Thursday, January 19, 2012 6:26:42 AM
Multicellular Life Evolves in Laboratory
At left, an original strain of brewer’s yeast. At right, the multicellular form.
(Ratcliff et al./PNAS)
By Brandon Keim
January 17, 2012 | 4:49 pm
An evolutionary transition that took several billion years to occur in nature has happened in a laboratory, and it needed just 60 days.
Under artificial pressure to become larger, single-celled yeast became multicellular creatures. That crucial step is responsible for life’s progression beyond algae and bacteria, and while the latest work doesn’t duplicate prehistoric transitions, it could help reveal the principles guiding them.
“This is actually simple. It doesn’t need mystical complexity or a lot of the things that people have hypothesized — special genes, a huge genome, very unnatural conditions,” said evolutionary biologist Michael Travisano of the University of Minnesota, co-author of a study Jan. 17 in the Proceedings of the National Academy of Sciences.
In the new study, researchers led by Travisano and William Ratcliff grew brewer’s yeast, a common single-celled organism, in flasks of nutrient-rich broth.
Once per day they shook the flasks, removed yeast that most rapidly settled to the bottom, and used it to start new cultures. Free-floating yeast were left behind, while yeast that gathered in heavy, fast-falling clumps survived to reproduce.
Within just a few weeks, individual yeast cells still retained their singular identities, but clumped together easily. At the end of two months, the clumps were a permanent arrangement. Each strain had evolved to be truly multicellular, displaying all the tendencies associated with “higher” forms of life: a division of labor between specialized cells, juvenile and adult life stages, and multicellular offspring.
Multicellular yeast reproduces itself; the offspring will not reproduce until it has grown.
(Ratcliff et al./PNAS)
“Multicellularity is the ultimate in cooperation,” said Travisano, who wants to understand how cooperation emerges in selfishly competing organisms. “Multiple cells make make up an individual that cooperates for the benefit of the whole. Sometimes cells give up their ability to reproduce for the benefit of close kin.”
Since the late 1990s, experimental evolution studies have attempted to induce multicellularity in laboratory settings. While some fascinating entities have evolved — Richard Lenski’s kaleidoscopically adapting E. coli [ https://en.wikipedia.org/wiki/E._coli_long-term_evolution_experiment ], Paul Rainey’s visible-to-the-naked-eye bacterial biofilms [ http://www.nature.com/nature/journal/v445/n7127/abs/nature05514.html ] — true multicellularity remained elusive.
According to Travisano, too much emphasis was placed on identifying some genetic essence of complexity. The new study suggests that environmental conditions are paramount: Give single-celled organisms reason to go multicellular, and they will.
Apart from insights into complexity’s origins, the findings could have implications for researchers in other fields. While multicellularity would have a hard time emerging now in nature, where existing animals have a competitive advantage, the underlying lesson of rapid, radical evolution is universal.
“That idea of easy transformability changes your perspective,” said Travisano. “I’m certain that rapid evolution occurs. We just don’t know to look for it.”
Targeted breeding of single-celled organisms into complex, multicellular forms could also become a biotechnological production technique.
“If you want to have some organism that makes ethanol or a novel compound, then — apart from using genetic engineering — you could do selection experiments” to shape their evolution, Travisano said. “What we’re doing right here, engineering via artificial selection, is something we’ve done for centuries with animals and agriculture.”
Citation: “Experimental evolution of multicellularity.” By William C. Ratcliff, R. Ford Denison, Mark Borrello, and Michael Travisano. Proceedings of the National Academy of Sciences, Jan. 17, 2012. [ http://www.pnas.org/content/early/2012/01/10/1115323109 , in full at http://www.pnas.org/content/early/2012/01/10/1115323109.full.pdf+html ( http://dx.doi.org/10.1073/pnas.1115323109 )]
*
Previously
570-Million-Year-Old Fossils Hint at Origins of Animal Kingdom
http://www.wired.com/wiredscience/2011/12/doubt-earliest-animal-fossils/
Origin of Life Chicken-and-Egg Problem Solved
http://www.wired.com/wiredscience/2010/04/pre-life-paradox/
Primordial Soup’s Missing Ingredient May Be Sulfur
http://www.wired.com/wiredscience/2011/03/sulfur-prebiotic-soup/
Forgotten Experiment May Explain Origins of Life
http://www.wired.com/wiredscience/2008/10/forgotten-exper/
Life’s First Spark Re-Created in the Laboratory
http://www.wired.com/wiredscience/2009/05/ribonucleotides/
*
Wired.com © Condé Nast Digital
http://www.wired.com/wiredscience/2012/01/evolution-of-multicellularity/ [with comments]
===
Researchers evolve a multicellular yeast in the lab in 2 months
By John Timmer | Published January 18, 2012
When we think of life on Earth, most of us think of multicellular organisms, like large mammals or massive trees. But we're only aware of three groups of complex, multicellular organisms, which suggested it might be a major hurdle. Now, a new study describes how researchers evolved a multicellular form of yeast (the same species that contributes to bread and beer), and were able to see specialized cell behaviors and reproduction in as little as 60 days.
