Darwin, From Birth to Death Carl Zimmer Provost Lecture Series Stony Brook University February 11, 2011

Thursday, February 17, 2011

Thursday, February 17, 2011

When  the  views  entertained  in  this   volume  on  the  origin  of  species,  or   when  analogous  views  are  generally   admitted,  we  can  dimly  foresee  that   there  will  be  a  considerable  revolution   in  natural  history.

Thursday, February 17, 2011

Light  will  be  thrown  on  the   origin  of  man  and  his  history.

Thursday, February 17, 2011

W. Eugene Robinson, http://images.google.com/hosted/life/ Thursday, February 17, 2011

“For over forty years Darwin suffered intermittently from various combinations of symptoms such as malaise, vertigo, dizziness, muscle spasms and tremors, vomiting, cramps and colics, bloating and nocturnal intestinal gas, headaches, alterations of vision, severe tiredness, nervous exhaustion, dyspnea, skin problems such as blisters all over the scalp and eczema, crying, anxiety, sensation of impending death and loss of consciousness, fainting, tachycardia, insomnia, tinnitus, and depression.” http://en.wikipedia.org/wiki/Charles_Darwin's_health

Thursday, February 17, 2011

Thursday, February 17, 2011

Randolph Nesse

Richard Dawkins Thursday, February 17, 2011

“Evolution: The Most Basic Science of Medicine”

George Williams 1926-2010

Thursday, February 17, 2011

Thursday, February 17, 2011

Life Branches Like A Tree

Thursday, February 17, 2011

Natural Selection

Based on Gregory, Evo Edu Outreach (2009) 2:156–175 Thursday, February 17, 2011

Natural Selection Generation 1

Based on Gregory, Evo Edu Outreach (2009) 2:156–175 Thursday, February 17, 2011

Non-random survival

X X X X Natural Selection

Generation 1

Based on Gregory, Evo Edu Outreach (2009) 2:156–175 Thursday, February 17, 2011

Reproduction and mutation Non-random survival

X X X X

Generation 2

Natural Selection Generation 1

Based on Gregory, Evo Edu Outreach (2009) 2:156–175 Thursday, February 17, 2011

Reproduction and mutation Non-random survival

X X X X Natural Selection

Generation 2

Non-random survival

X X X X

Generation 1

Based on Gregory, Evo Edu Outreach (2009) 2:156–175 Thursday, February 17, 2011

Reproduction and mutation Non-random survival

Generation 2

X X X X Natural Selection

Non-random survival

X X X X

Generation 1 Reproduction and mutation Generation 3 Based on Gregory, Evo Edu Outreach (2009) 2:156–175 Thursday, February 17, 2011

Reproduction and mutation Non-random survival

Generation 2

X X X X Natural Selection

Generation 1

Generation X

Non-random survival

Many generations of mutation and natural selection

X X X X

Reproduction and mutation Generation 3 Based on Gregory, Evo Edu Outreach (2009) 2:156–175

Thursday, February 17, 2011

Why do animals get old and die?

America Perales http://www.flickr.com/photos/americaperales/3505343905/ Thursday, February 17, 2011

Beneficial death?

Photo by jschinker: flickr.com/photos/63873121@N00/3889483525/ Thursday, February 17, 2011

American Am Nat. Jan;177(1):v-vii. Figure 1: Linda Cranford Lillegraven, now an artist Naturalist in Laramie,2011 Wyoming, was 2011 a graduate student at the University of Utah who drew caricatures of many seminar speakers. She drew this likeness of George (ca. 1976) when he gave a talk on eels at the university. George was fond of this image and used February it in correspondence for much of the last three decades. Thursday, 17, 2011

“My reaction was that if Emerson’s presentation was acceptable biology, I would prefer another calling.”

Figure 1: Linda Cranford Lillegraven, now an artist in Laramie, Wyoming, was a graduate student at the University of Utah who drew caricatures of many seminar speakers. She drew this likeness of George (ca. 1976) when he gave a talk on eels at the university. George was fond of this image and used it in correspondence for much of the last three decades.

century” hit Stony Brook, and the university shut down and sent everyone home at noon. The wind was fierce and snow was falling, but George and Doris insisted on walking home. Doris said she didn’t mind the walk because the conditions were not nearly so bad as they had been when she and George lived in Iceland. For many years, George spent most of the summer at his cabin in Ontario, Canada; Doris’s appointment only allowed her three weeks there each summer. It was set back about two miles off the Toronto-Ottawa highway, about equidistant between the two cities. It was fairly remote and had no phone. George and Doris liked to bathe in a cold stream about 100 yards from the cabin. Colleagues who visited George were treated to long talks, a little boating on the creek, and occasionally catching pike for dinner. George was gentle, quiet, and intensely thoughtful. He was also very approachable. It was not uncommon to sit down with him and toss out an initial idea for discussion. Then, when his ensuing silence grew uncomfortably long, one would toss out another idea, ultimately another, and then another. When he finally spoke, it became clear that he was still thinking deeply about the first or second idea.

Adaptation and Natural Selection, 1996, p. ix Thursday, February 17, 2011

He exhibited a modesty that was several standard deviations from the norm, at least the norm in academia. When Maynard Smith’s Evolution of Sex (1978, Cambridge University Press) appeared just three years after George’s book on the same topic (Sex and Evolution, Princeton University Press), George declared in his review that Maynard Smith’s book had made his own obsolete. In 2000, when George was invited to speak on Darwinian medicine at a symposium on applied evolution (George was the cofounder of the field, after all), he responded that the organizers should get someone who knew something about the topic. He did not hesitate to give credit to others for his inspiration. For example, he attributed his inspiration for Darwinian medicine to Paul Ewald and Margie Profet, and in the preface to the 1996 reprinting of his 1966 book, George gave credit to several people who had influenced his thinking. George also had a sense of humor about himself (fig. 1) and about his mistakes. For example, his prediction in 1966 that sex ratio was “solved” came just a year before the topic started on a meteoric rise in popularity, with Hamilton’s “extraordinary sex ratios” paper and the later

http://www.flickr.com/photos/roblee/102165753/ Thursday, February 17, 2011

Thursday, February 17, 2011

Figure 1: Linda Cranford Lillegraven, now an artist in Laramie, Wyoming, was a graduate student at the University of Utah who drew caricatures of many seminar speakers. She drew this likeness of George (ca. 1976) when he gave a talk on eels at the university. George was fond of this image and used it in correspondence for much of the last three decades.

