Fundamentals of Toxicologic Pathology

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Fundamentals of Toxicologic
Pathology

THIRD EDITION

EDITED BY

Matthew A. Wallig
University of Illinois at Urbana-Champaign, Urbana, IL, United States

Wanda M. Haschek
University of Illinois at Urbana-Champaign, Urbana, IL, United States

Colin G. Rousseaux
University of Ottawa, Ottawa, ON, Canada

Brad Bolon
GEMpath Inc., Longmont, CO, United States
Table of Contents

Cover image

Title page

Copyright

Frontispiece

Dedication

List of Contributors

Preface

Chapter 1. An Overview of Toxicologic Pathology

Introduction

What is Toxicologic Pathology?

The Basis of Toxicologic Pathology

The Challenges Facing Toxicologic Pathology

Training and Certification in Toxicologic Pathology

 The “Practitioner” of Toxicologic Pathology

Summary

Further Reading

Part I: Principles of Toxicologic Pathology

Chapter 2. Biochemical and Molecular Basis of Toxicity

Abstract

Introduction

General Principles of Xenobiotic Disposition

Interactions of Toxicants With Cellular and Molecular Targets

Protective Mechanisms, Repair Mechanisms, and Adaptation

Summary

Further Reading

Chapter 3. Pharmacokinetics and Toxicokinetics

Abstract

Introduction

Pharmacokinetics: Processes and Determinants

Pharmacokinetic (Time-Course) Data and Derivation of Dose Metrics

Pharmacokinetic Models and Computation of Dose Metrics

Pharmacokinetics and Its Use in Designing Toxicity Studies

Concluding Remarks

Further Reading
Chapter 4. Principles of Pharmacodynamics and Toxicodynamics
Abstract

Introduction: Pharmacodynamics and Toxicodynamics Defined

Relevance to Toxicological Pathology

Drug/Toxin–Receptor Interactions Lead to Signal Transduction and Effect

Selectivity and Safety: Therapeutic Versus Toxic Effect of Drugs

Quantal Dose–Effect Curves

Forms of Antagonism at the Tissue or Organism Level

Dynamic Receptor Phenomena Resulting in Altered Efficacy Following

Chronic Dosing

Variation in Drug/Toxin Responsiveness

Nonmonotonic Dose–Effect Curves

Principles of Pharmacokinetic/Pharmacodynamic Modeling

Further Reading

Chapter 5. Morphologic Manifestations of Toxic Cell Injury

Abstract

Introduction

Irreversible versus Reversible Cell Injury

Irreversible Cell Injury

Summary

Further Readings

Chapter 6. Carcinogenesis: Manifestation and Mechanisms

 Abstract

Overview of Carcinogenesis

Mechanisms of Carcinogenesis

Summary

Identifying Carcinogens

Overall Summary

Acknowledgments

Further Reading

Chapter 7. Design of Studies and Risk Management in Toxicologic

Pathology: Addressing Risks in Product Discovery and Development
Abstract

Introduction

Risk Assessment: The Process

Risk Assessment: The Elements

Toxicologic Pathology Issues in Risk Assessment

Risk Perception and Acceptability

Risk Communication

Risk Management

Financial Risk and Toxicologic Pathology

Summary

Acknowledgments

Further Reading
Part II: Systems Toxicologic Pathology
Chapter 8. Liver and Gall Bladder
Abstract

