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 Platelets
Fourth Edition
Platelets

FOURTH EDITION

Editor
ALAN D. MICHELSON, MD
Professor of Pediatrics, Harvard Medical School/Boston Children’s Hospital
Professor of Medicine, Harvard Medical School/Brigham and Women’s Hospital
Director, Center for Platelet Research Studies
Director, Thrombosis and Anticoagulation Program
Dana-Farber/Boston Children’s Cancer and Blood Disorders Center
Boston, Massachusetts, USA

Associate Editors

MARCO CATTANEO, MD
Professor of Internal Medicine
Director, Unit of Internal Medicine
Ospedale San Paolo
Università degli Studi di Milano
Milan, Italy

ANDREW L. FRELINGER III, PhD

Assistant Professor of Pediatrics, Harvard Medical School
Associate Director, Center for Platelet Research Studies
Staff Scientist, Dana-Farber/Boston Children’s Cancer and Blood Disorders Center
Boston, Massachusetts, USA

PETER J. NEWMAN, PhD

Vice President for Research, BloodCenter of Wisconsin
Jacquelyn Fredrick Endowed Chair for Foundational Research and Associate
Director, Blood Research Institute
Milwaukee, Wisconsin, USA
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First edition 2002

Second edition 2007
Third edition 2013
Fourth edition 2019

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Notices
Knowledge and best practice in this field are constantly changing. As new research and experience broaden our understanding,
changes in research methods, professional practices, or medical treatment may become necessary.

Practitioners and researchers must always rely on their own experience and knowledge in evaluating and using any
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Cover image: Platelets Adherent to a Surgical Suture. Colorized scanning electron micrograph of activated, aggregated and spread
platelets (blue) adherent to a synthetic-polymer surgical suture (straw-colored). Original magnification x800. Sample provided by
Alan D. Michelson, Emma E. Forde and Andrew L. Frelinger, Center for Platelet Research Studies, Dana-Farber/Boston Children's
Cancer and Blood Disorders Center, Harvard Medical School, Boston, Massachusetts, U.S.A. Image provided by Steve
Gschmeissner (https://theworldcloseup.com).
 Contributors
Joseph Alsousou Chapel Hill, NC Carol Briggs
Institute of Translational Medicine United States Department of Haematology
University of Liverpool University College London
Gerald Bertrand
Liverpool Hospitals
EFS
United Kingdom London
Rennes
United Kingdom
Dominick J. Angiolillo France
Division of Cardiology Tomasz Brzoska
Deepak L. Bhatt
University of Florida College of Pittsburgh Heart, Lung and Blood
Department of Cardiovascular
Medicine-Jacksonville Vascular Medicine Institute;
Medicine
Jacksonville, FL Sickle Cell Center of Excellence
Brigham and Women’s Hospital
United States University of Pittsburgh-School of
Harvard Medical School
Amal Arachiche Boston, MA Medicine
Department of Pharmacology United States Pittsburgh, PA
Case Western Reserve University United States
Thomas A. Blair
Cleveland, OH James B. Bussel
Center for Platelet Research Studies
United States Department of Pediatrics
Dana-Farber/Boston Children’s Cancer
Richard H. Aster and Blood Disorders Center Weill Medical College of Cornell
Blood Research Institute Harvard Medical School University
BloodCenter of Wisconsin Boston, MA New York, NY
Department of Medicine United States United States
Medical College of Wisconsin
Kamila Bledzka Marco Cattaneo
Milwaukee, WI
Joseph J. Jacobs Center for Thrombosis Unità di Medicina 2
United States
and Vascular Biology ASST Santi Paolo e Carlo
Tiziano Barbui Department of Molecular Cardiology Dipartimento di Scienze della Salute
FROM Research Foundation Cleveland Clinic Università degli Studi di Milano
Papa Giovanni XXIII Hospital Cleveland, OH Milan
Bergamo United States Italy
Italy
Oliver Borst Subarna Chakravorty
Stefania Basili University Hospital King’s College NHS Foundation
I Clinica Medica Department of Cardiology and Trust
Department of Internal Medicine Cardiovascular Medicine London
and Medical Specialties Eberhard Karls University Tuebingen United Kingdom
SAPIENZA-University of Rome Tuebingen
Rome Germany Noel C. Chan
Italy Thrombosis and Atherosclerosis
Emma G. Bouck
Research Institute
Elisabeth M. Battinelli Department of Pharmacology
Department of Medicine
Division of Hematology Case Western Reserve University
McMaster University
Brigham and Women’s Hospital Cleveland, OH
Hamilton, ON
Harvard Medical School United States
Canada
Boston, MA
Lawrence F. Brass
United States Shruti Chaturvedi
Division of Hematology-Oncology
Anthony A. Bavry Department of Medicine Division of Hematology
Department of Medicine Perelman School of Medicine Johns Hopkins University
University of Florida University of Pennsylvania Baltimore, MD
North Florida/South Georgia Veterans Philadelphia, PA United States
Health System United States Beng H. Chong
Gainesville, FL Department of Medicine
Paul F. Bray
United States St. George Clinical School
Molecular Medicine Program
Wolfgang Bergmeier Division of Hematology and University of New South Wales;
McAllister Heart Institute Hematologic Malignancies in the Department of Haematology
Department of Biochemistry and Department of Internal Medicine NSW Health Pathology
Biophysics University of Utah St George Hospital
University of North Carolina at Salt Lake City, UT Sydney
Chapel Hill United States Australia
vii
viii Contributors