The authors lay out the problem very simply in their introduction, stating that, "Multicellularity was one of the most significant innovations in the history of life, but its initial evolution remains poorly understood." There is some evidence that it can be a favorable trait—research shows that clusters of cells evolve when a single-celled organism is kept in culture with a predator that can only swallow one cell at a time.
But that's about as far as these experiments went. It wasn't clear how these clusters of cells formed, whether they were genetically related, or whether they engaged in any sort of specialized behavior. More significantly, it wasn't obvious whether these clusters took a sort of "every cell for itself" approach to reproduction. So, although this work showed a multicellular lifestyle could be selected for, the researchers didn't look into how far down the road towards specialization those cells would go.
The new study attempts to follow more the behavior of simple multicellular groups more closely. It uses baker's yeast (Saccharomyces cerevisiae), an organism that normally grows as single cells. The authors grew these in culture and, once a day, transferred them in a way that favored multicellular growth.
Their method was pretty simple. Normally, yeast are grown in a culture that's shaken, and the single cells will only slowly settle to the bottom when that's stopped. The authors only transferred the cells at the bottom of the culture to fresh food, so that they selected for those cells that settled to the bottom quickly. This favors large clusters of cells, instead of single ones.
With only 60 daily transfers, all of their experimental populations were dominated by yeast cells that grew as clusters, which the authors describe as "roughly spherical snowflake-like." These were formed because, instead of separating after they divided, cells would remain attached, expanding the cluster with each division. Although this comes at a cost compared to individual cells—the authors calculate that individual cells in the cluster are 10 percent less fit than their single-celled relatives when they're not selecting for things on the bottom. But, with the selection in place, the clusters had a huge advantage.
But the clusters didn't simply keep growing indefinitely. Instead, the yeast quickly evolved a form of reproduction by splitting off what the authors call "propagules," or smaller clusters that break off and go on to develop on their own.
With more generations, this form of reproduction began to include specialized cell behavior. A small percentage of cells in the cluster would start committing suicide through a process called apoptosis. This death would allow the propagule to split off cleanly at the site of the dead cell, improving the efficiency of reproduction. Normally, there's no evolutionary advantage to a cell ending up dead but, since the cells in the propagule are genetically identical, this behavior can be selected for.
This new form of growth and reproduction is still a long way off from the complex, specialized tissues found in most multicellular organisms. But the ease with which this behavior evolved suggests that the foundations of multicellularity may evolve very easily, and don't present the barrier to complexity that many people have assumed it was.
PNAS, 2012. DOI: 10.1073/pnas.1115323109 [ http://www.pnas.org/content/early/2012/01/10/1115323109 , in full at http://www.pnas.org/content/early/2012/01/10/1115323109.full.pdf+html ( http://dx.doi.org/10.1073/pnas.1115323109 )]
===
Yeast Experiment Hints at a Faster Evolution From Single Cells
Video [embedded]
Evolving Yeast Cells
By CARL ZIMMER
Published: January 16, 2012
Our ancestors were single-celled microbes for about three billion years before they evolved bodies. But in a laboratory at the University of Minnesota, brewer’s yeast cells can evolve primitive bodies in about two weeks.
The transition to multicellular life has long intrigued evolutionary biologists. The cells in our bodies have evolved to cooperate with exquisite precision. The human body has more than 200 types of cells, each dedicated to a different job. And a vast majority of the 100 trillion cells in our bodies sacrifice their own long-term legacy: Only eggs and sperm have a chance to survive our own death.
