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	<div class="center"><h1>Timeline of Discoveries in the Science of Aging</h1><div class="addthis_responsive_sharing"></div>A History on The Biology of Aging
<br><font size="-2"><a href="soa-timeline-references.html">Click for a collapsed view of the timeline</a></font><br><br><a href="gompertz-aging-human-mortality"><div class="timelineother"><div class="gallerycontainer"><a class="thumbnail" href="gompertz-aging-human-mortality.html"><b><font size="-1">June, 16,</font>1825 -
<div style="padding-right:15px; ">
<font size="-2">Benjamin Gompertz Finds that Mortality Increases Exponentially with Age</font>
<span><img style="float:left;" src="./images/Benjamin-Gompertz.jpg" width="80" height="100"><b>Benjamin Gompertz Finds that Mortality Increases Exponentially with Age</b>
<hr><b>Methods:</b> Empirically deriving the death rate from demographic data of England <br><b>Institution:</b> The Royal Society<br>
</a></span></div></div>
</b></div><a href="weismann-germ-line"><div class="timelinetwoother"><div class="gallerycontainer"><a class="thumbnailtwo" href="weismann-germ-line.html"><b>- 1892<font size="-1">, 01, January</font>
<div style="padding-left:15px;">
<font size="-2"> August Weismann Proposes a Difference Between Germ Line and Somatic Cells</font>
<span><img style="float:left;" src="./images/August-Weismann.jpg" width="80" height="100"><b> August Weismann Proposes a Difference Between Germ Line and Somatic Cells</b>
<hr><b>Methods:</b> By drawing upon current research and contrasting it to the current dogma under Darwin's theory of pangenises.<br><b>Institution:</b> University of Freiburg-in-Baden<br>
</a></span></div></div>
</b></div><a href="clive-mccay-prolonging-lifespan-caloric-restriction"><div class="timelineother"><div class="gallerycontainer"><a class="thumbnail" href="clive-mccay-prolonging-lifespan-caloric-restriction.html"><b><font size="-1">November, 01,</font>1934 -
<div style="padding-right:15px; ">
<font size="-2">Clive McCay Discovers that Calorie Restriction Extends Life-Span in Rats</font>
<span><img style="float:left;" src="./images/Clive-McCay.jpg" width="80" height="100"><b>Clive McCay Discovers that Calorie Restriction Extends Life-Span in Rats</b>
<hr><b>Methods:</b> Feeding one group of rats a lower calorie diet than another group, and observing that the under-fed group lived longer<br><b>Institution:</b> Cornell University<br>
</a></span></div></div>
</b></div><a href="medawar-theory-mutation-accumulation"><div class="timelinetwoother"><div class="gallerycontainer"><a class="thumbnailtwo" href="medawar-theory-mutation-accumulation.html"><b>- 1952<font size="-1">, 01, January</font>
<div style="padding-left:15px;">
<font size="-2">Peter Medawar proposes that aging evolved into existence since the force of natural selection diminishes after reproduction</font>
<span><img style="float:left;" src="./images/Peter-Medawar.jpg" width="80" height="100"><b>Peter Medawar proposes that aging evolved into existence since the force of natural selection diminishes after reproduction</b>
<hr><b>Institution:</b> University College London<br>
</a></span></div></div>
</b></div><a href="harman-free-radical-theory"><div class="timelineother"><div class="gallerycontainer"><a class="thumbnail" href="harman-free-radical-theory.html"><b><font size="-1">July, 1,</font>1956 -
<div style="padding-right:15px; ">
<font size="-2">Denham Harman proposes the free radical theory of aging</font>
<span><img style="float:left;" src="./images/Denham-Harman.jpg" width="80" height="100"><b>Denham Harman proposes the free radical theory of aging</b>
<hr><b>Methods:</b> Harman's logic proceeds from three observations: (1) irradiation causes premature aging; (2) irradiation creates oxygen radicals, which may mediate its effects; and (3) cells produce oxygen radicals under normal conditions. From these premises, he theorized that aging could be caused by endogenously generated oxygen radicals.<br><b>Institution:</b> University of California - Berkeley<br>
</a></span></div></div>
</b></div><a href="williams-antagonistic-pleiotropy"><div class="timelinetwoother"><div class="gallerycontainer"><a class="thumbnailtwo" href="williams-antagonistic-pleiotropy.html"><b>- 1957<font size="-1">, 26, February</font>
