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MANFRED MARTIN MAYER
June 15, 1916-September 18, 1984
BY K. FRANK AUSTEN
\~7HAT IS THE LEGACY of a scientist? A pioneer in the
~ ~ field of-immunochemistry, Manfred Mayer almost
singlehandeclly established the discipline of complement. He
contributed the one-hit theory of immune hemolysis. He un-
covered the first indications of the enzymatic cleavage of one
complement protein by another, leading to our eventual
understan(ling of the sequential interaction and function of
the eighteen proteins of the complement system. He appre-
ciatec} that cytolysis by complement is due to the insertion of
hydrophobic complement peptides into the lipid bilayer of
biomembranes and formation of transmembrane channels.
Finally, on a different tack, he and Robert Nelson developed
the Triponema pa111lidum immobilization test for syphilis. As a
teacher and mentor, his impeccable methodology and the
care he lavished on the members of his laboratory pro-
clucec! many distinguished intellectual descendants. Finally,
Manfred Mayer will always remain the model of a life lived
by the highest values, scientific and personal.
EDUCATION AND EARLY ElFE
Manfrec! was born in Frankfurt-am-Main, Germany, on
June 15, 1916, ant! stied in Baltimore, Maryland, on Septem-
ber IS, 1984. He received his primary and secondary school-
257
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258
BIOGRAPHICAL MEMOIRS
ing in Germany but was forced to leave that country in 1933,
at the age of seventeen, because of political events. He
worked his way through the City College of New York, re-
ceiving a B.S. in 193S, then entered a doctoral program at
Columbia University. His doctoral thesis was on the chemical
and immunologic properties of phosphorylate(1 serum al-
bumin. He received the Ph.D. degree in 1946.
From 1938 through 1942, Manfred supported himself
working as a laboratory assistant to Dr. Michael Heidel-
berger a founder of the cliscipline of immunochemistry—at
Columbia University. His background in physical chemistry
fit well with Heiclelberger's organic chemical background and
approach, and he was very comfortable in this laboratory that
also container! Forest Kendall and had just trained EIvin Ka-
bat. During his four years there, Manfrect progressed from
laboratory assistant to the role of distinguished graduate stu-
clent. He worker! on both the cross-reactions to Type Ill
pneumococcal capsular polysaccharides and the fixation of
the activity in immune complex reactions known as "comple-
ment." By 1946 Manfrecl was an accomplished immunochem-
ist with two unique interests of his own that would occupy his
subsequent scientific career: quantitative assessment of the
complement system and its components, ant! the elucida-
tion in biochemical terms of the reaction sequence.
The same year that Manfred receive(1 his Ph.D., Thomas
B. Turner, chairman of the Department of Bacteriology at
the Johns Hopkins School of Medicine, asked Michael
Heidelberger to recommenct someone in immunology.
Heiclelberger praised Mayer highly, and he was offered the
position of assistant professor. With his wife, Elinor, Manfrec!
proceeclec] to Baltimore, ant! within two years his contribu-
tions as a teacher and investigator had earned him promotion
to associate professor. In recognition of the quality of his
scholarship and his balances! approach to departmental
OCR for page 259
MANFRED MARTIN MAYER
259
issues, he was chosen acting head of the Department (though
not yet a full professor) when Thomas Turner left to become
dean. He served throughout 1957, when Barry Wooc} arrived
to take over the chairmanship, and was appointee} full pro-
fessor in ~ 960.
SCIENTIFIC CONTRIBUTION
Working with EIvin Kabat from 1942 to 1945, long before
his arrival at Hopkins, Mayer had completed Experimental
Immunochemistry ~ ~ 94S, ~ ), though this most important vol-
ume clid not appear in print until 1948. During that era,
everyone in the field! of immunochemistry hac! been in-
structed by Michael Heidelberger, either personally or
through his distinguished disciples, EIvin Kabat and Manfred
Mayer. The Heidelberger school tract developect techniques
for conducting quantitative precipitin reactions and agglutin-
ation determinations, and Kabat and Mayer cleciclec! it was
critical for the future of research in the field to produce a
textbook of quantitative immunochemistry that was both
conceptual and practical in content. For a number of years,
EIvin Kabat and Manfrect Mayer met virtually every weekend
in one another's apartments to react aloucl and revise every
word of the proposed text. Heiclelberger also react it and
ultimately prepared the introduction to the volume. These
were difficult times for the wives of immunochemists, but
Elinor supported Manfred throughout while at the same
time proceeding with her own substantial interests.
