Below are the first 10 and last 10 pages of uncorrected machine-read text (when available) of this chapter, followed by the top 30 algorithmically extracted key phrases from the chapter as a whole.
Intended to provide our own search engines and external engines with highly rich, chapter-representative searchable text on the opening pages of each chapter.
Because it is UNCORRECTED material, please consider the following text as a useful but insufficient proxy for the authoritative book pages.
Do not use for reproduction, copying, pasting, or reading; exclusively for search engines.
OCR for page 120
OCR for page 121
CLEMENT LAWRENCE MARKERT
April ]], 1917-October ], 1999
BY GERALD M. KIDDER
CLEMENT L. MARKERT DIED on October I, 1999, in Colorado
Springs, Colorado, at the age of 82. He and his wife,
Margaret, had been living there since his retirement from
North Carolina State University in 1993. Markert was born
in Las Animas, Colorado, and grew up in the area around
Pueblo. He and Margaret returned to "their mountain" near
Westcliffe, Colorado, each summer. It was in their beloved
mountain wilderness that he was laid to rest. Margaretjoined
him in death the following year. They are survived by three
children: Alan, Robert, and Samantha (Betsy) Schreck.
A MAN OF IDEALS AND ACTION
Clement Markert was a man of ideals whose devotion to
social causes was evident from early in his career. His father
had been a steel worker, and the family had suffered dur-
~ng the great Depression, when the mines and steel mills
were closing. This experience undoubtedly influenced the
· . . _~ . ~ ~
development of Markert's social conscience. Grateful for
scholarships awarded for his academic achievements, he
enrolled in the University of Colorado at Boulder to study
biology. At the same time, however, he was concerned about
events on the world stage, especially what he perceived to
121
OCR for page 122
122
B I O G RA P H I C A L
EMOIRS
be the failure of capitalistic economies to meet the neecis
of working-cIass people. He embraced socialism en cl even
organized a communist group at the university. He was very
soon presenter! with an opportunity to put his social icleals
into more direct practice. Responding to the threat of fas-
cist movements taking hoIcl in Europe in 193S, he inter-
ruptec! his studies and, along with his college roommate,
rocle freight trains to the East Coast, where the two men
stowocl away on a merchant ship bouncl for France. From
there they joiner! the famous Abraham Lincoin Brigacle,
which was fighting the forces of Generalissimo Franco in
Spain, hoping to prevent his toppling of the democratically
electec! government. Markert later explainer! that he hac!
been one of the few members of his combat unit to survive
the Spanish Civil War (his roommate was one of the casual-
ties). In an obituary for Markert in The New York Times
(October 10, 1999) he was quoted as having saicl in a 1986
interview, "I felt the most concrete thing I couIcl clo at the
time was to destroy fascism, en c! Spain was the battleground!
on which to clo that."
After the defeat of the anti-Franco forces Markert re-
turned to the University of Colorado to complete his un-
clergracluate studies. He was awarclecl a B.A. summa cum
laucle in ~ 940. In that same year he married Margaret
Rempfer, who was to be his partner for life. The couple
movecl to California so that Markert couIcl clo graduate work
at the University of California, Los Angeles, where he con-
cluctec! research in vertebrate embryology, however, woric!
events again intervened. The Unitecl States became involvecl
in WorIcl War II, en cl Markert chose to reactivate his per-
sonal fight against fascism. He took a master's degree in
1942 to terminate his graduate education en cl triecl to en-
list in the U.S. Army. Not surprisingly, given the political
climate of the time, his previous associations with American
OCR for page 123
CLEMENT LAWRENCE MARKERT
123
en c! Spanish communists, who hac! also been fighting against
Franco, macle him unacceptable for military service. In re-
sponse to this setback he movecl to San Diego to serve as a
clockworker until being acceptec! into the merchant ma-
rine. He served out the war as a raclio operator on a ship
supplying U.S. forces in the Pacific.