These demands for cooperation and sacrifice ought to make it hard for single-celled life to become multicellular. Yet animals, plants and other life forms have evolved bodies. “We know that multicellularity has evolved in different lineages at least 25 times in the history of life,” said William Ratcliff, a postdoctoral researcher at the University of Minnesota.
Dr. Ratcliff and his adviser, Michael Travisano, are experts in experimental evolution. They design experiments in which microbes can evolve interesting new traits within weeks.
“We were sitting in his office drinking coffee, talking about what would be the coolest thing you could do in the lab,” Dr. Ratcliff said. “O.K., the origin of life would be too hard. But other than the origin of life, what would be the coolest thing?” They decided it would be observing single-celled microbes evolving a primitive form of multicellularity.
The scientists designed an experiment with brewer’s yeast, which normally lives as single cells, feeding on sugar and budding off daughter cells to reproduce.
Dr. Ratcliff and his colleagues set up an experiment that might favor multicellularity in yeast. They reared lines of yeast, starting from a single cell, in 10 flasks of broth. They kept the flasks shaking for a day and then let the yeast settle. The scientists then took out a drop of the settled yeast cells and transferred it to a fresh flask, where the yeast could continue to grow. In this experiment, natural selection favored any new mutation that would let the yeast fall quickly. Yeast cells that were still floating high in the broth would not have a chance to be delivered to the next flask.
In a matter of weeks, Dr. Ratcliff noticed, the yeast was sinking fast, forming a cloudy layer at the bottom of the flasks. He put the yeast under a microscope and discovered that most of it was no longer growing as single cells. Instead, the broth was dominated by snowflake-shaped clusters of hundreds of cells stuck together.
These were not clumps of unrelated cells, he found. When he isolated individual cells and let them grow, they formed new snowflakes. Instead of drifting away, newly budded yeast cells remained stuck to their parents. By staying stuck together, these yeast clusters fell faster than individual cells.
A single cell needs a few hours to grow to “adult” size, Dr. Ratcliff found. After it matures, its growing branches start to press against one another until they snap apart. These broken branches are yeast versions of plant cuttings: Each one grows into a snowflake of its own, which then snaps apart in turn.
Dr. Ratcliff also found that this new form of reproduction is possible only because some of the yeast cells make the ultimate sacrifice. Once a snowflake reaches adult size, a fraction of the cells commit suicide. “The cells that kill themselves act as weak links,” he said.
The scientists describe their experiments in a paper [ http://www.pnas.org/content/early/2012/01/10/1115323109 , in full at http://www.pnas.org/content/early/2012/01/10/1115323109.full.pdf+html ( http://dx.doi.org/10.1073/pnas.1115323109 )] being published this week in Proceedings of the National Academy of Sciences [ http://www.pnas.org/ ].
“This is a really interesting and important study,” said Richard Lenski, a biologist at Michigan State University and the editor of the paper. “It shows that a major transition in evolution — going from unicellular to multicellular life forms — might not be as hard to achieve as most biologists have long thought.”
Dr. Ratcliff suspects that the transformation of the yeast in his lab may offer hints about how animals and other lineages became multicellular hundreds of millions of years ago. “Forming clusters isn’t a freaky yeast thing,” he said. The closest single-celled relatives of animals, called choanoflagellates [ http://www.nytimes.com/2010/12/14/science/14creatures.html ], also sometimes grow as clusters of cells.
Animals and plants did not evolve inside flasks, of course. But natural conditions could have favored clusters of cells. They might have been harder for predators to eat, for example. A cluster of cells might also be able to feed more efficiently in some cases.
Dr. Ratcliff and his colleagues are now examining 25 genomes of the evolved yeast, looking for the mutations that gave them snowflake bodies. Meanwhile, their yeast continues to evolve. Once the cells gain the ability to form snowflakes, they become better adapted to multicellular life. They snap off smaller branches, allowing them to reproduce faster.
Dr. Ratcliff would not go into detail about where the yeast evolution was going until he published the latest results. “We’re getting really interesting things happening now” was all he would say.
© 2012 The New York Times Company
http://www.nytimes.com/2012/01/17/science/yeast-reveals-how-fast-a-cell-can-form-a-body.html
===
from elsewhere this string, (linked in) http://investorshub.advfn.com/boards/read_msg.aspx?message_id=69760902 and following
At left, an original strain of brewer’s yeast. At right, the multicellular form.