By 1945 the unique anc} historically critical volume was
complete, only to be delayecl three years by the publishers-
allegecIly because of a paper shortage. The authors, however,
used the clelay to revise the manuscript extensively anct pro-
ducect a volume that went through three printings without
revision over the next ten years. In 1958 the authors began
work on a second edition in which Mayer's contribution on
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260
BIOGRAPHICAL MEMOIRS
the complement system was greatly expanded, largely due to
the findings of his laboratory. This edition (1961,5) went
through four printings before going out of print in 1984.
During the late 1940s and early 1950s Manfred had be-
gun to assemble an outstanding group committed to immu-
nochemistry in general and to complement research in spe-
cific—with startling results. During those years Lawrence
Levine demonstrated that the introduction of diisopropyl
fluorophosphate (DFP) would block enzymatic activity in the
first component of complement a critical step in the rec-
ognition of the biochemical events in the complement cas-
cade. Keith Cowan was studying how carbowax acted as a
substitute for specific antibody in mediating the hemolytic
action of complement. Al Marucci, with Manfred's guidance,
had begun to evaluate the use of radiolabeled antibody as an
analytic tool in defining immunochemical events. Finally,
Herbert Rapp was analyzing the different functions of rabbit
antibodies of different immunoglobulin classes and, with
Manfred, was beginning to develop a mathematical basis for
the analysis of the reaction sequence. Their work resulted in
the conclusion that the "third component of complement"
was not a single substance but, based upon its behavioral
characteristics as defined in mathematical terms, represented
multiple substances a conclusion subsequently substanti-
ated by the identification of five component proteins.
Manfred's definition of the cofactor functions of calcium
and magnesium made possible the singularly important dem-
onstration that the functional interactions of the components
of the complement system met the "one-hit" model of inter-
actions. By preparing, with his students, specific intermedi-
ates, he broke the reaction down into sequential events and
initially purified the components being analyzed. This "one-
hit" analysis permitted the measurement of complement
components or proteins in molecular terms with a level
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MANFRED MARTIN MAYER
261
of sensitivity that enabled the researchers, working with both
guinea pig and human sources, to isolate each individual
protein.
Effective molecule titration proved useful again some
years later when the alternative complement pathway or
properdin system- was rediscovered as a non-antibody-de-
pendent mechanism for recruiting the terminal capabilities
of the complement system. On this occasion, the method's
sensitivity and specificity enabled the researchers to isolate
and characterize the activating proteins rapidly.
The work of Mayer's laboratory on effective molecule ti-
tration of the components of the complement system also led
to the initial recognition that certain of the components had
multiple biologically-active sites. In the case of the second
complement component, these studies showed, the bincling
site to the fourth component was clearly distinguished from
the catalytic site, resulting in the cleavage activation of the
third component.
Mayer later turned his attention to the mechanism by
which the sequentially reacting proteins (at one time termed!
"C3") produced "holes" in the membrane of a target cell des-
tined to undergo lysis. He established that lysis was causer!
by a pentamolecular complex of the terminal five compo-
nents, C5-9, which formed a transmembrane channel iden-
tified (in earlier studies by English electron microscopists) as
discontinuities with an elevated border.
In addition to his unique contributions to the unclerstand-
ing of the sequential interaction and function of the eighteen
proteins of the complement system, Mayer ant! his colleague
Robert Nelson developed the Tr~ponema pallidum immobili-
zation test for syphilis an important contribution to clinical
medicine capable of eliminating false-positive reactions. At
that time the conventional test for syphilis yielder! false-
positive results in inclividuals with gamma globulin abnor-
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262
BIOGRAPHICAL MEMOIRS
malities, including those with autoimmune diseases who did
not have the antibody specific to the spirochete.
