During the twentieth century in the United States, Alfred Day Hershey studied phages, or viruses that infect bacteria, and experimentally verified that genes were made of deoxyribonucleic acid, or DNA. Genes are molecular, heritable instructions for how an organism develops. When Hershey started to study phages, scientists did not know if phages contained genes, or whether genes were made of DNA or protein. In 1952, Hershey and his research assistant, Martha Chase, conducted phage experiments that convinced scientists that genes were made of DNA. For his work with phages, Hershey shared the 1969 Nobel Prize in Physiology or Medicine with Max Delbrück and Salvador Luria. Hershey conducted experiments with results that connected DNA to the function of genes, thereby changing the way scientists studied molecular biology and the development of organisms.

In 2001, Yale University Press published Frederic Lawrence Holmes' book, Meselson, Stahl, and the Replication of DNA: A History of "The Most Beautiful Experiment in Biology" (Replication of DNA), which chronicles the 1950s debate about how DNA replicates. That experiment verified that DNA replicates semi-conservatively as originally proposed by Watson and Crick. Rather than focusing solely on experiments and findings, Holmes's book presents the investigative processes of scientists studying DNA replication. Based on personal accounts, letter correspondence, and preserved research documents, Replication of DNA serves as a detailed account of the initial issues surrounding DNA replication and the Meselson-Stahl experiment from a scientist's perspective.

On 6 May 1952, at King’s College London in London, England, Rosalind Franklin photographed her fifty-first X-ray diffraction pattern of deoxyribosenucleic acid, or DNA. Photograph 51, or Photo 51, revealed information about DNA’s three-dimensional structure by displaying the way a beam of X-rays scattered off a pure fiber of DNA. Franklin took Photo 51 after scientists confirmed that DNA contained genes. Maurice Wilkins, Franklin’s colleague showed James and Francis Crick Photo 51 without Franklin’s knowledge. Watson and Crick used that image to develop their structural model of DNA. In 1962, after Franklin’s death, Watson, Crick, and Wilkins shared the Nobel Prize in Physiology or Medicine for their findings about DNA. Franklin’s Photo 51 helped scientists learn more about the three-dimensional structure of DNA and enabled scientists to understand DNA’s role in heredity.

In May 1953, scientists James Watson and Francis Crick wrote the article “Genetical Implications of the Structure of Deoxyribonucleic Acid,” hereafter “Genetical Implications,” which was published in the journal Nature. In “Genetical Implications,” Watson and Crick suggest a possible explanation for deoxyribonucleic acid, or DNA, replication based on a structure of DNA they proposed prior to writing “Genetical Implications.” Watson and Crick proposed their theory about DNA replication at a time when scientists had recently reached the consensus that DNA contained genes, which scientists understood to carry information that determines an organism’s identity. Watson and Crick’s replication mechanism as presented in “Genetical Implications” contributed to the two scientists sharing a portion of the 1962 Nobel Prize in Physiology or Medicine. With their suggested DNA replication mechanism in “Genetical Implications,” Watson and Crick explained how genes are copied and passed along to new cells and organisms, thereby explaining how the information contained within genes is preserved through generations.

In April 1953, Rosalind Franklin and Raymond Gosling, published “Molecular Configuration in Sodium Thymonucleate,” in the scientific journal Nature. The article contained Franklin and Gosling’s analysis of their X-ray diffraction pattern of thymonucleate or deoxyribonucleic acid, known as DNA. In the early 1950s, scientists confirmed that genes, the heritable factors that control how organisms develop, contained DNA. However, at the time scientists had not determined how DNA functioned or its three-dimensional structure. In their 1953 paper, Franklin and Gosling interpret X-ray diffraction patterns of DNA fibers that they collected, which show the scattering of X-rays from the fibers. The patterns provided information about the three-dimensional structure of the molecule. “Molecular Configuration in Sodium Thymonucleate” shows the progress Franklin and Gosling made toward understanding the three-dimensional structure of DNA.

Franklin William Stahl studied DNA replication, bacteriophages, and genetic recombination in the US during the mid-twentieth and early twenty-first centuries. With his colleague Matthew Meselson, Stahl performed an experiment called the Meselson-Stahl experiment, which provided evidence for a process called semi-conservative DNA replication. Semi-conservative replication is a process in which each strand of a parental DNA double helix serves as a template for newly replicated daughter strands, so that one parental strand is conserved in every daughter double helix. Those findings supported the Watson-Crick Model for DNA replication proposed in 1953 by James Watson and Francis Crick, convincing many biologists about DNA’s structure and replication in the 1950s. Stahl’s genetics research, especially that of DNA replication, showed researchers how genetic information is distributed within a cell and is passed down from cell to cell.

Matthew Stanley Meselson conducted DNA and RNA research in the US during the twentieth and twenty-first centuries. He also influenced US policy regarding the use of chemical and biological weapons. Meselson and his colleague Franklin Stahl demonstrated that DNA replication is semi-conservative. Semi-conservative replication means that every newly replicated DNA double helix, which consists of two individual DNA strands wound together, contains one strand that was conserved from a parent double helix and that served as a template for the other strand. Meselson's work enabled researchers to better explain and control cellular development by showing how DNA are copied when a cell divides and interpreted when a cell makes proteins.

William Thomas Astbury studied the structures of fibrous materials, including fabrics, proteins, and deoxyribonucleic acid, or DNA, in England during the twentieth century. Astbury employed X-ray crystallography, a technique in which scientists use X-rays to learn about the molecular structures of materials. Astbury worked at a time when scientists had not yet identified DNA’s structure or function in genes, the genetic components responsible for how organisms develop and reproduce. He was one of the first scientists to use X-ray crystallography to study the structure of DNA. According to historians, Astbury helped establish the field of molecular biology as he connected microscopic changes in the structure of materials to changes in their large-scale properties. Astbury and his images helped scientists to understand the structure of DNA and its role in genetics.

In 1944, Oswald Avery, Colin MacLeod, and Maclyn McCarty published an article in which they concluded that genes, or molecules that dictate how organisms develop, are made of deoxyribonucleic acid, or DNA. The article is titled “Studies on the Chemical Nature of the Substance Inducing Transformation of Pneumococcal Types: Induction of Transformation by a Desoxyribonucleic Acid Fraction Isolated from Pneumococcus Type III,” hereafter “Transformation.” The authors isolated, purified, and characterized genes within bacteria and found evidence that those genes were made of DNA and not protein. Though scientists were initially skeptical that genes were made of DNA, they later recognized that the data reported in “Transformation” were clear evidence that DNA was genetic material, a revelation that furthered research about how organisms grow, develop, and pass on traits to offspring.

In February 1953, Linus Pauling and Robert Brainard Corey, two scientists working at the California Institute of Technology in Pasadena, California, proposed a structure for deoxyribonucleic acid, or DNA, in their article “A Proposed Structure for the Nucleic Acids,” henceforth “Nucleic Acids.” In the article, Pauling and Corey suggest a model for nucleic acids, including DNA, that consisted of three nucleic acid strands wound together in a triple helix. “Nucleic Acids” was published in Proceedings of the National Academy of Sciences shortly after scientists came to the consensus that genes, the biological factors that control how organisms develop, contained DNA. Though scientists proved Pauling and Corey’s model incorrect, “Nucleic Acids” helped scientists understand DNA’s structure and function as genetic material.

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