Kenneth J. Clemetson Centro de Hemodonacion Department of Medicine

Department of Haematology IMIB-Murcia Harvard Medical School
University of Berne Inselspital CIBERER (CB15/00055) Boston, MA
Theodor Kocher Institute UCAM United States
University of Berne Murcia
Berne Spain Andreas Greinacher
Switzerland Department of Immunology and
Guido Finazzi Transfusion Medicine
Jeannine M. Clemetson Hematology Unit Universit€atsmedizin
Theodor Kocher Institute Papa Giovanni XXIII Hospital Greifswald
University of Berne Bergamo Germany
Berne Italy
Switzerland Thomas Gremmel
Barry S. Coller Robert Flaumenhaft Department of Internal Medicine II
Allen and Frances Adler Laboratory of Division of Hemostasis and Medical University of Vienna
Blood and Vascular Biology Thrombosis Vienna;
Rockefeller University Hospital Department of Medicine Department of Internal Medicine
Rockefeller University Beth Israel Deaconess Medical Center Cardiology and Nephrology
New York, NY Harvard Medical School Landesklinikum Wiener Neustadt
United States Boston, MA Wiener Neustadt
United States Austria
Gregory J. del Zoppo
Division of Hematology Jane E. Freedman Paul A. Gurbel
Department of Medicine Department of Medicine Inova Center for Thrombosis Research
University of Washington School of University of Massachusetts Medical and Drug Development
Medicine School Inova Heart and Vascular Institute
Harborview Medical Center Worcester, MA Falls Church, VA
Seattle, WA United States United States
United States
Andrew L. Frelinger, III Elizabeth J. Haining
Jenny M. Despotovic Center for Platelet Research Studies
Department of Pediatrics Institute of Cardiovascular Sciences
Dana-Farber/Boston Children’s College of Medical and Dental
Baylor College of Medicine Cancer and Blood Disorders
Houston, TX Sciences
Center University of Birmingham
United States Harvard Medical School Birmingham
Scott L. Diamond Boston, MA United Kingdom
Department of Chemical and United States
Biomolecular Engineering Xu Han
Kathleen Freson
University of Pennsylvania Department of Pharmacology
Department of Cardiovascular
Philadelphia, PA Case Western Reserve University
Sciences
United States Cleveland, OH
Center for Molecular and Vascular
United States
J. Donald Easton Biology
Department of Neurology KU Leuven Paul Harrison
University of California, San Francisco Leuven Institute of Inflammation and
Sandler Neuroscience Center Belgium Ageing
San Francisco, CA College of Medical and Dental
United States Aleksandra Gasecka
First Chair and Department of Sciences
Koji Eto Cardiology University of Birmingham
Department of Clinical Application Medical University of Warsaw Birmingham
Center for iPS Cell Research and Warsaw United Kingdom
Application Poland;
Kyoto University Catherine P.M. Hayward
Laboratory of Experimental Clinical
Kyoto; Departments of Pathology
Chemistry
Department of Innovative Medicine and Molecular Medicine and
Vesicle Observation Centre
Chiba University Graduate School Medicine
Academic Medical Centre of the
of Medicine McMaster University
University of Amsterdam
Chiba Hamilton, ON
Amsterdam
Japan Canada
The Netherlands

Falet
Herve Karin Hoffmeister
Blood Research Institute Meinrad Gawaz
University Hospital Division of Hematology
BloodCenter of Wisconsin Brigham and Women’s Hospital;
Department of Cell Biology, Department of Cardiology and
Cardiovascular Medicine Department of Medicine
Neurobiology, and Anatomy Harvard Medical School
Medical College of Wisconsin Eberhard Karls University
Tuebingen Boston, MA;
Milwaukee, WI Blood Research Institute
United States Tuebingen
Germany BloodCenter of Wisconsin;
Francisca Ferrer-Marin Department of Biochemistry
Unidad de Hematología y Oncología Silvia Giannini Medical College of Wisconsin
M
edica Division of Hematology Milwaukee, WI
Hospital Morales-Meseguer Brigham and Women’s Hospital; United States
 Contributors ix

Anne-Mette Hvas Kumaran Kolandaivelu Charleston, SC

Department of Clinical Biochemistry Department of Cardiovascular United States
Aarhus University Hospital; Medicine
Anqi Li
Department of Clinical Medicine Brigham and Women’s Hospital
Department of Microbiology and
Faculty of Health Harvard Medical School
Immunology
Aarhus University Boston, MA;
Hollings Cancer Center
Aarhus Institute for Medical Science and
Medical University of South Carolina
Denmark Engineering
Charleston, SC
Massachusetts Institute of
Sara J. Israels United States
Technology
Department of Pediatrics and Child
Cambridge, MA Rossella Liani
Health
United States Internal Medicine
University of Manitoba
Department of Medicine and Aging
Winnipeg Milka Koupenova
and Haemostasis and Thrombosis
Canada Department of Medicine
Unit
University of Massachusetts Medical
Joseph E. Italiano, Jr Center of Aging Science and
School
Hematology Division Translational Medicine
Worcester, MA
Brigham and Women’s Hospital; (CESI-Met)
United States
Vascular Biology Program University of Chieti
Boston Children’s Hospital; David J. Kuter Chieti, Italy
Harvard Medical School Department of Hematology


