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Just, Ernest Everett 1883–1941

Ernest Everett Just 18831941


At a Glance

Won First Spingarn Medal From NAACP

Obtained His Ph.D. in Chicago

Established Professional Career

Encountered Prejudice at MBL

Europe Offered New Horizons

Sought a New World View

Returned to Howard Despite Poor Health

Selected writings


A brilliant marine biologist who made numerous significant discoveries concerning cell behavior, Ernest Everett Just faced many obstacles in his career because of racial prejudice. He overcame many of them, but they left their mark on him. Finding European scientists more humanitarian than their American counterparts, he went into a self-imposed exile in Europe during the last decade of his life.

From the time he was a young adult to the end of his life, Just was an outstanding student and academician. Born in 1883, he grew up in Charleston, South Carolina, and attended an all-black public elementary school. His father died when he was four years old, and his mother worked as a teacher to support her family. Just helped out by working in the fields after school.

Just and his mother agreed that he would receive a better education in the northern United States. After finishing public school in Charleston, he went to New York City at the age of 17 in order to earn enough money to attend the Kimball Academy in Meriden, New Hampshire. He was financially able to enroll in the school after only four weeks of working in the city. Eventually graduating with honors in 1903 at the age of 19, he took four years of classes in three and served as the editor of the school newspaper and president of the debating society.

Just went on to Dartmouth University, where he was the only black student in a class of 287. In his sophomore year, he was introduced to science as a new field of interest. He studied botany under Professor Lyman and biology under Professor William Patten, chairman of the biology department. Inspired by Patten, who was an esteemed scientist specializing in evolution, Just decided to major in science with a concentration in biology. By the time he graduated from Dartmouth, Just had taken all of the biology courses offered there.

In his junior year at Dartmouth, Just took three classes from the noted biologist and zoologist John H. Gerould, who had done several years of research at marine laboratories in Italy and France, countries in which Just himself would later work. By his senior year, Just was performing brilliantly. His superior grades earned him recognition as a Rufus Choate Scholar for the second year in a row, and he also received the Grimes Prize for general improvement.

Just earned his A.B. degree in 1907 and was elected to the

At a Glance

Born August 14, 1883, in Charleston, SC; son of Charles Fraser and Mary Just; died of cancer, October 27, 1941, in Washington, DC; married Ethel Highwarden (a high school teacher), June 26, 1912 (divorced, 1939); married Hedwig Schnetzler, August 11, 1939; children: (first marriage) Margaret, Highwarden, Maribel; (second marriage) Elisabeth. Education: Dartmouth College, A.B. (magna cum laude), 1907; graduate studies at Marine Biological Laboratory, Woods Hole, MA, 1909-15; University of Chicago, Ph.D., 1916.

Howard University, Washington DC, 1908-40, began as assistant professor, became professor of physiology and biology. Conducted research at the Marine Biological Laboratory, Woods Hole, MA, during summers from 1909-29, and at various European laboratories, including Stazione Zoologica, Naples, Italy, 1929; Kaiser Wilhelm Institut in Berlin, Germany, 1930 and 1931; and Station Biologique near Roscoff, France, 1939-40.

Awards: Spingarn Medal, National Association for the Advancement of Colored People (NAACP), 1915; grants from Julius Rosenwald Foundation, 1920 and 1928.

Member: American Society of Naturalists, American Society of Zoologists, American Association for the Advancement of Science, American Ecological Society, Societe nationale des sciences naturelles et mathematiques de Cherbourg.

honorary fraternity Phi Beta Kappa. He received special honors in zoology and history and was the only magna cum laude graduate in his class. In spite of his excellent academic performance, Just found the predominantly white professional world closed to him. Consequently, he chose to pursue a career in academia and accepted a position teaching English and rhetoric at Howard University in Washington, D.C. Eventually teaching more advanced English courses, his starting salary in 1907 was $400 per year.

In the summer of 1909, Just began his graduate training in biology at the Marine Biological Laboratory (MBL) in Woods Hole, Massachusetts. He had contacted one of his former teachers at Dartmouth, William Patten, about possible courses of graduate study in science. Patten referred him to Frank R. Lillie, head of the Department of Zoology at the University of Chicago and director of the MBL at Woods Hole. Lillie would become Justs mentor, and Just would become Lillies devoted student.

During his first summer at Woods Hole Just took invertebrate zoology, studying marine invertebrates. The next summer he studied embryology, and in 1911 Lillie arranged for Just to enroll in the University of Chicagos doctorate program in absentia by taking courses at the MBL. Just would only need to establish one year of residency in Chicago before formally applying for his Ph.D.