<div style="padding-left:15px;">
<font size="-2">George C. Williams expands on the theory of antagonistic pleiotropy</font>
<span><img style="float:left;" src="./images/George-Williams.jpg" width="80" height="100"><b>George C. Williams expands on the theory of antagonistic pleiotropy</b>
<hr><b>Methods:</b> By defining assumptions and providing set expectations<br><b>Institution:</b> Michigan State University<br>
</a></span></div></div>
</b></div><a href="hayflick-moorhead-senescence-telomeres"><div class="timelineother"><div class="gallerycontainer"><a class="thumbnail" href="hayflick-moorhead-senescence-telomeres.html"><b><font size="-1">May, 15,</font>1961 -
<div style="padding-right:15px; ">
<font size="-2">  Hayflick and Moorhead Characterize Cellular Senescence in Primary Human Cells</font>
<span><img style="float:left;" src="./images/Leonard-Hayflick.jpg" width="80" height="100"><b>  Hayflick and Moorhead Characterize Cellular Senescence in Primary Human Cells</b>
<hr><b>Methods:</b> The authors characterized 25 strains of human cells over the course of long-term growth in culture. <br><b>Institution:</b> Wistar Institute<br>
</a></span></div></div>
</b></div><a href="orgel-error-catastrophe"><div class="timelinetwoother"><div class="gallerycontainer"><a class="thumbnailtwo" href="orgel-error-catastrophe.html"><b>- 1963<font size="-1">, 15, February</font>
<div style="padding-left:15px;">
<font size="-2">   Leslie Orgel Proposes the "Error Catastrophe" Theory of Aging</font>
<span><img style="float:left;" src="./images/Leslie-Orgel.jpg" width="80" height="100"><b>   Leslie Orgel Proposes the "Error Catastrophe" Theory of Aging</b>
<hr><b>Methods:</b> Orgel hypothesizes that translation errors could decrease the fidelity of translation, setting in motion a vicious cycle ultimately causing a critical failure of the gene expression machinery; he further proposes that the resulting loss of cell viability could be a contributing factor in the aging process.<br><b>Institution:</b> University of Cambridge<br>
</a></span></div></div>
</b></div><a href="hayflick-cells-limited-lifespan"><div class="timelineother"><div class="gallerycontainer"><a class="thumbnail" href="hayflick-cells-limited-lifespan.html"><b><font size="-1">March, 01,</font>1965 -
<div style="padding-right:15px; ">
<font size="-2">Leonard Hayflick proposes the Cellular Theory of Aging based on the limited replicative lifespan of primary human cells</font>
<span><img style="float:left;" src="./images/Leonard-Hayflick.jpg" width="80" height="100"><b>Leonard Hayflick proposes the Cellular Theory of Aging based on the limited replicative lifespan of primary human cells</b>
<hr><b>Methods:</b> Normal diploid cells were serially passaged in culture until they stopped dividing<br><b>Institution:</b> Wistar Institute<br>
</a></span></div></div>
</b></div><a href="gershon-inactive-enzyme-molecules"><div class="timelinetwoother"><div class="gallerycontainer"><a class="thumbnailtwo" href="gershon-inactive-enzyme-molecules.html"><b>- 1970<font size="-1">, 19, September</font>
<div style="padding-left:15px;">
<font size="-2">   David and Harriet Gershon Find Accumulation of Non-active Enzymes in Older Nematodes</font>
<span><img style="float:left;" src="./images/David-Gershon.jpg" width="80" height="100"><b>   David and Harriet Gershon Find Accumulation of Non-active Enzymes in Older Nematodes</b>
<hr><b>Methods:</b> Using immunological techniques to detect catalytically altered or inactive enzyme molecules in the nematode Turbatrix aceti<br><b>Institution:</b> Israel Institute of Technology<br>
</a></span></div></div>
</b></div><a href="walford-middle-age-dietary-restriction-lifespan-cancer"><div class="timelineother"><div class="gallerycontainer"><a class="thumbnail" href="walford-middle-age-dietary-restriction-lifespan-cancer.html"><b><font size="-1">March, 12,</font>1982 -
<div style="padding-right:15px; ">
<font size="-2">Richard Weindruch finds that adult mice on dietary restriction live 20% longer and have lower cancer rates</font>
<span><img style="float:left;" src="./images/Richard-Weindruch.jpg" width="80" height="100"><b>Richard Weindruch finds that adult mice on dietary restriction live 20% longer and have lower cancer rates</b>