Dr. Mayer's contributions to the immunochemistry ant!
biochemistry of the complement field were recognized in
1969 by an honorary degree in medical science from the ~o-
hannes Gutenberg University of Mainz, Germany; in 1974 by
the Karl Lancisteiner Award of the American Association of
Blood Banks; in 1979 by election to the presidency of the
American Association of Immunologists; and in 1982 by the
Gairciner Foundation International Award. In ~ 953 he
shared with Robert Nelson the Kimble Award for MethocI-
ology for the development of the ~ pallidum immobilization
test. He was elected to the National Academy of Sciences in
1979.
TEACHER AND MENTOR
Most teachers of science provide their students with basic
skills and knowledge, but few can instill that additional in-
gredient: confidence to meet the challenges of independent
research. Manfred Mayer was an inspiring scholar who by
example, instruction, ant! wisdom made inclependent re-
searchers of many of his students. Well aware that Mayer's
own vision had uncovered the immunochemistry of comple-
ment Today a significant portion of the discipline of immu-
nology), they used his laboratory as the reference point for
all aspects of the field! of complement research and the mode!
for addressing with technical resourcefulness and appro-
priate critical analysis all clifficult scientific questions.
Dr. Mayer, politely but firmly, demanded technical mas-
tery of all the relevant immunologic methodologies before
he would trust a member of his laboratory to clear with critical
research questions. Technical competence, he maintained,
was the essential prerequisite for personal creativity. He ex-
amined each experiment with an open mincl, exploring the
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MANFRED MARTIN MAYER
263
established results ant! their implications. Several times a clay
he would go with colleagues to the blackboard to discuss
which ciata were secure and which required more work. He
frequently suggested an alternative hypothesis that required
the development of a new methoclology. If the new meth-
odology took months but was the only way to obtain an an-
swer, that was the direction the research took. Mayer's com-
mittec! belief that correct methodology was the prerequisite
for meaningful research meant that his laboratory's meth-
odologic development was continually in flux. His science was
state-of-the-art.
After a piece of work tract been completest, the researchers
had the remarkable experience of putting their results down
on paper for critique by other members of Mayer's labora-
tory. Manfred always treater! the literature of his field with
integrity, while discussing his own data with great imagina-
tion and insight.
What more can be sail! of a giant who cleveloped a whole
scientific field! not only in his personal research, but also
through the training he so generously gave to others? His
rocklike personal integrity became a part of his students' eclu-
cational environment. Never forgetting his own early years
as a refugee from Nazi oppression, he die] all in his power
for the displaced of any background. Truth not politics-
was his only goal, and in the search for truth he generously
shared new hypotheses to be tested with every student, mak-
ing sure that each had a part in the joy of discovery. His
hypotheses further stimulated those about him, generating
ever more definitive experiments. Not surprisingly, Mayer's
laboratory procluced a number of distinguishes! colleagues
ant! students who carry on his own high standards in a variety
of fields (immunochemistry, complement biology, cellular im-
munology), among them Teruko Ishizaka, Moon Shin, anc!
Hyun Shin.
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264
BIOGRAPHICAL MEMOIRS
Manfrec! was equally committed to the clevelopment of
new knowledge and to the education of those of us who in-
teractec! with him. He had no sense of status or rank, and
the friendships he formed with colleagues and students were
lifelong and meaningful. He felt the opportunity for a life in
research a rare privilege that obliged the researcher to strive
for the highest possible level of technical competence, re-
sourcefuIness, integrity, and commitment, both to research
ant! to education ant! he transferred these values to his stu-
dents. Manfred was conspicuously more concerned about the
development of the discipline of immunology and of com-
plement immunochemistry than about his own personal
fame.
Manfred's nonprofessional interests centered on his wife
and four children. Born into a musical family, he maintained
interests in music, languages, ant! art throughout his life.
Both he and Elinor were accomplished amateur pianists, as
well as collectors of art and archaeology.
An admirer of beauty in art, music, and science, Manfred
Mayer was a true role mode] of the scientist-teacher. He de-
veloped a major area of immunology and, with the aid of his
concepts and technologies, prepared those inclividuals who
now pursue it. He is sorely missed by everyone who trained
with him and or was influenced by his work. He will be re-
membered always as a scientist, a teacher, and the founder
of the cliscipline of complement immunochemistry.