Justs duties at Howard continued to increase during the regular school year. In 1910 the university opened a new science building, and Just was able to join the newly formed science faculty. He taught zoology and completed the switch from teaching English to science by acquiring the title of assistant professor of biology. To Howards scientific curricula he added an advanced course on animal histology that focused on cell structure.

According to Kenneth R. Manning in Black Apollo of Science: The Life of Ernest Everett Just, by 1911 Just was well on his way to instituting a rigorous and innovative academic program, and to establishing himself as a first-rate teacher of science. He was appointed associate professor at a salary of $1,500 per year. By 1912, he had extended his work to include the professional training of doctors. He became professor of physiology in the medical school as well as professor of biology in the regular college, earning a combined annual salary of $2,150.

Just was popular with the students at Howard. His extracurricular activities included organizing Howards Drama Club, which produced several plays each year. Manning observed that Just became known on campus as an energetic young man willing to offer advice and guidance to students. In 1912, he married Ethel Highwarden, a schoolteacher he had met in Washington, D.C.

Continuing to spend his summers at Woods Hole, Just was encouraged by Lillie to undertake scientific research. Lillie put Just to work on the problem of cell cleavage, or divisions, in the eggs of nereis, a type of sandworm, and arbacia. His research resulted in his first scientific publication, 1912s The Relation of the First Cleavage Plane to the Entrance Point of the Sperm, and it was the beginning of Justs lifelong interest in the study of marine eggs.

In his first paper, Just demonstrated that the sperm entrance point is critical in determining the line of cleavage of the egg. The importance of his discovery was generally recognized at the time, and years later the renowned geneticist and Nobel laureate T. H. Morgan would cite it as the fundamental and authoritative study on the subject, according to Manning. Just eventually became the foremost authority on the embryological resources of marine animals.

Won First Spingarn Medal From NAACP

In 1913 the National Association for the Advancement of Colored People (NAACP) initiated plans for an award to go to a person of African descent who had performed the foremost service to his race. The 30 nominees for the first Spingarn Medal reflected a predominant interest in art, politics, social work, business, literature, education, and athleticsthe fields in which black achievement had traditionally been concentrated, commented Manning.

The NAACP, Manning continued, also wanted science to be represented, feeling it would help place black people and the NAACP in particular in the mainstream of Western tradition. They asked Jacques Loeb, a noted biologist at the Rockefeller Institute for Medical Research, to recommend a science candidate. Loeb nominated Just, whom he had met at Woods Hole in 1912, for the important scientific research he had accomplished. Just was subsequently awarded the first Spingam Medal, and it was presented to him on February 12, 1915, by New Yorks governor, Charles Whitman. The award and its accompanying publicity made Just known to the American public.

Obtained His Ph.D. in Chicago

In June of 1915 Lillie decided that Just had completed the research requirements for his Ph.D. and agreed to accept previously published articles as a doctoral thesis. Just only needed to fulfill a few minor subject requirements and the residence requirement, which necessitated a leave of absence with pay from Howard University. According to Manning, Lillie persuaded the administration that the reputation of faculty members was an important consideration for any medical school, and university officials agreed to make sacrifices so Just could obtain his Ph.D.

Spending the 1915 to 1916 school year at the University of Chicago, Just took basic physiology classes to satisfy his minor subject requirements. He also registered for A. P. Mathewss advanced physiology course. This important course involved a detailed analysis of the chemistry of cell constituents, and, wrote Manning, it laid the foundation for Justs future work on the protoplasmic systems of marine invertebrates.

Just took courses in the fall, winter, and spring terms at the University of Chicago. At the end of the spring quarter, the department agreed to accept two of Justs articles as the main text for his doctoral thesis, which was titled Studies of Fertilization in Platynereis megalops. He received his diploma in June and took a two-month rest instead of going to Woods Hole that summer.

Established Professional Career

With his Ph.D. in hand, Just took his place among the scientists of the MBL at Woods Hole. By 1920 he was elected to several scientific societies, including the American Society of Naturalists, the American Society of Zoologists, the American Association for the Advancement of Science, the American Ecological Society, and the Societe nationale des sciences naturelles et mathematiques de Cherbourg.

In 1919 and 1920 Just published four articles in the Biological Bulletin based on four years of research on fertilization. His findings challenged Loebs previous work on artificial parthenogenesisor unfertilized reproductionand defended the theories of fertilization put forth by his mentor, Dr. Lillie. His published work on the fertilization reaction in Echinarachnius parma established Just as an outstanding scientist. Lillie asked him to contribute to the section on fertilization in the textbook General Cytology, by E. V. Cowdry.

