Embryo Project Encyclopedia

In the 1930s, George Beadle and Boris Ephrussi discovered factors that affect eye colors in developing fruit flies. They did so while working at the California Institute of Technology in Pasadena, California. (1) They took optic discs (colored fuchsia in the image) from fruit fly larvae in the third instar stage of development. Had the flies not been manipulated, they would have developed into adults with vermilion eyes. (2) Beadle and Ephrussi transplanted the donor optic discs into the bodies of several types of larvae, including those that would develop with normal colored eyes (brick red), and those that would develop eyes with other shades of red, such as claret, carmine, peach, and ruby (grouped together and colored black in the image). (3a) When implanted into normal hosts that would develop brick red eyes, the transplanted optic disc developed into an eye that also was brick red. (3b) When implanted into abnormal hosts that would develop eyes of some other shade of red, the transplanted optic discs developed into eyes that were vermilion. Beadle and Ephrussi concluded that there was a factor, such as an enzyme or some other protein, produced outside of the optic disc that influenced the color of the eye that developed from the disc.

Fruit flies of the species Drosophila melanogaster develop from eggs to adults in eight to ten days at 25 degrees Celsius. They develop through four primary stages: egg, larva, pupa, and adult. When in the wild, female flies lay their fertilized eggs in rotting fruit or other decomposing material that can serve as food for the larvae. In the lab, fruit flies lay their fertilized eggs in a mixture of agar, molasses, cornmeal, and yeast. After roughly a day, each egg hatches into a larva. The larva eats the material it finds itself in, and for four days it grows into stages of increasing size, called first-, second-, and third-instar stages. This figure shows a third-instar larva. Each larva has sections of tissue called imaginal discs, from which various parts of the adult anatomy develop. This figure shows the imaginal discs that will develop into antennae (colored purple), eyes (colored red), brain (colored blue), and wings (colored green). After four days, the larva turns into a pupa by making a casing, similar to caterpillars, and grows within the casing. After a four-day metamorphosis, the adult fly then emerges from its pupal casing. Adult males look somewhat different from adult females, as the males have darker rear abdomen segments than do females. The warmer the temperature around the eggs, the faster the flies develop to adults.

This diagram shows the life cycle of Neurospora crassa, a mold that grows on bread. N. crassa can reproduce through an asexual cycle or a sexual cycle. The asexual cycle (colored as a purple circle), begins in this figure with (1a) vegetative mycelium, which are strands of mature fungus. Some of the strands form bulbs (2a) in a process called conidiation. From those bulbs develop the conidia, which are spores. Next, (3a) a single conidium separates from its strand and elongates until it forms mycelium. The sexual cycle (colored as an orange circle) also starts with the (1b) vegetative mycelium. The strands develop into a structure called the proto-perithecium, and reproduction involves the proto-perithecium interacting with the conidia from a different mycelium. Reproduction also involves two mating types, called type A and type a. In reproduction, type A pairs with type a, and a conidium can be of either type, as can a proto-perithecium. A proto-perithecium fertilized by a conidium of the opposite mating type (2b) will develop into a perithecium. Inside the perithecium, croziers develop and mature into asci. (3b) In a maturing ascus, there are two nuclei (one represented as a white circle and one as a black circle), one of which comes from the conidium and the other from the proto-perithecium. Each nuclei has only one set of chromosomes (haploid). The two haploid nuclei fuse into a diploid nucleus (represented as a half black half white circle). The nucleus then divides, separating into two nuclei each with one set of chromosomes. Those nuclei duplicate themselves (represented as two white circles and two black circles), and then all the nuclei duplicate themselves again (represented as four white circles and four black circles). This process yields eight haploid ascospores within a mature ascus. Ascospores are spores, and function for the mold as do seeds for plants. The mature perithecium releases its ascospores (4b), which germinate and grow into mycelium. In the 1930s and 1940s, George Beadle and Ed Tatum collected the spores of irradiated N. crassa to study how genes produced enzymes.

In 1935, George Beadle and Boris Ephrussi developed a technique to transplant optic discs between fruit fly larvae. They developed it while at the California Institute of Technology in Pasedena, California. Optic discs are tissues from which the adult eyes develop. Beadle and Ephrussi used their technique to study the development of the eye and eye pigment. (1) The experimenter dissects a donor larva, which is in the third instar stage of development, and removes the optic disc (colored red) with a micropipette. Because the antenna disc is attached to the optic disc, they are often removed and transplanted together. (2) The experimenter then implants the optic disc into a host larva, in the part of the host that will develop into an adult abdomen. As the host larva matures to adulthood, the implanted optic disc develops into an eye inside the body cavity of the adult. (3) The adult host has an eye within its body, which Beadle and Ephrussi found by dissecting the adult hosts. If the antenna disc was also transplanted, sometimes the resulting eye developed with an antenna attached.

