'On the Permanent Life of Tissues outside of the Organism' reports Alexis Carrel's 1912 experiments on the maintenance of tissue in culture media. At the time, Carrel was a French surgeon and biologist working at the Rockefeller Institute in New York City. In his paper, Carrel reported that he had successfully maintained tissue cultures, which derived from connective tissues of developing chicks and other tissue sources, by serially culturing them. Among all the tissue cultures Carrel reported, one was maintained for more than two months, whereas previous efforts had only been able to keep tissues in vitro for three to fifteen days. Carrel’s experiments contributed to the development of long-term tissue culture techniques, which were useful in the study of embryology and eventually became instrumental in stem cell research. Despite later evidence to the contrary, Carrel believed that as long as the tissue culture method was accurately applied, tissues kept outside of the organisms should be able to divide indefinitely and have permanent life.

The Hayflick Limit is a concept that helps to explain the mechanisms behind cellular aging. The concept states that a normal human cell can only replicate and divide forty to sixty times before it cannot divide anymore, and will break down by programmed cell death or apoptosis. The concept of the Hayflick Limit revised Alexis Carrel's earlier theory, which stated that cells can replicate themselves infinitely. Leonard Hayflick developed the concept while at the Wistar Institute in Philadelphia, Pennsylvania, in 1965. In his 1974 book Intrinsic Mutagenesis, Frank Macfarlane Burnet named the concept after Hayflick. The concept of the Hayflick Limit helped scientists study the effects of cellular aging on human populations from embryonic development to death, including the discovery of the effects of shortening repetitive sequences of DNA, called telomeres, on the ends of chromosomes. Elizabeth Blackburn, Jack Szostak and Carol Greider received the Nobel Prize in Physiology or Medicine in 2009 for their work on genetic structures related to the Hayflick Limit.

Alexis Carrel was a doctor and researcher who studied tissue cultures. He continued Ross Granville Harrison's research and produced many improvements in the field of tissue culture and surgery. He was the recipient of the 1912 Nobel Prize in Physiology or Medicine for his development of surgical techniques to repair blood vessels. Carrel was born on 28 June 1873 in Sainte-Foy-les-Lyon, France, to Anne-Marie Ricard and Alexis Carrel Billiard. His father died when he was five years old. Carrel earned a bachelor's degree in letters in 1889 and another in science in 1890 from St. Joseph's Day School in Lyon, France. He entered medical school at the age of seventeen and was regarded as a good but not exceptional student. The assassination of Sadi Carnot, a French politician visiting Lyon who was stabbed in the abdomen and died from the loss of blood, further interested him in surgery.

In an effort to develop tissue culture techniques for long-term tissue cultivation, French surgeon and biologist Alexis Carrel, and his associates, produced and maintained a series of chick heart tissue cultures at the Rockefeller Institute in New York City. From 1912 to 1946, this series of chick heart tissue cultures remained alive and dividing. Since the duration of this culture greatly exceeded the normal chick life span, the cells were deemed immortal. Although this conclusion was challenged by further experiments in the 1960s, the publicity surrounding the immortal chick heart tissue significantly influenced the concept of cell immortality and cellular aging from the 1920s through the 1960s. Carrel's experiment convinced many biologists to accept immortality as an intrinsic property of all cells, not just the cell line through which genetic material is passed to offspring, called the germ line. Consequently, the phenomenon of cellular aging was regarded not as an intrinsic characteristic, but was attributed to external factors such as the accumulation of waste products within the cell.

In the 1910s, Alexis Carrel, a French surgeon and biologist, concluded that cells are intrinsically immortal. His claim was based on chick-heart tissue cultures in his laboratory that seemed to be able to proliferate forever. Carrel's ideas about cellular immortality convinced his many contemporaries that cells could be maintained indefinitely. In the 1960s, however, Carrel's thesis about cell immortality was put into question by the discovery that human diploid cells can only proliferate for a finite period. As it was gradually recognized that chick cells only have a finite proliferative life span in vitro as well, historians and scientists alike attempted to identify experimental errors that could have led to the extremely long life of Carrel's "immortal" chick-heart tissue cultures. Those reassessments not only point out potential experimental mistakes in pioneer tissue culture work in the early twentieth century, but are also relevant to current discussions about the different life spans of germ line cells, embryonic and adult stem cells, normal somatic cells, and cancer cells.

Alexis Carrel, the prominent French surgeon, biologist, and 1912 Nobel Prize laureate for Physiology or Medicine, was one of the pioneers in developing and modifying tissue culture techniques. The publicized work of Carrel and his associates at the Rockefeller Institute established the practice of long-term tissue culture for a wide variety of cells. At the same time, some aspects of their work complicated the operational procedures of tissue culture. Thus Carrel's legacy had a mixed influence on the development of tissue culture techniques, which have been widely used in the fields of embryology and stem cell research.

Leonard Hayflick studied the processes by which cells age during the twentieth and twenty-first centuries in the United States. In 1961 at the Wistar Institute in the US, Hayflick researched a phenomenon later called the Hayflick Limit, or the claim that normal human cells can only divide forty to sixty times before they cannot divide any further. Researchers later found that the cause of the Hayflick Limit is the shortening of telomeres, or portions of DNA at the ends of chromosomes that slowly degrade as cells replicate. Hayflick used his research on normal embryonic cells to develop a vaccine for polio, and from HayflickÕs published directions, scientists developed vaccines for rubella, rabies, adenovirus, measles, chickenpox and shingles.

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