<hr><b>Methods:</b> By feeding a control group a normal diet and another group a high nutrient calorie restricted diet and observing which lived longer<br><b>Institution:</b> University of California at Los Angeles<br>
</a></span></div></div>
</b></div><a href="cerami-maillard-aging"><div class="timelinetwoother"><div class="gallerycontainer"><a class="thumbnailtwo" href="cerami-maillard-aging.html"><b>- 1983<font size="-1">, 15, November</font>
<div style="padding-left:15px;">
<font size="-2">Anthony Cerami proposes the Maillard Reaction theory of aging stating that DNA accumulates damage due to nonenzymatic modification from sugars</font>
<span><b>Anthony Cerami proposes the Maillard Reaction theory of aging stating that DNA accumulates damage due to nonenzymatic modification from sugars</b>
<hr><b>Methods:</b> By incubating isolated DNA with high levels of glucose at 37 degrees Celsius, or body temperature, and recording changes in DNA through spectroscopy<br><b>Institution:</b> Rockefeller University<br>
</a></span></div></div>
</b></div><a href="rose-evolution-animal-senescence"><div class="timelineother"><div class="gallerycontainer"><a class="thumbnail" href="rose-evolution-animal-senescence.html"><b><font size="-1">February, 02,</font>1984 -
<div style="padding-right:15px; ">
<font size="-2">Michael R. Rose asserts that senescence can be postponed by natural selection</font>
<span><img style="float:left;" src="./images/Michael-Rose.jpg" width="80" height="100"><b>Michael R. Rose asserts that senescence can be postponed by natural selection</b>
<hr><b>Methods:</b> Reviewing past articles<br><b>Institution:</b> Dalhousie University<br>
</a></span></div></div>
</b></div><a href="greider-blackburn-discover-telomerase"><div class="timelinetwoother"><div class="gallerycontainer"><a class="thumbnailtwo" href="greider-blackburn-discover-telomerase.html"><b>- 1985<font size="-1">, 01, December</font>
<div style="padding-left:15px;">
<font size="-2">     Carol W. Greider and Elizabeth H. Blackburn Discover A Specific Telomere Terminal
Transferase Later Known as Telomerase</font>
<span><img style="float:left;" src="./images/Elizabeth-Blackburn.jpg" width="80" height="100"><b>     Carol W. Greider and Elizabeth H. Blackburn Discover A Specific Telomere Terminal
Transferase Later Known as Telomerase</b>
<hr><b>Methods:</b> By obtaining extracts of Tetrahymena during macronuclear development when new telomeres are generated<br><b>Institution:</b> University of California - Berkeley<br>
</a></span></div></div>
</b></div><a href="glucocorticoid-cascade-hypothesis"><div class="timelineother"><div class="gallerycontainer"><a class="thumbnail" href="glucocorticoid-cascade-hypothesis.html"><b><font size="-1">January, 01,</font>1986 -
<div style="padding-right:15px; ">
<font size="-2">         Robert M. Sapolsky, Lewis C. Krey, and Bruce S. McEwen Propose the Glucocorticoid Cascade Hypothesis of Aging</font>
<span><img style="float:left;" src="./images/Robert-Sapolsky.jpg" width="80" height="100"><b>         Robert M. Sapolsky, Lewis C. Krey, and Bruce S. McEwen Propose the Glucocorticoid Cascade Hypothesis of Aging</b>
<hr><b>Methods:</b> Citing past research indicating that older mice release greater amounts of glucocorticoids in response to stress, and that this release precipitates a cascade effect of hypersecretion<br><b>Institution:</b> The Salk Institute<br>
</a></span></div></div>
</b></div><a href="johnson-friedman-define-age-1-gene"><div class="timelinetwoother"><div class="gallerycontainer"><a class="thumbnailtwo" href="johnson-friedman-define-age-1-gene.html"><b>- 1988<font size="-1">, 05, February</font>
<div style="padding-left:15px;">
<font size="-2">     Thomas Johnson and David Friedman Define the AGE-1 gene</font>
<span><img style="float:left;" src="./images/Thomas-Johnson.jpg" width="80" height="100"><b>     Thomas Johnson and David Friedman Define the AGE-1 gene</b>
<hr><b>Methods:</b> By obtaining long-lived mutants of C. elegans and crossing them to screen which alleles resulted in longer lifespans<br><b>Institution:</b> University of California - Irvine<br>
</a></span></div></div>
</b></div><a href="kenyon-elegans-mutant-lives-twice-as-long"><div class="timelineother"><div class="gallerycontainer"><a class="thumbnail" href="kenyon-elegans-mutant-lives-twice-as-long.html"><b><font size="-1">September, 15,</font>1993 -