When his war years were finally behinc! him, Markert
enrollecl in the cloctoral program in biology at Johns Hopkins
University, where he concluctecl research uncler the
mentorship of one of the country's foremost clevelopmen-
tal biologists at the time, Benjamin H. Willier. After earn-
ing the doctorate in 1948 he completecl his research train-
ing as a Merck-NRC postcloctoral fellow at the California
Institute of Technology. There he specializecl in biochemi-
cal genetics uncler the influence of George W. BeacIle, the
foremost proponent of that emerging fielcI.
A MAN OF INTEGRITY AND COURAGE
Markert's first independent academic appointment was
at the University of Michigan in Ann Arbor, where he ac-
cepted an assistant professorship in the Zoology Depart-
ment in 1950. He became the intellectual leacler of a group
of junior faculty who were in tune with the recent advances
in biochemistry en cl genetics that lecl in 1953 to Watson
en c! Crick's publication of the structure of DNA. The Markert
family, which by this time incluclecl all three chilciren, settlecl
into the pleasant life of the Ann Arbor academic commu-
nity, en c! it seemec! that Markert's earlier life as a social
activist was a thing of the past.
This notion was shattered in 1954 when Markert en cl
two colleagues were called before a subcommittee of the
House Un-American Activities Committee meeting in East
Lansing, Michigan. The subcommittee, chaired by Michi-
gan Representative Kit CIarcly, was manciatec! to identify
OCR for page 124
124
B I O G RA P H I C A L
EMOIRS
and root out communists from academia. Those who were
targeted by the committee were threatened with being ex-
posed as communists unless they named their former asso-
ciates who might be considered communists or sympathiz-
ers. The three men declined to cooperate, refusing to name
anyone. As a consequence all three were suspended from
their positions with the university, which set up review com-
mittees at various levels to examine their cases. Markert was
the only one of the three who was reinstated. According to
David Nanney, a departmental colleague of Markert's at
the time,~~ ~ ~ ~
Markert survived the ordeal because of support
from his academic colleagues, who were convinced of his
personal integrity, as well as scientists elsewhere (including
George Beadle) who were impressed with Markert's scien-
tific acumen and wrote letters on his behalf. Markert would
later relate this experience to his students to emphasize the
importance of standing up for one's convictions, whether
scientific or political, regardless of the cost. Years later the
university invited the three men back to Ann Arbor to re-
ceive an apology for the way they had been treated.
The controversy surrounding Markert's youthful social-
ist activism did not end there. In 1957 he applied for the
position in developmental biology at Johns Hopkins that
was being vacated by his retiring mentor, Professor Willier.
When the search committee recommended Markert's ap-
pointment, administrative resistance developed. Markert had
made no secret of his past, indeed, he was proud of it! The
impasse was resolved when, after interviewing Markert him-
self, the president of the university, Milton Eisenhower
(brother of the President), recommended Markert's appoint-
ment as a full professor and threatened to resign if the
appointment was not confirmed. Markert accepted the po-
sition and remained at Hopkins until moving to Yale Uni-
versity in 1965 to become chair of the Department of Biol-
OCR for page 125
CLEMENT LAWRENCE MARKERT
125
ogy. Once again Markert took pains to ensure that YaTe's
president at the time, Kingman Brewster, was fully aware of
his past en cl his intention to remain involvecl in social causes.
During the late 1960s Markert was an outspoken opponent
of the government's continuing involvement in the Viet-
nam conflict en cl took an active role in public protests.
Other causes that receiver! his outspoken support incluclec!
affirmative action to promote women in academia en cl the
"Zero Population Growth" campaign (his car license plate
for a time was ZPG). Through this entire advocacy, as al-
ways, Margaret was by his sicle. IncleecI, Markert attributed
much of his confidence cluring those clifficult times in Michi-
gan to the knowlecige that his wife wouic! be able to cope
with whatever hardship his political activism brought upon
them.
MOST NOTABLE SCIENTIFIC CONTRIBUTIONS
Throughout his career Markert aimed high: He wan tell
to tackle the big questions in biological science, questions
like how genes control clevelopment en cl how the genome
of an organism can be manipulated to bring about genetic
improvement. In many cases answering such questions re-
quirecl the clevelopment of new research techniques. His
scientific contributions covered a wicle range from biochem-
istry through clevelopmental en c! reproductive genetics.
