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1 Chapter 11 Evolution by Gene Duplication Chau-Ti Ting [email protected] Unless noted, the course materials are licensed under Creative Commons Attribution- NonCommercial - ShareAlike 3.0 Taiwan (CC BY-NC-SA 3.0)
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Page 2: 1 Chapter 11 Evolution by Gene Duplication Chau-Ti Ting ctting@ntu.edu.tw Unless noted, the course materials are licensed under Creative Commons Attribution-NonCommercial-ShareAlike.

National Taiwan University Chau-Ti Ting

The Bar Locus in Drosophila

+/+, wild-type B/+, heterozygous Bar B/B, homozygous Bar

Page 3: 1 Chapter 11 Evolution by Gene Duplication Chau-Ti Ting ctting@ntu.edu.tw Unless noted, the course materials are licensed under Creative Commons Attribution-NonCommercial-ShareAlike.

Unexpected Observations

Occasionally the homozygous stock gives rise to wild-type appearance.More rarely the stock gives rise to a more extreme reduction in eye-size, Ultra-Bar.Sturtevant and Morgan • both wild-type and Ultra-Bar flies recovered

from the Bar stock were associated with crossing over.

Page 4: 1 Chapter 11 Evolution by Gene Duplication Chau-Ti Ting ctting@ntu.edu.tw Unless noted, the course materials are licensed under Creative Commons Attribution-NonCommercial-ShareAlike.

Hypothesis: Unequal Crossing-over

Source: W. H. Freeman and P. Benjamin2005. Genetics: A Conceptual Approach, 2nd ed., p. 239. W. H. Freeman and Company. New York, USA.

Page 5: 1 Chapter 11 Evolution by Gene Duplication Chau-Ti Ting ctting@ntu.edu.tw Unless noted, the course materials are licensed under Creative Commons Attribution-NonCommercial-ShareAlike.

Cytological Evidence(Bridges 1936)

Source: C. B. Bridges1936. The Bar “gene” a duplication. Science 83: p. 210.

Page 7: 1 Chapter 11 Evolution by Gene Duplication Chau-Ti Ting ctting@ntu.edu.tw Unless noted, the course materials are licensed under Creative Commons Attribution-NonCommercial-ShareAlike.

Genome duplication• Kuwada (1911)

– Maize (Zea mays) was an ancient tetraploid• two sets of paralogous chromosomes in its karyotype.

Diploid Parent (2N) Tetraploid Parent (4N)National Taiwan University Chau-Ti Ting

Page 8: 1 Chapter 11 Evolution by Gene Duplication Chau-Ti Ting ctting@ntu.edu.tw Unless noted, the course materials are licensed under Creative Commons Attribution-NonCommercial-ShareAlike.

=

2N Parent 4N Parent

Meiosis

N Gamete 2N Gamete

3N Zygote

Mating

Uneven Chromosome Number National Taiwan University Chau-Ti Ting

Page 9: 1 Chapter 11 Evolution by Gene Duplication Chau-Ti Ting ctting@ntu.edu.tw Unless noted, the course materials are licensed under Creative Commons Attribution-NonCommercial-ShareAlike.

Autopolyploidy

Mitosis(Failure of cell division)

2N 4N

Meiosis

2N 4N

Fertilization

National Taiwan University Chau-Ti Ting

Page 10: 1 Chapter 11 Evolution by Gene Duplication Chau-Ti Ting ctting@ntu.edu.tw Unless noted, the course materials are licensed under Creative Commons Attribution-NonCommercial-ShareAlike.

Allopolypoidy

Species A2N = 6

Species B2N = 4

Failure of cell division

4 chromosome gameteHybrid

Chromosome = 7

Normal gameteChromosome = 3

7 chromosome gamete

Fertile hybrid Chromosome = 10

National Taiwan University Chau-Ti Ting

Page 11: 1 Chapter 11 Evolution by Gene Duplication Chau-Ti Ting ctting@ntu.edu.tw Unless noted, the course materials are licensed under Creative Commons Attribution-NonCommercial-ShareAlike.

Evolution by Gene duplication(Historical View I)

In 1918, Bridges wrote that the main interest in duplications lay in their offering a method for evolutionary increase in lengths of chromosomes with identical genes, which could subsequently mutate separately and diversify their effects

Muller further proposed that the redundant loci produced by the duplication of chromosomes parts could experience divergent mutations and eventually be regarded as nonhomologous genes

Source: J. S. Taylor and J. Raes2004. Duplication and divergence: The evolutionof new genes and old Ideas. Annual Review of Genetics 38: p. 615.