<div style="padding-right:15px; ">
<font size="-2">Cynthia Kenyon finds that mutations in daf-2 and daf-16 genes cause C. elegans worms to live over twice as long</font>
<span><img style="float:left;" src="./images/Cynthia-Kenyon.jpg" width="80" height="100"><b>Cynthia Kenyon finds that mutations in daf-2 and daf-16 genes cause C. elegans worms to live over twice as long</b>
<hr><b>Methods:</b> By regulating the formation of a dauer state, and observing how long the daf-2 mutated strains lived compared to other strains.<br><b>Institution:</b> University of California at San Francisco<br>
</a></span></div></div>
</b></div><a href="yeast-daughter-cells-shortened-lifespan"><div class="timelinetwoother"><div class="gallerycontainer"><a class="thumbnailtwo" href="yeast-daughter-cells-shortened-lifespan.html"><b>- 1994<font size="-1">, 01, December</font>
<div style="padding-left:15px;">
<font size="-2">  Brian K. Kennedy et al. Discover that the Daughter Cells of Old Yeast have Shorter Life-Spans than those of Younger Strains</font>
<span><img style="float:left;" src="./images/Brian-Kennedy.jpg" width="80" height="100"><b>  Brian K. Kennedy et al. Discover that the Daughter Cells of Old Yeast have Shorter Life-Spans than those of Younger Strains</b>
<hr><b>Methods:</b> By growing yeast and manipulating away their progeny, then counting how long both the first strain and the progeny lived<br><b>Institution:</b> Massachusetts Institute of Technology<br>
</a></span></div></div>
</b></div><a href="guarente-sir4-delay-yeast-aging"><div class="timelineother"><div class="gallerycontainer"><a class="thumbnail" href="guarente-sir4-delay-yeast-aging.html"><b><font size="-1">February, 10,</font>1995 -
<div style="padding-right:15px; ">
<font size="-2"> Brian K. Kennedy et al. Find that SIR4-42 Extends the Life Span of Yeast by 30% and May Play a Key Role in Regulating Aging</font>
<span><img style="float:left;" src="./images/Brian-Kennedy.jpg" width="80" height="100"><b> Brian K. Kennedy et al. Find that SIR4-42 Extends the Life Span of Yeast by 30% and May Play a Key Role in Regulating Aging</b>
<hr><b>Methods:</b> By culturing various strains of yeast under stressful conditions and analyzing the longest lived strains for key mutations<br><b>Institution:</b> Massachusetts Institute of Technology<br>
</a></span></div></div>
</b></div><a href="no-sensescence-in-hydra"><div class="timelinetwoother"><div class="gallerycontainer"><a class="thumbnailtwo" href="no-sensescence-in-hydra.html"><b>- 1997<font size="-1">, 04, August</font>
<div style="padding-left:15px;">
<font size="-2"> Daniel E. Mart�nez finds that Hydra do not show signs of Biological Aging</font>
<span><b> Daniel E. Mart�nez finds that Hydra do not show signs of Biological Aging</b>
<hr><b>Methods:</b> By keeping hydra alive for four years and plotting their mortality rates vs. that of similar sized organisms like Drosophila melanogaster (fruit fly)<br><b>Institution:</b> Pomona College<br>
</a></span></div></div>
</b></div><a href="extension-of-life-span-telomerase"><div class="timelineother"><div class="gallerycontainer"><a class="thumbnail" href="extension-of-life-span-telomerase.html"><b><font size="-1">January, 16,</font>1998 -
<div style="padding-right:15px; ">
<font size="-2">Andrea Bodnar, Michel Ouellette, et al. Find An Extension of Human Cell Life-Span with the Introduction of Telomerase</font>
<span><img style="float:left;" src="./images/Andrea-Bodnar.jpg" width="80" height="100"><b>Andrea Bodnar, Michel Ouellette, et al. Find An Extension of Human Cell Life-Span with the Introduction of Telomerase</b>
<hr><b>Methods:</b> By transfecting cells with vectors encoding the human telomerase catalytic subunit and noticing that the cells exceeded normal replicative lifespan<br><b>Institution:</b> Geron<br>
</a></span></div></div>
</b></div><a href="murphy-downstream-daf-16"><div class="timelinetwoother"><div class="gallerycontainer"><a class="thumbnailtwo" href="murphy-downstream-daf-16.html"><b>- 2003<font size="-1">, 17, July</font>
<div style="padding-left:15px;">
<font size="-2">Coleen Murphy et al. find genes downstream of daf-16 which regulate aging in C.elegans nematodes</font>