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MANFRED MARTIN MAYER
HONORS AND DISTINCTIONS
PROFESSIONAL AND ACADEMIC POSITIONS
265
1938-1942 Laboratory Assistant in Immunochemistry, Columbia
University
1942-1945 Member of the Scientific Staff, Project of the Office
of Scientific Research and Development, Columbia
University
1946 Instructor in Biochemistry, Columbia University
1946-1947 Assistant Professor of Bacteriology, Johns Hopkins
University School of Hygiene and Public Health
1948
1957
1960
LEARNED SOCIETIES
Associate Professor of Microbiology, Johns Hopkins
University School of Hygiene and Public Health
Acting Chairman, Department of Microbiology,
Johns Hopkins University School of Hygiene and
Public Health
Professor of Microbiology, Department of Micro-
biology, Johns Hopkins University of Medicine
American Association for the Advancement of Science
American Association of Immunologists
American Chemical Society
American Society of Biological Chemists
Biochemical Society
National Academy of Sciences
Society for Experimental Biology and Medicine
HONORARY MEMBERSHIPS
Phi Beta Kappa
Sigma Xi
Collegium Internationale Allergologicum
OTHER PROFESSIONAL ACTIVITIES
Consultant, United States Public Health Service
Consultant, National Science Foundation
Consultant, Office of Naval Research
Consultant, Plum Island Animal Disease Laboratory, Department
of Agriculture
Associate Editor, Biological Abstracts
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266
BIOGRAPHICAL MEMOIRS
Associate Editor, Journal of Immunology
Associate Editor, Analytical Biochemistry
Associate Editor, Immunochemistry
President, American Association of Immunologists
Editorial Board, journal of Immunology
PRIZES AND AWARDS
1945 Citation, Columbia University, for work in the Division of
War Research during World War II
1953 Kimble Award for Methodology
1957 Selman Waksman Lectureship Award
1969 Honorary Doctor of Medical Science, tohannes Gutenberg
University, Mainz, Germany
1974 Karl Landsteiner Award, American Association of Blood
Banks
1976 Albion 0. Bernstein Award, Medical Society of the State of
New York
1982 Gairdner Foundation International Award, Toronto,
Canada
OCR for page 271
MANFRED MARTIN MAYER
271
1958
Studies on the mechanism of hemolysis by antibody and comple-
ment. Prog. Allergy, 5:215.
With B. Roizman and W. Hopken. Immunochemical studies of
poliovirus. II. Kinetics of the formation of infectious and non-
infectious type I poliovirus in three cell strains of human deri-
vation. I. Immunol., 80:386.
With B. Roizman and H. I. Rapp. Immunochemical studies of
poliovirus. III. Further studies on the immunological and phys-
ical properties of poliovirus particles produced in tissue culture.
I. Immunol., 81:419.
1959
With B. Roizman and P. R. Roane, in Immunochemical studies of
poliovirus. IV. Alteration of the immunological specificity of
purified poliovirus by heat and ultraviolet light. I. Immunol.,
82:119.
With L. G. Hoffmann, H. J. Rapp, and J. R. Vinas. A kinetic flow
technique for study of immune hemolysis. Proc. Soc. Exp. Biol.
Med., 100:211.
1961
Development of the one-hit theory of immune hemolysis. In: Im-
munochemical Approaches to Problems in Microbiology. New Bruns-
wick, N.~.: Rutgers University Press.
With T. Borsos and H. J. Rapp. Studies on the second component
of complement. II. The reaction between EAC'1,4 and C'2:
Evidence on the single site mechanism of immune hemolysis
and determination of C'2 on an absolute molecular basis. I.
Immunol., 87:310.
With T. Borsos and H. J. Rapp. Studies on the second component
of complement. II. The nature of the decay of EAC'1,4,2. I.
Immunol., 87:326.
On the destruction of erythrocytes and other cells by antibody and
complement. Cancer Res., 21:1262.
With E. A. Kabat. Experimental Immunochemistry, 2d ed. Springfield,
Ill.: C. C. Thomas.