Markert was best known early in his career for eluciciat-
ing the importance and structural basis of isozymes, mul-
tiple molecular forms of enzymes. The stage was set for that
work when in 1957 Markert en cl his University of Michigan
colleague, Robert L. Hunter, combined enzyme histochem-
istry with the starch gel electrophoresis technique newly
clevelopecl by Oliver Smithies to show that there are more
than 10 separable forms of esterases in mouse liver (Hunter
en c! Markert, 1957~. Using different substrates or inhibitors
OCR for page 126
126
B I O G RA P H I C A L
EMOIRS
in the histochemical staining reaction, they obtained evi-
clence that the different esterase bancis in the gel were en-
zymatically distinct. The same technique but using cliffer-
ent histochemical reagents also revealer! multiple forms of
other enzymes, demonstrating that this phenomenon is not
limitecl to esterases. The investigators termed their stained
gel, showing multiple bancis representing the same enzy-
matic function, a zymogram. In a subsequent paper com-
municatecl to Proceedings of the National Academy of Sci-
ences by Benjamin Willier, Markert en c! Frecicly Mailer user!
the zymogram technique to show that the number of mo-
lecular forms of lactate clehycirogenase (LDH) in mamma-
lian tissues is greater than hac! been appreciates! en c! pro-
posecl the term isozyme to denote these forms (Markert
en cl Mailer, 1959~. They also showocl that tissues cliffer in
the number of LDH isozymes they contain en c! their rela-
tive proportions. Most importantly, their ciata macle it clear
that the isozyme patterns of embryonic tissues change through
ontogeny until the tissue-appropriate aclult pattern is
achieved, a phenomenon that was interpreted as indicating
changes in gene expression relatecl to cell differentiation.
This insight into the utility of isozyme studies for under-
stancling clevelopmental mechanisms was to influence
Markert's research for years to come. Mailer, a Dane who
hac! been trainee! in veterinary medicine before joining
Markert's lab (by then at Hopkins), later creclitecl the ex-
citement of those clays with his decision to make research
his career. Markert's insights into the importance of cliffer-
ential gene activation cluring clevelopment proviclecl a new
way of looking at abnormal clevelopment as well, en cl he
was one of the first to point out that diseases such as cancer
can be viewocl as cell differentiation gone awry (Markert,
1 968~.
As important as it was, Markert en c! M011er's 1959 paper
OCR for page 127
CLEMENT LAWRENCE MARKERT
127
left unexplained! the molecular basis of isozymes. There was
little appreciation at the time of the existence of gene fami-
lies, evolutionarily relatecl genes encocling proteins of simi-
lar or overlapping function. Yet Markert en c! Mailer clic!
offer that as one explanation, citing the multiplicity of genes
encocling fetal en cl aclult hemoglobins. They also suggested
that a single gene might somehow encocle an array of isozymes
differing in "structural variations," a concept that seems to
presage our current unclerstancling of alternative mRNA
splicing en c! post-transTational protein mollification. It was
several years later, through the efforts of Ettore AppelIa, an
Italian postcloc, that the Markert laboratory finally came to
a clear unclerstancling of the molecular basis of LDH isozymes.
By treating the enzyme with denaturing agents it was Earned
that LDH is a tetramer of two types of polypepticle chains
(Appella and Markert, 1961~. Thus the multiple-gene hy-
pothesis was partially correct: Two different LDH subunits,
each encoclecl by a distinct gene, re-sort themselves in vari-
ous tetrameric combinations to give rise to five different
isozymes (Markert, ~ 963 ) . During the succeeding years
Markert en cl his students en cl postclocs continual to stucly
the molecular basis and biological significance of isozymes
en cl showocl how the stucly of isozymes couIcl contribute to
our unclerstancling of the biochemical variation that uncler-
lies cell differentiation and evolution. The culmination of
this work was the new perspective presented in a Science
paper (Markert et al., 1975) entitled "Evolution of a Gene,"
coauthored! with former graduate students James B. Shaklee
en cl Gregory S. Whitt. Markert took particular pricle in his
role in eluciciating the isozyme concept, not least because
this was a case of a clevelopmental biologist teaching some-
thing about biochemistry to the biochemists. For several
years he served as editor or coeditor of the multivolume
OCR for page 128
128
B I O G RA P H I C A L
EMOIRS
series "Isozymes" that emanates! from the annual Interna-
tional Congress on Isozymes.
