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HANDBOOK
OF
PLATELET
PHYSIOLOGY
AND
PHARMACOLOGY
Edited
by
Gundu
H. R. Rao
University
of
Minnesota
KLUWER
ACADEMIC PUBLISHERS
Boston
/
Dordrecht
/
London
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for
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Library
of
Congress
Cataloging-in-Publication
Data
Handbook
of
platelet physiology
and
pharmacology
/
edited
by
Gundu
H.R.
Rao.
p. cm.
Includes bibliographical references
and
index.
ISBN
0-7923-8538-1
(alk.
paper)
1.
Blood platelets Handbooks,
manuals,
etc.
I.
Rao,
Gundu
H.R
1938-
.
QP97.H36
1999
612.ri7-dc21
99-27962
CIP
Copyright
©
1999
by
Kluwer Academic
Publishers
All
rights reserved.
No
part
of
this publication
may be
reproduced, stored
in a
retrieval system
or
transmitted
in any
form
or by any
means, mechanical, photo-
copying, recording,
or
otherwise, without
the
prior written permission
of the
publisher,
Kluwer Academic Publishers,
101
Philip Drive, Assinippi Park, Norwell,
Massachusetts
02061
Printed
on
acid-free
paper.
Printed
in the
United States
of
America
List
of
Contributors
1
.
Kailash
C.
Agarwal,
Ph. D.
Department
of
Molecular Pharmacology,
Brown
University
Providence,
RI
02912,USA
2.
Colin
N.
Baigent,
Ph. D.
ATT
Collaboration
CTSU
Harkness
Building
Radcliffe
Infirmary
Woodstock Road
Oxford
0X2
6HE
United
Kingdom
3.
Rodger
L.
Bick,
M.
D.,
Ph. D.
Departments
of
Pathology
&
Pharmacology
Loyola
University
Med.
Center
21260 South First Ave.
Maywood,
IL
60153,USA
4.
David
C.
Calverley,
M. D.
USC
School
of
Medicine
1441EastlakeAve
NOR MS 34
Los
Angeles,
CA
90033,
USA
5.
Thomas
Chandy,
Ph. D.
Chemical
Engineering
and
Material
Sciences
University
of
Minnesota
Minneapolis,
MN
55455,
USA
6.
Kenneth
J.
Clemetson,
Ph. D.
Theodor
Kocher
Institut
Der
Universitat Bern
Freiestrasse
1,
Ch-3012
Berne
Switzerland
7.
Robert
W.
Colman,
M. D.
Thrombosis Research Center
Temple
Univ. Sch.
of
Medicine
3400
N.
Broad
Street
Philadelphia,
PA
19140,USA
8.
Maribel
Diaz-Ricart,
Ph. D.
Servicio
de
Hemoterapia
Hospital
Clinico
Provincial
Villarroel
170,
Barcelona
08036,
Spain
9.
Gines
Escolar,
M. D. Ph. D.
Servicio
de
Hemoterapia
Hospital Clinico Provincial
Villarroel
170,
Barcelona
08036,
Spain.
10.
Daniel
Fareed,
B.Sc.
Departments
of
Pathology
&
Pharmacology
Loyola
University Med. Center
2
1260
South First Ave.
Maywood,
IL
60153,USA
1 1
.
Jawed Fareed,
Ph. D.
Departments
of
Pathology
&
Pharmacology
Loyola
University Med. Center
2
1
260
South
First
Ave.
Maywood,
IL
60153,USA
12.
Deborah French,
M. D.
Department
of
Medicine
Mount
Sinai Hospital
&
Medical
School
One
Gustave
L.
Levy
Place
New
York,
NY
10029-6574,USA
1
3.
Mony
M.
Frojmovic,
Ph. D.
Mclntyre
Medical
Science Building
McGiIl
University
3655
Drummond
Street
Montreal,
QB
Canada,
H3GIY6
14.
Nicholas J.Greco,
Ph. D.
Platelet Biology Laboratory
American
Red
cross
1
5601
Crabbs Branch
Way
Rockville,
MD
20855,USA
15.
HolmHolmsen,Ph.D.
Department
of
Biochemistry
and
Molecular Biology
University
of
Bergen
Astradveien
19,
Bergen
N5009,
Norway
1
6.
Debra Hoppensteadt,
Ph. D.
Departments
of
Pathology
&
Pharmacology
Loyola
University Med. Center
2
1260
South First Ave.