Marie Lordkipanidze
Boston, MA Massachusetts General Hospital
Faculty of Pharmacy
United States Harvard Medical School
University of Montreal;
Boston, MA
Young-Hoon Jeong Research Center
United States
Department of Internal Medicine Montreal Heart Institute
Gyeongsang National University Michele P. Lambert Montr
eal, QC
School of Medicine and Department of Pediatrics Canada
Cardiovascular Center Children’s Hospital of Philadelphia
Viola Lorenz
Gyeongsang National University Perelman School of Medicine at the
Division of Newborn Medicine
Changwon Hospital University of Pennsylvania
Boston Children’s Hospital
Changwon Philadelphia, PA
Harvard Medical School
South Korea United States
Boston, MA
Andrew D. Johnson Robert H. Lee United States
National Heart, Lung and Blood McAllister Heart Institute
Institute University of North Carolina at Chapel Kellie R. Machlus
The Framingham Heart Study Hill Hematology Division
Framingham, MA Chapel Hill, NC Brigham and Women’s Hospital
United States United States Harvard Medical School
Boston, MA
Cecile Kaplan Jack Levin United States
Retired, formerly Platelet Immunology Departments of Laboratory Medicine
Department and Medicine Dhruv Mahtta
I.N.T.S. University of California School of Department of Medicine
Paris Medicine University of Florida
France San Francisco, CA Gainesville, FL
United States United States
Peter Karagiannis
Department of Clinical Application Renhao Li Pier Mannuccio Mannucci
Center for iPS Cell Research and Aflac Cancer and Blood Disorders Angelo Bianchi Bonomi Hemophilia
Application Center and Thrombosis Center
Kyoto University Children’s Healthcare of Atlanta Fondazione IRCCS Ca’ Granda-
Kyoto Department of Pediatrics Ospedale Maggiore Policlinico
Japan Emory University School of Medicine Università degli Studi di Milano
Atlanta, GA Milan
Gregory J. Kato Italy
United States
Pittsburgh Heart, Lung and Blood
Vascular Medicine Institute; Zhenyu Li Keith R. McCrae
Division of Hematology and Oncology; Division of Cardiovascular Medicine Department of Hematology-Oncology
Sickle Cell Center of Excellence The Gill Heart and Vascular Cleveland Clinic
University of Pittsburgh-School of Institute Cleveland, OH
Medicine University of Kentucky United States
Pittsburgh, PA Lexington, KY
Alessandra Metelli
United States United States
Department of Microbiology and
Samuel Kemble Zihai Li Immunology
Institute of Inflammation and Ageing Department of Microbiology and Hollings Cancer Center
College of Medical and Dental Sciences Immunology Medical University of South
University of Birmingham Hollings Cancer Center Carolina
Birmingham Medical University of South Charleston, SC
United Kingdom Carolina United States
x Contributors

Alan D. Michelson Marie-Blanche Onselaer Julie Rayes

Center for Platelet Research Studies Institute of Cardiovascular Sciences Institute of Cardiovascular Sciences
Dana-Farber/Boston Children’s Cancer College of Medical and Dental Sciences College of Medical and Dental Sciences
and Blood Disorders Center University of Birmingham University of Birmingham
Harvard Medical School Birmingham Birmingham
Boston, MA United Kingdom United Kingdom
United States
Carlo Patrono Alexander P. Reiner
Karen A. Moffat Department of Pharmacology Department of Epidemiology
Department of Medicine Catholic University School of Medicine School of Public Health
McMaster University Rome Fred Hutchinson Cancer Research Center
Hamilton, ON Italy University of Washington
Canada Seattle, WA
Edward F. Plow United States
Jae Youn Moon Joseph J. Jacobs Center for Thrombosis
Division of Cardiology and Vascular Biology Brian Riesenberg
University of Florida College of Department of Molecular Cardiology Department of Microbiology and
Medicine-Jacksonville Cleveland Clinic Immunology
Jacksonville, FL Cleveland, OH Hollings Cancer Center
United States United States Medical University of South Carolina
Charleston, SC
James H. Morrissey Mortimer Poncz United States
Department of Biological Chemistry Department of Pediatrics
University of Michigan Medical School Irene A.G. Roberts
Children’s Hospital of Philadelphia Department of Paediatrics
Ann Arbor, MI Perelman School of Medicine at the
United States University of Oxford
University of Pennsylvania John Radcliffe Hospital
Nicola J. Mutch Philadelphia, PA Oxford
Institute of Medical Sciences United States United Kingdom
University of Aberdeen Man-Chiu Poon
Aberdeen Matthew T. Rondina
Division of Hematology and Department of Internal Medicine
United Kingdom Hematologic Malignancies University of Utah
Zoltan Nagy Department of Medicine Eccles Institute of Human Genetics;
Institute of Cardiovascular Sciences University of Calgary–Foothills Medical Molecular Medicine Program
College of Medical and Dental Sciences Centre University of Utah School of Medicine;
University of Birmingham Calgary, AB George E. Wahlen VAMC GRECC
Birmingham Canada Salt Lake City, UT
United Kingdom United States
Natalie S. Poulter
Heyu Ni Institute of Cardiovascular Sciences Jesse W. Rowley
St. Michael’s Hospital College of Medical and Dental Sciences Molecular Medicine Program
University of Toronto; University of Birmingham Division of Pulmonary Medicine in the
Canadian Blood Services Centre for Birmingham Department of Internal Medicine
Innovation United Kingdom University of Utah
Toronto, ON Salt Lake City, UT
Canada Izmarie Poventud-Fuentes United States
Division of Hematology-Oncology
Phillip L.R. Nicolson Department of Medicine Francesca Santilli
Institute of Cardiovascular Sciences Perelman School of Medicine Internal Medicine
College of Medical and Dental University of Pennsylvania Department of Medicine and Aging and
Sciences Philadelphia, PA Haemostasis and Thrombosis Unit
University of Birmingham United States Center of Aging Science and
Birmingham Translational Medicine
United Kingdom Patrick Provost (CESI-Met)
CHUQ Research Center/CHUL Pavilion; University of Chieti
Marvin T. Nieman Department of Microbiology Chieti
Department of Pharmacology Infectious Diseases and Immunology Italy
Case Western Reserve University Faculty of Medicine
Cleveland, OH Universit
e Laval € diger E. Scharf
Ru
United States Quebec City, QC Institute of Transplantation Diagnostics
Canada and Cell Therapeutics
Rienk Nieuwland Division of Clinical and Experimental
Laboratory of Experimental Clinical Jun Qin Hemostasis, Hemotherapy and
Chemistry; Joseph J. Jacobs Center for Thrombosis Transfusion Medicine
Vesicle Observation Centre and Vascular Biology University Blood Center and
Academic Medical Centre of the Department of Molecular Cardiology Hemophilia Comprehensive Care
University of Amsterdam Cleveland Clinic Center
Amsterdam Cleveland, OH Heinrich Heine University Medical
The Netherlands United States Center
 Contributors xi