Markert's preclilection for tackling the big questions some-
times causer! problems for the person in his laboratory who
was taking the leacl on a project. For example, one iclea
that was tested cluring Markert's Hopkins years was that the
program of gene expression within a cell is clictatec! by the
constellation of nuclear proteins interacting with its DNA.
If so, then it was hypothesized that introducing nuclear
proteins from another source shouic! reprogram a cell's ge-
netic machinery. This was tested by injecting liver nuclear
proteins into fertilized frog eggs with the expectation that
the embryos wouic! clevelop characteristics of liver cells. In-
steacl the embryos arrested their clevelopment, en cl little
was Earned from the experiment despite exhaustive attempts
to analyze the embryos using the techniques of the clay.
Markert was later criticized for investing resources en cl stu-
clent time in such a simple-minclecl approach to a very com-
plex problem, but if the experiment hac! worker! at least
partially, it wouIcl have been a major step forward.
Shortly after moving to Yale, Markert's laboratory be-
came involves! in a new research topic that was to have an
impact at least as important as that of the isozyme concept.
Yoshio Masui, a young scientist from Konan University in
Japan, arrived at Yale in 1966 on sabbatical leave to study
biochemical aspects of cell differentiation and clevelopment.
Masui had become intrigued by Markert's view of develop-
ment as emanating from clifferential gene activation en c!
wan tell to contribute to the eluciciation of that concept. He
began working on LDH isozymes in penguin embryos, char-
acterizing their changing expression patterns cluring clevel-
opment. After less than a year, however, Masui came to the
conclusion that the complexity of regulation of even a single
enzymatic function during development was too great to be
OCR for page 129
CLEMENT LAWRENCE MARKERT
129
eTuciciatec! by the technology available in the ~ 960s. He
wan tell a more tractable problem to work on. Markert en-
couragecl him to choose a project of his own interest, one
that he conic! continue working on after returning to
Japan.
Masui cleciclecl that to unclerstancl cell differentiation it
wouic! be advantageous to stucly an unambiguous cell change
inclucecl by a well-clefinecl external signal. Remembering
the classical experiment by Heilbrunn et al. (1939) in which
oocytes were inclucec! to be releaser! from frog ovaries treater!
in vitro with a pituitary glancl suspension, Masui reasoned
that this must be an example of a clevelopmental incluction
evokes! by a hormone. He was impresser! that a hormone
couIcl act clirectly on its target tissue in vitro. Furthermore,
Masui realizecl that hormonal incluction of melotic matura-
tion en c! ovulation of the frog oocyte conic! provicle a highly
advantageous system for studying the control of cell cycle
events: It wouIcl allow the investigator to use distinct stimuli
to incluce oocyte maturation (response to the hormone)
en cl egg activation (cleavage in response to fertilization),
thus separating the signals that drive the cell cycle from G2
to M phase and from M to G! phase, respectively. He hoped
in this way to develop a research program in nucleocyto-
plasmic interactions that he couIcl continue in Japan, where
research resources were not as plentiful at the time, taking
advantage of the ability to obtain large numbers of synchro-
nous frog oocytes for biochemical analysis. For his part
Markert was enthusiastic about that line of investigation,
because he hacl often musecl about the possibility of sup-
pressing meiosis in oocytes as a route to parthenogenesis.
Masui's proposer! experiments were seen as an early step
along that roacI, since they couIcl reveal how meiosis is con-
trolled (Masui, 2001~.