(Ratcliff et al./PNAS)
By Brandon Keim
January 17, 2012 | 4:49 pm
An evolutionary transition that took several billion years to occur in nature has happened in a laboratory, and it needed just 60 days.
Under artificial pressure to become larger, single-celled yeast became multicellular creatures. That crucial step is responsible for life’s progression beyond algae and bacteria, and while the latest work doesn’t duplicate prehistoric transitions, it could help reveal the principles guiding them.
“This is actually simple. It doesn’t need mystical complexity or a lot of the things that people have hypothesized — special genes, a huge genome, very unnatural conditions,” said evolutionary biologist Michael Travisano of the University of Minnesota, co-author of a study Jan. 17 in the Proceedings of the National Academy of Sciences.
In the new study, researchers led by Travisano and William Ratcliff grew brewer’s yeast, a common single-celled organism, in flasks of nutrient-rich broth.
Once per day they shook the flasks, removed yeast that most rapidly settled to the bottom, and used it to start new cultures. Free-floating yeast were left behind, while yeast that gathered in heavy, fast-falling clumps survived to reproduce.
Within just a few weeks, individual yeast cells still retained their singular identities, but clumped together easily. At the end of two months, the clumps were a permanent arrangement. Each strain had evolved to be truly multicellular, displaying all the tendencies associated with “higher” forms of life: a division of labor between specialized cells, juvenile and adult life stages, and multicellular offspring.
Multicellular yeast reproduces itself; the offspring will not reproduce until it has grown.
(Ratcliff et al./PNAS)
“Multicellularity is the ultimate in cooperation,” said Travisano, who wants to understand how cooperation emerges in selfishly competing organisms. “Multiple cells make make up an individual that cooperates for the benefit of the whole. Sometimes cells give up their ability to reproduce for the benefit of close kin.”
Since the late 1990s, experimental evolution studies have attempted to induce multicellularity in laboratory settings. While some fascinating entities have evolved — Richard Lenski’s kaleidoscopically adapting E. coli [ https://en.wikipedia.org/wiki/E._coli_long-term_evolution_experiment ], Paul Rainey’s visible-to-the-naked-eye bacterial biofilms [ http://www.nature.com/nature/journal/v445/n7127/abs/nature05514.html ] — true multicellularity remained elusive.
According to Travisano, too much emphasis was placed on identifying some genetic essence of complexity. The new study suggests that environmental conditions are paramount: Give single-celled organisms reason to go multicellular, and they will.
Apart from insights into complexity’s origins, the findings could have implications for researchers in other fields. While multicellularity would have a hard time emerging now in nature, where existing animals have a competitive advantage, the underlying lesson of rapid, radical evolution is universal.
“That idea of easy transformability changes your perspective,” said Travisano. “I’m certain that rapid evolution occurs. We just don’t know to look for it.”
Targeted breeding of single-celled organisms into complex, multicellular forms could also become a biotechnological production technique.
“If you want to have some organism that makes ethanol or a novel compound, then — apart from using genetic engineering — you could do selection experiments” to shape their evolution, Travisano said. “What we’re doing right here, engineering via artificial selection, is something we’ve done for centuries with animals and agriculture.”
Citation: “Experimental evolution of multicellularity.” By William C. Ratcliff, R. Ford Denison, Mark Borrello, and Michael Travisano. Proceedings of the National Academy of Sciences, Jan. 17, 2012. [ http://www.pnas.org/content/early/2012/01/10/1115323109 , in full at http://www.pnas.org/content/early/2012/01/10/1115323109.full.pdf+html ( http://dx.doi.org/10.1073/pnas.1115323109 )]
*
Previously
570-Million-Year-Old Fossils Hint at Origins of Animal Kingdom
http://www.wired.com/wiredscience/2011/12/doubt-earliest-animal-fossils/
Origin of Life Chicken-and-Egg Problem Solved
http://www.wired.com/wiredscience/2010/04/pre-life-paradox/
Primordial Soup’s Missing Ingredient May Be Sulfur
http://www.wired.com/wiredscience/2011/03/sulfur-prebiotic-soup/
Forgotten Experiment May Explain Origins of Life
http://www.wired.com/wiredscience/2008/10/forgotten-exper/
Life’s First Spark Re-Created in the Laboratory
http://www.wired.com/wiredscience/2009/05/ribonucleotides/
*
Wired.com © Condé Nast Digital
http://www.wired.com/wiredscience/2012/01/evolution-of-multicellularity/ [with comments]
===
Researchers evolve a multicellular yeast in the lab in 2 months
By John Timmer | Published January 18, 2012
When we think of life on Earth, most of us think of multicellular organisms, like large mammals or massive trees. But we're only aware of three groups of complex, multicellular organisms, which suggested it might be a major hurdle. Now, a new study describes how researchers evolved a multicellular form of yeast (the same species that contributes to bread and beer), and were able to see specialized cell behaviors and reproduction in as little as 60 days.