Thursday, February 17, 2011

century” hit Stony Brook, and the university shut down and sent everyone home at noon. The wind was fierce and snow was falling, but George and Doris insisted on walking home. Doris said she didn’t mind the walk because the conditions were not nearly so bad as they had been when she and George lived in Iceland. For many years, George spent most of the summer at his cabin in Ontario, Canada; Doris’s appointment only allowed her three weeks there each summer. It was set back about two miles off the Toronto-Ottawa highway, about equidistant between the two cities. It was fairly remote and had no phone. George and Doris liked to bathe in a cold stream about 100 yards from the cabin. Colleagues who visited George were treated to long talks, a little boating on the creek, and occasionally catching pike for dinner. George was gentle, quiet, and intensely thoughtful. He was also very approachable. It was not uncommon to sit down with him and toss out an initial idea for discussion. Then, when his ensuing silence grew uncomfortably long, one would toss out another idea, ultimately another, and then another. When he finally spoke, it became clear that

He exhibited a modesty that was several standard deviations from the norm, at least the norm in academia. When Maynard Smith’s Evolution of Sex (1978, Cambridge University Press) appeared just three years after George’s book on the same topic (Sex and Evolution, Princeton University Press), George declared in his review that Maynard Smith’s book had made his own obsolete. In 2000, when George was invited to speak on Darwinian medicine at a symposium on applied evolution (George was the cofounder of the field, after all), he responded that the organizers should get someone who knew something about the topic. He did not hesitate to give credit to others for his inspiration. For example, he attributed his inspiration for Darwinian medicine to Paul Ewald and Margie Profet, and in the preface to the 1996 reprinting of his 1966 book, George gave credit to several people who had influenced his thinking. George also had a sense of humor about himself (fig. 1) and about his mistakes. For example, his prediction in 1966 that sex ratio was “solved” came just a year before the topic started on a meteoric rise in popularity, with

America Perales http://www.flickr.com/photos/americaperales/3505343905/ Thursday, February 17, 2011

“Walking home from the lecture with my wife, Doris, I regaled her with my unhappiness about the lecture and proposed the obvious idea...

Adaptation and Natural Selection, 1996, p. ix America Perales http://www.flickr.com/photos/americaperales/3505343905/ Thursday, February 17, 2011

...that selection among indivivuals in any population would be biased in favor of the young, as long as the likelihood of living to age X was greater than to age X+1”

Adaptation and Natural Selection, 1996, p. ix America Perales http://www.flickr.com/photos/americaperales/3505343905/ Thursday, February 17, 2011

The Williams eel, the fish that never gets old Figure 1: Linda Cranford Lillegraven, now an artist in Laramie, Wyoming, was a graduate student at the University of Utah who drew caricatures of many seminar speakers. She drew this likeness of George (ca. 1976) when he gave a talk on eels at the university. George was fond of this image and used it in correspondence for much of the last three decades.

century” hit Stony Brook, and the university shut down and sent everyone home at noon. The wind was fierce and snow was falling, but George and Doris insisted on walking home. Doris said she didn’t mind the walk because the conditions were not nearly so bad as they had been when she and George lived in Iceland. For many years, George spent most of the summer at his cabin in Ontario, Canada; Doris’s appointment only allowed her three weeks there each summer. It was set back about two miles off the Toronto-Ottawa highway, about equidistant between the two cities. It was fairly remote and had no phone. George and Doris liked to bathe in a cold stream about 100 yards from the cabin. Colleagues who visited George were treated to long talks, a little boating on the creek, and occasionally catching pike for dinner. George was gentle, quiet, and intensely thoughtful. He was also very approachable. It was not uncommon to sit down with him and toss out an initial idea for discussion. Then, when his ensuing silence grew uncomfortably long, one would toss out another idea, ultimately another, and then another. When he finally spoke, it became clear that he was still thinking deeply about the first or second idea.

Thursday, February 17, 2011

He exhibited a modesty that was several standard deviations from the norm, at least the norm in academia. When Maynard Smith’s Evolution of Sex (1978, Cambridge University Press) appeared just three years after George’s book on the same topic (Sex and Evolution, Princeton University Press), George declared in his review that Maynard Smith’s book had made his own obsolete. In 2000, when George was invited to speak on Darwinian medicine at a symposium on applied evolution (George was the cofounder of the field, after all), he responded that the organizers should get someone who knew something about the topic. He did not hesitate to give credit to others for his inspiration. For example, he attributed his inspiration for Darwinian medicine to Paul Ewald and Margie Profet, and in the preface to the 1996 reprinting of his 1966 book, George gave credit to several people who had influenced his thinking. George also had a sense of humor about himself (fig. 1) and about his mistakes. For example, his prediction in 1966 that sex ratio was “solved” came just a year before the topic started on a meteoric rise in popularity, with Hamilton’s “extraordinary sex ratios” paper and the later