Encountered Prejudice at MBL

Justs role at the MBL became more prominent. In 1922 he gave a series of guest lectures in the embryology course and taught other investigators his own techniques and skills for handling marine invertebrates. Lillie wanted to set him up with a permanent position at MBL, but he knew relations between Just and some of the MBL community were strained as a result of racial prejudice. In one instance, Just was asked not to attend a social gathering celebrating a new clubhouse for the MBL Club. While he had a circle of friends that helped protect him from prejudice at Woods Hole, he was left with feelings of bitterness from the incidents he had experienced.

Justs important research articles eventually brought him to the attention of officials from various science foundations and won financial support for both his own work and Howard University. He knew Julius Rosenwald personally and in 1920 received a Rosenwald grant that lasted for seven years. In 1927, Just met Edwin R. Embree, who was leaving the Rockefeller Foundation to assume the presidency of the Julius Rosenwald Fund. The Rosenwald Fund was expanding its operations, and its assets had dramatically increased. Just initially proposed funding to build a graduate department and laboratory at Howard University, but Embree preferred to fund Justs work as an individual scientist.

In 1928 Embree prepared a proposal for Just, and with the help of a glowing evaluation from Ralph Lillie, the brother of Justs mentor, Frank Lillie, the Rosenwald trustees approved a major resolution in favor of Just. He received an $80,000 grant for a five-year period that would enable him to work and travel in Europe as well as provide support for Howards biology department.

Europe Offered New Horizons

Accompanied by his daughter Margaret, Just boarded the Dresden in January of 1929 for a trip to Europe. After spending a week in Paris, France, the two went to Naples, Italy, where Just would research European species of marine invertebrates at the Stazione Zoologica. Freed from the demands of students and young investigators, Just enjoyed spending the majority of his time on research. The purpose of his studies at the Stazione Zoologica was to prove his hypothesis that the European seaworm Mereis dumerilii and the American seaworm Platynereis megalops were not the same species. He also repeated his Woods Hole experiments in fertilization, supporting Lillie by discovering that fertilization capacity and the level of the glutinous substance Lillie called fertilizin were directly proportional to each other in certain European species as well, according to Manning in Black Apollo of Science.

After his stay in Italy, Just developed a strong kinship with European scientists and acted as an ambassador on their behalf to the American science community. He found, however, that most American scientists were not concerned with their European counterparts. He returned to Europe in 1930 to work at the Kaiser Wilhelm Institut in Berlin, Germany, where he studied a new species, Amoeba proteus, a unicellular freshwater organism, to test his theory that cortical, or ectoplasmic, behaviorthe function of the outer layer of an organism or cellwas the key to all problems in biology.

Just returned to the United States for a short time in 1930 to attend the celebration of Frank Lillies 60th birthday at Woods Hole. He spoke on behalf of Lillies work on the problems of fertilization in his formal presentation. Then, stepping down from the podium, he announced, as quoted by Manning, I have received more in the way of fraternity and assistance in my one year at the Kaiser Wilhelm Institut than in all my other years at Woods Hole put together. When Just left, he never returned to Woods Hole, conducting the rest of his research in Europe.

Just was under pressure in 1930 and 1931 to reorganize the zoology department at Howard University. He took a personal leave and spent his own money to accept an invitation to be a keynote speaker at the International Congress on Zoology in Padua, Italy. After submitting his report on the zoology department, Just returned to the Kaiser Wilhelm Institut in the spring of 1931. His social circle there included Margaret Boveri, whom he had met in Naples, and the former crown prince of Germany, the son of Kaiser Wilhelm II.

The 1930s were a decade of trauma and excitement for Just. After the summer of 1931, he made seven trips to Europe. He courted and in 1939 married Hedwig Schnetzler, a philosophy student he had met in Berlin. His first wife, Ethel, whom he divorced, and their children faded out of his life. Many of his childhood dreams resurfaced; his underlying awareness of his German ancestry, his religious intensity, his penchant for fantasy, noted Manning. Though approaching middle age, he was not too old to change. Europe made him a different person.

Sought a New World View

Europe had opened up [Justs] science, had given him the confidence to take daring steps in his scientific work, wrote Manning. He moved away from experimental details to larger philosophical views; he searched for a new world view through some grand alliance of the various fields of science, history, and philosophy. By 1938 Just was almost penniless when he began a self-imposed exile in Europe. The scientist went to France, where he and Hedwig worked at the Station Biologique in the Finistere district and completed what Just considered his crowning achievement, the book The Biology of the Cell Surface.