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272
BIOGRAPHICAL MEMOIRS
1962
With T. Borsos. Mechanism of action of guinea pig complement.
In: Mechanism of Cell and Tissue Damage Produced by Immune Re-
actions (Second International Symposium on Immunopathol-
ogy, Brook Lodge, Michigan). Basel: Benno Schwabe & Co.
1963
Enzymatic cleavage of C'2 by EAC'la,4: Fixation of C'2a on the
cell and release of C'2i. Science, 141 :738.
With H. I. Rapp and T. Borsos. Complement. National Cancer
Institute Workshop, February 28-March 1, 1963. Bethesda,
Maryland.
1965
With W. F. Willoughby. Antibody-complement complexes. Science,
150:907. Mechanism of hemolysis by complement. In: CIBA
Foundation Symposium on Complement, eds. G. E. W. Woltsten-
holme and l. Knight, London: I. & A. Churchill, Ltd., p. 4.
With L. G. Hoffman and A. T. McKenzie. The steady state system
in immune hemolysis. Description and analysis; Application to
the enumeration of SAC'4. Immunochemistry, 2: 13.
With J. A. Miller. Inhibition of guinea pig C'2 by rabbit antibody,
quantitative measurement of inhibition, discrimination between
immune inhibition and complement fixation, specificity of in-
hibition and demonstration of uptake of C'2 by EAC' la,4. Im-
munochemistry, 2:71.
With R. M. Stroud and K. F. Austen. Catalysis of C'2 fixation by
C' la. Reaction kinetics, competitive inhibition by TAMe, and
transferase hypothesis of the enzymatic action of C' la on C'2,
one of its natural substrates. Immunochemistry, 2:219.
1966
With G. Sitomer and R. M. Stroud. Reversible adsorption of C'2
by EAC'4: role of Mg++, enumeration of competent SAC'4,
two-step nature of C'2a fixation and estimation of its efficiency.
Immunochemistry, 3:57.
With R. M. Stroud, J. A. Miller, and A. T. McKenzie. C'2a&, an
inactive derivative of C'2 released during decay of EAC'4,2a.
Immunochemistry, 3:163.
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MANFRED MARTIN MAYER
273
1967
With H. S. Shin and l. A. Miller. Fragmentation of guinea pig
complement components C'2 and C'3c. In: Protides of the Bio-
log~cal Fluids ( 1 5th Annual Colloquium), Amsterdam: Elsevier
Publishing Co., p. 411.
1968
With H. S. Shin. The third component of the guinea pig comple-
ment system. I. Purification and characterization. Biochemistry,
7:2991.
With H. S. Shin. The third component of the guinea pig comple-
ment system. II. Kinetic study of the reaction of EAC'4,2a with
guinea pig C'3. Enzymatic nature of C'3 consumption, multi-
phasic character of fixation, and hemolytic titration of C'3. Bio-
chemistry, 7:2997.
With H. S. Shin. The third component of the guinea pig comple-
ment system. III. Effect of inhibitors. Biochemistry, 7:3003.
With I. A. Miller. On the cleavage of C'2 by Cla: Immunological
and physical comparisons of C'2a~ and C'2a/i. Proc. Soc. Exp.
Biol. Med., 129: 127.
With H. S. Shin, R. Snyderman, E. B. Friedman, and A. I. Mellors.
A chemotactic and anaphylatoxic fragment cleaved from the
fifth component of guinea pig complement. Science, 162:361.
1969
With D. I. Hingson and R. K. Massengill. The kinetics of release
of 86rubidium and hemoglobin from erythrocytes damaged by
antibody and complement. Immunochemistry, 6:295.
1970
With I. A. Miller. Photometric analysis of proteins and peptides at
191-194 m. Analyt. Biochem., 36:91.
With F. A. Rommel, M. B. Goldlust, F. C. Bancroft, and A. H.
Tashjian, Jr. Synthesis of the ninth component of complement
by a clonal strain of rat hepatoma cells. J. Immunol., 105:396.
With I. A. Miller and H. S. Shin. A specific method for purification
of the second component of guinea pig complement and a
chemical evaluation of the one-hit theory. J. Immunol.,
105:327.