Most Williams eels will have relatively young parents Figure 1: Linda Cranford Lillegraven, now an artist in Laramie, Wyoming, was a graduate student at the University of Utah who drew caricatures of many seminar speakers. She drew this likeness of George (ca. 1976) when he gave a talk on eels at the university. George was fond of this image and used it in correspondence for much of the last three decades.

century” hit Stony Brook, and the university shut down and sent everyone home at noon. The wind was fierce and snow was falling, but George and Doris insisted on walking home. Doris said she didn’t mind the walk because the conditions were not nearly so bad as they had been when she and George lived in Iceland. For many years, George spent most of the summer at his cabin in Ontario, Canada; Doris’s appointment only allowed her three weeks there each summer. It was set back about two miles off the Toronto-Ottawa highway, about equidistant between the two cities. It was fairly remote and had no phone. George and Doris liked to bathe in a cold stream about 100 yards from the cabin. Colleagues who visited George were treated to long talks, a little boating on the creek, and occasionally catching pike for dinner. George was gentle, quiet, and intensely thoughtful. He was also very approachable. It was not uncommon to sit down with him and toss out an initial idea for discussion. Then, when his ensuing silence grew uncomfortably long, one would toss out another idea, ultimately another, and then another. When he finally spoke, it became clear that he was still thinking deeply about the first or second idea.

Thursday, February 17, 2011

He exhibited a modesty that was several standard deviations from the norm, at least the norm in academia. When Maynard Smith’s Evolution of Sex (1978, Cambridge University Press) appeared just three years after George’s book on the same topic (Sex and Evolution, Princeton University Press), George declared in his review that Maynard Smith’s book had made his own obsolete. In 2000, when George was invited to speak on Darwinian medicine at a symposium on applied evolution (George was the cofounder of the field, after all), he responded that the organizers should get someone who knew something about the topic. He did not hesitate to give credit to others for his inspiration. For example, he attributed his inspiration for Darwinian medicine to Paul Ewald and Margie Profet, and in the preface to the 1996 reprinting of his 1966 book, George gave credit to several people who had influenced his thinking. George also had a sense of humor about himself (fig. 1) and about his mistakes. For example, his prediction in 1966 that sex ratio was “solved” came just a year before the topic started on a meteoric rise in popularity, with Hamilton’s “extraordinary sex ratios” paper and the later

A beneficial mutation will have a big effect if it helps young eels survive and have more offspring

Figure 1: Linda Cranford Lillegraven, now an artist in Laramie, Wyoming, was a graduate student at the University of Utah who drew caricatures of many seminar speakers. She drew this likeness of George (ca. 1976) when he gave a talk on eels at the university. George was fond of this image and used it in correspondence for much of the last three decades.

century” hit Stony Brook, and the university shut down and sent everyone home at noon. The wind was fierce and snow was falling, but George and Doris insisted on walking home. Doris said she didn’t mind the walk because the conditions were not nearly so bad as they had been when she and George lived in Iceland. For many years, George spent most of the summer at his cabin in Ontario, Canada; Doris’s appointment only allowed her three weeks there each summer. It was set back about two miles off the Toronto-Ottawa highway, about equidistant between the two cities. It was fairly remote and had no phone. George and Doris liked to bathe in a cold stream about 100 yards from the cabin. Colleagues who visited George were treated to long talks, a little boating on the creek, and occasionally catching pike for dinner. George was gentle, quiet, and intensely thoughtful. He was also very approachable. It was not uncommon to sit down with him and toss out an initial idea for discussion. Then, when his ensuing silence grew uncomfortably long, one would toss out another idea, ultimately another, and then another. When he finally spoke, it became clear that he was still thinking deeply about the first or second idea.

Thursday, February 17, 2011

He exhibited a modesty that was several standard deviations from the norm, at least the norm in academia. When Maynard Smith’s Evolution of Sex (1978, Cambridge University Press) appeared just three years after George’s book on the same topic (Sex and Evolution, Princeton University Press), George declared in his review that Maynard Smith’s book had made his own obsolete. In 2000, when George was invited to speak on Darwinian medicine at a symposium on applied evolution (George was the cofounder of the field, after all), he responded that the organizers should get someone who knew something about the topic. He did not hesitate to give credit to others for his inspiration. For example, he attributed his inspiration for Darwinian medicine to Paul Ewald and Margie Profet, and in the preface to the 1996 reprinting of his 1966 book, George gave credit to several people who had influenced his thinking. George also had a sense of humor about himself (fig. 1) and about his mistakes. For example, his prediction in 1966 that sex ratio was “solved” came just a year before the topic started on a meteoric rise in popularity, with Hamilton’s “extraordinary sex ratios” paper and the later

A beneficial mutation that affects older eels will have less effect because most eels are dead by then Figure 1: Linda Cranford Lillegraven, now an artist in Laramie, Wyoming, was a graduate student at the University of Utah who drew caricatures of many seminar speakers. She drew this likeness of George (ca. 1976) when he gave a talk on eels at the university. George was fond of this image and used it in correspondence for much of the last three decades.

century” hit Stony Brook, and the university shut down and sent everyone home at noon. The wind was fierce and snow was falling, but George and Doris insisted on walking home. Doris said she didn’t mind the walk because the conditions were not nearly so bad as they had been when she and George lived in Iceland. For many years, George spent most of the summer at his cabin in Ontario, Canada; Doris’s appointment only allowed her three weeks there each summer. It was set back about two miles off the Toronto-Ottawa highway, about equidistant between the two cities. It was fairly remote and had no phone. George and Doris liked to bathe in a cold stream about 100 yards from the cabin. Colleagues who visited George were treated to long talks, a little boating on the creek, and occasionally catching pike for dinner. George was gentle, quiet, and intensely thoughtful. He was also very approachable. It was not uncommon to sit down with him and toss out an initial idea for discussion. Then, when his ensuing silence grew uncomfortably long, one would toss out another idea, ultimately another, and then another. When he finally spoke, it became clear that he was still thinking deeply about the first or second idea.