Introduction

Structure, Function, and Cell Biology

Evaluation of Liver Toxicity

Response to Injury

Mechanisms of Toxicity in Liver

Summary

Further Reading

Chapter 9. Cardiovascular System

Abstract

Introduction

Heart

Blood Vessels

Summary

Further Reading

Chapter 10. Skeletal Muscle System

Abstract

Structure and Function

Evaluation of Skeletal Muscle Toxicity

Response to Injury

Mechanisms of Toxicity
 Summary

Further Reading

Chapter 11. Urinary System

Abstract

Part 1: Kidney

Part 2: Lower Urinary Tract

Acknowledgments

Further Reading

Chapter 12. Immune System

Abstract

Introduction

Mechanisms and Evaluation of Toxicity

Response to Injury

Summary

Further Reading

Chapter 13. Hematopoietic System

Abstract

Introduction

Phylogenesis and Ontogenesis

Structure, Function, and Cell Biology

Evaluation of Hematotoxicity
 Mechanisms of Toxicity

Responses of Hematopoietic Tissues to Injury

Summary

Further Reading

Chapter 14. Respiratory System

Abstract

Introduction

Structure, Function, and Cell Biology

Testing for Toxicity

Response to Injury

Mechanisms of Toxicity

Summary

Acknowledgment

Further Reading

Chapter 15. Digestive System

Abstract

Introduction

Structure and Function of the Gastrointestinal Tract

Evaluation of Gastrointestinal Toxicity

Response of the Gastrointestinal Tract to Injury

Mechanisms of Gastrointestinal Toxicity

Summary
 Further Reading

Chapter 16. Exocrine Pancreas

Abstract

Introduction

Normal Structure and Function of the Exocrine Pancreas

Evaluation of Toxicity

Responses of Exocrine Pancreas to Injury

Mechanisms of Exocrine Pancreatic Toxicity

Summary

Further Reading

Chapter 17. Male Reproductive System

Abstract

Introduction

Structure, Function, and Cell Biology

Evaluation of Toxicity

Response to Injury

Mechanisms of Toxicity

Further Reading

Chapter 18. Female Reproductive System

Abstract

Introduction
 Structure, Function, and Cell Biology

Evaluation of Female Reproductive System Toxicity

Mechanisms of Toxicity in the Female Reproductive System

Carcinogenesis in the Female Reproductive System

Further Reading

Chapter 19. The Mammary Gland

Abstract

Introduction

Mammary Gland Structure and Function

Morphologic Evaluation

Evaluation of Toxicity

Response to Injury

Mechanisms of Toxicity

Summary

Further Reading

Chapter 20. Endocrine System

Abstract

Introduction

Part 1: Adrenal Cortex

Part 2: Adrenal Medulla

Part 3: Pituitary Gland

Part 4: Thyroid C Cells

 Part 5: Thyroid Follicular Cells

Part 6: Parathyroid Gland

Part 7: Endocrine Pancreas

Summary

Acknowledgments

Further Reading

Chapter 21. Nervous System

Abstract

Introduction

Neuroanatomic Considerations in Toxicologic Neuropathology

Evaluation of Neurotoxicity

Nervous System Responses to Neurotoxic Injury

Mechanisms of Nervous System Injury

Summary

Further Reading

Chapter 22. Special Senses

Chapter 22a. Special Senses—Eye

Introduction

Structure and Function

Evaluation of Toxicity

Response to Injury by Specific Ocular Tissues

Mechanisms of Ocular Toxicity

 Summary

Further Reading

Chapter 22b. Special Senses—Ear

Introduction

Structure and Function

Testing for Otic Toxicity

Responses to Injury and Mechanisms of Injury

Summary

Further Reading

Chapter 23. Bone and Joints

Abstract

Introduction

Structure, Function, and Cell Biology of Bone and Cartilage

Evaluation of Toxicity

Responses to Injury

Mechanisms of Toxicity

Summary

Further Reading

Chapter 24. The Integumentary System

Abstract

Introduction

Structure and Function

 Response to Injury

Mechanisms of Toxicity

Summary

Further Reading

Chapter 25. Embryo, Fetus, and Placenta

Abstract

Introduction

Normal Morphologic Development

Developmental Toxicity Testing and Risk Assessment

Responses to Injury

Principles of Developmental Toxicity

Mechanisms of Developmental Toxicity

Summary

Further Reading

Index
Copyright
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Frontispiece
“To teach is to learn…” Japanese Proverb
Dedication

We are profoundly grateful to those individuals

responsible for our love of pathology and who
mentored us during our careers as well as to our
colleagues and students who inspired and supported
us along this journey.