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274
BIOGRAPHICAL MEMOIRS
Highlights of complement research during the past twenty-five
years. Immunochemistry, 7:485.
1971
With C. T. Cook, H. S. Shin, and K. Laudenslayer. The fifth com-
ponent of the guinea pig complement system. I. Purification
and characterization. I. Immunol., 106:467.
With H. S. Shin and R. l. Pickering. The fifth component of the
guinea pig complement system. II. Reaction of C5 with
EAC' 1,4,2,3. I. Immunol., 106:473.
With H. S. Shin and R. I. Pickering. The fifth component of the
guinea pig complement system. III. The properties of
EAC 1,4,2,3,5b. I. Immunol., 106:480.
With M. B. Goldlust and H. S. Shin. Elution of guinea pig "C5b/6"
activity from EAC 1,4,2a,3b,5b,6. J. Immunol (abstract).,
107:318.
With R. L. Marcus and H. S. Shin. An alternate pathway: Dem-
onstration of C3 cleaving activity, other than C4,2a, on endo-
toxic lipopolysaccharide after treatment with guinea pig serum.
Relation to the properdin system. Proc. Natl. Acad. Sci. USA,
68:1351.
With C. S. Henney. Specific cytolytic activity of lymphocytes: Effect
of antibodies against complement components C2, C3, and C5.
Cell. Immunol., 2:702.
1972
Mechanism of cytolysis by complement. Proc. Natl. Acad. Sci. USA,
69:2954. With M. K. Gately. The effect of antibodies to comple-
ment components C2, C3, and C5 on the production and action
of lymphotoxin. J. Immunol., 109:728.
With V. Brade, C. T. Cook, and H. S. Shin. Studies on the proper-
din system: Isolation of a heat-labile factor from guinea pig
serum related to a human glycine-rich beta-glycoprotein (GBC
or factor B). I. Immunol., 109: 1174.
1973
With F. A. Rommel. Studies of guinea pig complement component
C9: Reaction kinetics and evidence that lysis of EAC1-8 results
from a single membrane lesion caused by one molecule of C9.
J. Immunol., 110:637.
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MANFRED MARTIN MAYER
275
With A. Eden, C. Bianco, and V. Nussenzweig. C3 split products
inhibit the binding of antigen-antibody-complement com-
plexes to B lymphocytes. J. Immunol., 110: 1452.
The complement system. Sci. Am., 229:54.
With V. Brade, G. D. Lee, A. Nicholson, and H. S. Shin. The re-
action of zymosan with the properdin system in normal and C4-
deficient guinea pig serum: Demonstration of C3- and C5-
cleaving and multi-unit enzymes, both containing factor B. and
acceleration of their formation by the classical complement
pathway. I. Immunol., 111: 1389.
1974
With A. Nicholson, V. Brade, G. D. Lee, and H. S. Shin. Kinetic
studies of the formation of the properdin system enzymes on
zymosan. Evidence that nascent C3b controls the rate of as-
sembly. l. Immunol., 112:1115.
With M. K. Gately. The molecular dimensions of guinea pig lym-
photoxin. I. Immunol., 112: 168.
With V. Brade, A. Nicholson, and G. D. Lee. The reaction of zym-
osan with the properdin system. Isolation of purified factor D
from guinea pig serum and study of its reaction characteristics.
I. Immunol., 112: 1845.
With M. B. Goldlust, H. S. Shin, and C. H. Hammer. Studies of
complement complex C5b,6 eluted from EAC-6: Reaction of
C5b,6 with EAC4b,3b and evidence on the role of C2a and C3b
in the activation of C5. I. Immunol., 113:998.
1975
Complement. An immunological and pathological mediator sys-
tem. Medizin. Prisma, (May):2.
With C. L. Gately and M. K. Gately. The molecular dimensions of
mitogenic factor from guinea pig lymph node cells. I. Immu-
nol., 114:10.
With C. H. Hammer and A. Nicholson. On the mechanism of cy-
tolysis by complement. Evidence on insertion of the C5b and
C7 subunits of the C5b,6,7 complex into the phospholipid bi-
layer of the erythrocyte membrane. Proc. Natl. Acad. Sci. USA,
72:5076.