Maywood,
IL
60153,USA
17.
Huzoor-Akbar,
Ph. D.
Molecular
and
Cellular Biology
Department
of
Biological
Sciences,
Irvine Hall
Athens,
OH
45701,USA
1
8.
G.
A.
Jamieson,
Ph. D, D. Sc.
Platelet Biology Laboratory
American
Red
Cross
15601 Crabbs Branch
Way
Rockville,
MD
20855,USA
1
9.
Gerhard
J.
Johnson,
M. D.
Veterans
Affairs
Medical Center
One
Veterans
Way
Minneapolis,
MN
55417,USA
20.
BeateKehrel,Ph.D.
Experimental
and
Clinical
Haemostaseology
Department
of
Anaesthesiology
and
Intensive Care Medicine
University
of
Muenster
D-48149
Muenster, Germany
2
1
.
Bruce
R.Lester,
Ph.
D,.
Knowledge
Frontiers
3989
Central Ave,
N.
E.,
# 625
Minneapolis,
MN
55421,USA
22.
Mahadev
Murthy,
Ph. D.
Division
Endocrinolgy,
Metabolism
&
Nutrition
Department
of
Medicine
Hennepin
County
Medical Center
914
South Eighth Street,
D-3
Minneapolis,
MN
55404.
USA
23.
Ellinor
I.
Peerschke,
Ph. D.
Cornell Medical Center
New
York University
525 E
68th Street,
Rm
F51
1
J
New
York,
NY
10021,USA
24.
Anna
S.
Radomski
Division
of R and D
Lacer,
S.A.
08025
Barcelona
Spain
25.
Marek
W.
Radomski,
M.D,D.Sc.
Division
of R and D
Lacer, S.A.,
08025
Barcelona
Spain
26.
Gundu
H. R.
Rao,
Ph. D.
Departments
of
Lab.
Med.
&
Pathol.
and
Biomed. Engineering
P.B.
609
UMHC
Academic Health Center
University
of
Minnesota
Minneapolis,
MN
55455,USA
27. A.
Koneti Rao,
M. D.
Department
of
Medicine
Temple University School
of
Medicine
3400
N.
Broad
St Rm
300-OMS
Philadelphia,
PA
19140,USA
28.
Gerald
J.
Roth,
M. D.
Division
of
Hematology
V.A.
Medical Center
1660
South Columbian
Way
Seattle,
WA
98108,USA
29.
Anita
Ryningen,
Ph. D.
Department
of
Biochemistry
and
Molecular Biology
University
of
Bergen
Astradveien
19
Bergen
N-5009
Norway
30.
Shivendra
D.
Shukla,
Ph. D.
University
of
Missouri-
Columbia
517B
Medical
Science
Building
One
Hospital Drive
Columbia,
MO
65212,USA
3
1
.
Cathie Sudlow,
Ph. D.
ATT
Collaboration
CTSU
Harkness
Building
Radcliffe
Infirmary
Woodstock Road
Oxford
OX2
6HE
United Kingdom
32.
Narendra
N.
Tandon,
Ph. D.
Thrombosis
&
Vascular Biology
Otsuka
America Pharmaceutical
9900
Medical Center Drive
Rockville,
Maryland
20850,
USA
33.
Jeanine
M.
Walenga,
Ph. D.
Departments
of
Pathology
&
Pharmacology
Loyola
University
Medical Center
2
1260
South
First
Ave.
Maywood,
IL
60153,USA
34.
Douglas
J.
Weiss,
D.
V.M.,
Ph. D.
Department
of
Pathobiology
and
Veterinary
Sciences
University
of
Minnesota
St.
Paul,
MN
55108,USA
35.
Helmut
Wolf,
M. D, Ph. D.
Departments
of
Pathology
&
Pharmacology
Loyola
University
Medical Center
2
1260
South
First
Ave.
Maywood,
IL
60153,USA
PREFACE
Despite
my
many years
of
research
and
teaching
in
platelet physiology
and
pharmacology
at the
University
of
Minnesota,
I am
often
confronted with conflicting
opinions
as to the
relevance
of
nonnucleated
platelets
in
human health
and
disease.