Source: J. S. Taylor and J. Raes2004. Duplication and divergence: The evolutionof new genes and old Ideas. Annual Review of Genetics 38: p. 615.

Page 12: 1 Chapter 11 Evolution by Gene Duplication Chau-Ti Ting ctting@ntu.edu.tw Unless noted, the course materials are licensed under Creative Commons Attribution-NonCommercial-ShareAlike.

Evolution by Gene duplication(Historical View II)

In 1932, Haldane suggested that duplication events might be favorable because they produce genes that could be altered without disadvantage to the organism

In 1938, Serebrovsky proposed that selection could be relaxed in genes that occur in duplicate (achaete-scute complex)

Source: J. S. Taylor and J. Raes2004. Duplication and divergence: The evolutionof new genes and old Ideas. Annual Review of Genetics 38: p. 615.

Source: J. S. Taylor and J. Raes2004. Duplication and divergence: The evolutionof new genes and old Ideas. Annual Review of Genetics 38: p. 615.

Page 13: 1 Chapter 11 Evolution by Gene Duplication Chau-Ti Ting ctting@ntu.edu.tw Unless noted, the course materials are licensed under Creative Commons Attribution-NonCommercial-ShareAlike.

Evolution by Gene duplication(Historical View III)

Stephens (1951) proposed the only way of achieving “evolutionary progress” would be by increasing the number of genetic loci, either by the synthesis of new loci from nongenic material or by the duplication and subsequent differentiation of existing loci via genome duplication or unequal recombination

Source: J. S. Taylor and J. Raes2004. Duplication and divergence: The evolutionof new genes and old Ideas. Annual Review of Genetics 38: p. 615.

Page 14: 1 Chapter 11 Evolution by Gene Duplication Chau-Ti Ting ctting@ntu.edu.tw Unless noted, the course materials are licensed under Creative Commons Attribution-NonCommercial-ShareAlike.

Gene Duplication and DNA Content

Britten and Davidson (1969) summarized DNA content from widely diverse species, with a plot showing that a great increase in DNA is “a necessary concomitant to increased complexity of organization.” Ohno noted a twofold to threefold increase in genome size between Ciona intestinalis and placental mammals and concluded that this difference was due to extensive gene duplication, possibly polyploidization Source: J. S. Taylor and J. Raes

2004. Duplication and divergence: The evolutionof new genes and old Ideas. Annual Review of Genetics 38: p. 615.

Source: J. S. Taylor and J. Raes2004. Duplication and divergence: The evolutionof new genes and old Ideas. Annual Review of Genetics 38: p. 615.

Page 15: 1 Chapter 11 Evolution by Gene Duplication Chau-Ti Ting ctting@ntu.edu.tw Unless noted, the course materials are licensed under Creative Commons Attribution-NonCommercial-ShareAlike.

GENE DUPLICATION AND ISOZYME ELECTROPHORESIS

Duplicated isozyme loci produced more bands than single-copy genes, and many isozyme electrophoresis studies uncovered gene duplicates in polyploidsData from isozyme studies not only uncovered new examples of gene duplication, it also shed light on the consequences of gene duplication (PGI in fish)

Source: J. S. Taylor and J. Raes2004. Duplication and divergence: The evolutionof new genes and old Ideas. Annual Review of Genetics 38: p. 615.

Source: J. S. Taylor and J. Raes2004. Duplication and divergence: The evolutionof new genes and old Ideas. Annual Review of Genetics 38: p. 615.

Page 16: 1 Chapter 11 Evolution by Gene Duplication Chau-Ti Ting ctting@ntu.edu.tw Unless noted, the course materials are licensed under Creative Commons Attribution-NonCommercial-ShareAlike.

Susumu Ohno ( 大野 乾 )1928-2000

http://www.nap.edu/html/biomems/sohno.html

Page 17: 1 Chapter 11 Evolution by Gene Duplication Chau-Ti Ting ctting@ntu.edu.tw Unless noted, the course materials are licensed under Creative Commons Attribution-NonCommercial-ShareAlike.