We also thank our partners, who encouraged and

sustained us during our efforts to complete this
revision.
List of Contributors
Carl L. Alden, Mayflower Consulting, Wrenceburg, IN, United
States
Brian R. Berridge, GlaxoSmithKline Research & Development,
King of Prussia, PA, United States
Brad Bolon, GEMpath, Inc., Longmont, CO, United States
Denise I. Bounous, Bristol-Myers Squibb Co., Princeton, NJ,
United States
Mark T. Butt, Tox Path Specialists, LLC, Frederick, MD, United
States
Russell C. Cattley, Auburn University, Auburn, AL, United States
Robert E. Chapin, Retired, Pfizer Global R&D, Groton, CT, United
States
Sandrine F. Chebekoue, University of Montreal, Montreal, QC,
Canada
Dianne M. Creasy, Dianne Creasy Consulting, Norfolk, United
Kingdom
John M. Cullen, North Carolina State University, Raleigh, NC,
United States
Dimitry M. Danilenko, Genentech, South San Francisco, CA,
United States
Barbara Davis, Innogenics, Inc., Harvard, MA, United States
Kelly L. Diegel, GlaxoSmithKline Research & Development, King
of Prussia, PA, United States
David C. Dorman, North Carolina State University, Raleigh, NC,
United States
Richard R. Dubielzig, University of Wisconsin-Madison,
Madison, WI, United States
Susan A. Elmore, National Toxicology Program/NIEHS, Research
Triangle Park, NC, United States
Suzanne E. Fenton, National Institute of Environmental Health
Sciences, Research Triangle Park, NC, United States
Duncan C. Ferguson, University of Illinois at Urbana-Champaign,
Urbana, IL, United States
George L. Foley, Abbvie, Chicago, IL, United States
Robert H. Garman, Consultants in Veterinary Pathology, Inc.,
Murrysville, PA, United States
Kathryn E. Gropp, Pfizer Inc., Groton, CT, United States
Diane Gunson, Novartis Pharmaceuticals Corporation, East
Hanover, NJ, United States
Gordon C. Hard, Toxicologic Pathology Consulting, Tairua, New
Zealand
Jack R. Harkema, Michigan State University, East Lansing, MI,
United States
Wanda M. Haschek, University of Illinois at Urbana-Champaign,
Urbana, IL, United States
Eugene Herman, National Cancer Institute, Bethesda, MD, United
States
Mark J. Hoenerhoff, University of Michigan, Ann Arbor, MI,
United States
Evan B. Janovitz, Bristol-Meyer Squibb Company, Princeton, NJ,
United States
Kanwar N.M. Khan, Worldwide R&D, Pfizer, Groton, CT, United
States
Kannan Krishnan
Risk Sciences International, Ottawa, ON, Canada
University of Montreal, Montreal, QC, Canada
C. Frieke Kuper, TNO, Zeist, The Netherlands
Lois D. Lehman-McKeeman, Bristol-Myers Squibb Company,
Princeton, NJ, United States
Xiantang Li, Worldwide R&D, Pfizer, Groton, CT, United States
David E. Malarkey, National Institute of Environmental Health
Sciences, Research Triangle Park, NC, United States
Robert R. Maronpot, Maronpot Consulting LLC, Raleigh, NC,
United States
Kristen J. Nikula, Seventh Wave Laboratories, LLC, Maryland
Heights, MO, United States
Ricardo Ochoa, Pre-Clinical Safety, Inc., Niantic, CT, United States
George A. Parker, Charles River Laboratories, Inc., Hillsborough,
NC, United States
Lila Ramaiah, Bristol-Myers Squibb Co., New Brunswick, NJ,