In October of 1939 the Station Biologique received orders from the French government to close its facilities to foreigners; Just and Hedwig decided to stay in the French seaport town of Roscoff anyway. Just wrote an article, Unsolved Problems of General Biology, for the periodical Physiological Zoology, in which he expressed what he felt was the most pressing need in biology: to find what in any protoplasmic system, cell or otherwise, is the living substance. A reflective piece on biologys unanswered questions, the article featured Justs synthesized research findings, suggested fruitful paths for future research, and called for collaboration among scientists from different disciplineschemists, biologists, and physiciststo solve the dilemma of how life begins, how it is continued, and how transmitted. It was his last full-length paper, and in it he justified his lifes work, writing that in the final analysis, chemistry and physics were dependent on biology to establish, beyond question, criteria of normality, the range of normal processes, and the extent of normal variability.

Returned to Howard Despite Poor Health

Just and his wife had difficulty leaving France. In August of 1940 the German Nazis interned Just in a camp, but he was released through the influence of Hedwigs father. With Hedwig several months pregnant, the couple made their way out of France through Spain and Portugal, boarding the S.S. Excambion in Lisbon, Portugal. Just had to convince the reluctant ships captain that Hedwig would be able to manage the voyage in her condition.

Despite his deteriorating health, Just went back to teaching at Howard University. Hedwig gave birth to their daughter, who was named Elisabeth after her mother. Tired and thin, Just suffered severe stomach trouble in February of 1941. He continued his work and was preparing his final manuscript, Ethics and the Struggle for Existence, which promised to be a synthesis of biology, history, sociology, and anthropology. His illness, though, made it difficult for him to concentrate, and the resulting paper was fragmentary.

Just and Hedwig planned to spend the summer of 1941 by the seaside in Portland, Maine, but Just became so ill in early July that he had to return to New York City, where he was admitted to Mount Sinai Hospital. It turned out that the trouble he was having with spinal nerves was a symptom of a more serious problem, pancreatic cancer. After a decorated career in academia marked by numerous significant contributions to the field of biology, Just succumbed to cancer on October 27, 1941.

Selected writings


Basic Methods for Experiments on Eggs of Marine Animals, Blakistons, 1939.

The Biology of the Cell Surface, Blakistons, 1939.


The Relation of the First Cleavage Plane to the Entrance Point of the Sperm, Biological Bulletin,1912.

The Fertilization Reaction in Echinarachnius parma, Parts I-IV, Biological Bulletin,1919-20.

On the orgin of Mutations, American Naturalist, 1932.

Unsolved Problems of General Biology, Physiological Zoology,1940.


American Black Scientists and Inventors, National Science Teachers Association, 1975.

Haber, Louis, Black Pioneers of Science and Invention, Harcourt, Brace & World, 1970.

Manning, Kenneth R., Black Apollo of Science: The Life of Ernest Everett Just, Oxford University Press, 1983.

David Bianco

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Just, Ernest Everett


(b. Charleston, South Carolina, 14 August 1883; d. Washington, D.C., 27 October 1941),

embryology, developmental biology, cell biology.

Best known for his discovery of the “wave of negativity” that sweeps over the surface of the marine invertebrate egg upon fertilization, a wave that correlates with what has become known as the “fast block to polyspermy,” Just more generally showed that the egg cell surface plays an important role in fertilization and development. He was the first to associate cell surface changes with stages of embryonic development experimentally.

As an African American in early-to-mid-twentieth-century America, Just lived and worked in a social and cultural milieu that was often hostile to his research efforts. He succeeded in obtaining a prestigious research fellowship from philanthropist Julius Rosenwald through the National Research Council in the middle of his career (1920–1930). Later on, however, he had considerable difficulty in obtaining financial support and spent much of his time asking for help from potential donors, including various American foundations, wealthy individuals—and even Italian dictator Benito Mussolini. Despite these potentially crippling challenges, Just made lasting contributions to biology.

Formative Years . Just was the son Charles Frazier Just Jr. and Mary Matthews Just. In 1887, after Charles’s death, the family moved to James Island, just off the coast of

Charleston, South Carolina. In his early years, young Ernest was educated mainly by his mother, a strong-willed, independent woman who established the first school and church on the island. At age twelve, he left James Island to attend the Colored Normal Industrial Agricultural and Mechanics College at Orangeburg (now South Carolina State College), and at fifteen he left the South for New England. He entered Kimball Union Academy in Meriden, New Hampshire, at age seventeen and then went to Dartmouth College in Hanover, New Hampshire, where he studied biology, history, literature, and the classics. He graduated magna cum laude from Dartmouth in 1907 as an esteemed Rufus Choate scholar.