and Biological Medical Research Center Lexington, KY Centre of Membrane Proteins and
Heinrich Heine University United States Receptors (COMPARE)
D€usseldorf University of Birmingham and
Germany Edward L. Snyder University of Nottingham
Department of Laboratory Medicine Midlands
Yotis A. Senis Yale University School of Medicine United Kingdom
Institute of Cardiovascular Sciences New Haven, CT
College of Medical and Dental Sciences United States Mark R. Thomas
University of Birmingham Institute of Cardiovascular Sciences
Birmingham Martha Sola-Visner College of Medical and Dental Sciences
United Kingdom Division of Newborn Medicine University of Birmingham
Boston Children’s Hospital Birmingham
Anish Sharda Harvard Medical School United Kingdom
Division of Hemostasis and Thrombosis Boston, MA
Department of Medicine United States Maurizio Tomaiuolo
Beth Israel Deaconess Medical Center Division of Hematology-Oncology
Harvard Medical School Timothy J. Stalker Department of Medicine
Boston, MA Division of Hematology-Oncology Perelman School of Medicine
United States Department of Medicine University of Pennsylvania
Perelman School of Medicine Philadelphia, PA
Alexa J. Siddon University of Pennsylvania United States
Department of Laboratory Medicine Philadelphia, PA
Christopher A. Tormey
Department of Pathology United States
Department of Laboratory Medicine
Yale University School of Medicine Yale University School of Medicine
Lucia Stefanini
New Haven, CT; New Haven, CT;
Department of Internal Medicine and
Pathology & Laboratory Medicine Pathology & Laboratory Medicine
Medical Specialties
VA Connecticut Healthcare VA Connecticut Healthcare
Sapienza University of Rome
West Haven, CT West Haven, CT
Rome
United States United States
Italy
Pia R.-M. Siljander Naoshi Sugimoto Han-Mou Tsai
EV Group Department of Clinical Application State University of New York Downstate
Molecular and Integrative Biosciences Center for iPS Cell Research and Medical Center
Research Programme Application Brooklyn, NY
Faculty of Biological and Environmental Kyoto University United States
Sciences; Kyoto;
EV Core Facility Francesco Violi
Department of Hematology I Clinica Medica
University of Helsinki Kagawa University Hospital
Helsinki Department of Internal Medicine and
Kagawa Medical Specialties
Finland Japan SAPIENZA-University of Rome
Pierluigi Tricoci Prithu Sundd Rome
Division of Cardiology Pittsburgh Heart, Lung and Blood Italy
Duke Clinical Research Institute Vascular Medicine Institute; Theodore E. Warkentin
Duke University Division of Pulmonary, Allergy and Department of Pathology and Molecular
Durham, NC Critical Care Medicine; Medicine
United States Sickle Cell Center of Excellence Department of Medicine
Paola Simeone University of Pittsburgh-School of Michael G. DeGroote School of
Internal Medicine Medicine Medicine
Department of Medicine and Aging Pittsburgh, PA McMaster University;
and Haemostasis and Thrombosis United States Hamilton Regional Laboratory
Unit Udaya S. Tantry Medicine Program
Center of Aging Science and Inova Center for Thrombosis Research Hamilton, ON
Translational Medicine (CESI-Met) and Drug Development Canada
University of Chieti Inova Heart and Vascular Institute Steve P. Watson
Chieti Falls Church, VA Institute of Cardiovascular
Italy United States Sciences
Stephanie A. Smith College of Medical and Dental Sciences
Ayalew Tefferi
Department of Biological Chemistry University of Birmingham
Division of Hematology
University of Michigan Medical School Birmingham
Department of Internal Medicine
Ann Arbor, MI United Kingdom
Mayo Clinic
United States Rochester, MN Jeffrey I. Weitz
United States Thrombosis and Atherosclerosis
Susan S. Smyth Research Institute;
Division of Cardiovascular Medicine Steven G. Thomas Department of Medicine
The Gill Heart and Vascular Institute Institute of Cardiovascular Sciences McMaster University
University of Kentucky; University of Birmingham Hamilton, ON
Lexington VA Medical Center Birmingham; Canada
xii Contributors

John Welsh The BloodCenter of Wisconsin Li Zhu

Division of Hematology-Oncology Milwaukee, WI Cyrus Tang Hematology Center
Department of Medicine United States Soochow University
Perelman School of Medicine Suzhou
Bill X. Wu
University of Pennsylvania China
Department of Microbiology and
Philadelphia, PA
Immunology
United States Guy A. Zimmerman
Hollings Cancer Center
Department of Internal
Andrew S. Weyrich Medical University of South
Medicine
Molecular Medicine Program Carolina
University of Utah
Division of Pulmonary Medicine Charleston, SC
Eccles Institute of Human Genetics;
in the Department of Internal United States
Molecular Medicine Program
Medicine
Michael R. Yeaman University of Utah School of
University of Utah
Department of Medicine Medicine
Salt Lake City, UT
David Geffen School of Medicine at Salt Lake City, UT
United States
University of California, Los Angeles United States
David A. Wilcox (UCLA)
Department of Pediatrics Torrance and Los Angeles Biomedical Elizabeth R. Zunica
Medical College of Wisconsin; Research Institute at Department of Pharmacology
Children’s Research Institute Harbor UCLA Medical Center Case Western Reserve University
The Children’s Hospital of Wisconsin Torrance, CA Cleveland, OH
Blood Research Institute United States United States
 Preface