United States
Colin G. Rousseaux, University of Ottawa, Ottawa, ON, Canada
Daniel G. Rudmann, Flagship Biosciences, Westminster, CO,
United States
Christine Ruehl-Fehlert, Bayer AG, Wuppertal, Germany
Stefan U. Ruepp, Bristol-Myers Squibb Company, Princeton, NJ,
United States
Kenneth A. Schafer, Vet Path Services, Inc., Mason, OH, United
States
John M. Sullivan, Eli Lilly & Company, Indianapolis, IN, United
States
Leandro Teixeira, University of Wisconsin-Madison, Madison, WI,
United States
John F. Van Vleet†, Purdue University, West Lafayette, IN, United
States
Aurore Varela, Charles River Laboratories, Inc., Senneville, QC,
Canada
Matthew A. Wallig, University of Illinois at Urbana-Champaign,
Urbana, IL, United States
Zbigniew W. Wojcinski, Drug Development Preclinical Services,
LLC, Ann Arbor, MI, United States
†In Memoriam
Preface
Matthew A. Wallig, Brad Bolon, Wanda M. Haschek, Colin G.
Rousseaux and Beth W. Mahler
Fundamentals of Toxicologic Pathology, as stated in the preface of its
first edition, “examines the interface between toxicology and
pathology, providing an overview of structural alterations caused by
toxicants and the mechanisms which result in those changes.” It is
designed as a “textbook” with easy and ready access to core
information about toxicologic pathology. It is divided into two
sections, Part I containing basic information pertaining to toxicology
and pathology, and Part II in an organ systems format. The basis for
this textbook is the Handbook of Toxicologic Pathology, a much larger
reference work that comprehensively addresses virtually all aspects
of toxicologic pathology and its multiple interfaces with numerous-
related scientific disciplines. This edition updates and expands
information presented in the second edition of Fundamentals of
Toxicologic Pathology, not only to provide additional necessary core
information regarding essential toxicologic principles, core
mechanisms of injury, and basic tissue responses to toxic injury (Part
I) but also to provide organ system-specific mechanisms of toxicity,
responses to toxic injury, and basic methods of evaluating injury
(Part II). The mammary gland and special senses have been added to
the organ systems section.
As with the first and second editions, this third edition is focused
toward entry-level professionals in the field of toxicologic pathology,
mainly pathology/toxicology residents and graduate students, who
require a brief but comprehensive overview that addresses the
integration of structural and functional changes, which occur with
toxic injury. In addition, this textbook is also a useful reference for
more experienced pathologists, toxicologists, and other health
professionals. Whether you are new to the field or already an
accomplished practitioner, the editors and chapter authors hope that
this volume serves you well.
The editors wish to acknowledge the efforts of the original authors
of the chapters in the Handbook of Toxicologic Pathology, third edition,
many of whom actively participated in updating and revising the
chapters for this book.
CHAPTER 1