In early 1967 Masui starter! research on oocyte matura-
OCR for page 130
30
B I O G RA P H I C A L
EMOIRS
tion by repeating Heilbrunn's classical experiment using
Ran a pipiens. His experiments eventually lecl him to con-
clucle that pituitary gonadotropin acts on the follicle cells
of the ovary to stimulate them to release a progesterone-
like hormone that clirectly acts on the oocyte. Further work
revealecl that progesterone couIcl have an effect only when
it actec! from the outsicle of the oocyte or on the cell sur-
face, leacling him to propose that the oocyte cytoplasm car-
ries the hormonal signal to the oocyte nucleus to incluce
the first meiotic division. To test this hypothesis Masui in-
jectecl the cytoplasm of oocytes inclucecl to mature by proges-
terone into immature oocytes en cl found that these oocytes
were inclucec! to mature without hormone treatment. That
was the now famous experiment that clemonstratecl the pres-
ence of a cytoplasmic factor, which Masui en cl Markert callecl
maturation promoting factor (MPF), that caused oocyte
maturation by triggering meiosis (Masui and Markert, 1971~.
Using the same bioassay it was shown that MPF appears
before the oocyte enters M phase, but clecTines when the
oocyte proceeds to G] phase after fertilization. Masui also
clemonstratecl that maturing oocytes contain another fac-
tor, namer! cytostatic factor (CSF), that is responsible for
the arrest of oocyte meiosis until fertilization. The manu-
script reporting these exciting results (Masui and Markert,
1971) was published shortly after Masui moved to the Uni-
versity of Toronto, where he is still working. It was the first
significant step in unclerstancling how cell division is con-
trollecI. That work was follower! by research in other labora-
tories studying cell cycle regulation in yeasts, where the
power of genetics was used to identify specific molecules
having the properties of MPF and CSF. Today we know that
MPF, more generally known as M-phase promoting factor,
is a complex of cyclin B2, a regulatory protein that is syn-
thesizec! en c! then clestroyoc! in each cell cycle, en c! Crick, a
OCR for page 131
CLEMENT LAWRENCE MARKERT
131
catalytic protein that promotes entry into M phase. CSF is a
Mos protein-containing complex that acts to prevent cyclin
B2 clegraciation, thus maintaining the cell in M phase
(Duesbery en c! Van cle Woucle, 2002~. The importance of
Masui and Markert's 1971 paper was recognized in 1992
with the awarding of the prestigious Gairciner Awarcl (
OCR for page 132
132
B I O G RA P H I C A L
EMOIRS
blastocysts were obtainer! by this procedure, but none sur-
vivecl to term after transfer to foster mothers (Markert en cl
Petters, 1977~. Soon afterward another team of researchers
ciaimec! to have succeeclec! with the same procedure (Hoppe
en cl IlImensee, 1977~. Those results have not been repli-
catecI. The current view, basecl on a large bocly of ciata, is
that clifferential epigenetic mollification of sperm en c! egg
genomes preclucles normal post-blastocyst clevelopment when
the embryonic genome is clerivecl from a single parent.
Markert liver! just Tong enough to see mammalian cloning,
now performed by nuclear transfer into enucleatecl oocytes,
become a reality (Wilmut et al., 1997~.
Markert en c! postcloc Robert Petters clic! succeec! with
another technically clemancling experiment: the procluction
of hexaparental chimeras, mice macle up of cells from three
different embryos having different genotypes (Markert en c!
Petters, 1978~. They then repeated the experiment with
four different embryos, producing octaparental mice (Petters
en c! Markert, 1980~. This result prover! that at least four
embryonic stem cells of the early embryo give rise to the
fetus. Pictures of those hexaparental mice are still featured
in clevelopmental biology textbooks. At the same time, a
graduate student in Markert's laboratory, Vijay Thadani,
was showing that rat oocytes can be fertilized by sperm of
other mammalian species if the sperm are injected! clirectly
into the oocytes (Thadani, 1980~. This experiment demon-
stratecl that fertilization can occur without the normal pro-
cesses of sperm activation, penetration of the zone pellu-
cida, and sperm-oocyte binding, processes that are sometimes
defective in infertile men, it presaged the now widely used
technique of intracytoplasmic sperm injection (ICSI) as a
treatment for male infertility.
