Guest post by Pamela K. Mulligan and Susanne M. Gollin.

Robert C. King, Professor of genetics at Northwestern University for over four decades and a pioneer in studies of Drosophila oogenesis, passed away on June 25th. He was 89.

Among scientists in the field, Bob (as he was generally known) was recognized as a world-renowned geneticist, cell and developmental biologist, educator, and author. However, those who knew him well saw in Bob not only an outstanding researcher and academic, but also a congenial colleague, a supportive mentor, a generous and loyal friend, and an affectionate husband, father, and grandfather.

Bob King is widely known for his definitive work on the morphological characteristics and classification of the fourteen developmental stages of Drosophila oogenesis. It was through cytological observations and painstaking three-dimensional reconstructions of electron microscope images that he elucidated the formation and development of the basic structural and functional unit of the ovary, the egg chamber.

This work included identifying and describing germline stem cells at the tip of the ovary; the pattern of asymmetric cell divisions that gives rise to a 16-cell germline syncytium from which one cell becomes the oocyte; the formation of the fusome, an organelle required for the proper formation of the germline syncytium; the existence of cytoplasmic bridges, or ring canals, connecting the germline cells; the cytology and migratory behaviour of somatic cells surrounding the germline cluster; and other developmental events that culminate in the production of a mature, functional oocyte.

The cytological and ultrastructural work on the formation and differentiation of the egg chamber was combined with studies of female-sterile mutations that caused defects in normal oogenesis. Through this work, genes whose products are required for various cellular processes during oogenesis were identified.  These studies laid the earliest foundation for understanding the genetic control of Drosophila oogenesis and helped advance the burgeoning field of developmental genetics.

This basic foundation has been built upon by later generations of scientists using new and advanced tools, and Drosophila oogenesis is now widely recognized as an excellent model system in which to study developmental genetics and cell biology in the minutest molecular detail. For example, the ovarian tumor (otu) gene, one of the most extensively characterized by Bob, was subsequently isolated using hybrid dysgenesis-induced mutations and its role in the proliferation and differentiation of the germline described. The morphogenesis and molecular components of the fusome and its role in oocyte specification have been advanced. Studies of mutations that disrupt normal ring canal development have led to the identification of genes that encode ring canal proteins, and the highly ordered process of ring canal assembly has been elucidated in striking detail. Time-lapse confocal microscopy has described intercellular transport of distinct biomolecules through the ring canals. And outside of this system, Bob’s investigations were the inspiration for the recent finding that oocyte formation in mammals occurs in a manner similar to that in the Drosophila ovary.

In addition to publishing research papers Bob was also the author and editor of books that have contributed significantly to the field of genetics. These include Genetics (Oxford University Press), the multi-volume Handbook of Genetics (Plenum Publishing), Ovarian Development in Drosophila melanogaster (Academic Press), Insect Ultrastructure, Volumes 1 and 2 (Plenum Publishing) and eight editions of A Dictionary of Genetics (Oxford University Press). These books and reference works continue to educate budding scientists and experts around the globe. At the time of his death Bob was working on the ninth edition of his beloved Dictionary, an interdisciplinary book translated into five languages, that has become an invaluable reference work for students of classical and molecular genetics.

Bob King was born in New York City in 1928 and received his PhD in Zoology from Yale University in 1952 at the early age of 24. He then spent a few years at Brookhaven National Laboratory and in 1956 accepted an Assistant Professorship in the Department of Biology at Northwestern University. At Northwestern he taught undergraduate and graduate courses in genetics, developmental genetics, cell biology and cytology, and established his now renowned research and writing career. He became a Full Professor in 1964 and retired in 2000 as Emeritus Professor in the Department of Biochemistry, Molecular Biology and Cell Biology.

Although Bob King was a leader in his chosen field, he was generally content to remain in the background due to his selfless nature. Bob’s quiet, intangible characteristics are what his students and associates will most miss: his strong, silent mentoring and guidance through the process of higher education and scientific research; his strict adherence to the highest ethical standards; his insistence on near-perfection in writing and scientific endeavor; his genuine love for science; and his fatherly caring about his students and their futures. Many who benefited from Bob’s tutelage continue his legacy of high standards in scientific pursuit and scientific writing.

We, as former students, remember Bob’s lab as a hub of exciting research and cheerfulness. Bob was always the first to arrive for work. Long before even the earliest risers amongst us entered the lab, he would have been at his desk for an hour or more, and one would open the lab door to the smell of freshly-brewed coffee, with the sound of Bach, Beethoven, or Stravinsky in the background. There was an ethnically diverse population of students and research associates, and his children were frequent visitors. There were parties to celebrate publication of papers and funding of grants. Bob loved to travel, presenting papers and spending sabbaticals abroad. It was in Korea that he met his future wife and loving companion, Suja. After retirement, Bob indulged in his insatiable interest in books (including murder mysteries), music, and cinema. And he continued to write, producing new editions of his Dictionary every few years.

The passing of Bob King brings into sharp focus the life of an extraordinary man who loved life and who leaves behind a legacy of academic excellence, hard work, generosity in friendship, and a sincere desire to make life better for others. As we mourn his loss, we are deeply grateful for his vital contributions to science and for his humanity.

For additional information about Robert C. King’s work and life, see:

Robert C. King: An Appreciation of His Work 

Pamela K. Mulligan


For an introduction to Drosophila oogenesis, see this GENETICS Primer article:

Drosophila Oocytes as a Model for Understanding Meiosis: An Educational Primer to Accompany “Corolla Is a Novel Protein That Contributes to the Architecture of the Synaptonemal Complex of Drosophila

Elizabeth T. Ables

GENETICS January 2015 199: 17-23;

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  1. King’s Genetics text shaped my early thinking in Genetics. Very nice to rad about the man and see how great he was to his lab.