An Overview of Toxicologic
Pathology
Colin G. Rousseaux1, Matthew A. Wallig2, Wanda M. Haschek2 and Brad
Bolon3, 1University of Ottawa, Ottawa, ON, Canada, 2University of Illinois at Urbana-Champaign,
Urbana, IL, United States, 3GEMpath, Inc., Longmont, CO, United States

OUTLINE

Introduction 1
What Is Toxicologic Pathology? 3
The Basis of Toxicologic Pathology 3
The Challenges Facing Toxicologic Pathology 4
Training and Certification in Toxicologic Pathology 7
The “Practitioner” of Toxicologic Pathology 7
Regulatory (Industrial) Toxicologic Pathology 8
Toxicologic Pathology Related to the Environment and Food
Safety 9
Diagnostic Toxicologic Pathology 9
Investigative Toxicologic Pathology 10
Management Roles for Toxicologic Pathologists 10
Summary 11
Further Reading 12
Introduction
Humans, animals, and/or the environment are exposed to dozens of
xenobiotics (exogenous compounds or materials) each day, and
thousands of xenobiotics over the course of time. These agents may
be encountered as a single substance or in complex mixtures, in
doses large or small, and for a limited period, intermittently or
continuously over time. The impact of xenobiotics ranges from no
detectable effect to toxicity of various severity to death. Toxicologic
pathologists are instrumental in protecting the well-being of
humans, animals, and the environment.
Even though toxicologic pathology was practiced for decades
prior to the 1970s, widespread use of toxicologic pathology did not
begin until the declaration of the “War on Cancer” in the United
States by President Richard Nixon under the National Cancer Act in
1971. Over the past 50 years, this effort led to a major effort to
identify carcinogens and toxicants that occur in the environment and
workplace. At this time the National Cancer Institute (NCI), a
subdivision of the US National Institutes of Health (NIH), initiated
the first large-scale testing program, known as the bioassay program.
In 1978 this program was transferred to the National Toxicology
Program (NTP), located within the National Institute of
Environmental Health Sciences (NIEHS), another subdivision of the
NIH. To date, over 500 two-year rodent toxicity and carcinogenicity
studies have been conducted by the NCI/NTP, resulting in
identification of numerous environmental and workplace hazards.
Other federal agencies in the United States participating in this
effort through the use of animal models, in vitro toxicology and
molecular biology include the National Institute for Occupational
Safety and Health (NIOSH) and the Environmental Protection
Agency (EPA). In addition the US Food and Drug Administration
(FDA) is responsible for ensuring the safety and efficacy of human
and veterinary drugs, biological products, and medical devices as
well as the safety of the food supply, cosmetics, products that emit
radiation, and tobacco products. Similar programs to address
concerns about the impact of environmental toxicants on human
health also have been created in other countries, with well-known
laboratories including the Fraunhofer Institute for Toxicology and
Experimental Medicine in Hannover, Germany; the Institute for
Applied Scientific Research (TNO) in the Netherlands; the Maltoni
Institute in Bologna, Italy; and the British Industrial Biological
Research Association (BIBRA). Together, these programs have
substantially decreased human exposure to potentially harmful
agents and thus have improved public health throughout the world.
Societies in developed countries have responded to concerns
raised by focus groups and the general public by introducing
legislation to address a number of issues regarding not only the
effects of chemical dispersal on the environment but also the safety
of products developed for industrial use. The increasing demands
placed on industry to demonstrate that their products are “safe”
have catalyzed an expanded interest in toxicologic pathology as a
key means to develop better assessments of safety.
Industry, whether chemical or pharmaceutical, must provide
safety assessment of their products to appropriate government
agencies for approval prior to marketing. To address this
requirement, an increasing number of pathologists have been
recruited to evaluate anatomic pathology (morphologic) and clinical
pathology (biochemical and hematologic) changes following
exposure to xenobiotics of concern under controlled experimental
conditions (i.e., safety testing). Since long-term exposure is
anticipated with many environmental contaminants, a major focus of
testing has been carcinogenesis using the 2-year rodent bioassay.
Good Laboratory Practices (GLP) regulations, central to the
industrial toxicologic pathologist’s work, have been developed and
implemented following exposure of fraud in laboratory testing of
chemicals in the late 1970s. These regulations ensure the uniformity,
reliability, and integrity of the data produced during safety
assessment. Periodically, GLP regulations are revised to ensure that
animal-derived data used to predict potential human responses are
of the highest possible quality.
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5. The Shaft, and its Summit magnified.
 ERICA verticillata.

CHARACTER SPECIFICUS.

Erica, antheris aristatis, inclusis; corollis pollicaribus, cernuis, ventricoso-

cylindracis, saturate rubris, pedunculis longissimis; foliis quaternis.

DESCRIPTIO.

Caulis erectus, bipedalis, basi simplicissimus, rami verticillati,

subsimplices, erecto-patentes.
Folia quaterna, glabra, lineari, acuta, subtus sulcata, recta.
Flores in apicibus ramorum verticillati, nutantes, conferti; pedunculi
longissimi, bracteis tribus ad basin instructi.
Calyx. Perianthium tetraphyllum, foliolis lato-ovatis, acuminatis, apice
dilute rubris, basi viridis.
Corolla ventricoso-cylindrica, pollicaris, ad basin-tetragona, profunde
costata; ore arctata, tenuissime laciniata; saturate rubentia.
Stamina. Filamenta octo capillaria antheræ aristatæ, inclusæ.
Pistillum. Germen globosum, sulcatum. Stylus filiformis longitudine
corollæ. Stigma tetragonum.
Habitat ad Caput Bonæ Spei.
Floret a mensi Julii, in Novembrem.

REFERENTIA.

1. Calyx, et Corolla.
2. Calyx, lente auctus.
3. Stamina, et Pistillum.
4. Stamina a Pistillo diducta, anthera una lente aucta.
5. Stylus, et Stigma, lente aucta.
 SPECIFIC CHARACTER.

Heath, with bearded tips, within the blossoms, which are an inch long,
hanging downward, of the shape of an inflated cylinder, deep red, and very
long foot-stalks; leaves growing by fours.

DESCRIPTION.

