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8Muller’s Lineage from Morgan’s Second Mentor, William Keith Brooks

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H. J. Muller’s undergraduate years were inspired by the courses taught by Edmund Beecher Wilson. His graduate years were spent, however, in the laboratory of Thomas Hunt Morgan. Both Wilson and Morgan received their PhDs from the same mentor, William Keith Brooks. Hence Wilson’s pedigree follows the same lineage through Brooks as does Morgan.

Edmund Beecher Wilson (1856–1939) was born in Geneva, Illinois, and died in New York City, NY. In his youth he considered becoming a cellist. His father was a judge. Instead, Wilson went to Yale and loved learning about nature. He went to Johns Hopkins for his PhD studying with William Keith Brooks. He chose embryology and cell biology as his special interest and taught first at Bryn Mawr and then at Columbia University. As chair of that department, he recruited T. H. Morgan who came to Johns Hopkins several years after Wilson had graduated with his PhD. In 1896 Wilson published the first edition of The Cell in Development and Inheritance. It summed up the latest findings in cell biology and Wilson predicted that the nucleic acids might be the source of the hereditary component of the chromosomes. His laboratory work focused on the chromosomes and he used beetles to work out the sex chromosomes, which he designated as X and Y with XX as female and XY as male (with some species having a single X male and two X females and no Y chromosome which he designated as XO male and XX female). Independently, Nettie Stevens found sex chromosomes in Diptera that she studied. His student Walter Sutton was the first to apply Mendelian inheritance to meiosis and with Wilson and Theodor Boveri they called this the chromosome theory of heredity.

William Keith Brooks (1848–1908) was born in Cleveland and died in Baltimore. He had frail health from a congenital heart defect. He was an avid scholar and read widely in classics and philosophy before committing himself to natural history after reading Darwin’s Origin of Species. At Harvard he got his PhD with Louis and Alexander Agassiz. He studied embryology of tunicates (the genus Salpa) and invertebrates, especially mollusks. He believed a study of tunicates would contribute to the understanding of the evolution of vertebrates from their invertebrate ancestors. He was a gifted teacher and among his PhD and postdoctoral students were William Bateson, T. H. Morgan, Edmund B. Wilson, H. V. Wilson, and Ross Harrison. He wrote seven influential books, including The Law of Heredity (1883). His views were Lamarckian. He told Bateson that the most interesting area he could enter was heredity. He instructed his students to work with living specimens and study their living functions. Bateson followed that advice but rejected Brook’s theoretical approach.

Louis Agassiz (1807–1873) was born in Môtier, Switzerland, and died in Boston, Massachusetts. He studied medicine at Zurich, Heidelberg, and Munich and he studied natural history at Erlanger, Paris, and Munich. He did his research with Alexander Cuvier and Alexander von Humboldt. Cuvier inspired him to become an ichthyologist and Humboldt convinced him of the importance of geology for a study of natural history. Agassiz gained fame by descending through a vertical tunnel he dug into a glacier to examine if water flowed under it. He wrote numerous volumes on fresh water fish and fossil fish. From their distribution he concluded that Europe was covered by glaciers that he described as the Ice Age. When he visited the United States to study North American fish, he fell in love with American society and accepted an appointment at Harvard. His students included David Starr Jordan and William Keith Brooks. He established a forerunner of Woods Hole laboratories at Penikese Island in Buzzard’s Bay, inspiring Brooks to set up a similar, but portable, summer laboratory by the sea in Beaufort, Maryland. He opposed natural selection but never criticized his students who were all Darwinians. At Woods Hole his motto greets visitors: “Study Nature, Not Books.”

Georges Cuvier (1769–1832) was born in Montbéliard, and died in Paris, both in France. He was raised as a Lutheran, a faith he kept throughout his life. His father was in the Swiss Guard. Young Cuvier had a near photographic memory and remembered what he read in prodigious amounts including Buffon’s volumes on natural history. He studied at Stuttgart and then in Paris where he gained attention for his studies of living and fossil elephants. He described Asian and African elephants as separate species and the fossil elephant (that he named mastodon) also a separate species. He received his doctorate from Ignaz Döllinger. Cuvier is a founder of the field of comparative anatomy and made numerous contributions to taxonomy, the use of stratigraphy to represent layers of animals that became fossilized. He explained this with a theory of sudden extinctions (attributed to massive tsunamis) and replacements with new creations. He opposed vehemently Lamarck’s theory of evolution and modification by use and disuse and wrote a scathing eulogy in 1832 denouncing Lamarck’s character. Cuvier’s most famous student was Louis Agassiz.

Ignaz Döllinger (1770–1841) was born in Bamberg, Germany, where his father was a Professor and physician. He died in Munich. He got his doctorate in 1794 and studied in Würzburg, Padua, and Vienna before settling in to Munich where he was Professor of Anatomy. His medical degree was from Padua and his doctoral advisor was Antonio Scarpa. Döllinger taught medicine as a natural science. He devoted most of his research to embryonic development. His students included Louis Agassiz, Karl Ernst von Baer, Christian Pander, Lorenz Oken, and Johann Lukas Schönlein. Pander and von Baer helped establish the field of embryology. Pander introduced the idea of three germ layers in the early embryo — ectoderm, mesoderm, and endoderm. It was von Baer who described development as epigenetic with organs formed not by enlargement but by differentiation of embryonic germ layers. Schönlein was one of the first German Professors to lecture in German rather than in Latin. Oken was a founder of a holistic, transcendental approach to biology called naturphilosophie that was extended by Goethe. It tried to make sense of homologous structures and how they came to be modified.