It
is
fascinating
to
think that
how
cells with
no
apparent nucleus, have such
a
towering
impact
on
concepts, dealing with
often
overlapping physiological (i.e.
hemostasis,
wound
healing, etc.)
and
pathophysiological
(i.e. thrombosis, stroke, atherosclerosis,
wound
healing, diabetes,
inflammation
and
cancer) components. Although
the
idea
of
compiling
new frontiers of
platelet research
in the
form
of a
book
was
quite simple
at
the
beginning,
the
project turned
out to be a
major
undertaking
from my
part.
At the
end,
I am
elated
that
the
contributors
to
this book were gracious enough
to
write chapters
in
their area
of
research expertise despite their pressing
and
highly valuable time.
For
me,
it has
been
an
humbling experience
as the
chapters that
I
have compiled,
are
written
by
people with incredible recognition
for
their relentless contributions over
the
years
to
strengthen
the
understanding
of
platelet physiology
and
pharmacology.
In my
opinion,
this
has
added
an
immense value
to the
book.
I am
proud
to
have been involved
in
this
undertaking
despite several unexpected problems
and
delays during this project.
I am
confident
that this book would
be
highly
useful
to the
community
of
scientists, including
graduate
students, researchers, academicians, physicians
and
other health care
professionals,
and
pharmaceutical industry scientists.
Circulating
platelets which lack nucleus neither adhere
to the
vessel wall
nor
aggregate
unless they encounter
a
zone
of
injury.
Upon encountering such
a
zone
of
injury,
they
become almost instantly activated, which leads
to
their adhesion
and
aggregation, both
reactions
are of
fundamental
importance
to
hemostasis
and
thrombosis. Because
of
this
reason, platelet research
has
clearly
led the way in the
continuing development
of new
strategies
and
drugs that
can
help prevent
and
treat arterial thrombosis, stroke
and
atherosclerosis. Unquestionably, platelet research
has
also impacted concepts dealing
with
many
other
diseases.
Nevertheless, considerable progress
has
been made
in the
development
of new
antiplatelet
agents
in
recent years. These newer agents
are
aimed
at
interrupting specific sites
and
pathways
of
platelet activation. Inhibitors
of
specific
platelet
agonist-receptor interactions include
antithrombins,
thromboxane
A
2
receptor
antagonists,
and
adenosine
diphosphate receptor
blockers
(i.e.
ticlopidine,
clopidogrel).
In
addition, inhibitors
of
arachidonic
acid metabolism
and
thromboxane
A
2
include
aspirin, newer COX-2 inhibitors, other NSAIDs, thromboxane
A
2
synthase
inhibitors
and
o>-3
fatty
acids. Moreover, long awaiting drugs that block
ligand
binding
to the
platelet
glycoprotein
Ilb/IIIa
complex (i.e.
tirofiban)
have
now
entered
the
market.
In
this book,
the
chapters
are
organized into
six
major
sections, including
Introduction,
Receptor
Biology,
Platelet
Biochemistry,
Experimental
Physiology,
Platelet Pathology
and
Platelet
Pharmacology. Authoritative chapters
in
each section have provided
a
collective strength
to our
initial philosophy
of
accomplishing
a
comprehensive review
of
current
concepts
in
each discipline. Although every attempt
has
been made
to
provide
an
interdisciplinary discussion
on the
subject
of
platelets
in
this book, there
may
still
be
some
gaps
and
lapses
for
which readers
are
urged
to
consult other articles
and
reviews.
I
have deliberately avoided going into
any
specific comments
on
reviews
in
order
to let
the
imagination
of the
readers
flow freely. I
believe that
the
readers
are
intelligent
enough
to
judge
and
form
their
own
critical opinion.
I
must humbly express
my
deep gratitude
to
thirty
five
scientists
in the field for
their
invaluable contributions.
I now
honestly believe that this publication would
not
have
been possible without their meritorious contributions.
I
am
deeply indebted
to my
dear
friend and
close
research collaborator,
Mahadev
Murthy,
Ph.
D.,
Director
of
Research, Division
of
Endocrinology, Metabolism
and
Nutrition, Department
of
Medicine,
Hennepin
County Medical Center, Minneapolis,
MN,
USA,
for his
commitment
and
contribution
to
this project.
He has
spent countless
hours during this project
in
reviewing
and
preparing camera ready manuscripts
for final
submission
to the
Kluwer Academic Publisher.
In
addition,
he has
written
two
excellent
chapters
for the
book.
I
must confess that this publication would
not
have been
completed without
his
generous
and
truly dedicated
efforts.