Stem upright, grows two feet high, simple at the bottom, the branches
growing in whorls, nearly simple, upright, and spreading.
Leaves grow by fours, smooth, linear, sharp-pointed, furrowed on the
under part, and growing straight out from the stem.
Flowers grow in whorls, at the upper part of the branches, hanging
downward, and close together, with long foot-stalks, that have three floral
leaves just at their base.
Empalement. Cup four-leaved, which are of a broad-oval shape, sharp-
pointed, the upper part of a light red, the bottom of a green colour.
Blossom is of a swelled-cylindrical shape, an inch long, four-cornered at
the base, and deeply ribbed; the mouth is narrowed, and very slightly cut; of
a deep red colour.
Chives. Eight hair-like threads; the tips are bearded, and within the
blossom.
Pointal. Seed-vessel round, and furrowed. Shaft thread-shaped, the
length of the blossom. Summit four-cornered.
Native of the Cape of Good Hope.
Flowers from July, till November.

REFERENCE.

1. The Empalement, and Blossom.

2. The Empalement, magnified.
3. The Chives, and Pointal.
4. The Chives detached from the Pointal; one tip magnified.
5. The Shaft, and its Summit, magnified.
 ERICA versicolor.

CHARACTER SPECIFICUS.

Erica, antheris muticis, subinclusis; stylo exserto; corollis subcylindraceis,

costatis, variè coloratis; foliis ternis.

DESCRIPTIO.

Caulis fruticosus, bipedalis, erectus, attenuatus in apicem; ramis

simplicibus.
Folia terna, linearia, subulata, supra plana, subtus sulcata, glabra,
saturate viridia; petiolis brevissimis, adpressis.
Flores plures, subcernui, tres quatuorve ramuli terminantes prope caulis
summitatem; pedunculi brevissimi.
Calyx. Perianthium duplex, interius tetraphyllum, foliolis aurantiis,
apice sulcatis, adpressis; exterius triphyllum, priori brevior, apice virescens.
Corolla subcylindracea, medio paulula arctata, costata, ima parte
aurantia, apice virescens; laciniis reflexis.
Stamina. Filamenta octo, ima parte spathulata, summa linearia,
receptaculo inserta. Antheræ muticæ, subinclusæ.
Pistillum. Germen orbiculatum, sulcatum. Stylus filiformis, exsertus.
Stigma tetragonum, virescens.
Habitat ad Caput Bonæ Spei.
Floret a mense Octobris, in Aprilem.

REFERENTIA.

1. Calyx, et Corolla.
2. Calyx lente auctus.
3. Stamina et Pistillum.
 4. Stamina a Pistillo diducta; et stamen unum lente auctum.
5. Stylus, et Stigma, lente aucta.

SPECIFIC CHARACTER.

Heath with beardless tips, just within the blossom, the pointal without; the
blossoms are nearly cylindrical, ribbed, and variously coloured; leaves grow
by threes.

DESCRIPTION.

Stem shrubby, grows two feet high, erect, tapering to the top; the
branches are nearly simple.
Leaves grow by threes, linear, awl-shaped, flat on the upper, and
furrowed on the under surface, smooth, and of a deep green; with very short
foot-stalks pressed to the branches.
The Flowers are numerous, bending a little downward, terminating the
small branches by threes or fours, near the top of the stem; the foot-stalks
being very short.
Empalement. The Cup is double, the inner is four-leaved, which are
orange-coloured, furrowed at the point, and pressed to the blossom; the
outer is three-leaved, shorter than the inner, and greenish at the point.
The Blossom is nearly cylindrical, a little narrowed at the middle,
ribbed, the lower part orange colour, the mouth greenish, whose segments
are reflexed.
Chives. Eight threads, whose lower parts are spatula-shaped, and upper
linear, fixed into the receptacle. Tips beardless, and almost within the
blossom.
Pointal. Seed-vessel globe-shaped, and furrowed. Shaft thread-shaped,
and without the blossom. Summit four-cornered and greenish.
Native of the Cape of Good Hope.
Flowers from October, till April.

REFERENCE.
1. The Empalement, and Blossom.
2. The Empalement magnified.
3. The Chives and Pointal.
4. The Chives detached from the Pointal; one tip with its thread, magnified.
5. The Shaft, and its Summit, magnified.
 ERICA vestita alba.

CHARACTER SPECIFICUS.