In the autumn of 1907, Just accepted a faculty position at Howard University, an African American school in Washington, D.C. His initial appointment was in English, but in 1910 he moved to the Biology Department and soon became the first head of the Department of Zoology. With funding from the Rosenwald Fund, he established a master’s program in zoology. In 1915 Just was chosen from among a group of distinguished nominees to receive the first NAACP (National Association for the Advancement of Colored People) Spingarn Medal for his research excellence and his promotion of medical education at Howard. Despite a sometimes rocky relationship with the university administration and many trips to Europe, he remained a Howard faculty member until his death in 1941.

Woods Hole . In 1909 Just began making summer excursions to the Marine Biological Laboratory (MBL) in Woods Hole, Massachusetts, where he worked as an assistant to its director, the eminent embryologist Frank R. Lillie of the University of Chicago. Under Lillie’s supervision, Just received his PhD in experimental embryology in 1916. His early studies at Woods Hole on the fertilization of the marine annelid Platynereis megalops formed the basis of his PhD thesis.

At Woods Hole, using a light microscope, Just recorded in detail the changes that take place within the egg during fertilization. In one particularly innovative study, Just exposed eggs of the sand dollar Echinarachnius parma to dilute seawater at various precisely timed intervals after insemination, and he carefully measured the position of membrane elevation relative to the point of sperm contact along with the time it took for each egg to rupture at this position. He found that the fertilization envelope forms as the result of a wave of structural instability that moves from the point of sperm entry to the opposite side. It has since become known that this wave of instability is actually a wave of cortical granule exocytosis that forms the fertilization envelope. As early as 1919, Just had observed that a “wave of negativity” moves from the point of sperm contact around the surface to the egg’s opposite side. He noticed that the wave was associated with an immediate block to polyspermy (aberrant fertilization by multiple sperm), a block that preceded the liftoff of the fertilization membrane. He also observed the slow, mechanical block that occurred after membrane separation. Together, these two phenomena constitute what became known as the fast and slow blocks to polyspermy.

While at Woods Hole, Just studied the effects of a range of conditions—hypotonic and hypertonic seawater, temperature, degree of hydration, and ultraviolet irradiation, for example—on both normal development and parthenogenesis (egg activation without sperm) in a number of marine animals, including E. parma, the parchment worm Chaetopterus pergamentaceus, the clam worm Nereis limbata, and the sea urchin Arbacia punctulata. He also performed experiments that demonstrated the validity of Lillie’s “fertilizin” hypothesis of fertilization. As early as 1906, Lillie had proposed that eggs release a diffusible substance (fertilizin) that, when it contacts spermatozoa, causes them to agglutinate. Fertilizin was hypothesized to have two ends, one that interacted with a receptor on the egg, and another that interacted with a receptor on the sperm. In this way, the fertilizin molecule allowed sperm and egg to come together. Lillie’s hypothesis was in conflict with a rival idea of Jacques Loeb’s known as the lysin hypothesis. Based on Loeb’s observations of experimental parthenogenesis (see below), the hypothesis proposed that a cytolytic factor (lysin) in the sperm activates the egg, thereby initiating development. Throughout his career, Just sought experimental support for Lillie’s hypothesis, proving that it was valid for a number of marine animals. From 1919 onward, in published papers and at scientific conferences, Just mounted a sustained attack on Loeb’s rival hypothesis as well as on his work on experimental parthenogenesis. In 1930 Just wrote a long defense of the fertilizin hypothesis.

In the course of his work, Just discovered the important effects that environmental factors have on embryonic development. Based on these discoveries, he came to believe strongly that in order for laboratory experiments to be valid, the conditions in the laboratory must match as closely as possible those in nature. In marine settings at the MBL and, later on, at marine stations in Italy and France, he observed and carefully recorded the breeding behavior of the animals whose eggs he studied. Through his intimate knowledge of marine invertebrate natural history, and by testing the effects of different variables on development, he was able to formulate specific indices of normal development for a range of species. These indices, based for the most part on when and under what conditions fertilization envelope separation occurs, allowed him to predict with a high degree of certainty whether or not a particular egg, upon fertilization, would develop normally. So deep was Just’s knowledge of marine invertebrate natural history, and so great was his ability to coax marine embryos to develop normally when other researchers failed, that he was often consulted on matters related to the proper handling of marine eggs. This led him to write and publish his first book, Basic Methods for Experiments on Eggs of Marine Animals (1939). He was widely regarded at Woods Hole and beyond as a brilliant experimental embryologist and a leading expert on embryo handling and culture.