I
would like
to
take this opportunity
to
thank Charles
W.
Schmieg,
Jr., Acquisitions
Editor, Kluwer Academic Publishers,
101
Philip Drive,
Assinippi
Park,
Norwell,
MA,
02061, USA,
for
facilitating
the
publication
of
this book.
I am
specially
thankful
for his
cooperation
and
patience even
though
this project
was
delayed
by
about
four
months.
Finally,
I
would
not be in
this
field
today without
my
mentor, James
G.
White,
M.
D.,
Regents'
Professor
&
Associate Dean, Academic Health Center, University
of
Minnesota,
Minneapolis,
MN,
USA.
I
humbly
dedicate this publication
to
James
G.
White,
M.
D.,
who has
been
my
mentor, teacher, associate
and
dear
friend,
during
my
long
career
in
platelet research.
In the
end,
my
academic success
and
accomplishments
over
the
years, would
not
have been possible without
the
support
of my
wife
Yashoda,
my
daughter
Aupama
and my son
Prashanth.
I
sincerely acknowledge
and
appreciate
their
patience
and
support throughout
my
career.
Gundu
H. R. Rao
University
of
Minnesota
Professor
Minneapolis,
MN
55455
v
This page has been reformatted by Knovel to provide easier navigation.
Contents
Contributors
ix
Preface
xiii
Introduction 1
1. Platelet Physiology & Pharmacology: an Overview 1
1.1 Introduction 1
1.2 Role of Platelets in Hemostasis and
Thrombosis 2
1.3 Platelet Morphology and Biochemistry 2
1.4 Platelet Physiology 5
1.5 Altered Physiology and Function 6
1.6 Platelet Pharmacology 8
1.7 Platelet Function Inhibitory Drugs 9
1.8 Acknowledgements 14
References 15
Receptor Biology 21
2. Human Platelet Thrombin Receptors and the Two
Receptor Model for Platelet Activation 21
2.1 Introduction 21
2.2 Binding Studies 22
2.3 Membrane Microviscosity 24
2.4 Candidate Receptors 26
vi Contents
This page has been reformatted by Knovel to provide easier navigation.
2.5 The GPIb-IX-V Complex 27
2.6 Two Receptor Model 31
References 33
3. Platelet Thromboxane Receptors: Biology and
Function 38
3.1 Introduction 38
3.2 Biological Effects of TP Receptor Activation 39
3.3 Smooth Muscle Contraction 39
3.4 TP Receptor Structure 41
3.5 TP Receptor Function 49
3.6 Altered TP Receptor Function 58
References 66
4. Collagen Receptors: Biology and Functions 80
4.1 Introduction 80
4.2 Collagens 82
4.3 Von-Willebrand-Factor 83
4.4 P65 84
4.5 CD36 84
4.6 a
2
b
1
-Integrin (GPIa/IIa, VLA
2
, ECMRII) 87
4.7 GPVI/FcRg 89
4.8 Collagen-Induced Signal Transduction 90
References 92
5. Adenosine Receptors: Biology and Function 102
5.1 Introduction 102
5.2 Adenosine Receptors 103
5.3 Antiplatelet Action of Adenosine 104
5.4 Adenosine Production and Platelet
Inactivation 106
5.5 Agents Affecting Adenosine Actions 109
Contents vii
This page has been reformatted by Knovel to provide easier navigation.