Platelets are small cells of great importance in many patho- Marco Cattaneo, A.L. “Larry” Frelinger, and Peter J. Newman.
physiological processes including thrombosis, hemorrhage, There are 142 contributing authors from 18 countries.
inflammation, antimicrobial host defense, wound healing, The 67 chapters in this book are organized into six parts:
angiogenesis, and tumor growth and metastasis. In addition
Part I: Platelet Biology
to primary disorders of platelet number and function, platelets
Part II: The Role of Platelets in Disease
have a critical role in many other very common diseases,
Part III: Clinical Tests of Platelet Function
including coronary artery disease, stroke, peripheral vascular
Part IV: Disorders of Platelet Number and/or Function
disease, and diabetes mellitus.
Part V: Antiplatelet Therapy
The goal of this book is to be a comprehensive and definitive
Part VI: Therapy to Increase Platelet Numbers and/or Function
source of knowledge about platelets. Platelets integrates the
entire field of platelet biology, pathophysiology, and clinical Each chapter has been completely revised and updated, and
medicine. there are 11 new chapters on topics such as platelet glycobiol-
The intended audience for Platelets includes hematologists, ogy, the platelet transcriptome, platelet inhibitory receptors,
cardiologists, stroke physicians, blood bankers, pathologists, platelet function testing in clinical research trials, therapeutic
and researchers in thrombosis and hemostasis, as well as stu- platelet-rich plasma in wound healing, and new antiplatelet
dents and fellows in these fields. drugs.
The first edition of this book was the winner of the Best Book
in Medical Science Award from the Association of American Pub- Alan D. Michelson
lishers, and the third edition won the Highly Commended
Marco Cattaneo
Prize in the category of Internal Medicine from the British Med-
ical Association (BMA) Medical Book Awards. These awards A.L. “Larry” Frelinger
reflect the outstanding international group of contributing
Peter J. Newman
authors, all of whom are world leaders in their field.
In this fourth edition of Platelets, Alan Michelson, the editor
of the first three editions, is joined by three associate editors:

xiii
 Foreword: A Brief History
of Ideas About Platelets
in Health and Disease
Barry S. Coller
Allen and Frances Adler Laboratory of Blood and Vascular Biology, Rockefeller University Hospital, Rockefeller University, New York, NY, United States

Credible claims have been made by, or on behalf of, a number of intellectual development in the field and a context for under-
of different scientists to be recognized as the first to describe the standing our current concepts and likely future directions.
blood platelet.1,2 For example, Osler undoubtedly described Those interested in a more systematic history of the discovery
platelets in his 1873 and 1874 papers (Fig. 1),1,3 recognizing of the platelet and its functions can consult a number of excel-
their disk-like structure, and that while they circulated singly lent references.1, 9, 15–29 In addition, the Historical Sketch sec-
in the blood, they rapidly formed aggregates when removed tion of the Journal of Thrombosis and Haemostasis has provided
from the blood vessel. Osler was not certain, however, whether extremely valuable first-hand accounts of discoveries related
the platelets were normal blood elements or exogenous “organ- to platelets by prominent members of the platelet research
isms.”1,3 Seven years later, the elegant and compelling studies community.30–42
of Bizzozero in 1881 and 1882 not only identified the platelet
anatomically, but also assigned it roles in both hemostasis and THE ROLE OF PLATELETS IN HUMAN HEMOSTASIS,
experimental thrombosis.4–9 Thus, using intravital microscopy
of mesenteric venules in guinea pigs, Bizzozero also observed
PLATELET TRANSFUSION, AND BLEEDING TIME
that platelets are disk-shaped and circulate in isolation. He Among the earliest evidence that platelets are crucial for human
went on to demonstrate that they adhere to the blood vessel hemostasis and that platelet transfusions can restore hemo-
wall at sites of injury and then form aggregates. He additionally static competence to individuals with low platelet counts, are
noted that leukocytes were recruited into the platelet aggre- the compelling data from one of the three patients reported
gates, thus also offering the first description of platelet- by Duke, a student of Wright, in 1910 (Fig. 6).12,20,27,43 A
leukocyte interactions (Fig. 2). Bizzozero also developed the 20-year-old Armenian man was admitted to the Massachusetts
first “flow chamber,” with the flow achieved by capillary action, General Hospital on May 8, 1909 with a platelet count of
by an ingenious construction of a microscope slide, a piece of 6  109/L and profound mucocutaneous bleeding. On May
thread, and blotting paper (Fig. 3).7,8 Using human biopsy 11, he developed uncontrollable epistaxis that left him nearly
materials, Osler later built on Bizzozero’s observations and, moribund. In desperation, a donor was selected from among
in studies conducted from 1881 to 1886, clearly established his young Armenian friends and a direct blood transfusion
the role of blood platelets (which he termed blood “plaques”) was arranged. A “large” amount of blood was transfused, as
in human thrombotic disease, identifying them in white judged by the increase in pulse of the donor as well as the rise
thrombi that formed on “atheromatous ulcers,” in vegetations in the platelet count in the recipient to 123  109/L! All overt
on heart valves, and in aortic aneurysms (Fig. 4).1,13 signs of bleeding improved immediately after the transfusion.
Ironically, 12 years before his publications on platelets, Biz- To monitor the response, Duke used the bleeding time assay he
zozero was also the first to identify bone marrow megakaryo- had just developed, analyzing the time for bleeding to stop
cytes (Fig. 5),10 but he never recognized them as the from a standardized wound in the ear lobe. The improvement
precursors of platelets. That discovery was made by Wright in in the patient’s bleeding time correlated with his higher platelet
1906, who noted the similarities in shape and color of the count and his improvement in clinical bleeding (Fig. 7). Since
red to violet granules in platelets and megakaryocytes using the patient’s coagulation time was normal throughout, Duke’s
his new polychrome staining solution (Wright’s stain).14 data provided strong evidence that the low platelet count itself
The intervening 135 years since Bizzozero’s pioneering stud- was responsible for both the clinical hemorrhage and the pro-
ies have seen a remarkable increase in our understanding of longed bleeding time, and that normal coagulation tests do not
hemostasis and thrombosis, and the important roles that plate- insure a normal bleeding time or adequate hemostasis in the
lets play in both of these phenomena. Thus, it is timely that face of thrombocytopenia.
Dr. Alan Michelson, who himself has made many important Despite this remarkable success, many obstacles to routine
discoveries regarding platelet function and pioneered the inno- platelet transfusion remained. These began to be systematically
vative application of flow cytometry to the study of platelets, addressed 35 years later when U.S. government support of
has once again brought together contributions from the leaders research in this area was spurred by the observation that throm-
in this discipline to provide a comprehensive review of the bocytopenic purpura was a major cause of death after radiation
topic in this fourth edition of Platelets. In this Foreword, I offer exposure from atomic weapons (Fig. 8).44,45 Further impetus
a brief and highly selective historical analysis of topics in plate- for improving platelet transfusion support came soon thereaf-
let history, with the hope of providing both a sense of the sweep ter when it became apparent that thrombocytopenia was a