<span><img style="float:left;" src="./images/Coleen-Murphy.jpg" width="80" height="100"><b>Coleen Murphy et al. find genes downstream of daf-16 which regulate aging in C.elegans nematodes</b>
<hr><b>Methods:</b> By comparing transcriptional profiles of long lived c.elegans mutants vs. wild-type and identifying genes that were upregulated or downregulated<br><b>Institution:</b> University of California - San Francisco<br>
</a></span></div></div>
</b></div><a href="sinclair-howitz-resveratrol"><div class="timelineother"><div class="gallerycontainer"><a class="thumbnail" href="sinclair-howitz-resveratrol.html"><b><font size="-1">August, 24,</font>2003 -
<div style="padding-right:15px; ">
<font size="-2">     David A. Sinclair et al. Discover Three Classes of Molecules that Activate Sirtuins (Including Resveratrol)</font>
<span><img style="float:left;" src="./images/David-Sinclair.jpg" width="80" height="100"><b>     David A. Sinclair et al. Discover Three Classes of Molecules that Activate Sirtuins (Including Resveratrol)</b>
<hr><b>Methods:</b> By screening small molecule libraries for molecules which could modulate sirtuin activity<br><b>Institution:</b> Harvard Medical School<br>
</a></span></div></div>
</b></div><a href="sharpless-ink4a-arf-locus-senescence"><div class="timelinetwoother"><div class="gallerycontainer"><a class="thumbnailtwo" href="sharpless-ink4a-arf-locus-senescence.html"><b>- 2004<font size="-1">, 01, November</font>
<div style="padding-left:15px;">
<font size="-2">Norman E. Sharpless finds that expression of the Ink4a/Arf locus is a biomarker of aging</font>
<span><img style="float:left;" src="./images/Norman-Sharpless.jpg" width="80" height="100"><b>Norman E. Sharpless finds that expression of the Ink4a/Arf locus is a biomarker of aging</b>
<hr><b>Methods:</b> Using PCR assays on the expression of cyclin-dependent kinase inhibitor family members to find variations in expression of the gene p16<sup><font size="-2">INK4a</font></sup><br><b>Institution:</b> UNC Chapel Hill<br>
</a></span></div></div>
</b></div><a href="Sharpless-DePinho-stem-cell-aging"><div class="timelineother"><div class="gallerycontainer"><a class="thumbnail" href="Sharpless-DePinho-stem-cell-aging.html"><b><font size="-1">January, 01,</font>2004 -
<div style="padding-right:15px; ">
<font size="-2">       Norman Sharpless and Ronald Pinho Propose the Stem-Cell Theory of Aging</font>
<span><img style="float:left;" src="./images/Norman-Sharpless.jpg" width="80" height="100"><b>       Norman Sharpless and Ronald Pinho Propose the Stem-Cell Theory of Aging</b>
<hr><b>Methods:</b> By citing past evidence their reasoning follows that a cell with genomic instability will either undergo senescence/apoptosis, or proliferate into cancer, depending on the integrity and ability of various cell cycle check points.<br><b>Institution:</b> UNC Chapel Hill<br>
</a></span></div></div>
</b></div><a href="life-span-tor-sch9"><div class="timelinetwoother"><div class="gallerycontainer"><a class="thumbnailtwo" href="life-span-tor-sch9.html"><b>- 2005<font size="-1">, 18, November</font>
<div style="padding-left:15px;">
<font size="-2">   Matt Kaeberlein et al. propose that the TOR and Sch9 kinases regulate longevity in yeast</font>
<span><img style="float:left;" src="./images/Matt-Kaeberlein.jpg" width="80" height="100"><b>   Matt Kaeberlein et al. propose that the TOR and Sch9 kinases regulate longevity in yeast</b>
<hr><b>Methods:</b> Large-scale analysis of 564 gene deletions which yielded 6 genes corresponding to the TOR and Sch9 pathways<br>
</a></span></div></div>
</b></div><a href="bodnar-telomerase-sea-urchins"><div class="timelineother"><div class="gallerycontainer"><a class="thumbnail" href="bodnar-telomerase-sea-urchins.html"><b><font size="-1">July, 24,</font>2006 -
<div style="padding-right:15px; ">
<font size="-2"> Andrea Bodnar Finds a Lack of Age-Associated Telomere Shortening in Long and Short Lived Sea Urchins</font>
<span><img style="float:left;" src="./images/Andrea-Bodnar.jpg" width="80" height="100"><b> Andrea Bodnar Finds a Lack of Age-Associated Telomere Shortening in Long and Short Lived Sea Urchins</b>
<hr><b>Institution:</b> Bermuda Institute of Ocean Sciences<br>
</a></span></div></div>
</b></div><br><br>
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