Europe . Just’s first European trip was to the Stazione Zoologica in Naples, Italy, in 1929, where for six months he studied cortical reactions in sea urchin eggs in order to test Lillie’s fertilizin hypothesis further. Following up on an idea he had conceived earlier, he also showed that P. megalops of Woods Hole was not the same species as the Mediterranean annelid Nereis dumerilii. Just’s second trip came only a year after his first. He received an invitation, exceedingly rare for an American at that time, to visit the Kaiser Wilhelm Institut für Biologie in Berlin for six months beginning in January 1930. While there, Just met and became friends with such eminent German embryologists as Max Hartmann (who had invited Just), Otto Mangold, and Richard Goldschmidt. He became particularly close to Johannes Holtfreter, Mangold’s assistant, who in subsequent years published groundbreaking papers on the role of the cell surface in adhesion during amphibian embryo morphogenesis. Holtfreter’s papers heavily cited Just’s second book The Biology of the Cell Surface (1939) but, curiously, Just’s work, though groundbreaking in its own right, was not cited much after that. After World War II, his work was almost completely forgotten.

Altogether, from his first trip in 1929 to his last in 1938, Just made nine visits to Europe to pursue research interests. Some of these were to Berlin, some were to Naples, and some were to Paris and Roscoff, a French village on the English Channel that had a small marine biological station. Although he had conceived of it earlier (around 1931), while at the Stazione Zoologica in Naples in 1934 Just began to work in earnest on The Biology of the Cell Surface, which was to be a fusion of his scientific work and his philosophical views on biology.

When at Woods Hole, Just had gained international recognition for the quality of his experimental technique and for his significant discoveries regarding the fertilization process, but he had not had the opportunity to explore more general theories about biology. At the Kaiser Wilhelm Institut für Biologie, however, he was encouraged to develop his ideas about the importance of the cell surface (the ectoplasm) and extend them to species other than marine invertebrates, such as amoebae. Emboldened by his European experience, Just underwent a transformation: he began to display increasing confidence in tackling the larger problems of biology. After 1936, except for one paper of medical significance and three papers on his work on fertilization of marine animals at Roscoff, Just’s writings turned strongly in a philosophical direction. His papers had bold titles such as “A Single Theory for the Physiology of Development and Genetics” (1936) and “Phenomena of Embryogenesis and Their Significance for a Theory of Development and Heredity” (1937).

Philosophical Leanings . It is likely that Just’s first paper, “The Relation of the First Cleavage Plane to the Entrance Point of the Sperm” (1912), in which he showed that the first cleavage plane of the fertilized Nereis egg is determined by the point of sperm entry, planted the seed of his later view that the organism is a holistic system having properties that emerge out of its complexity and organization. Just reasoned that if the plane of first cleavage is not preformed, but is contingent on the sperm entry point, then development must be an epigenetic rather than a preformationistic process. This holistic view, which considers living systems to be more than the sum of their parts, is becoming more prevalent at the beginning of the twenty-first century as biologists increasingly embrace systems biology. It was also common among embryologists at the beginning of the twentieth century. What set Just apart from his contemporaries, however, was his willingness to articulate openly the embryologist’s view in the face of the opposing reductionistic views of such prominent biologists as Loeb and Thomas Hunt Morgan.

By 1900 Loeb had discovered that he could partheno-genetically activate sea urchin and marine annelid eggs through an experimental two-step method involving treatment with butyric acid followed by hypertonic seawater. This became known as Loeb’s “superficial-cytolysis-corrective-factor” method. In papers, at scientific conferences, and in The Biology of the Cell Surface, Just challenged Loeb’s method. He demonstrated not only that the order of treatment—butyric acid, then hypertonic seawater—was inconsequential, but that only the butyric acid was required to activate the egg. He showed that Loeb’s conclusions were based on a mishandling of the eggs and that the cytolytic effect of the butyric acid was due to overexposure of the eggs to the acid. In sharp contrast to Loeb, a staunch mechanist who believed that he had created life through chemical means, Just believed that experimental parthenogenesis (and fertilization) showed the egg to be a dynamic system poised to respond to external agents of various kinds. It was not a passive physico-chemical object upon which external agents acted to “cause” the development that ensued. Parthenogenesis and fertilization revealed what he called the independent irritability of the egg cell surface, its ability to respond to external stimuli. Thus, for Just, it was the cell surface and the structured layer just below it, the ectoplasm, that mediated fertilization and all subsequent developmental events. Likewise, it was the cytoplasmic surface, which was in contact with the environment, that was critical in the evolution of the first living thing, the ancestor of all life.

Just’s focus on the cell cytoplasm (ectoplasm), as opposed to genes in the nucleus, led him to challenge Nobel laureate Morgan at the 1935 meeting of the American Society of Zoologists in Princeton, New Jersey. Morgan had proposed the gene theory, which postulated that genes located in linear arrays on chromosomes in the nucleus were the units of inheritance. Morgan believed that the purpose of the cytoplasm was only to execute the orders of genes in the nucleus. In contrast, Just believed that genes and chromosomes played secondary roles, and that the units of inheritance were located in the cytoplasm, not the nucleus.