xv
xvi Foreword

Fig. 1 Drawings by Osler (1874)1, 3 of “corpuscles” from rat

blood (5–7) and within a vein in the loose connective tissue of a
rat (8 and 9). (From Osler,3 with permission.)

Fig. 2 Drawing by Bizzozero (1882) of intravital microscopic image

of platelet and leukocyte deposition on damaged blood vessel in Fig. 4 Figs. 5 and 6 from Osler (1886)13 demonstrating “white
guinea pig mesentery.5 (From Bizzozero,5 with permission.) thrombi” in the aorta of a patient who died with cancer of the
stomach, and the “plaques” (platelets) that could be retrieved from
the thrombi. (From Osler,13 with permission.)

Fig. 3 Bizzozero’s “flow chamber” in which he watched platelets

deposit on a piece of thread, which was purposely frayed at both
ends (z). The chamber was formed by placing a coverslip (x) over two
strips of regular paper (a) and a strip of blotting paper (b) connected
to a wad of blotting paper. To encourage blood flow, the blood entry side
of the slide (o’) was slightly raised. (From Brewer,8 with permission.)

major cause of death from the then new chemotherapeutic

agents used to treat malignancies. In fact, successful platelet
transfusion therapy was absolutely crucial for the development
of modern chemotherapy.45,46
Many dramatic improvements in platelet transfusion ther-
apy were made over the next decades, including: improved anti-
coagulants; development of a differential centrifugation Fig. 5 Drawings by Bizzozero (1869)10 of giant bone marrow cells with
technique to produce platelet concentrates; recognition of the multilobular nuclei, now recognizable as megakaryocytes. (From
benefits of room temperature storage; improved plastics and Baserga A,11 with permission.)
 Foreword xvii

been ongoing for more than 50 years,18, 60–71 this goal has not yet
been achieved, despite the advances in our understanding of
platelet physiology. The need for such an agent(s) grew more
apparent in the 1980s and 1990s as a result of the dramatic
increase in the demand for platelet transfusions, especially with
the growth of bone marrow and stem cell transplantation and
autologous stem cell reconstitution, reaching 8,330,000 units
of platelets transfused in the United States in 1992.72 Since then
the growth rate has slowed substantially, with just over 9,000,000
units transfused in both 1997 and 1999,73 and 10,388,000 units
transfused in 2006.74 The number of units transfused per year
more recently has shown a downward trend as more restrictive
transfusion guidelines have been broadly adopted.75
There is also an increased appreciation of the seriousness of
platelet transfusion-transmitted bacterial infections, a conse-
quence of room temperature platelet storage, and estimated
to cause life-threatening sepsis in one in 100,000 recipients
in 2004.76 As a result, new methods to prevent bacterial con-
tamination, including diversion of the first blood drawn, and
to detect bacterial contamination before releasing platelets
for transfusion have been implemented, but their efficacy is
not absolute.77 Photochemical pathogen-reduction methods
have been introduced to lower the risk of transmitting infec-
tion, but current evidence suggests that such treatments may
increase the risk of developing refractoriness to platelet transfu-
sions and increase the number of platelet units transfused.78
Storage of platelets at 4°C would dramatically reduce the risk
of bacterial contamination, but it has been known for more
than 50 years, since the studies of Zucker and Borelli,79 that
exposure to cold temperature affects platelet morphology. Sub-
sequent studies demonstrated that exposure to cold results in
platelet activation,80,81 seriously compromising in vivo survival
and hemostatic effectiveness of the platelets.82 More recently,
basic studies led by Hoffmeister provided insights into the
mechanism responsible for the rapid clearance of platelets
incubated in the cold, identifying the lectin-like interaction
between integrin αMβ2 on hepatic macrophages and β-N-
acetylglucosamine residues on clustered platelet glycoprotein
Fig. 6 Bleeding time results from a patient described by Duke in (GP) Ib molecules as playing a central role in platelets sub-
1910.12 The series from rows A–D are blots from the ear wound made jected to acute cooling and the interaction between exposed
when the patient’s platelet count was 3000/μL. The blots in series A galactose residues and hepatocyte asialoglycoprotein
were taken immediately after the ear was pricked; series B began at (Ashwell-Morell) receptors in platelets refrigerated for
20 minutes; series C began at 40 minutes; series D began at 60 minutes; 48 hours.83 Refrigeration also increases the binding of von
and series E began at 80 minutes. The bleeding time was 90minutes. In Willebrand factor to GPIb and, in animal models, preventing
row F are the blots obtained after transfusion (platelet count 110,000/μL), this binding improves in vivo survival of the platelets.84 Loss
demonstrating a bleeding time of 3 minutes. (From Duke,12 with of sialic acid during normal platelet aging can also lead to plate-
permission.) let clearance via Ashwell-Morell receptors and may stimulate
thrombopoietin production (see Chapter 4).85
conditions of bag storage to facilitate gas exchange and the At present, a number of different factors are influencing the
maintenance of pH; cryopreservation; HLA matching or partial field of platelet transfusion therapy (see Chapter 64), includ-
matching to improve platelet responses in patients refractory to ing: the tendency to transfuse platelets only at lower platelet
random donor platelets; platelet “cross-matching;” single counts and with smaller numbers of platelets than in the
donor platelet collection via continuous flow centrifugation; past53,59,86; the questioning of the need for prophylactic plate-
blood screening assays to prevent viral transmission; hepatitis let transfusions87; the emergence of cord blood as a source of
B vaccination; prestorage leukoreduction to decrease febrile stem cells since megakaryocyte reconstitution is often delayed
transfusion reactions; ultraviolet irradiation and leukoreduc- with this source of stem cells88–90; the development of new con-
tion to decrease immunogenicity; development of synthetic ditioning regimens for stem cell reconstitution that produce
storage solutions; new methods to eliminate viable pathogens much less myelosuppression than in the past91; the availability
by damaging their DNA; and attempts to define better the most of agents that can activate the thrombopoietin receptor (mpl)
appropriate platelet count at which to transfuse platelets (see (see Chapter 61)92,93; attempts to expand hematopoietic stem
Chapter 64).23, 47–58 Ironically, one of the most important cells ex vivo to increase the likelihood and speed of megakaryo-
advances related to platelet transfusion therapy had nothing cytic engraftment after transplantation with cytokines, growth
to do with the platelet product, but rather concerned the recog- factors, and a small molecule CD34+ expander, as well as extra-
nition that aspirin inhibits platelet function and thus aspirin cellular proteins in the hematopoietic bone marrow niche, such
should not be used as an antipyretic or analgesic in patients as fibronectin94–100; the increasing concern about the compli-
who are thrombocytopenic.59 cations of platelet transfusion therapy, including thrombosis,
Although attempts to develop substitutes for fresh platelets transfusion-related acute lung injury (TRALI), and immuno-
that could secure hemostasis in thrombocytopenic patients have modulation101; the renewed emphasis on ABO blood group
xviii Foreword

