Embryo Project Encyclopedia

Carl Richard Moore was a professor and researcher at the University of Chicago in Chicago, Illinois who studied sex hormones in animals from 1916 until his death in 1955. Moore focused on the role of hormones on sex differentiation in offspring, the optimal conditions for sperm production, and the effects of vasectomy or testicular implants on male sex hormone production. Moore's experiments to create hermaphrodites in the laboratory contributed to the theory of a feedback loop between the pituitary and fetal gonadal hormones to control sex differentiation. Moore showed that the scrotal sac controls the temperature for the testes, which is necessary for sperm production. He also helped distinguish the hormones testosterone, and androsterone from testicular extracts.

Frank Rattray Lillie's research on freemartins from 1914 to 1920 in the US led to the theory that hormones partly caused for sex differentiation in mammals. Although sometimes applied to sheep, goats, and pigs, the term freemartin most often refers to a sterile cow that has external female genitalia and internal male gonads and was born with a normal male twin. Lillie theorized that a freemartin is a genetic female whose process of sexual development from an undifferentiated zygote was suppressed or antagonized by her twin's release of male hormones via their shared blood circulation in utero. Despite publications of similar findings by physician Julius Tandler in Vienna, Austria, in 1910 and physician Karl Keller in Wiesensteig, Germany in 1916 prior to Lillie's research, Lillie often receives credit for the hormonal theory of sex differentiation in the freemartin. Lillie's study of freemartins, and the subsequent research by graduate students in Lillie's laboratory at the University of Chicago in Chicago, Illinois, prompted many embryologists to research sex differentiation and hermaphroditism in mammals.

Ontogeny and Phylogeny is a book published in 1977, in which the author Stephen J. Gould, who worked in the US, tells a history of the theory of recapitulation. A theory of recapitulation aims to explain the relationship between the embryonic development of an organism (ontogeny) and the evolution of that organism's species (phylogeny). Although there are several variations of recapitulationist theories, most claim that during embryonic development an organism repeats the adult stages of organisms from those species in it's evolutionary history. Gould suggests that, although fewer biologists invoked recapitulation theories in the twentieth century compared to those in the nineteenth and eighteenth centuries, some aspects of the theory of recapitulation remained important for understanding evolution. Gould notes that the concepts of acceleration and retardation during development entail that changes in developmental timing (heterochrony) can result in a trait appearing either earlier or later than normal in developmental processes. Gould argues that these changes in the timing of embryonic development provide the raw materials or novelties upon which natural selection acts.

The Spandrels of San Marco and the Panglossian Paradigm:
A Critique of the Adaptationist Programme, hereafter called
The Spandrels, is an article written by Stephen J. Gould and
Richard C. Lewontin published in the Proceedings of the Royal
Society of London in 1979. The paper emphasizes issues with
what the two authors call adaptationism or the adaptationist
programme as a framework to explain how species and traits evolved. The paper
is one in a series of works in which Gould emphasized the
role of development in evolutionary theories. The article suggests
that constraints on how organisms can develop and constraints on how species can evolve from others play a
central role in explaining the how species and traits evolve. The
authors note that organisms from different species develop as
embryos through stages similar across species, genera, and higher
classes. Gould and Lewontin hypothesize that those stages
constrained the possible pathways of evolution and has therefore
guided the history of life. Throughout the paper, the authors rely on analogy of some parts of organisms to architectural structures called spandrels, marked in this image as 'a'."

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.

The Law of Acceleration of Growth is a theory proposed by Edward Drinker Cope in the US during the nineteenth century. Cope developed it in an attempt to explain the evolution of genera by appealing to changes in the developmental timelines of organisms. Cope proposed this law as an additional theory to natural selection. He argued that the evolution of genera, the more general groups within which biologists group species, occurs when the individual in a species move through developmental stages faster than did their ancestors, but within the same fixed period of gestation, and thus can undergo new developmental stages and develop new traits. The Law of Acceleration compliments Cope's Law of Retardation of Growth. He described the later law as the process by which organisms revert to an ancestral stage. In these cases, forces suppress the most recent traits or stages common to the development of individuals from different species within the same genus. Cope described evolution as progressive, following a predetermined path, a perspective about evolution sometimes called orthogenetic. Cope's was one among many orthogenic theories in the second half of the nineteenth century. Furthermore, the theory was part of a trend in nineteenth century in which some biologists claimed that the changes in developmental timing of organisms could explain large changes in biological forms throughout natural history.