At the Princeton meeting, Just presented his own theory of genetic restriction in opposition to Morgan’s theory. The theory was founded on Just’s detailed experiments showing that the cell nuclei increased in mass during the cleavage process that follows fertilization. It was an attempt to explain how a single-celled fertilized egg could differentiate into a multicellular organism; it proposed that the nuclei, and the genes within them, act only to remove selectively obstacles from the cytoplasm so that the “activity of the cytoplasm” can be released in one direction as opposed to another. This would lead to differentiation of cells into different types. Thus, although Morgan’s purely nucleocentric view was at one extreme, Just’s purely cytoplasmic view was clearly at the other. Morgan’s gene theory was incorrect in attributing almost unlimited power to genes, but Just’s genetic restriction theory was even more off the mark. Both were too onesided, and neither succeeded in bridging the divide that separated the new geneticists (represented by Morgan) and the traditional embryologists (such as Just). Indeed, it is only at the beginning of the twenty-first century, with the rise of epigenetics and systems biology, that these two perspectives on development—the nucleocentric and the cytoplasmic—are becoming reconciled. The genecentric view, which has dominated biology in the latter half of the twentieth century, is giving way to a recognition of the importance of the gene in context.

Endings . In 1938 Just initiated a self-imposed exile in Europe and began working at the Station Biologique at Roscoff. In May 1940, however, Nazi Germany invaded France. All foreigners were ordered to leave the country, but Just decided to stay behind to finish writing his paper, “Unsolved Problems in General Biology” (1940). In June the Nazis seized control of Paris and the surrounding countryside, including Roscoff. Just was briefly imprisoned, but he was released with the help of friends. Although he had expected to reside permanently in Europe following his exile attempt, he was forced to return to the United States. In October 1941, having become gravely ill with pancreatic cancer, he died. He was fifty-eight years old.

Just was, by all accounts, a brilliant experimental embryologist who made important contributions to our understanding of fertilization and development. His experiments on the importance of the cell surface during development influenced others such as Holtfreter, who extended Just’s work. His holistic view of the embryo is increasingly relevant as the organism is more and more viewed as an integrated system with properties that emerge from its complexity and organization. Just made these contributions despite the racist social and cultural milieu of his time, a milieu that influenced how his work was perceived as well as his ability to secure research support. The words of Just’s friend and mentor, Lillie, expressed in his obituary of Just in the journal Science, are particularly poignant:

An element of tragedy ran through all Just’s scientific career due to the limitations imposed by being a Negro in America, to which he could make no lasting psychological adjustment in spite of earnest efforts on his part.… In Europe he was received with universal kindness, and made to feel at home in every way; he did not experience social discrimination on account of his race, and this contributed greatly to his happiness there. Hence, at least in part, his prolonged self-imposed exile on many occasions. That a man of his ability, scientific devotion, and of such strong personal loyalties as he gave and received, should have been warped in the land of his birth must remain a matter for regret. (1942, p. 11)



“The Relation of the First Cleavage Plane to the Entrance Point of the Sperm.” Biological Bulletin 22 (1912): 239–252.

With Frank R. Lillie. “Breeding Habits of the Heteronereis Form of Nereis limbata at Woods Hole, Mass.” Biological Bulletin 27 (1913): 147–169.

“The Present Status of the Fertilizin Theory of Fertilization.” Protoplasma 10 (1930): 300–342.

“A Single Theory for the Physiology of Development and Genetics.” American Naturalist Supplement 70 (1936): 267–312.

“Phenomena of Embryogenesis and Their Significance for a Theory of Development and Heredity.” American Naturalist 71 (1937): 97–112.

Basic Methods for Experiments on Eggs of Marine Animals. Philadelphia: Blakiston’s Son, 1939.

The Biology of the Cell Surface. New York: Garland Publishing, 1939.

“Unsolved Problems in General Biology.” Physiological Zoology 13 (1940): 23–142.


Byrnes, W. Malcolm, and William R. Eckberg. “Ernest Everett Just (1883–1941)—An Early Ecological Developmental Biologist.” Developmental Biology 296 (2006): 1–11. Highlights Just’s appreciation of the role of environmental factors in development and ties this in with the emerging field of ecological developmental biology.

Gilbert, Scott F. “Cellular Politics: Ernest Everett Just, Richard B. Goldschmidt, and the Attempt to Reconcile Embryology and Genetics.” In The American Development of Biology, edited by Ronald Rainger, Keith R. Benson, and Jane Maienschein. Philadelphia: University of Pennsylvania Press, 1988. Compares and contrasts Just with Goldschmidt, arguing that Just had a more egalitarian, decentralized view of the cell, while Goldschmidt had a more imperialistic view, with the nucleus controlling everything.