Fig. 7 The clinical course of the patient reported by Duke.12 The transfusion was performed at 2 a.m. on May 12. (From Duke,12 with permission.)

matching102; and continued attempts to develop platelet microenvironment, including the shear forces experienced by
substitutes.60–71,103 megakaryocytes as they extend into bone marrow sinu-
The advances in understanding stem cell biology, the basic soids.108,111–121 Transfusion of megakaryocyte precursors is
mechanisms of megakaryocyte differentiation, and platelet an alternative approach that avoids the need to accomplish
production offer the potential to improve stem cell replace- the final, and most technically difficult, steps in producing
ment therapy and platelet transfusion therapy (see platelets in vitro.120–123 Platelets produced from a potential
Chapter 66). Megakaryocytes have been generated from both recipient’s iPS cells provide hope of eliminating the immuno-
mouse and human embryonic stem cells when cultured logic responses that make some patients refractory to platelet
in vitro under various conditions, but the final step, platelet transfusions, although recent studies in other systems have
production, has been difficult to achieve with high efficiency raised questions about the immune recognition of iPS cells.124
or yield.104–109 The technology of induced pluripotent stem There is good reason for platelets to lead the way in the use of
cells (iPS) has been applied to developing human megakaryo- iPS-derived cells, since the concerns about the teratogenicity of
cytes from human dermal fibroblasts, with demonstrated iPS cells can be eliminated by transfusing anucleate, irradiated
success in obtaining platelets that both circulate in immunode- platelets. The search for an artificial platelet substitute will cer-
ficient mice for more than 24 hours and participate in throm- tainly continue, and this quest may be significantly advanced if
bus formation.110 Several groups are now actively engaged in nanotechnology is able to develop small biocompatible matri-
developing large-scale production of platelets from iPS cells, ces, since the platelet’s small size is an important feature of its
cord blood, or other sources for transfusion using a variety of biology, reducing the shear force it experiences relative to large
ingenious devices to simulate the megakaryocyte bone marrow cells at comparable shear rates.
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 Oceanography - Workbook
Summer 2024 - Department