Ever the adventurer, Markert was eager until the end of
his active research career to tackle the most clifficult and.
OCR for page 133
CLEMENT LAWRENCE MARKERT
133
as he saw it, the most important biological questions. After
retiring from Yale he finished his career as Distinguished
University Research Professor in Animal Science en cl Ge-
netics at North Carolina State University.
SERVICE TO SCIENCE
Markert believer! that scientists have an obligation to clo
their share of administrative work en cl to serve on volun-
teer boards en cl committees for the goocl of the scientific
enterprise. In aciclition to serving as chairman of the De-
partment of Biology at Yale (1965-71) he was director of
the Center for Reproductive Biology (1974-85~. He was man-
aging editor of The Jo urn a] of Experimen tad Zoology ~ ~ 963-
85) en cl coeditor (with John G. Scancialios) of Developmen-
ta] Genetics (1979-92~. He served terms as president of the
American Institute of Biological Sciences ~966), American
Society of Zoologists (1967), en cl the Society for Develop-
mental Biology ~ ~ 973-74) . Agencies en cl institutions that
benefited from Markert's advice as a board member included
the Bermuda Biological Station ~959-83), Presiclent's Bio-
meclical Research Panel (1975), American Cancer Society
(1976-78), Bioscience Information Service (1976-81 ), La Jolla
Cancer Research Fund (1977-86), National Research Coun-
cil ~ ~ 979-83), lane Coffin Fund for Meclical Research ~ ~ 979-
87), American Academy of Arts en c! Sciences (1981-84), en c!
the Fecleration of American Societies for Experimental Bi-
ology (1987-93~. As a member of the National Academy of
Sciences he server! on several committees en c! was elected!
to the Acaclemy's governing board, the Council.
MARKERT AS TEACHER AND MENTOR
Markert was a superb teacher whose lectures were leg-
endary among undergraduates. Like his research interests,
his lectures emphasizer! the big questions. He taught a course
OCR for page 134
34
B I O G RA P H I C A L
EMOIRS
at Yale entitles! "Biology of Reproduction" that covered, in
aciclition to the important biological principles, hot-button
issues of the time such as overpopulation en cl abortion rights.
The course also presenter! cutting-ecige reproductive tech-
nology, inclucling actual procluction of chimeric mice. Markert
returned to Yale for several years after his mandatory retire-
ment to give lectures in the course that he hac! pioneered.
For many of us who trained with Clem Markert, our
memories are as much about the culture of his laboratory
as about the science that was clone. Graduate students even
more so than postclocs were given free reign to choose their
own research topics en cl to pursue them more or less incle-
penclently, the only requirement being that any project
neeclecl to fit within the broacl scope of Markert's research
interests, which was certainly not clifficult. In the late 1960s,
for example, research in the Markert laboratory ranger! from
LDH isozymes in various species through maturation en cl
fertilization of frog and mouse oocytes to ribosomal gene
redundancy and the molecular biology of molluscan devel-
opment. Given the inclepenclence with which graduate stu-
dents pursued their research, Markert usually declined to
alit! his name to their publications, he clicI, however, re-
ceive explicit acknowlecigement for financial support of the
work and his mentorship. Despite his heavy administrative
responsibilities he was often available to talk with incliviclual
trainees without prior appointment and, unless he was trav-
eling, couIcl be expected to sit clown to a bag lunch with
laboratory members on a tinily basis. In aciclition to science,
the conversations often focused on history (the American
and Russian revolutions, for example), politics (a topic on
which Markert was never hesitant to share his views), en c!
even religion. An avowed atheist, Markert nonetheless was
knowlecigeable about en cl respected the beliefs of his train-
ees en c! colleagues. Whether his trainees agrees! with him
OCR for page 135
CLEMENT LAWRENCE MARKERT
135
or not they knew they were being mentorec! by a man of
superior intellect en cl strong social convictions who was willing
to put his life en cl career on the line for what he believecl
in. That, most of all, was Markert's legacy.