The authors lay out the problem very simply in their introduction, stating that, "Multicellularity was one of the most significant innovations in the history of life, but its initial evolution remains poorly understood." There is some evidence that it can be a favorable trait—research shows that clusters of cells evolve when a single-celled organism is kept in culture with a predator that can only swallow one cell at a time.
But that's about as far as these experiments went. It wasn't clear how these clusters of cells formed, whether they were genetically related, or whether they engaged in any sort of specialized behavior. More significantly, it wasn't obvious whether these clusters took a sort of "every cell for itself" approach to reproduction. So, although this work showed a multicellular lifestyle could be selected for, the researchers didn't look into how far down the road towards specialization those cells would go.
The new study attempts to follow more the behavior of simple multicellular groups more closely. It uses baker's yeast (Saccharomyces cerevisiae), an organism that normally grows as single cells. The authors grew these in culture and, once a day, transferred them in a way that favored multicellular growth.
Their method was pretty simple. Normally, yeast are grown in a culture that's shaken, and the single cells will only slowly settle to the bottom when that's stopped. The authors only transferred the cells at the bottom of the culture to fresh food, so that they selected for those cells that settled to the bottom quickly. This favors large clusters of cells, instead of single ones.
With only 60 daily transfers, all of their experimental populations were dominated by yeast cells that grew as clusters, which the authors describe as "roughly spherical snowflake-like." These were formed because, instead of separating after they divided, cells would remain attached, expanding the cluster with each division. Although this comes at a cost compared to individual cells—the authors calculate that individual cells in the cluster are 10 percent less fit than their single-celled relatives when they're not selecting for things on the bottom. But, with the selection in place, the clusters had a huge advantage.
But the clusters didn't simply keep growing indefinitely. Instead, the yeast quickly evolved a form of reproduction by splitting off what the authors call "propagules," or smaller clusters that break off and go on to develop on their own.
With more generations, this form of reproduction began to include specialized cell behavior. A small percentage of cells in the cluster would start committing suicide through a process called apoptosis. This death would allow the propagule to split off cleanly at the site of the dead cell, improving the efficiency of reproduction. Normally, there's no evolutionary advantage to a cell ending up dead but, since the cells in the propagule are genetically identical, this behavior can be selected for.
This new form of growth and reproduction is still a long way off from the complex, specialized tissues found in most multicellular organisms. But the ease with which this behavior evolved suggests that the foundations of multicellularity may evolve very easily, and don't present the barrier to complexity that many people have assumed it was.
PNAS, 2012. DOI: 10.1073/pnas.1115323109 [ http://www.pnas.org/content/early/2012/01/10/1115323109 , in full at http://www.pnas.org/content/early/2012/01/10/1115323109.full.pdf+html ( http://dx.doi.org/10.1073/pnas.1115323109 )]
===
Yeast Experiment Hints at a Faster Evolution From Single Cells
Video [embedded]
Evolving Yeast Cells
By CARL ZIMMER
Published: January 16, 2012
Our ancestors were single-celled microbes for about three billion years before they evolved bodies. But in a laboratory at the University of Minnesota, brewer’s yeast cells can evolve primitive bodies in about two weeks.
The transition to multicellular life has long intrigued evolutionary biologists. The cells in our bodies have evolved to cooperate with exquisite precision. The human body has more than 200 types of cells, each dedicated to a different job. And a vast majority of the 100 trillion cells in our bodies sacrifice their own long-term legacy: Only eggs and sperm have a chance to survive our own death.