Gould, Stephen J. “Just in the Middle: A Solution to the Mechanist-Vitalist Controversy.” In The Flamingo’s Smile: Reflections in Natural History. New York: Norton, 1985. Makes the argument that Just’s organicist views were “in the middle” between those of vitalism on the one hand and mechanism on the other.

Lillie, Frank R. “Obituary of Ernest Everett Just.” Science 95 (1942): 11.

Manning, Kenneth R. Black Apollo of Science: The Life of Ernest Everett Just. New York: Oxford University Press, 1983. An excellent, well-researched biography of Just.

W. Malcolm Byrnes

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Ernest Just

Ernest Just

Ernest Just (1883-1941) was a prominent African American biologist who was noted for his contributions to marine biology.

Ernest Everett Just was born on August 14, 1883, in Charleston, South Carolina. He had a rough childhood; his father died when he was very young, leaving his mother to fend for herself and her family. He was educated by his mother until the age of 13, when he entered the Colored Normal, Industrial, Agricultural and Mechanical College. He received a Licentiate of Instruction that permitted him to teach in the black public schools of South Carolina.

The prospects for advancement in the school system were small, however, and Just wanted to continue his education. He thus obtained a scholarship to attend the Kimball Union Academy in Vermont, where he was the only African American in a group of 170 students. There he received a broad education, and he was able to continue his schooling at Dartmouth, where he won a degree in biology with a minor in history in 1907. Just soon obtained a teaching position at Howard University, the most prestigious African American institution in the country at the time. He helped to develop the science curriculum there, largely by teaching zoology and setting up laboratories.

Determined to pursue the same type of career that a white man would have in science, Just started spending his summers as research assistant at the famed Marine Biology Laboratory in Woods Hole, Massachusetts. There he worked under Frank Lillie, the head of the Zoology Department of the University of Chicago. Lillie helped him enroll "in absentia" in the Ph.D. program at the University of Chicago, and after considerable delay, due mostly to his teaching responsibilities at Howard, he received his Ph.D. in 1916, becoming one of the first African Americans to do so.

Throughout his life Just had problems obtaining funding for his work, partly because of racial discrimination and partly due to the general lack of funding for science during the Great Depression of the 1930s. He continued to teach on and off at Howard for the remainder of his life. He also went to Europe several times, having achieved greater recognition there than in his own country. Just preferred Europe to the United States. In 1938 he went to France with the intention of staying for good. In the middle of 1940, however, the Nazis drove him out, and he was forced to return to Washington, D.C., where he died on October 27, 1941.

During the earlier stages of his career Just was primarily concerned with collecting a mass of verifiable data on marine eggs. He concentrated on some of the fundamental problems of cell biology, and in particular dealt with the problem of parthenogenesis, or the ability of certain types of eggs to reproduce without sperm. He developed a precise and much respected style of experimentation and was considered an authority on experimentation with marine invertebrates. He wrote a number of well-received articles on experimental methods that was later collected in a volume entitled Basic Methods for Experiments on Eggs of Marine Animals (1939).

Once Just became an established scientist with a worldwide reputation he started concentrating on a theory that had been brewing in his mind for several years. He theorized that the outer part of the cell (the cortical cytoplasm) was more significant in vital life processes than had been previously recognized. He thought, in fact, that it was a crucial link between an animal and its environment. One way he used to show this was by showing that the fertilization of an egg was independent of how mature it was and that the cytoplasm thus became important in the fertilization process.

While in France, Just wrote a book with Hedwig Schnetzler, his second wife and research assistant, entitled The Biology of the Cell Surface (1939). He explained this theory, along with many of his past scientific achievements, in this book. He also categorized biological experimentation and claimed that it fell into three categories: experiments done on living systems, those done on killed living systems, and those done on nonliving systems. He showed the importance of the purity of the system being tested, which was an aspect that had not been emphasized enough by most biologists. Finally, the book was also somewhat of a philosophical treatise, endeavoring to answer the question "what is life?"

Just was also known for his work on cell morphology (the form and the changes in form of a cell), and in particular cell division, the process by which living cells reproduce themselves. Because of his problems with funding, with certain members of the American scientific community, and with racial discrimination, Just was never able to undertake a major research project. However, the accumulation of his work is important to the field of marine biology. His achievements were particularly significant and encouraging to the African American scientists that succeeded him.

Further Reading

The complete story of Ernest Just's life—both the personal and the professional aspects of it—is presented in the biography by Kenneth R. Manning, Black Apollo of Science: the Life of Ernest Everett Just (1983). □

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