Conrad Hal Waddington's "Experiments on Embryonic Induction III," published in 1934 in the Journal of Experimental Biology, describes the discovery that the primitive streak induces the mammalian embryo. Waddington's hypothesis was that a transplanted primitive streak could induce neural tissue in the ectoderm of the rabbit embryo. The primitive streak defines the axis of an embryo and is capable of inducing the differentiation of various tissues in a developing embryo during gastrulation. In this experiment Waddington was, in fact, able to induce neural differentiation. Waddington noted that the tissue is "competent"; for a chick organizer, and by deduction a mammalian organizer must exist. Competence refers to a cell's ability to respond to an inducing signal, which is temporally limited to certain developmental stages. Waddington's initial work laid the foundation for many decades of research to follow, including further experiments by Waddington with the mammalian organizer.

Rosalind Elsie Franklin worked with X-ray crystallography at King's College London, UK, and she helped determine the helical structure of DNA in the early 1950s. Franklin's research helped establish molecular genetics, a field that investigates how heredity works on the molecular level. The discovery of the structure of DNA also made future research possible into the molecular basis of embryonic development, genetic disorders, and gene manipulation.

Endoderm is one of the germ layers-- aggregates of cells that organize early during embryonic life and from which all organs and tissues develop. All animals, with the exception of sponges, form either two or three germ layers through a process known as gastrulation. During gastrulation, a ball of cells transforms into a two-layered embryo made of an inner layer of endoderm and an outer layer of ectoderm. In more complex organisms, like vertebrates, these two primary germ layers interact to give rise to a third germ layer, called mesoderm. Regardless of the presence of two or three layers, endoderm is always the inner-most layer. Endoderm forms the epithelium-- a type of tissue in which the cells are tightly linked together to form sheets-- that lines the primitive gut. From this epithelial lining of the primitive gut, organs like the digestive tract, liver, pancreas, and lungs develop.

The sex of a reptile embryo partly results from the production of sex hormones during development, and one process to produce those hormones depends on the temperature of the embryo's environment. The production of sex hormones can result solely from genetics or from genetics in combination with the influence of environmental factors. In genotypic sex determination, also called genetic or chromosomal sex determination, an organism's genes determine which hormones are produced. Non-genetic sex determination occurs when the sex of an organism can be altered during a sensitive period of development due to external factors such as temperature, humidity, or social interactions. Temperature-dependent sex determination (TSD), where the temperature of the embryo's environment influences its sex development, is a widespread non-genetic process of sex determination among vertebrates, including reptiles. All crocodilians, most turtles, many fish, and some lizards exhibit TSD.

Wilhelm Johannsen in Denmark first proposed the distinction between genotype and phenotype in the study of heredity in 1909. This distinction is between the hereditary dispositions of organisms (their genotypes) and the ways in which those dispositions manifest themselves in the physical characteristics of those organisms (their phenotypes). This distinction was an outgrowth of Johannsen's experiments concerning heritable variation in plants, and it influenced his pure line theory of heredity. While the meaning and significance of the genotype-phenotype distinction has been a topic of debate-among Johannsen's contemporaries, later biological theorists, and historians of science-many consider the distinction one of the conceptual pillars of twentieth century genetics. Moreover some have used it to characterize the relationships between studies of development, genetics, and evolution.

The General Embryological Information Service (GEIS) was an annual report published by the Hubrecht Laboratory in Utrecht, The Netherlands from 1949 to 1981 that disseminated contemporary research information to developmental biologists. The purpose of the annual report was to catalog the names, addresses, and associated research of every developmental biologist in the world. Pieter Nieuwkoop edited each issue from 1949 until 1964, when Job Faber began assisting Nieuwkoop. Bert Z. Salome joined the editing team in 1968 before Nieuwkoop ceased editing duties in 1971. Faber and Salome remained the editors from 1971 until the periodical's final year of circulation in 1981. The Hubrecht Laboratory, a national laboratory created to house a large collection of comparative embryological materials and loan them to interested researchers, sponsored the publication after World War II to facilitate international collaboration and prevent unnecessary duplication of work. The catalog of researchers and the scientific topics grew in number and variety as the field of developmental biology changed during the publication's thirty-two year history.