Thursday, February 17, 2011

He exhibited a modesty that was several standard deviations from the norm, at least the norm in academia. When Maynard Smith’s Evolution of Sex (1978, Cambridge University Press) appeared just three years after George’s book on the same topic (Sex and Evolution, Princeton University Press), George declared in his review that Maynard Smith’s book had made his own obsolete. In 2000, when George was invited to speak on Darwinian medicine at a symposium on applied evolution (George was the cofounder of the field, after all), he responded that the organizers should get someone who knew something about the topic. He did not hesitate to give credit to others for his inspiration. For example, he attributed his inspiration for Darwinian medicine to Paul Ewald and Margie Profet, and in the preface to the 1996 reprinting of his 1966 book, George gave credit to several people who had influenced his thinking. George also had a sense of humor about himself (fig. 1) and about his mistakes. For example, his prediction in 1966 that sex ratio was “solved” came just a year before the topic started on a meteoric rise in popularity, with Hamilton’s “extraordinary sex ratios” paper and the later

The same is true for harmful mutations

Figure 1: Linda Cranford Lillegraven, now an artist in Laramie, Wyoming, was a graduate student at the University of Utah who drew caricatures of many seminar speakers. She drew this likeness of George (ca. 1976) when he gave a talk on eels at the university. George was fond of this image and used it in correspondence for much of the last three decades.

century” hit Stony Brook, and the university shut down and sent everyone home at noon. The wind was fierce and snow was falling, but George and Doris insisted on walking home. Doris said she didn’t mind the walk because the conditions were not nearly so bad as they had been when she and George lived in Iceland. For many years, George spent most of the summer at his cabin in Ontario, Canada; Doris’s appointment only allowed her three weeks there each summer. It was set back about two miles off the Toronto-Ottawa highway, about equidistant between the two cities. It was fairly remote and had no phone. George and Doris liked to bathe in a cold stream about 100 yards from the cabin. Colleagues who visited George were treated to long talks, a little boating on the creek, and occasionally catching pike for dinner. George was gentle, quiet, and intensely thoughtful. He was also very approachable. It was not uncommon to sit down with him and toss out an initial idea for discussion. Then, when his ensuing silence grew uncomfortably long, one would toss out another idea, ultimately another, and then another. When he finally spoke, it became clear that he was still thinking deeply about the first or second idea.

Thursday, February 17, 2011

He exhibited a modesty that was several standard deviations from the norm, at least the norm in academia. When Maynard Smith’s Evolution of Sex (1978, Cambridge University Press) appeared just three years after George’s book on the same topic (Sex and Evolution, Princeton University Press), George declared in his review that Maynard Smith’s book had made his own obsolete. In 2000, when George was invited to speak on Darwinian medicine at a symposium on applied evolution (George was the cofounder of the field, after all), he responded that the organizers should get someone who knew something about the topic. He did not hesitate to give credit to others for his inspiration. For example, he attributed his inspiration for Darwinian medicine to Paul Ewald and Margie Profet, and in the preface to the 1996 reprinting of his 1966 book, George gave credit to several people who had influenced his thinking. George also had a sense of humor about himself (fig. 1) and about his mistakes. For example, his prediction in 1966 that sex ratio was “solved” came just a year before the topic started on a meteoric rise in popularity, with Hamilton’s “extraordinary sex ratios” paper and the later

The same is true for harmful mutations

Figure 1: Linda Cranford Lillegraven, now an artist in Laramie, Wyoming, was a graduate student at the University of Utah who drew caricatures of many seminar speakers. She drew this likeness of George (ca. 1976) when he gave a talk on eels at the university. George was fond of this image and used it in correspondence for much of the last three decades.

century” hit Stony Brook, and the university shut down and sent everyone home at noon. The wind was fierce and snow was falling, but George and Doris insisted on walking home. Doris said she didn’t mind the walk because the conditions were not nearly so bad as they had been when she and George lived in Iceland. For many years, George spent most of the summer at his cabin in Ontario, Canada; Doris’s appointment only allowed her three weeks there each summer. It was set back about two miles off the Toronto-Ottawa highway, about equidistant between the two cities. It was fairly remote and had no phone. George and Doris liked to bathe in a cold stream about 100 yards from the cabin. Colleagues who visited George were treated to long talks, a little boating on the creek, and occasionally catching pike for dinner. George was gentle, quiet, and intensely thoughtful. He was also very approachable. It was not uncommon to sit down with him and toss out an initial idea for discussion. Then, when his ensuing silence grew uncomfortably long, one would toss out another idea, ultimately another, and then another. When he finally spoke, it became clear that he was still thinking deeply about the first or second idea.

Thursday, February 17, 2011

He exhibited a modesty that was several standard deviations from the norm, at least the norm in academia. When Maynard Smith’s Evolution of Sex (1978, Cambridge University Press) appeared just three years after George’s book on the same topic (Sex and Evolution, Princeton University Press), George declared in his review that Maynard Smith’s book had made his own obsolete. In 2000, when George was invited to speak on Darwinian medicine at a symposium on applied evolution (George was the cofounder of the field, after all), he responded that the organizers should get someone who knew something about the topic. He did not hesitate to give credit to others for his inspiration. For example, he attributed his inspiration for Darwinian medicine to Paul Ewald and Margie Profet, and in the preface to the 1996 reprinting of his 1966 book, George gave credit to several people who had influenced his thinking. George also had a sense of humor about himself (fig. 1) and about his mistakes. For example, his prediction in 1966 that sex ratio was “solved” came just a year before the topic started on a meteoric rise in popularity, with Hamilton’s “extraordinary sex ratios” paper and the later