Erica, antheris muticis, sub-inclusis; folia sena, tremulenta, linearia,

conferta, attenuata in petiolos capillares; floribus clavatis, albidis.

DESCRIPTIO.

Caulis tripedalis, laxus, erectus, basi simplicissimus, ramis simplicibus

adscendentibus.
Folia sena, conferta, glabra, linearia, tremulenta, subtus sulcata,
attenuata in petioles capillares semiunguiculares.
Flores in ramulis verticillati, erecto-patentes; pedunculis brevibus
bracteis tribus instructis.
Calyx. Perianthium tetraphyllum, foliolis subulatis, adpressis, glabris,
viscosis.
Corolla clavata, pollicaris, alba, pubescens; oris laciniis patentibus.
Stamina. Filamenta octo capillaria; antheræ muticæ, subinclusæ.
Pistillum. Germen turbinatum, summum pilis longis coronatum. Stylus
filiformis, exsertus. Stigma tetragonum.
Habitat ad Caput Bonæ Spei.
Floret a mense Julii in Decembrem.

REFERENTIA.

1. Calyx et Corolla.
2. Calyx lente auctus.
3. Stamina et Pistillum.
4. Stamina a Pistillo diducta; anthera una lente aucta.
5. Stylus et Stigma lente aucti.
 SPECIFIC CHARACTER.

Heath, with beardless tips, just within the blossoms; leaves grow by sixes,
are tremulous, linear, and crowded together, tapering into hair-like foot-
stalks; the flowers are club-shaped, and white.

DESCRIPTION.

Stem grows three feet high, loose and upright, simple at the base, with
simple ascending branches.
Leaves grow by sixes, crowded together, smooth, linear, tremulous,
furrowed underneath, and tapering into hair-like foot-stalks a quarter of an
inch in length.
Flowers grow in whorls upon the smaller branches, upright, and
spreading; with short foot-stalks having three floral leaves.
Empalement. Cup four-leaved, which are awl-shaped, pressed to the
blossom, smooth, and clammy.
Blossom club-shaped, an inch long, white and downy, the segments of
the mouth are spread outward.
Chives. Eight hair-like threads, tips beardless, and just within the
blossom.
Pointal. Seed-vessel turban-shaped, the top-being crowned with long
hairs. Shaft thread-shaped, and without the blossom. Summit four-cornered.
Native of the Cape of Good Hope.
Flowers from July till December.

REFERENCE.

1. The Empalement and Blossom.

2. The Empalement magnified.
3. The Chives and Pointal.
4. The Chives detached from the Pointal; one tip magnified.
5. The Shaft and its Summit magnified.
 ERICA vestita purpurea.

CHARACTER SPECIFICUS.

Erica, antheris muticis, subexsertis; foliis senis, linearibus, tremulentibus,

attenuata in petiolos capillares, semiunguiculares; floribus axillaribus,
clavatis, purpureis.

DESCRIPTIO.

Caulis fruticosus, bipedalis, erectus, basi simplicissimus, dein

verticillatim ramosus, ramis simplicibus, adscendentibus.
Folia sena, linearia, tremulanta, acuta, glabra, subtus leviter sulcata,
attenuata in petiolos capillares, semiunguiculares.
Flores in ramulis mediis verticillati, recti; pedunculis brevibus, bracteis
tribus instructis.
Calyx. Perianthium tetraphyllum, foliolis lanciolatis, basi serratis,
viridis, glabris.
Corolla clavata, pollicaria, obtusa, recta, purpurea, ore arctato, limbo
quadrifido, reflexo.
Stamina. Filamenta octo, capillaria, receptaculo inserta. Antheræ
muticæ, subexsertæ.
Pistillum. Germen turbinatum, apice, pilis longis, plumosis, octofarium
coronatum. Stylus filiformis, exsertus. Stigma tetragonum.
Habitat ad Caput Bonæ Spei.
Floret a mensi Julii in Januarium.

REFERENTIA.

1. Folium unum cum petiolo.

2. Calyx et Corolla.
 3. Calyx lente auctus.
4. Stamina et Pistillum.
5. Stamina a Pistillo diducta; anthera una lente aucta.
6. Stylus et Stigma lente aucta.

SPECIFIC CHARACTER.