The complex complement system. Inflo, 8: 1.
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276
BIOGRAPHICAL MEMOIRS
1976
With C. L. Gately and M. K. Gately. Separation of lymphocyte mi-
togen from lymphotoxin and experiments on the production
of lymphotoxin by lymphoid cells stimulated with the partially
purified mitogen: A possible amplification mechanism of cel-
lular immunity and allergy. I. Immunol., 116:669.
With D. W. Michaels and A. S. Abramovitz. Increased ion perme-
ability of planar lipid bilayer membranes after treatment with
the C5b-9 cytolytic attack mechanism of complement. Proc.
Natl. Acad. Sci. USA, 73:2852.
With C. H. Hammer and A. S. Abramovitz. A new activity of com-
plement component C3: Cell-bound C3b potentiates lysis of
erythrocytes by C5b,6 and terminal components. I. Immunol.,
117:830.
On the mechanism of cytolysis by complement: Experimental stud-
ies of the transmembrane channel hypothesis. In: The Nature
and Significance of Complement Activation (An international sym-
posium sponsored by Ortho Research Institute of Medical Sci-
ence September, 1976, in Raritan, New Jersey).
With M. K. Gately and C. S. Henney. Effect of anti-lymphotoxin
on cell-mediated cytotoxicity. Evidence for two pathways, one
involving lymphotoxin and the other requiring intimate contact
between the plasma membranes of killer and target cells. Cell.
Immunol., 27:82.
1977
The cytolytic attack mechanism of complement. In: Mediators of the
Immediate Type Inflammatory Reaction. Mono. Allergy 12, Basel:
S. Karger.
With C. H. Hammer, M. L. Shin, and A. S. Abramovitz. On the
mechanism of cell membrane damage by complement: Evi-
dence on insertion on polypeptide chains from C8 and C9 into
the lipid of erythrocytes. l. Immunol., 119: 1.
With M. L. Shin, W. A. Paznekas, and A. S. Abramovitz. On the
mechanism of cell membrane damage by complement: Expo-
sure of hydrophobic sites on activated complement protein.
J. Immunol., 119: 1358.
On the mechanism of cytolysis by lymphocytes: A comparison with
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MANFRED MARTIN MAYER
277
complement. (Presidential address to the American Association
of Immunologists, April, 1977.) J. Immunol., 119: 1195.
With H. I. Muller-Eberhard and L. G. Hoffmann. Complement.
In: Methods in Immunology and Immunochemistry, eds. A. W. Curtis
and M.W.Chase,vol.4,p. 1Y7.
With S. Cohen, J. David, M. Feldmann, P. R. Glade, J. J. Oppen-
heim, et al. Current state of studies of mediators of cellular
immunity: A progress report. Cell. Immunol., 33:233.
1978
Complement, past and present. In: The Harvey Lect., 72, 1976-
1977. New York: Academic Press.
With M. Okamoto. Studies on the mechanism of action of guinea
pig lymphotoxin. I. Membrane active substances prevent target
cell lysis by lymphotoxin. J. Immunol., 120:272.
With M. Okamoto. Studies on the mechanism of action of guinea
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damaged target cells. I. Immunol., 120:279.
With M. L. Shin and W. A. Paznekas. On the mechanism of mem-
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uration of the acyl chains in liposomal bilayers and the effect of
cholesterol concentration in sheep erythrocytes and liposomal
membranes. I. Immunol., 120: 1996.
With M. K. Gately. Purification and characterization of lympho-
kines: An approach to the study of molecular mechanisms of
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With C. H. Hammer, D. W. Michaels, and M. L. Shin. Immunolog-
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Dlement action and the similarity of lymphocyte-mediated cy-
. ~ —~—
totoxicity. Transplant. Prox., ~ u:-/ u-/ .
1979
With C. H. Hammer, D. W. Michaels, and M. L. Shin. Immunolog-
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totoxicity. I mmunochemistry, 15 :813.
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brane lipid composition in a liposome system. Biochim. Bio-
phys. Acta, 55:79.
With M. K. Gately, M. Okamoto, M. L. Shin, and I. B. Willoughby.