Pleiotropy: two effects from one gene

Figure 1: Linda Cranford Lillegraven, now an artist in Laramie, Wyoming, was a graduate student at the University of Utah who drew caricatures of many seminar speakers. She drew this likeness of George (ca. 1976) when he gave a talk on eels at the university. George was fond of this image and used it in correspondence for much of the last three decades.

century” hit Stony Brook, and the university shut down and sent everyone home at noon. The wind was fierce and snow was falling, but George and Doris insisted on walking home. Doris said she didn’t mind the walk because the conditions were not nearly so bad as they had been when she and George lived in Iceland. For many years, George spent most of the summer at his cabin in Ontario, Canada; Doris’s appointment only allowed her three weeks there each summer. It was set back about two miles off the Toronto-Ottawa highway, about equidistant between the two cities. It was fairly remote and had no phone. George and Doris liked to bathe in a cold stream about 100 yards from the cabin. Colleagues who visited George were treated to long talks, a little boating on the creek, and occasionally catching pike for dinner. George was gentle, quiet, and intensely thoughtful. He was also very approachable. It was not uncommon to sit down with him and toss out an initial idea for discussion. Then, when his ensuing silence grew uncomfortably long, one would toss out another idea, ultimately another, and then another. When he finally spoke, it became clear that he was still thinking deeply about the first or second idea.

Thursday, February 17, 2011

He exhibited a modesty that was several standard deviations from the norm, at least the norm in academia. When Maynard Smith’s Evolution of Sex (1978, Cambridge University Press) appeared just three years after George’s book on the same topic (Sex and Evolution, Princeton University Press), George declared in his review that Maynard Smith’s book had made his own obsolete. In 2000, when George was invited to speak on Darwinian medicine at a symposium on applied evolution (George was the cofounder of the field, after all), he responded that the organizers should get someone who knew something about the topic. He did not hesitate to give credit to others for his inspiration. For example, he attributed his inspiration for Darwinian medicine to Paul Ewald and Margie Profet, and in the preface to the 1996 reprinting of his 1966 book, George gave credit to several people who had influenced his thinking. George also had a sense of humor about himself (fig. 1) and about his mistakes. For example, his prediction in 1966 that sex ratio was “solved” came just a year before the topic started on a meteoric rise in popularity, with Hamilton’s “extraordinary sex ratios” paper and the later

Antagonistic pleiotropy: one effect is positive, and one is negative

Figure 1: Linda Cranford Lillegraven, now an artist in Laramie, Wyoming, was a graduate student at the University of Utah who drew caricatures of many seminar speakers. She drew this likeness of George (ca. 1976) when he gave a talk on eels at the university. George was fond of this image and used it in correspondence for much of the last three decades.

century” hit Stony Brook, and the university shut down and sent everyone home at noon. The wind was fierce and snow was falling, but George and Doris insisted on walking home. Doris said she didn’t mind the walk because the conditions were not nearly so bad as they had been when she and George lived in Iceland. For many years, George spent most of the summer at his cabin in Ontario, Canada; Doris’s appointment only allowed her three weeks there each summer. It was set back about two miles off the Toronto-Ottawa highway, about equidistant between the two cities. It was fairly remote and had no phone. George and Doris liked to bathe in a cold stream about 100 yards from the cabin. Colleagues who visited George were treated to long talks, a little boating on the creek, and occasionally catching pike for dinner. George was gentle, quiet, and intensely thoughtful. He was also very approachable. It was not uncommon to sit down with him and toss out an initial idea for discussion. Then, when his ensuing silence grew uncomfortably long, one would toss out another idea, ultimately another, and then another. When he finally spoke, it became clear that he was still thinking deeply about the first or second idea.

Thursday, February 17, 2011

He exhibited a modesty that was several standard deviations from the norm, at least the norm in academia. When Maynard Smith’s Evolution of Sex (1978, Cambridge University Press) appeared just three years after George’s book on the same topic (Sex and Evolution, Princeton University Press), George declared in his review that Maynard Smith’s book had made his own obsolete. In 2000, when George was invited to speak on Darwinian medicine at a symposium on applied evolution (George was the cofounder of the field, after all), he responded that the organizers should get someone who knew something about the topic. He did not hesitate to give credit to others for his inspiration. For example, he attributed his inspiration for Darwinian medicine to Paul Ewald and Margie Profet, and in the preface to the 1996 reprinting of his 1966 book, George gave credit to several people who had influenced his thinking. George also had a sense of humor about himself (fig. 1) and about his mistakes. For example, his prediction in 1966 that sex ratio was “solved” came just a year before the topic started on a meteoric rise in popularity, with Hamilton’s “extraordinary sex ratios” paper and the later

Figure 1: Linda Cranford Lillegraven, now an artist in Laramie, Wyoming, was a graduate student at the University of Utah who drew caricatures of many seminar speakers. She drew this likeness of George (ca. 1976) when he gave a talk on eels at the university. George was fond of this image and used it in correspondence for much of the last three decades.

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century” hit Stony Brook, and the university shut down and sent everyone home at noon. The wind was fierce and snow was falling, but George and Doris insisted on walking home. Doris said she didn’t mind the walk because the conditions were not nearly so bad as they had been when she and George lived in Iceland. For many years, George spent most of the summer at his cabin in Ontario, Canada; Doris’s appointment only allowed her three weeks there each summer. It was set back about two miles off the Toronto-Ottawa highway, about equidistant between the two cities. It was fairly remote and had no phone. George and Doris liked to bathe in a cold stream about 100 yards from the cabin. Colleagues who visited George were treated to long talks, a little boating on the creek, and occasionally catching pike for dinner. George was gentle, quiet, and intensely thoughtful. He was also very approachable. It was not uncommon to sit down with him and toss out an initial idea for discussion. Then, when his ensuing silence grew uncomfortably long, one would toss out another idea, ultimately another, and then another. When he finally spoke, it became clear that he was still thinking deeply about the first or second idea.