Mutual Affinities of Organic Beings: Morphology: Embryology: Rudimentary Organs is the thirteenth chapter of Charles Darwin's book The Origin of Species, first published in England in 1859. The book details part of Darwin's argument for the common ancestry of life and natural selection as the cause of speciation. In this chapter, Darwin summarizes the evidence for evolution by connecting observations of development in organisms to the processes of natural selection. Darwin shows how the theory of special creation, which claims that God directly created all organisms in their current form, is inferior to the theory of natural selection for its ability to explain the diversity of life. In this chapter, Darwin also discusses classification and homology as they relate to natural selection.

Stephen Jay Gould studied snail fossils and worked at Harvard University in Cambridge, Massachusetts during the latter half of the twentieth century. He contributed to philosophical, historical, and scientific ideas in paleontology, evolutionary theory, and developmental biology. Gould, with Niles Eldredge, proposed the theory of punctuated equilibrium, a view of evolution by which species undergo long periods of stasis followed by rapid changes over relatively short periods instead of continually accumulating slow changes over millions of years. In his 1977 book, Ontogeny and Phylogeny, Gould reconstructed a history of developmental biology and stressed the importance of development to evolutionary biology. In a 1979 paper coauthored with Richard Lewontin, Gould and Lewonitn criticized many evolutionary bioligists for relying solely on adaptive evolution as an explanation for morphological change, and for failing to consider other explanations, such as developmental constraints.

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.

This illustration shows George Beadle and Edward Tatum's experiments with Neurospora crassa that indicated that single genes produce single enzymes. The pair conducted the experiments at Stanford University in Palo Alto, California. Enzymes are types of proteins that can catalyze reactions inside cells, reactions that produce a number of things, including nutrients that the cell needs. Neurospora crassa is a species of mold that grows on bread. In the early 1940s, Beadle and Tatum conducted an experiment to discover the abnormal genes in Neurospora mutants, which failed to produce specific nutrients needed to survive. (1) Beadle and Tatum used X-rays to cause mutations in the DNA of Neurospora, and then they grew the mutated Neurospora cells in glassware. (2) They grew several strains, represented in four groups of paired test tubes. For each group, Neurospora was grown in one of two types of growth media. One medium contained all the essential nutrients that the Neurospora needed to survive, which Beadle and Tatum called a complete medium. The second medium was a minimal medium and lacked nutrients that Neurospora needed to survive. If functioning normally and in the right conditions, however, Neurospora can produce these absent nutrients. (3) When Beadle and Tatum grew the mutated mold strains on both the complete and on the minimal media, all of the molds survived on the complete media, but not all of the molds survived on the minimal media (strain highlighted in yellow). (4) For the next step, the researchers added nutrients to the minimal media such that some glassware received an amino acid mixture (represented as colored squares) and other glassware received a vitamin mixture (represented as colored triangles) in an attempt to figure out which kind of nutrients the mutated molds needed. The researchers then took mold from the mutant mold strain that had survived on a complete medium and added that mold to the supplemented minimal media. They found that in some cases the mutated mold grew on media supplemented only with vitamins but not on media supplemented only with amino acids. (5) To discover which vitamins the mutant molds needed, Beadle and Tatum used several tubes with the minimal media, supplementing each one with a different vitamin, and then they attempted to grow the mutant mold in each tube. They found that different mutant strains of the mold grew only on media supplemented with different kinds of vitamins, for instance vitamin B6 for one strain, and vitamin B1 for another. In experiments not pictured, Beadle and Tatum found in step (4) that other strains of mutant mold grew on minimal media supplemented only with amino acids but not on minimal media supplemented only with vitamins. When they repeated step (5) on those strains and with specific kinds of amino acids in the different test tubes, they found that the some mutated mold strains grew on minimal media supplemented solely with one kind of amino acid, and others strains grew only on minimal media supplemented with other kinds of amino acids. For both the vitamins and amino acid cases, Beadle and Tatum concluded that the X-rays had mutated different genes in Neurospora, resulting in different mutant strains of Neurospora cells. In a cell of a given strain, the X-rays had changed the gene normally responsible for producing an enzyme that catalyzed a vitamin or an amino acid. As a result, the Neurospora cell could no longer produce that enzyme, and thus couldn't catalyze a specific nutrient.