Two mechanisms of cell-mediated cytotoxicity: (1) Ca++ trans-
port modulation by lymphotoxin, and (2) transmembrane chan-
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With L. E. Ramm. Life-span and size of the trans-membrane chan-
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With I. B. Willoughby. On the channel hypothesis of antibody-
dependent cell-mediated cytotoxicity (ADCC): Evaluation of a
liposome model system. In: Biochemical Characterization of Lym-
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York: Academic Press.
Trans-membrane channels produced by complement proteins.
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1981
With M. L. Shin and G. M. Hansch. Effect of agents that produce
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Hopkins Med. }, 148:243.
With G. M. Hansch, C. H. Hammer, and M. L. Shin. Activation of
the fifth and sixth component of the complement system: Sim-
ilarities between C5b6 and C(56)a with respect to lytic enhance-
ment by cell-bound C3b or A2C, and species preferences of
target cell. J. Immunol., 127:999.
With D. W. Michaels, L. E. Ramm, M. B. Whitlow, J. B. Willoughby,
and M. L. Shin. Membrane damage by complement. Crit. Rev.
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Membrane attack by complement (with comments on cell-mediated
cytotoxicity). In: Mechanisms of Cell-Mediated Cytotoxicity, eds.
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With L. E. Ramm and M. B. Whitlow. Size of the trans-membrane
channels produced by complement proteins C5b-8. I. Immu-
nol., 129:1143.
With L. E. Ramm and M. B. Whitlow. Trans-membrane channel
formation by complement. Functional analysis of the number
of C5b6, C7, C8, and C9 molecules required for a single chan-
nel. Proc. Natl. Acad. Sci. USA, 79:4751.
1983
With L. E. Ramm and M. B. Whitlow. Size distribution and stability
of the trans-membrane channels formed by complement com-
plex C5b-9. Mol. Immunol., 20: 155.
With L. E. Ramm, D. W. Michaels, and M. B. Whitlow. On the size,
heterogeneity and molecular composition of the trans-
membrane channels produced by complement. In: Biological
Response Mediators and Modulators, ed. l. T. August, New York:
Academic Press.
With C. L. Koski, L. E. Ramm, C. H. Hammer, and M. L. Shin.
Cytolysis of nucleated cells by complement: Cell death displays
multi-hit characteristics. Proc. Natl. Acad. Sci. USA, 80:3816.
With L. E. Ramm, M. B. Whitlow, C. L. Koski, and M. L. Shin.
Elimination of complement channels from the plasma mem-
branes of U937, a nucleated mammalian cell line: Temperature
dependence of the elimination rate. I. Immunol., 121:1411.
With D. K. Imagawa, L. E. Ramm, and M. B. Whitlow. Membrane
attack by complement and its consequences. In: Progress in Im-
munology, eds. Y. Yamamura and T. Tada, Tokyo: Academic
Press Japan, p. 427.
With D. K. Imagawa, N. E. Osilchin, W. A. Paznekas, and M. L.
Shin. Consequences of cell membrane attack by complement:
Release of arachidonate and formation of inflammatory deriv-
atives. Proc. Natl. Acad. Sci. USA, 80:6647.
Complement. Historical perspectives and some current issues.
Complement, 1:2.
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With L. E. Ramm and M. B. Whitlow. Complement lysis of nu-
cleated cells: Effect of temperature and puromycin on the
number of channels required from cytolysis. Mol. Immunol.,
21:1015.
With M. B. Whitlow and L. E. Ramm. Penetration of C8 and C9 in
the C5~9 complex across the erythrocyte membrane into the
cytoplasmic space. I. Biol. Chem., 260:998.
With L. E. Ramm and M. B. Whitlow. The relationship between
channel size and the number of C9 molecules in the C5~9
complex. I. Immunol., 134:2594.
1987
With D. K.Imagawa, N. E. Osifchin, L. E. Ramm, P. G. Koga, C.
H. Hammer, and H. S. Shin. Release of arachidonic acid and
formation of oxygenated derivatives following complement at-
tack on macrophages: Role of channel formation. }. Immunol.
OCR for page 281
Representative terms from entire chapter:
complement fixation