Natural selection can favor mutations that are harmful in old age

Thursday, February 17, 2011

He exhibited a modesty that was several standard deviations from the norm, at least the norm in academia. When Maynard Smith’s Evolution of Sex (1978, Cambridge University Press) appeared just three years after George’s book on the same topic (Sex and Evolution, Princeton University Press), George declared in his review that Maynard Smith’s book had made his own obsolete. In 2000, when George was invited to speak on Darwinian medicine at a symposium on applied evolution (George was the cofounder of the field, after all), he responded that the organizers should get someone who knew something about the topic. He did not hesitate to give credit to others for his inspiration. For example, he attributed his inspiration for Darwinian medicine to Paul Ewald and Margie Profet, and in the preface to the 1996 reprinting of his 1966 book, George gave credit to several people who had influenced his thinking. George also had a sense of humor about himself (fig. 1) and about his mistakes. For example, his prediction in 1966 that sex ratio was “solved” came just a year before the topic started on a meteoric rise in popularity, with Hamilton’s “extraordinary sex ratios” paper and the later

The vigor of youth.... Figure 1: Linda Cranford Lillegraven, now an artist in Laramie, Wyoming, was a graduate student at the University of Utah who drew caricatures of many seminar speakers. She drew this likeness of George (ca. 1976) when he gave a talk on eels at the university. George was fond of this image and used it in correspondence for much of the last three decades.

century” hit Stony Brook, and the university shut down and sent everyone home at noon. The wind was fierce and snow was falling, but George and Doris insisted on walking home. Doris said she didn’t mind the walk because the conditions were not nearly so bad as they had been when she and George lived in Iceland. For many years, George spent most of the summer at his cabin in Ontario, Canada; Doris’s appointment only allowed her three weeks there each summer. It was set back about two miles off the Toronto-Ottawa highway, about equidistant between the two cities. It was fairly remote and had no phone. George and Doris liked to bathe in a cold stream about 100 yards from the cabin. Colleagues who visited George were treated to long talks, a little boating on the creek, and occasionally catching pike for dinner. George was gentle, quiet, and intensely thoughtful. He was also very approachable. It was not uncommon to sit down with him and toss out an initial idea for discussion. Then, when his ensuing silence grew uncomfortably long, one would toss out another idea, ultimately another, and then another. When he finally spoke, it became clear that he was still thinking deeply about the first or second idea.

Thursday, February 17, 2011

He exhibited a modesty that was several standard deviations from the norm, at least the norm in academia. When Maynard Smith’s Evolution of Sex (1978, Cambridge University Press) appeared just three years after George’s book on the same topic (Sex and Evolution, Princeton University Press), George declared in his review that Maynard Smith’s book had made his own obsolete. In 2000, when George was invited to speak on Darwinian medicine at a symposium on applied evolution (George was the cofounder of the field, after all), he responded that the organizers should get someone who knew something about the topic. He did not hesitate to give credit to others for his inspiration. For example, he attributed his inspiration for Darwinian medicine to Paul Ewald and Margie Profet, and in the preface to the 1996 reprinting of his 1966 book, George gave credit to several people who had influenced his thinking. George also had a sense of humor about himself (fig. 1) and about his mistakes. For example, his prediction in 1966 that sex ratio was “solved” came just a year before the topic started on a meteoric rise in popularity, with Hamilton’s “extraordinary sex ratios” paper and the later

The vigor of youth.... causes old age Figure 1: Linda Cranford Lillegraven, now an artist in Laramie, Wyoming, was a graduate student at the University of Utah who drew caricatures of many seminar speakers. She drew this likeness of George (ca. 1976) when he gave a talk on eels at the university. George was fond of this image and used it in correspondence for much of the last three decades.

century” hit Stony Brook, and the university shut down and sent everyone home at noon. The wind was fierce and snow was falling, but George and Doris insisted on walking home. Doris said she didn’t mind the walk because the conditions were not nearly so bad as they had been when she and George lived in Iceland. For many years, George spent most of the summer at his cabin in Ontario, Canada; Doris’s appointment only allowed her three weeks there each summer. It was set back about two miles off the Toronto-Ottawa highway, about equidistant between the two cities. It was fairly remote and had no phone. George and Doris liked to bathe in a cold stream about 100 yards from the cabin. Colleagues who visited George were treated to long talks, a little boating on the creek, and occasionally catching pike for dinner. George was gentle, quiet, and intensely thoughtful. He was also very approachable. It was not uncommon to sit down with him and toss out an initial idea for discussion. Then, when his ensuing silence grew uncomfortably long, one would toss out another idea, ultimately another, and then another. When he finally spoke, it became clear that he was still thinking deeply about the first or second idea.

