads, few of them
appear to be abundant.

Leptoplana tremellaris is frequently associated with colonies of

Botryllus, and if separated soon perishes, whereas the free-living
individuals are distinctly hardy (Hallez). A closely allied but possibly
distinct form lives upon the surface of the Polyzoon Schizoporella, on
the French side of the Channel, and cannot long endure separation
from its natural habitat, to which it is adaptively coloured. A striking
case of protective mimicry is exhibited by Cycloporus papillosus, on
the British coasts. This species, eminently variable in colour and in
the presence or absence of dorsal papillae, is usually a quarter of an
inch in length and of a firm consistency. Fixed by its sucker to
Polyclinid and other Ascidians, Cycloporus appears part and parcel
of the substratum, an interesting parallel to Lamellaria perspicua,[29]
though we are not justified in calling the Polyclad parasitic. Indeed,
though a few cases of association between Polyclads and large
Gasteropods, Holothurians, and Echinids are known,[30] there is only
one case, that of Planocera inquilina,[31] in the branchial chamber of
the Gasteropod Sycotypus canaliculatus, which would seem to bear
the interpretation of parasitism. The jet-black Pseudoceros velutinus
and the orange Yungia aurantiaca of the Mediterranean, are large
conspicuous forms with no attempt at concealment, but their taste,
which is not known, may protect them. Other habits, curiously
analogous with devices employed by Nudibranch Mollusca (compare
Thysanozoon brocchii with Aeolis papillosa), emphasise the
conclusion that the struggle for existence in the littoral zone has
adapted almost each Polyclad to its particular habitat.

As regards the vertical distribution of this group on the British coasts,

Leptoplana tremellaris has an extensive range, and appears to come
from deeper to shallower water to breed.[32] In the upper part of the
Laminarian zone, Cycloporus papillosus, and, among brown weeds,
Stylochoplana maculata are found. At and below lowest water-mark
Prostheceraeus vittatus, P. argus, and Eurylepta cornuta occur.
Stylostomum variabile and Oligocladus sanguinolentus, though
occasionally found between tide-marks, especially in the Channel
Islands, are characteristic, along with Leptoplana droebachensis and
L. fallax, of dredge material from 10 to 20 fathoms.
Locomotion.—Locomotion is generally performed by Polyclads at
night when in search of food, and two methods, creeping and
swimming, are usually employed—creeping by the cilia, aided
possibly, as in the case of some Gasteropod Mollusca, by the
longitudinal muscles of the ventral surface; and swimming, by
undulations of the expanded margins of the body. In the former case
the cilia work in a glandular secretion which bathes the body, and
enables them to effect their purpose equally well on different
substrata. The anterior region is generally lifted up, exploring the
surroundings by the aid of the tentacles, which are here usually
present. The rest of the body is closely appressed to the ground.

Swimming is particularly well performed by the Pseudoceridae,

certain species of Prostheceraeus, the large Planoceridae, some
Stylochoplana, Discocelis, and Leptoplana, and in the same manner
as in Leptoplana tremellaris (p. 9). In Cryptocelis, Leptoplana alcinoi,
and L. pallida, however, the whole body executes serpentine
movements like an active leech (e.g. Nephelis); a cross section of
the body would thus present the same appearance during the whole
movement. Many Polyclads, notably Anonymus (Lang), if irritated,
spread out in all directions, becoming exceeding thin and
transparent.

Fig. 9.—Discocelis lichenoides Mert. (after Mertens), creeping on the

inner side of a glass vessel by means of the lobes of the extended
and exceedingly mobile pharynx (ph). These lobes also serve to
enclose Crustacea (a), and one lobe may then be withdrawn
independently of the rest, back into the body (b). The brain (br)
and shell-gland (sg) are shown by transparency.
Discocelis lichenoides, Planocera graffii, and Anonymus virilis have
peculiar modes of progression. The first, according to Mertens, will
climb up the sides of a vessel by means of the expanded lobes of
the pharynx (Fig. 9, ph), a habit of considerable interest, since we
know that certain Ctenophores—Lampetia, for instance—progress
when not swimming on the expanded lobes of their "stomach."[33]
Planocera and Anonymus creep by extending parts of the anterior
margin and dragging the rest of the body behind. In consequence,
the brain and dorsal tentacles may come to lie actually behind the
middle of the body, and thus no definite anterior end or "head"
advances first. Along with this curious habit it may be noticed (Lang)
that the radial symmetry of the body is well marked; but even without
accepting this author's suggestion of the concurrent development of
a "head" with locomotion in a definite direction, the facts, whether
these two forms are primitive or not, are highly interesting.

Food.—Though we are probably right in calling Polyclads a

carnivorous group, the food of very few forms has been ascertained.
Those which possess a large frilled pharynx (most Acotylea)
probably enclose and digest large, and, it may be, powerful prey, as
appears to be the case in Leptoplana tremellaris. Cryptocelis alba
has been seen by Lang with the pharynx so distended, owing to a
large Drepanophorus (Nemertine) which it contained, as to resemble
a yolk-sac projecting from the under surface of an embryo. The
Cotylea such as Thysanozoon, with a bell- or trumpet-shaped
pharynx, are fond of fixing this to the side of the aquarium, but
whether they thus obtain minute organisms is not clear.
Prosthiostomum shoots out its long pharynx with great vehemence
(Fig. 8, F) and snaps up small Annelids by its aid (Lang). Those
Polyclads which, as Cycloporus and others, are definitely associated
with other organisms are not certainly known to feed upon the latter,
though "Planaria velellae" has been seen by Lesson[34] devouring
the fleshy parts of its host. The salivary glands which open on the
lips and the inner surface of the pharynx powerfully disintegrate the
flesh of the prey. Digestion takes place in the main-gut, and the
circulation of the food is accomplished by the sphinctral musculature
of the intestinal branches (conf. Leptoplana, p. 13).

Fig. 10.—Diagram of the musculature, causing peristaltic movements

of the intestinal branches of Polyclads. (After Lang.)

A distinct vent or anus is always absent. After a meal the faecal

matter collects in the main-gut, and is discharged violently by the
pharynx into the water. In a few species, however, the intestinal
branches open to the exterior (Lang). Yungia aurantiaca, a large and
abundant Neapolitan form, possesses such openings over the
greater part of the dorsal surface; Cycloporus papillosus has
marginal pores; Oligocladus sanguinolentus apparently possesses
an opening at the posterior end of the main-gut; and Thysanozoon
brocchii frequently rends at this point, in consequence of the
accumulation of food.

Respiration.—The oxygen of the atmosphere dissolved in the sea-

water is, in default of a special circulatory fluid, brought to the tissues
of Polyclads in two ways. The ciliated epidermis provides a constant
change of the surrounding water, by which the superficial organs
may obtain their supply; and the peristaltic movements of the
digestive system, aided by the cilia of the endoderm cells, ensure a
rough circulation of the sea-water, which enters along with the food,
to the internal organs. The papillae of Thysanozoon brocchii,
containing outgrowths of the intestinal branches, are possibly so
much additional respiratory surface, although still larger forms (other
Pseudoceridae) are devoid of such outgrowths.

Excretion.—The excretory system of only one Polyclad

(Thysanozoon brocchii) is accurately known. Lang, by compressing
light-coloured specimens, found the three parts of the system known