Antonio Scarpa (1752–1832) was born in Liorenzaga, Italy, and died in Padua, Italy. He received his medical degree at the University of Padua, studying with Giovanni Battista Morgani. He studied the anatomy of the inner ear and cardiac nerves. He taught at the University of Modena. He was a bachelor and had several children out of wedlock. Born poor, he became wealthy, and a collector of art. Scarpa’s most noted student was Ignaz Döllinger .

Giovanni Battista Morgagni (1682–1771) was born in Foril, Italy, and died in Padua, Italy. He was a professor of anatomical pathology at the University of Padua. He rejected the prevailing view of the time that disease was systemic (influenced by vital humors or toxins). Instead he argued that it had a localized origin and was usually organ specific. He wrote a five-volume treatise, On the Seat and Causes of Diseases based on 640 dissections he carried out. His most famous student was Antonio Scarpa. His mentor was Antonio Maria Valsalva.

Antonio Maria Valsalva (1666–1723) was born in Imola, Italy. He studied the anatomy of the throat and named the Eustachian tube. His mentor was Marcello Malpighi.

Marcello Malpighi (1628–1694) was born near Bologna, Italy. He has been described as “the father of microscopic anatomy, histology, physiology, and embryology.” He discovered the trachea network in insects and associated them with breathing. He discovered the Malpighian tubes in kidneys and demonstrated that the pigment of Africans was in a lower layer of the skin. He identified the capillaries as the structures that connected arterial blood to venous blood flow. His colleague Giovanni Alfonso Borelli at Pisa introduced him to experimental science and Malpighi chose the microscope for his studies. He was an excellent artist and drew careful illustrations of his microscopic specimens. He demonstrated that galls found in plants were caused by insects that laid eggs in the plant tissue. The Royal Society in London published many of his findings in plant and animal cellular anatomy.

Giovanni Alfonso Borelli (1608–1679) was born in Naples of a Spanish father and Italian mother. He took an interest in experimental physics and mathematics but became a physician. For his medical studies, he took an interest in animal motion and worked out the relation of muscles and bones to limb motion. He was also the first to note the existence of stomata in plant leaves. He is considered the founder of the field of biophysics. He was mentored by Benedetto Castelli with whom he studied the detailed orbits of Jupiter’s moons.

Benedetto Castelli (1578–1643) was born in Brescia, Italy, and became a mathematician and taught at the University of Padua before becoming the Abbott of Monte Casino. He worked on sunspots with his mentor Galileo Galilei.

Galileo Galilei (1564–1642) was born in Pisa, Italy. His father was a composer and played the lute. He wanted Galileo to become a physician but while in medical school Galileo found courses on astronomy and science more interesting and convinced his father to let him change his field. He studied mathematics and made contributions to the physics of falling bodies, using experiments to demonstrate these laws. He moved to the University of Padua where he taught and wrote most of his books. He also made a telescope and applied it to the skies. He discovered the moon had craters, Venus had phases like the moon, Jupiter had four moons, and the sun had sunspots that followed its rotation. He also described Saturn as having “ears” because the rings at that time were tilted and the lenses were not as sharp as later lenses that revealed these were rings. He felt he had the evidence for the Copernican theory and began a series of disputes with his fellow astronomers. He replied in polemic style ridiculing his opponents, many of them Jesuits or influential in the Church. This led to his eventual trial and conviction as a heretic forcing recantation and household arrest for the rest of his life. He is considered one of the greatest scientists of all time and he helped launch the scientific revolution of the later Renaissance. His mentor in Pisa was Ostilio Ricci.

Ostilio Ricci (1540–1603) was a mathematician who taught at the University of Pisa. Galileo took his courses and was converted by Ricci to become a scientist and mathematician. He taught Galileo Euclidian and Archimedean mathematics. Ricci believed that mathematics was not a science itself but a tool that could be used for applied science. He was mentored by Niccolò Fontana Tartaglia.

Niccolò Fontana Tartaglia (1499–1557) was born in Brescia. When the French defeated the Italians in that region they massacred most of its inhabitants. Niccolò was a child and was hit by a saber that sliced his jaw and palate. His mother nursed him back to health, but it made speech difficult and his nickname (the stammerer) became his last name. He never shaved, believing his beard would hide his wounds. He learned engineering and wrote books on applied mathematics. He worked out the mathematics for ballistics. He wrote a treatise on salvaging sunken ships. He translated Euclid into Italian. He solved the mathematics for cubic equations. His treatise on mathematics was a sixteenth century best seller. His most famous student was Ostilio Ricci.

How to Construct Your Intellectual Pedigree

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