Heath, with beardless tips, almost without the blossom; leaves grow by
sixes, linear, tremulous, and tapering into hair-like foot-stalks a quarter of an
inch long; the flowers grow from the base of the leaves, are club-shaped,
and purple.

DESCRIPTION.

Stem shrubby, grows two feet high, and upright, simple at the base, then
branching out from a whorl into simple ascending branches.
Leaves grow by sixes, are linear, tremulous, sharp-pointed, smooth,
lightly furrowed on the under part, and tapering into hair-like leaf-stems a
quarter of an inch in length.
Flowers are in whorls about the middle of the branches, straight out,
having short foot-stalks with three floral leaves.
Empalement. Cup four-leaved, which are lance-shaped, sawed at the
base, green, and smooth.
Blossom club-shaped, an inch long, blunt, straight, and purple,
compressed at the mouth, with a four-lobed, reflexed border.
Chives. Eight hair-like threads fixed into the receptacle. Tips beardless,
and nearly of a length with the blossom.
Pointal. Seed-vessel turban-shaped, the top crowned by eight plumes of
long hairs. Shaft thread-shaped, and without the blossom. Summit four-
cornered.
Native of the Cape of Good Hope.
Flowers from July till January.
 REFERENCE.

1. A leaf with its foot-stalk.

2. The Empalement and Blossom.
3. The Empalement magnified.
4. The Chives and Pointal.
5. The Chives detached from the Pointal; one tip magnified.
6. The Shaft and its Summit magnified.
 ERICA vestita coccinea.

CHARACTER SPECIFICUS.

Erica, antheris muticis, sub-exsertis; foliis senis, tremulis, linearibus,

attenuatis in petiolos capillares; corollis clavatis, pollicaribus, coccineis.

DESCRIPTIO.

Caulis erectus, sesquipedalis, parum ramosus, ramulis simplicibus, foliis

tectis.
Folia sena, linearia, tremula, glabra, subtus sulcata, attenuata in petiolos
capillares semiunguiculares.
Flores in ramulis mediis conferti, numerosi, erecto-patentes, pedunculis
brevibus, bracteis tribus linearibus adpressis.
Calyx. Perianthium tetraphyllum, foliolis subulatis adpressis glabris.
Corolla clavata, pollicaris, coccinea, oris laciniis revolutis.
Stamina. Filamenta octo capillaria, receptaculo inserta; antheris muticis,
sub-exsertis.
Pistillum. Germen turbinatum, sulcatum. Stylus filiformis. Stigma
tetragonum.
Habitat ad Caput Bonæ Spæi.
Floret a mense Augusto in Novembrem.

REFERENTIA.

1. Calyx et Corolla.
2. Calyx lente auctus.
3. Stamina et Pistillum.
4. Stamina a Pistillo diducta; antherâ unâ lente auctâ.
5. Stylus et Stigma lente aucti.
 SPECIFIC CHARACTER.

Heath, with beardless tips just without the blossom; the leaves grow by
sixes, are tremulous, linear, and taper into hair-like foot-stalks; the blossoms
are club-shaped, an inch long, and deep scarlet.

DESCRIPTION.

Stem upright, grows a foot and a half high, branching but little; the
smaller branches are simple, and covered with leaves.
Leaves grow by sixes, are linear, tremulous, smooth, furrowed
underneath, and tapering into hair-like foot-stalks half an inch long.
Flowers are clustered together about the middle of the branches, are
numerous, spreading upwards, having short foot-stalks, with three linear
floral leaves pressed to the cup.
Empalement. Cup four-leaved, which are awl-shaped, pressed to the
blossom, and smooth.
Blossom club-shaped, an inch long, and of a deep scarlet colour; the
segments of the mouth are rolled back.
Chives. Eight hair-like threads, fixed into the receptacle; tips beardless,
and just without the blossom.
Pointal. Seed-vessel turban-shaped, and furrowed. Shaft thread-shaped.
Summit four-cornered.
Native of the Cape of Good Hope.
Flowers from August till November.

REFERENCE.

1. The Empalement and Blossom.

2. The Empalement magnified.
3. The Chives and Pointal.
4. The Chives detached from the Pointal; one tip magnified.
5. The Shaft and its Summit magnified.