Thursday, February 17, 2011

He exhibited a modesty that was several standard deviations from the norm, at least the norm in academia. When Maynard Smith’s Evolution of Sex (1978, Cambridge University Press) appeared just three years after George’s book on the same topic (Sex and Evolution, Princeton University Press), George declared in his review that Maynard Smith’s book had made his own obsolete. In 2000, when George was invited to speak on Darwinian medicine at a symposium on applied evolution (George was the cofounder of the field, after all), he responded that the organizers should get someone who knew something about the topic. He did not hesitate to give credit to others for his inspiration. For example, he attributed his inspiration for Darwinian medicine to Paul Ewald and Margie Profet, and in the preface to the 1996 reprinting of his 1966 book, George gave credit to several people who had influenced his thinking. George also had a sense of humor about himself (fig. 1) and about his mistakes. For example, his prediction in 1966 that sex ratio was “solved” came just a year before the topic started on a meteoric rise in popularity, with Hamilton’s “extraordinary sex ratios” paper and the later

The steeper the curve, the more intense the selection Figure 1: Linda Cranford Lillegraven, now an artist in Laramie, Wyoming, was a graduate student at the University of Utah who drew caricatures of many seminar speakers. She drew this likeness of George (ca. 1976) when he gave a talk on eels at the university. George was fond of this image and used it in correspondence for much of the last three decades.

century” hit Stony Brook, and the university shut down and sent everyone home at noon. The wind was fierce and snow was falling, but George and Doris insisted on walking home. Doris said she didn’t mind the walk because the conditions were not nearly so bad as they had been when she and George lived in Iceland. For many years, George spent most of the summer at his cabin in Ontario, Canada; Doris’s appointment only allowed her three weeks there each summer. It was set back about two miles off the Toronto-Ottawa highway, about equidistant between the two cities. It was fairly remote and had no phone. George and Doris liked to bathe in a cold stream about 100 yards from the cabin. Colleagues who visited George were treated to long talks, a little boating on the creek, and occasionally catching pike for dinner. George was gentle, quiet, and intensely thoughtful. He was also very approachable. It was not uncommon to sit down with him and toss out an initial idea for discussion. Then, when his ensuing silence grew uncomfortably long, one would toss out another idea, ultimately another, and then another. When he finally spoke, it became clear that he was still thinking deeply about the first or second idea.

Thursday, February 17, 2011

He exhibited a modesty that was several standard deviations from the norm, at least the norm in academia. When Maynard Smith’s Evolution of Sex (1978, Cambridge University Press) appeared just three years after George’s book on the same topic (Sex and Evolution, Princeton University Press), George declared in his review that Maynard Smith’s book had made his own obsolete. In 2000, when George was invited to speak on Darwinian medicine at a symposium on applied evolution (George was the cofounder of the field, after all), he responded that the organizers should get someone who knew something about the topic. He did not hesitate to give credit to others for his inspiration. For example, he attributed his inspiration for Darwinian medicine to Paul Ewald and Margie Profet, and in the preface to the 1996 reprinting of his 1966 book, George gave credit to several people who had influenced his thinking. George also had a sense of humor about himself (fig. 1) and about his mistakes. For example, his prediction in 1966 that sex ratio was “solved” came just a year before the topic started on a meteoric rise in popularity, with Hamilton’s “extraordinary sex ratios” paper and the later

Williams (1957) Prediction “Low adult death rates [from extrinsic causes] should be associated with low rates of senescence…”

Thursday, February 17, 2011

Steven Austad, University of Texas

Thursday, February 17, 2011

Mainland site

Sapelo Island

Thursday, February 17, 2011

Mainland site ~80% of opossums killed by predators

Sapelo Island

Thursday, February 17, 2011

Mainland site ~80% of opossums killed by predators

• Sapelo Island lacks mammalian opossum predators • Sapelo opossums have evolved in isolation for ~5,000 years Thursday, February 17, 2011

Sapelo Island

Thursday, February 17, 2011

Mainland

Island

Longevity (mo)

20.0

24.5

Max. longevity (mo)

31

45

Age 1st Repro (mo)

10.6

11.6

Litter size (1st yr)

7.6

5.9

Litter size (2nd yr)

7.6

5.4

SN Austad J. Zool. London 1993 229:695-708 Thursday, February 17, 2011

Death rates climb more slowly on the island

SN Austad J. Zool. London 1993 229:695-708 Thursday, February 17, 2011

Tendons get old: old fibers break faster

Thursday, February 17, 2011

Island opossums have slower-aging tendons

SN Austad J. Zool. London 1993 229:695-708 Thursday, February 17, 2011

Thursday, February 17, 2011

Charles with son William Anne Darwin Thursday, February 17, 2011

Thursday, February 17, 2011

What’s true for animals... Figure 1: Linda Cranford Lillegraven, now an artist in Laramie, Wyoming, was a graduate student at the University of Utah who drew caricatures of many seminar speakers. She drew this likeness of George (ca. 1976) when he gave a talk on eels at the university. George was fond of this image and used it in correspondence for much of the last three decades.

century” hit Stony Brook, and the university shut down and sent everyone home at noon. The wind was fierce and snow was falling, but George and Doris insisted on walking home. Doris said she didn’t mind the walk because the conditions were not nearly so bad as they had been when she and George lived in Iceland. For many years, George spent most of the summer at his cabin in Ontario, Canada; Doris’s appointment only allowed her three weeks there each summer. It was set back about two miles off the Toronto-Ottawa highway, about equidistant between the two cities. It was fairly remote and had no phone. George and Doris liked to bathe in a cold stream about 100 yards from the cabin. Colleagues who visited George were treated to long talks, a little boating on the creek, and occasionally catching pike for dinner. George was gentle, quiet, and intensely thoughtful. He was also very approachable. It was not uncommon to sit down with him and toss out an initial idea for discussion. Then, when his ensuing silence grew uncomfortably long, one would toss out another idea, ultimately another, and then another. When he finally spoke, it became clear that he was still thinking deeply about the first or second idea.