THIS MEMOIR COULD not have been compiled without the assistance of
former students and colleagues of Clem Markert. I am especially
indebted to Richard Elinson, Yoshio Masui, David Nanney, Robert
Petters, George Seidel, Vijay Thadani, and Gregory Whitt for pro-
viding documentation and commentaries pertaining to Markert's
life and career.
REFERENCES
Appella, E., and C. L. Markert. 1961. Dissociation of lactate dehy-
drogenase into subunits with guanidine hydrochloride. Biochem.
Biophys. Res. Comm. 6:171-76.
Duesbery, N. S., and G. F. Vande Woude. 2002. Developmental biol-
ogy: an arresting activity. Nature 416:804-805.
Heilbrunn, L.V., K. Daugherty, and K. M. Wilbur. 1939. Initiation
of maturation in the frog egg. Physiol. Zool. 12:97-100.
Hoppe, P. C., and K. Illmensee. 1977. Microsurgically produced
homozygous-diploid uniparental mice. Proc. Natl. Acad. Sci. U. S.
A 74:5657-61.
Hunter, R. L., and C. L. Markert. 1957. Histochemical demonstra-
tion of enzymes separated by zone electrophoresis in starch gels.
Science 125:1294-95.
Markert, C. L. 1963. Lactate dehydrogenase isozymes: Dissociation
and recombination of subunits. Science 140:1329-30.
Markert, C. L. 1968. Neoplasia: A disease of cell differentiation?
Canc. Res. 28:1908-14.
Markert, C. L., and F. M011er. 1959. Multiple forms of enzymes:
tissue, ontogenetic, and species specific patterns. Proc. Natl. A cad.
Sci. U. S. A. 45:753-63.
Markert, C. L., and R. M. Petters. 1977. Homozygous mouse em-
bryos produced by microsurgery. J. Exp. Zool. 201:295-302.
Markert, C. L., and R. M. Petters. 1978. Manufactured hexaparental
mice show that adults are derived from three embryonic cells.
Science 202:56-58.
OCR for page 136
136
B I O G RA P H I C A L
EMOIRS
Markert, C. L., T. B. Shaklee, and G. S. Whitt. 1975. Evolution of a
gene. Multiple genes for LDH isozymes provide a model of the
evolution of gene structure, function, and regulation. Science 189: 102-
14.
Masui, Y. 2001. From oocyte maturation to the in vitro cell cycle:
the history of discoveries of Maturation-Promoting Factor (MPF)
and Cytostatic Factor (CSF). Differentiation 69:1-17.
Masui, Y., and C. L. Markert. 1971. Cytoplasmic control of nuclear
behavior during meiotic maturation of frog oocytes. 7. Exp. Zool.
177:129-45.
McGrath, T., and D. Solter. 1984. Inability of mouse blastomere
nuclei transferred to enucleated zygotes to support development
in vitro. Science 226:1317-19.
Petters, R. M., and C. L. Markert. 1980. Production and reproduc-
tive performance of hexaparental and octaparental mice. 7. Hered.
71 :70-74.
Thadani, V. M. 1980. A study of hetero-specific sperm-egg interac-
tions in the rat, mouse, and deer mouse using in vitro fertiliza-
tion and sperm injection. 7. Exp. Zool. 212:435-53.
Wilmut, I., A. E. Schnieke, T. McWhir, A. T. Kind, and K. H. Campbell.
1997. Viable offspring derived from fetal and adult mammalian
cells. Nature 385:810-13.
OCR for page 137
CLEMENT LAWRENCE MARKERT
SELECTED BIBLIOGRAPHY
1948
137
The effects of thyroxine and anti-thyroid compounds on the synthe-
sis of pigment granules in chick melanoblasts cultured in vitro.
Physiol. Zool. 21: 309-27.
1950
The effects of genetic changes on tyrosinase activity in Glomerella.
Genetics 35: 60-75.
