Benjamin Harrison Willier is considered one of the most versatile embryologists to have ever practiced in the US. His research spanned most of the twentieth century, a time when the field of embryology evolved from being a purely descriptive pursuit to one of experimental research, to that of incorporating molecular biology into the research lab. Willier was born on 2 November 1890 near Weston, Ohio to Mary Alice Ricard. He spent his childhood doing farming chores and running the farm while his father, David Willier worked as a banker. Willier graduated from high school with no immediate desire to attend college; his goal was to become a public school teacher. After a year of special study he was certified to teach. But this desire took a turn when his cousin and public school teacher, Alice Ricard, convinced him to attend a six-week summer session of the State Normal College of Miami University (Ohio). It was here that Willier was introduced to nature study and where he made his decision to pursue a scholarly life.
In his 1907 paper, "Experiments in Transplanting Limbs and Their Bearing Upon the Problems of the Development of Nerves," in the Journal of Experimental Zoology that he edited, Ross Granville Harrison tested the development of nerves in transplanted tissue. He studied neural development by examining two competing theories. Victor Hensen proposed a syncytial theory as a way to explain neural development, suggesting that all the nerves of an embryo were connected directly by cytoplasm laid down early in development, and leaving no room for later modification. Santiago Ramón y Cajal proposed a competing outgrowth theory that nerves develop from the central nervous system, pushing through tissues by growing nerve fibers and growth cones. Harrison's experiment refuted many of the claims of the syncytial theory although it did not produce evidence that could directly prove the outgrowth theory.
In 1952 Robert Briggs and Thomas J. King published their article, "Transplantation of Living Nuclei from Blastula Cells into Enucleated Frogs' Eggs," in the Proceedings of the National Academy of Sciences, the culmination of a series of experiments conducted at the Institute for Cancer Research and Lankenau Hospital Research Institute in Philadelphia, Pennsylvania. In this paper Briggs and King examined whether nuclei of embryonic cells are differentiated, and by doing so, were the first to conduct a successful nuclear transplantation with amphibian embryos. Previously nuclear transplantation had only been performed using amoebae cells. Briggs and King believed that by removing the egg nucleus and replacing it with a differentiated cell, they could study nuclear differentiation. During the experiment, they used two different species of frogs, Rana pipiens and Rana catesbeiana, to study and test whether the nucleus is differentiated. The nuclear transplantations performed in the experiment would later be referred to as cloning.
Gustav Jacob Born was an experimental embryologist whose original work with amphibians served as the platform for his wax-plate method of embryo modeling, heteroblastic (different tissues) and xenoplastic (similar species) transplantation methods, environmental influences on sex ratio studies, and proposed function of the corpus luteum. He was born 22 April 1851 in Kempen, Prussia, but his family moved to the larger city of Görlitz within a year after Born's birth. His father was Marcus Born, a physician and public health officer who practiced in the town of Görlitz. Born began his formal education at the Gymnasium in Görlitz and later attended the universities of Breslau, Bonn, Strassburg, and Berlin. During the Franco-Prussian War (1870-1871), Born enlisted and served as an orderly and physician's assistant. He returned to the University of Breslau at war's end and graduated with a MD in 1874. In 1877 Born became a Privatdozent at the University of Breslau. In 1886 he was made assistant professor and in 1898 he became professor of histology and comparative anatomy. During his tenure at Breslau, Born worked with Karl Hasse and reported being under constant scrutiny by Hasse who held widely known anti-semitic views.
Leo Loeb developed an experimental approach to studying cancer and pioneered techniques for tissue culture and in vitro tissue transplantation which impacted early-to-mid twentieth century experimental embryology. Loeb received his medical degree from the University of Zurich in 1897. As part of his doctorate, he completed a thesis on the outcomes of tissue transplantation in guinea pigs. Loeb's thesis inspired a life-long interest in tissue transplantation. His research culminated in greater than 400 publications, including a book called The Biological Basis of Individuality, in which he demonstrated the potential immortality of certain mammalian tissues.
In "Versuche zur Analyse der Induktionsmittel in der Embryonalentwicklung," published in Naturwissenschaften in 1932, Hermann Bautzmann, Johannes Holtfreter, Otto Mangold, and Hans Spemann jointly reported on experiments each had conducted testing the activity of organizers killed by boiling, freezing, alcohol, and drying. Each of the authors had been independently conducting similar experiments, when Holtfreter made a breakthrough allowing him to produce many more successful transplantations. When he told Spemann the news, Spemann suggested that he tell Bautzmann, who had been working extensively on the question. Spemann then coordinated the joint paper to avoid a conflict among the researchers.
In this paper Viktor Hamburger and Rita Levi-Montalcini collaborated to examine the effects of limb transplantation and explantation on neural development. In 1947 Hamburger invited Levi-Montalcini to his lab at Washington University in St. Louis to examine this question. Independently, each had previously arrived at opposing conclusions based on the same data. Hamburger concluded that limb transplantations caused the ganglia to develop more connections and grow larger while Levi-Montalcini concluded that the ganglia first produce a large amount of neurons, then degenerate the unsuccessful neurons. She concluded that larger ganglia must be due to the increase in successful connections. This joint paper, published in the Journal of Experimental Zoology in 1949, corroborated the findings reported by Levi-Montalcini and established that nerve degeneration is an integral part of development.
Samuel Randall Detwiler was an embryologist who studied neural development in embryos and vertebrate retinas. He discovered evidence for the relationship between somites and spinal ganglia, that transplanted limbs can be controlled by foreign ganglia, and the plasticity of ganglia in response to limb transplantations. He also extensively studied vertebrate retinas during and after embryonic development. Detwiler's work established many principles studied in later limb transplantation experiments and was identified by Viktor Hamburger as an important bridge between his and Ross Granville Harrison's research.
Hans Spemann was an experimental embryologist best known for his transplantation studies and as the originator of the "organizer" concept. One of his earliest experiments involved constricting the blastomeres of a fertilized salamander egg with a noose of fine baby hair, resulting in a partially double embryo with two heads and one tail. Spemann continued changing variables such as the amount of time the embryo was constricted and the degree of constriction, all of which added more empirical evidence to Hans Driesch's studies showing that embryonic cells could self-regulate to varying degrees. Spemann's long list of "simple" experiments and significant findings were mainly carried out at his laboratory, the Spemann School at the University of Freiburg, Germany, where numerous graduate students collaborated with Spemann to investigate embryonic induction.