Thursday, February 17, 2011

He exhibited a modesty that was several standard deviations from the norm, at least the norm in academia. When Maynard Smith’s Evolution of Sex (1978, Cambridge University Press) appeared just three years after George’s book on the same topic (Sex and Evolution, Princeton University Press), George declared in his review that Maynard Smith’s book had made his own obsolete. In 2000, when George was invited to speak on Darwinian medicine at a symposium on applied evolution (George was the cofounder of the field, after all), he responded that the organizers should get someone who knew something about the topic. He did not hesitate to give credit to others for his inspiration. For example, he attributed his inspiration for Darwinian medicine to Paul Ewald and Margie Profet, and in the preface to the 1996 reprinting of his 1966 book, George gave credit to several people who had influenced his thinking. George also had a sense of humor about himself (fig. 1) and about his mistakes. For example, his prediction in 1966 that sex ratio was “solved” came just a year before the topic started on a meteoric rise in popularity, with Hamilton’s “extraordinary sex ratios” paper and the later

...is true for us, too

Human populations living under poor hygiene with little access to medicine compared to wild chimpanzees Caleb Finch, PNAS 2010 Thursday, February 17, 2011

Caleb Finch, PNAS 2010 Thursday, February 17, 2011

Antagonistic pleiotropy?

Caleb Finch, PNAS 2010 Thursday, February 17, 2011

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6SHHGLVPHDVXUHGDVWKHUHFLSURFDORIWKHPHDQUHTXLUHGWLPHLQPLQXWHVIRUDJLYHQ\HDU The Tithonus Error in Modern Gerontology George C. Williams

LQWKHSRS ILWQHVV JHQHUDO TheRWKHU Quarterly ReviewFDXVHLPSDLUV of Biology Vol.XODWLRQRIWHQ 74, No. 4 (Dec., 1999), pp. 405-415 FRXQ HQRXJKWREHHIIHFWLYHO\ Thursday, February 17, 2011

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Breast cancer cells

Thursday, February 17, 2011

Death rate from breast cancer (per 100,00 women) in US

Data: http://seer.cancer.gov/csr/1975_2007/ Graph: http://commons.wikimedia.org/wiki/File:Age-Specific_SEER_Incidence_Rates,_2003-2007_Breast_Cancer.svg Thursday, February 17, 2011

DNA damage, abnormal growth

Adapted from Coppe et al, Annu. Rev. Pathol. Mech. Dis. 2010. 5:99–118 Thursday, February 17, 2011

DNA damage, abnormal growth

p53 pathway activated

Adapted from Coppe et al, Annu. Rev. Pathol. Mech. Dis. 2010. 5:99–118 Thursday, February 17, 2011

DNA damage, abnormal growth

p53 pathway activated

Adapted from Coppe et al, Annu. Rev. Pathol. Mech. Dis. 2010. 5:99–118 Thursday, February 17, 2011

Senescence, cell suicide

DNA damage, abnormal growth

p53 pathway activated

Adapted from Coppe et al, Annu. Rev. Pathol. Mech. Dis. 2010. 5:99–118 Thursday, February 17, 2011

Senescence, cell suicide

CANCER STOPPED!

DNA damage, abnormal growth

p53 pathway activated

Senescence, cell suicide

CANCER STOPPED!

Tissue damage

Adapted from Coppe et al, Annu. Rev. Pathol. Mech. Dis. 2010. 5:99–118 Thursday, February 17, 2011

DNA damage, abnormal growth

p53 pathway activated

Adapted from Coppe et al, Annu. Rev. Pathol. Mech. Dis. 2010. 5:99–118 Thursday, February 17, 2011

Senescence, cell suicide

CANCER STOPPED!

Less regeneration in stem cells

Tissue damage

DNA damage, abnormal growth

p53 pathway activated

Inflammation, DNA damage

Adapted from Coppe et al, Annu. Rev. Pathol. Mech. Dis. 2010. 5:99–118 Thursday, February 17, 2011

Senescence, cell suicide

CANCER STOPPED!

Less regeneration in stem cells

Tissue damage

DNA damage, abnormal growth

p53 pathway activated

Inflammation, DNA damage

Senescence, cell suicide

CANCER STOPPED!

Less regeneration in stem cells

Tissue damage

Aging, late-life cancer, and other diseases Adapted from Coppe et al, Annu. Rev. Pathol. Mech. Dis. 2010. 5:99–118 Thursday, February 17, 2011

Delaying cancer in youth..... ...leads to cancer late in life?

Data: http://seer.cancer.gov/csr/1975_2007/ Graph: http://commons.wikimedia.org/wiki/File:Age-Specific_SEER_Incidence_Rates,_2003-2007_Breast_Cancer.svg Thursday, February 17, 2011

Antagonistic pleiotropy in the brain

Mo Morgan http://www.flickr.com/photos/momorgan/1483612005/ Thursday, February 17, 2011

The Two Faces of ApoE4

Trotter et al, Functional Ecology 2011, 25, 40–47 Thursday, February 17, 2011

The Two Faces of ApoE4 Benefits early in life Infection resistance Memory recall Enhanced learning

Trotter et al, Functional Ecology 2011, 25, 40–47 Thursday, February 17, 2011

The Two Faces of ApoE4 Benefits early in life

Costs late in life

Infection resistance

Heart disease

Memory recall Enhanced learning

Alzheimer’s disease

Trotter et al, Functional Ecology 2011, 25, 40–47 Thursday, February 17, 2011

Can aging be reversed, or at least slowed?

Thursday, February 17, 2011

Daf-2 mutants: The Methuselah Worms --Live twice as long

Chen et al, J Gerontol A Biol Sci Med Sci. 2007 Feb;62(2):126-35. Image: http://www.nematode.net/Images/c_elegans.jpg Thursday, February 17, 2011

Daf-2 mutants: The Methuselah Worms --Live twice as long

--Have fewer offspring Chen et al, J Gerontol A Biol Sci Med Sci. 2007 Feb;62(2):126-35. Image: http://www.nematode.net/Images/c_elegans.jpg Thursday, February 17, 2011

Thursday, February 17, 2011

Thanks to Doris Williams and Steven Austad

For more information, visit carlzimmer.com Thursday, February 17, 2011

Darwin Day 2011 Stony Brook-for pdf.pdf

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