Prepared by: Associate Prof. Brown

Date: July 28, 2025

Review 1: Study tips and learning strategies

Learning Objective 1: Learning outcomes and objectives
• Interdisciplinary approaches
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Learning Objective 2: Comparative analysis and synthesis
• Best practices and recommendations
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Learning Objective 3: Best practices and recommendations
• Theoretical framework and methodology
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Learning Objective 4: Case studies and real-world applications
• Current trends and future directions
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Learning Objective 5: Ethical considerations and implications
• Best practices and recommendations
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Note: Practical applications and examples
• Theoretical framework and methodology
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Example 6: Problem-solving strategies and techniques
• Case studies and real-world applications
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Important: Historical development and evolution
• Historical development and evolution
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
[Figure 8: Diagram/Chart/Graph]
Practice Problem 8: Key terms and definitions
• Research findings and conclusions
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Practice Problem 9: Statistical analysis and interpretation
• Comparative analysis and synthesis
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Formula: [Mathematical expression or equation]
Chapter 2: Interdisciplinary approaches
Definition: Problem-solving strategies and techniques
• Study tips and learning strategies
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Formula: [Mathematical expression or equation]
Important: Historical development and evolution
• Case studies and real-world applications
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Remember: Case studies and real-world applications
• Key terms and definitions
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Key Concept: Critical analysis and evaluation
• Problem-solving strategies and techniques
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Definition: Research findings and conclusions
• Historical development and evolution
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Key Concept: Study tips and learning strategies
• Theoretical framework and methodology
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Note: Historical development and evolution
• Ethical considerations and implications
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Key Concept: Experimental procedures and results
• Fundamental concepts and principles
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Remember: Case studies and real-world applications
• Experimental procedures and results
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Practice Problem 19: Comparative analysis and synthesis
• Learning outcomes and objectives
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
[Figure 20: Diagram/Chart/Graph]
Part 3: Interdisciplinary approaches
Definition: Current trends and future directions
• Literature review and discussion
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Remember: Ethical considerations and implications
• Statistical analysis and interpretation
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Important: Case studies and real-world applications
• Current trends and future directions
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Important: Study tips and learning strategies
• Ethical considerations and implications
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Practice Problem 24: Learning outcomes and objectives
• Key terms and definitions
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Formula: [Mathematical expression or equation]
[Figure 25: Diagram/Chart/Graph]
Important: Critical analysis and evaluation
• Historical development and evolution
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Example 26: Case studies and real-world applications
• Assessment criteria and rubrics
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Example 27: Best practices and recommendations
• Research findings and conclusions
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Practice Problem 28: Comparative analysis and synthesis
• Interdisciplinary approaches
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Important: Key terms and definitions
• Interdisciplinary approaches
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Section 4: Current trends and future directions
Key Concept: Learning outcomes and objectives
• Interdisciplinary approaches
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Practice Problem 31: Ethical considerations and implications
• Problem-solving strategies and techniques
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Key Concept: Learning outcomes and objectives
• Learning outcomes and objectives
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
[Figure 33: Diagram/Chart/Graph]
Definition: Critical analysis and evaluation
• Fundamental concepts and principles
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Definition: Research findings and conclusions
• Research findings and conclusions
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Important: Comparative analysis and synthesis
• Ethical considerations and implications
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Important: Practical applications and examples
• Practical applications and examples
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Remember: Interdisciplinary approaches
• Practical applications and examples
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Example 38: Research findings and conclusions
• Research findings and conclusions
- Sub-point: Additional details and explanations
- Example: Practical application scenario
[Figure 39: Diagram/Chart/Graph]
Important: Current trends and future directions
• Ethical considerations and implications
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Background 5: Ethical considerations and implications
Important: Study tips and learning strategies
• Experimental procedures and results
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Example 41: Study tips and learning strategies
• Study tips and learning strategies
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
[Figure 42: Diagram/Chart/Graph]
Definition: Ethical considerations and implications
• Problem-solving strategies and techniques
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Example 43: Literature review and discussion
• Key terms and definitions
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Note: Key terms and definitions
• Practical applications and examples
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Note: Critical analysis and evaluation
• Study tips and learning strategies
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Remember: Case studies and real-world applications
• Historical development and evolution
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Example 47: Case studies and real-world applications
• Study tips and learning strategies
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Formula: [Mathematical expression or equation]
Example 48: Statistical analysis and interpretation
• Study tips and learning strategies
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Important: Practical applications and examples
• Study tips and learning strategies
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Exercise 6: Current trends and future directions
Key Concept: Learning outcomes and objectives
• Fundamental concepts and principles
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
[Figure 51: Diagram/Chart/Graph]
Important: Statistical analysis and interpretation
• Current trends and future directions
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Practice Problem 52: Literature review and discussion
• Assessment criteria and rubrics
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Note: Historical development and evolution
• Key terms and definitions
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Important: Ethical considerations and implications
• Study tips and learning strategies
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Note: Fundamental concepts and principles
• Assessment criteria and rubrics
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Important: Experimental procedures and results
• Study tips and learning strategies
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Important: Statistical analysis and interpretation
• Ethical considerations and implications
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Remember: Interdisciplinary approaches
• Research findings and conclusions
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Important: Experimental procedures and results
• Problem-solving strategies and techniques
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Abstract 7: Literature review and discussion
Definition: Experimental procedures and results
• Literature review and discussion
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Example 61: Interdisciplinary approaches
• Fundamental concepts and principles
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Important: Learning outcomes and objectives
• Study tips and learning strategies
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Important: Theoretical framework and methodology
• Current trends and future directions
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Note: Historical development and evolution
• Theoretical framework and methodology
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Definition: Research findings and conclusions
• Research findings and conclusions
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Note: Comparative analysis and synthesis
• Research findings and conclusions
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Definition: Theoretical framework and methodology
• Historical development and evolution
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Practice Problem 68: Historical development and evolution
• Research findings and conclusions
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Practice Problem 69: Research findings and conclusions
• Key terms and definitions
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Discussion 8: Problem-solving strategies and techniques
Remember: Problem-solving strategies and techniques
• Practical applications and examples
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Key Concept: Case studies and real-world applications
• Assessment criteria and rubrics
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Example 72: Interdisciplinary approaches
• Experimental procedures and results
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Formula: [Mathematical expression or equation]
Practice Problem 73: Comparative analysis and synthesis
• Ethical considerations and implications
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Key Concept: Study tips and learning strategies
• Assessment criteria and rubrics
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
[Figure 75: Diagram/Chart/Graph]
Definition: Interdisciplinary approaches
• Research findings and conclusions
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Practice Problem 76: Problem-solving strategies and techniques
• Ethical considerations and implications
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Practice Problem 77: Interdisciplinary approaches
• Historical development and evolution
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Definition: Current trends and future directions
• Theoretical framework and methodology
- Sub-point: Additional details and explanations
- Example: Practical application scenario
[Figure 79: Diagram/Chart/Graph]
Remember: Statistical analysis and interpretation
• Experimental procedures and results
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Test 9: Literature review and discussion
Example 80: Key terms and definitions
• Best practices and recommendations
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Definition: Statistical analysis and interpretation
• Comparative analysis and synthesis
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Important: Interdisciplinary approaches
• Comparative analysis and synthesis
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Note: Theoretical framework and methodology
• Problem-solving strategies and techniques
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Note: Research findings and conclusions
• Practical applications and examples
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Example 85: Case studies and real-world applications
• Current trends and future directions
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Practice Problem 86: Assessment criteria and rubrics
• Best practices and recommendations
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Note: Practical applications and examples
• Learning outcomes and objectives
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
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