These demands for cooperation and sacrifice ought to make it hard for single-celled life to become multicellular. Yet animals, plants and other life forms have evolved bodies. “We know that multicellularity has evolved in different lineages at least 25 times in the history of life,” said William Ratcliff, a postdoctoral researcher at the University of Minnesota.
Dr. Ratcliff and his adviser, Michael Travisano, are experts in experimental evolution. They design experiments in which microbes can evolve interesting new traits within weeks.
“We were sitting in his office drinking coffee, talking about what would be the coolest thing you could do in the lab,” Dr. Ratcliff said. “O.K., the origin of life would be too hard. But other than the origin of life, what would be the coolest thing?” They decided it would be observing single-celled microbes evolving a primitive form of multicellularity.
The scientists designed an experiment with brewer’s yeast, which normally lives as single cells, feeding on sugar and budding off daughter cells to reproduce.
Dr. Ratcliff and his colleagues set up an experiment that might favor multicellularity in yeast. They reared lines of yeast, starting from a single cell, in 10 flasks of broth. They kept the flasks shaking for a day and then let the yeast settle. The scientists then took out a drop of the settled yeast cells and transferred it to a fresh flask, where the yeast could continue to grow. In this experiment, natural selection favored any new mutation that would let the yeast fall quickly. Yeast cells that were still floating high in the broth would not have a chance to be delivered to the next flask.
In a matter of weeks, Dr. Ratcliff noticed, the yeast was sinking fast, forming a cloudy layer at the bottom of the flasks. He put the yeast under a microscope and discovered that most of it was no longer growing as single cells. Instead, the broth was dominated by snowflake-shaped clusters of hundreds of cells stuck together.
These were not clumps of unrelated cells, he found. When he isolated individual cells and let them grow, they formed new snowflakes. Instead of drifting away, newly budded yeast cells remained stuck to their parents. By staying stuck together, these yeast clusters fell faster than individual cells.
A single cell needs a few hours to grow to “adult” size, Dr. Ratcliff found. After it matures, its growing branches start to press against one another until they snap apart. These broken branches are yeast versions of plant cuttings: Each one grows into a snowflake of its own, which then snaps apart in turn.
Dr. Ratcliff also found that this new form of reproduction is possible only because some of the yeast cells make the ultimate sacrifice. Once a snowflake reaches adult size, a fraction of the cells commit suicide. “The cells that kill themselves act as weak links,” he said.
The scientists describe their experiments in a paper [ http://www.pnas.org/content/early/2012/01/10/1115323109 , in full at http://www.pnas.org/content/early/2012/01/10/1115323109.full.pdf+html ( http://dx.doi.org/10.1073/pnas.1115323109 )] being published this week in Proceedings of the National Academy of Sciences [ http://www.pnas.org/ ].
“This is a really interesting and important study,” said Richard Lenski, a biologist at Michigan State University and the editor of the paper. “It shows that a major transition in evolution — going from unicellular to multicellular life forms — might not be as hard to achieve as most biologists have long thought.”
Dr. Ratcliff suspects that the transformation of the yeast in his lab may offer hints about how animals and other lineages became multicellular hundreds of millions of years ago. “Forming clusters isn’t a freaky yeast thing,” he said. The closest single-celled relatives of animals, called choanoflagellates [ http://www.nytimes.com/2010/12/14/science/14creatures.html ], also sometimes grow as clusters of cells.
Animals and plants did not evolve inside flasks, of course. But natural conditions could have favored clusters of cells. They might have been harder for predators to eat, for example. A cluster of cells might also be able to feed more efficiently in some cases.
Dr. Ratcliff and his colleagues are now examining 25 genomes of the evolved yeast, looking for the mutations that gave them snowflake bodies. Meanwhile, their yeast continues to evolve. Once the cells gain the ability to form snowflakes, they become better adapted to multicellular life. They snap off smaller branches, allowing them to reproduce faster.
Dr. Ratcliff would not go into detail about where the yeast evolution was going until he published the latest results. “We’re getting really interesting things happening now” was all he would say.
© 2012 The New York Times Company
http://www.nytimes.com/2012/01/17/science/yeast-reveals-how-fast-a-cell-can-form-a-body.html
===
from elsewhere this string, (linked in) http://investorshub.advfn.com/boards/read_msg.aspx?message_id=69760902 and following
Greensburg, KS - 5/4/07
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upon the Right of Election, 1790
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