1956
With W. K. Silvers: The effects of genotype and cell environment on
melanoblast differentiation in the house mouse. Genetics 41:429-
50.
1957
With R. L. Hunter: Histochemical demonstration of enzymes sepa-
rated by zone electrophoresis in starch gels. Science 125:1294-95.
1959
With F. M011er: Multiple forms of enzymes: tissue, ontogenetic, and
species specific patterns. Proc. Natl. Acad. Sci. U. S. A. 45:753-63.
1961
With E. Appella: Dissociation of lactate dehydrogenase into sub-
units with guanidine hydrochloride. Biochem. Biophys. Res. Comm.
6:171-76.
1963
Lactate dehydrogenase isozymes: Dissociation and recombination
of subunits. Science 140: 1329-30.
1965
With I. Faulhaber: Lactate dehydrogenase isozyme patterns of fish.
J. Exp. Zool. 159:319-32.
OCR for page 138
38
B I O G RA P H I C A L
1966
EMOIRS
With W. J. Sladen: Stability of lactate dehydrogenase isozyme pat-
terns in penguins. Nature 210: 948-49.
1968
With E. J. Massaro: Lactate dehydrogenase isozymes: dissociation
and denaturation by dilution. Science 162:695-97.
1969
With R. S. Holmes: Immunochemical homologies among subunits
of trout lactate dehydrogenase isozymes. Proc. Natl. Acad. Sci. U.
S. A. 64:205-10.
1971
With H. Ursprung: Developmental Genetics. Englewood Cliffs, N. J .:
Prentice-Hall.
With Y. Masui: Cytoplasmic control of nuclear behavior during mei-
otic maturation of frog oocytes. J. Exp. Zool. 177:129-45.
1975
With J. B. Shaklee and G. S. Whitt: Evolution of a gene. Multiple
genes for LDH isozymes provide a model of the evolution of
gene structure, function, and regulation. Science 189:102-14.
1977
With R. M. Petters: Homozygous mouse embryos produced by mi-
crosurgery. J. Exp. Zool. 201 :295-302.
1978
With R. M. Petters: Manufactured hexaparental mice show that adults
are derived from three embryonic cells. Science 202:56-58.
1979
With K. I. Yamamura and Z. I. Ogita: Epigenetic formation of lac-
tate dehydrogenase isozymes in the house mouse, Mus musculus.
J. Exp. Zool. 208:271-80.
With G. L. Hammond and E. Wieben: Molecular signals for initiat-
ing protein synthesis in organ hypertrophy. Proc. Natl. Acad. Sci.
U. S. A. 76:2455-59.
OCR for page 139
CLEMENT LAWRENCE MARKERT
1980
139
With T. Y. Lu: Manufacture of diploid/tetraploid chimeric mice.
Proc. Natl. Acad. Sci. U. S. A. 77:6012-16.
1982
With G. L. Hammond and Y. K. Lai: The molecules that initiate
cardiac hypertrophy are not species-specific. Science 216:529-31.
1983
Fertilization of mammalian eggs by sperm injection. J. Exp. Zool.
228:195-201.
1986
With C. Anderegg: Successful rescue of microsurgically produced
homozygous uniparental mouse embryos via production of ag-
gregation chimeras. Proc. Natl. Acad. Sci. U. S. A. 83:6509-13.
1990
With K. Momoi and E. Goldberg: Expression of the human lactate
dehydrogenase-C gene in transgenic mice. Prog. Clin. Biol. Res.
344:441-52.
1993
With K. Salehi-Ashtiani, R. J. Widrow, and E. Goldberg: Testis-spe-
cific expression of a metallothionein I-driven transgene corre-
lates with undermethylation of the locus in testicular DNA. Proc.
Natl. A cad. Sci. U. S. A. 90:8886-90.
1998
With T. M. Amet and E. Goldberg: Human testis-specific lactate
dehydrogenase-C promoter drives overexpression of mouse lac-
tate dehydrogenase-1 cDNA in testes of transgenic mice. J. Exp.
Zool. 282:171-78.
Representative terms from entire chapter:
lawrence markert