5.6 Adenosine Effects on Intracellular Ca
2+
Mobilization 113
5.7 Conclusions 114
References 115
6. Platelet Activating Factor and Platelets 120
6.1 PAF Discovery, Structure and Heterogeneity 120
6.2 PAF Biosynthesis in Platelets 121
6.3 Responses of Platelets to PAF 122
6.4 PAF Receptor and Signal Transduction
Pathways in Platelets 123
6.5 Antagonist 124
6.6 PAF Receptor 125
6.7 Phospholipases 126
6.8 Platelet and PAF in Pathophysiological and
Disease States 129
6.9 Acknowledgement 133
References 133
7. Platelet Glycoprotein Ib-V-IX: Biology and Function 142
7.1 Introduction 142
7.2 Structure 143
7.3 Post-Translational Modification of GPIb-V-IX 145
7.4 Basic Functions 146
7.5 Signal Transduction 148
7.6 GPIb-V-IX as a Target for Pharmacological
Inhibition 149
7.7 Genetic Disorders Affecting GPIb-V-IX 151
7.8 Tissue Specific Expression of GPIb-V-IX
Subunits 153
7.9 Future Developments 154
References 155
[...]... both human and animal models Furthermore, a careful correlation between signal-driven responses and platelet function will lead to a better understanding of platelet pathophysiology (27,31, also discussed in other chapters) 1.6 Platelet pharmacology Earlier studies on platelet biochemistry, physiology and function suggest that release of granule contents is essential for recruitment of platelets, irreversible... provide a comprehensive review on various aspects of platelet physiology and pharmacology 1.2 Role of platelets in hemostasis and thrombosis Platelets contribute significantly to the normal hemostatic process (1-6) They play a critical role in the recognition of injured vasculature, formation of hemostatic plugs, prevention of bleeding, retraction of clots and wound healing (6-9) When they develop severe... of Platelet Function 460 21.5 Mechanisms of NO Action on Platelets 461 21.6 Peroxynitrite 462 21.7 Nitric Oxide and Vascular Disorders 463 21.8 Nitric Oxide, Platelets and Septicemia 464 21.9 Pharmacology of NO Generation and Action in the Platelet Microenvironment 465 Acknowledgements 469 References 469 22 Aspirin, Prostaglandins and Platelet Function: Pharmacology. .. aggregation and secretion of granule contents Platelet activation is a prerequisite for the formation of a hemostatic plug and arrest of bleeding at the injured site Although formation of a hemostatic plug is a natural response to injury, the role of platelets in atherosclerosis, thrombosis and stroke are pathological manifestations (1-19) 1.3 Platelet morphology and biochemistry Blood platelets have... discussed in other chapters of this book) Binding results in the activation and stimulation of various effector enzymes and formation of second messengers, leading to aggregation and secretion of granule contents Specific mechanisms involved in the process of centralization of granules and release of their contents are poorly understood(26,27, 31) 1.5 Altered physiology and function Many investigators... easier navigation 1 PLATELET PHYSIOLOGY AND PHARMACOLOGY: AN OVERVIEW Gundu H R Rao, Ph D Departments of Laboratory Medicine & Pathology and Biomedical Engineering Institute Academic Health Center University of Minnesota, Minneapolis, MN 55455, USA 1.1 INTRODUCTION Dr Gundu Rao has spent the last 25 years studying and teaching platelet physiology and pharmacology at the University of Minnesota Over the... antiplatelet and antithrombotic drugs have provided a great deal of excitement and have led to the discovery of many specific antagonists Although the biochemistry of platelets remains highly complex, we now have well defined pathways that could impact platelet functional responses It is very likely that further understanding of the role of these pathways and the specific mechanisms that underlie platelet. .. Activation and Aggregation in Flow 316 14.3 Range of Shear Rates in Normal and Pathological Settings 317 14.4 Flow Regimes and Corresponding Devices Used to Study in Vitro Platelet Aggregation 318 14.2 Ligands and Receptors Involved in Platelet Aggregation 319 14.3 Quantitation of Aggregation: Theoretical and Experimental Approaches 322 14.4 Platelet Aggregation in Non-Stirred Platelet. .. Acid and Platelets 302 13.8 PAF and Phospholipase A2 303 13.9 LDL and Platelet Function 304 Concluding Remarks 304 Acknowledgements 305 References 305 Experimental Physiology 315 14 Platelet Biorheology: Adhesive Interactions in Flow 315 14.1 Introduction: General Overview for Flow Studies of Platelet Aggregation 315 14.2 General Physiology of Platelet. .. Medicine & Pathology and Biomedical Engineering Institute References 1 Marcus AJ, Zucker MB The physiology of blood platelets New York: Crime and Scrutton, 1965 2 Kowalski E, Niewiarowski S Biochemistry of blood platelets New York: Academic Press, 1966 3 Caen J Platelet aggregation Paris: Masson and Cie, 1971 4 Brinkhous KM, Sherman RW, Mostof FK The Platelet Baltimore: Williams and Wilkson Co, 1971 . HANDBOOK
OF
PLATELET
PHYSIOLOGY
AND
PHARMACOLOGY
Edited
by
Gundu
H. R. Rao
University
of
Minnesota
KLUWER
ACADEMIC. <http://www.wkap.nl>
Library
of
Congress
Cataloging-in-Publication
Data
Handbook
of
platelet physiology
and
pharmacology
/
edited
by
Gundu
H.R.
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