• J. Zhang2003. Evolution by gene duplication: an update. Trends in Ecology & Evolution 18: p. 292-298.• M. Long, E. Betran, K. Thornton, and W. Wang2003. The origin of new genes: glimpses from the young and old. Nature Reviews Genetics 4: p. 865-875.

Page 18: 1 Chapter 11 Evolution by Gene Duplication Chau-Ti Ting ctting@ntu.edu.tw Unless noted, the course materials are licensed under Creative Commons Attribution-NonCommercial-ShareAlike.

Source: J. Zhang2003. Evolution by gene duplication: an update. Trends in Ecology & Evolution 18: p. 292.

Page 19: 1 Chapter 11 Evolution by Gene Duplication Chau-Ti Ting ctting@ntu.edu.tw Unless noted, the course materials are licensed under Creative Commons Attribution-NonCommercial-ShareAlike.

Mechanisms of Gene Duplication

Source: J. Zhang2003. Evolution by gene duplication: an update. Trends in Ecology & Evolution 18: p. 292.

Page 20: 1 Chapter 11 Evolution by Gene Duplication Chau-Ti Ting ctting@ntu.edu.tw Unless noted, the course materials are licensed under Creative Commons Attribution-NonCommercial-ShareAlike.

(Pseudogenization)

Fates of Gene Duplication

Source: A. Force, M. Lynch, F. B. Pickett, A. Amores, Y. Yan, and J. Postlethwait1999. Preservation of duplicate genes by complementary, degenerative mutations. Genetics 151: p. 1531.

Page 21: 1 Chapter 11 Evolution by Gene Duplication Chau-Ti Ting ctting@ntu.edu.tw Unless noted, the course materials are licensed under Creative Commons Attribution-NonCommercial-ShareAlike.

Division of expression after gene duplication

Source: J. Zhang2003. Evolution by gene duplication: an update. Trends in Ecology & Evolution 18: p. 292.

Page 22: 1 Chapter 11 Evolution by Gene Duplication Chau-Ti Ting ctting@ntu.edu.tw Unless noted, the course materials are licensed under Creative Commons Attribution-NonCommercial-ShareAlike.

Source: M. Long, E. Betran, K. Thornton, and W. Wang2003. The origin of new genes: glimpses from the young and old. Nature Reviews Genetics 4: p. 865.

Page 23: 1 Chapter 11 Evolution by Gene Duplication Chau-Ti Ting ctting@ntu.edu.tw Unless noted, the course materials are licensed under Creative Commons Attribution-NonCommercial-ShareAlike.

The origin of jingwei

Source: M. Long, E. Betran, K. Thornton, and W. Wang2003. The origin of new genes: glimpses from the young and old. Nature Reviews Genetics 4: p. 865.

Page 24: 1 Chapter 11 Evolution by Gene Duplication Chau-Ti Ting ctting@ntu.edu.tw Unless noted, the course materials are licensed under Creative Commons Attribution-NonCommercial-ShareAlike.

Accelerated evolution of new genes

Source: M. Long, E. Betran, K. Thornton, and W. Wang2003. The origin of new genes: glimpses from the young and old. Nature Reviews Genetics 4: p. 865.

Page 25: 1 Chapter 11 Evolution by Gene Duplication Chau-Ti Ting ctting@ntu.edu.tw Unless noted, the course materials are licensed under Creative Commons Attribution-NonCommercial-ShareAlike.

Source: M. Long, E. Betran, K. Thornton, and W. Wang2003. The origin of new genes: glimpses from the young and old. Nature Reviews Genetics 4: p. 865.

The biased distribution of retroposition events

Page 26: 1 Chapter 11 Evolution by Gene Duplication Chau-Ti Ting ctting@ntu.edu.tw Unless noted, the course materials are licensed under Creative Commons Attribution-NonCommercial-ShareAlike.

Source: H. Innan and F. Kondrashov2010. The evolution of gene duplications: classifying and distinguishing between models. Nature Review Genetics 11: p. 97.

Page 27: 1 Chapter 11 Evolution by Gene Duplication Chau-Ti Ting ctting@ntu.edu.tw Unless noted, the course materials are licensed under Creative Commons Attribution-NonCommercial-ShareAlike.

Outstanding questions about the evolution of duplicate genes (I)

1. What is the relative importance of positive darwinian selection and relaxation of purifying selection in functional divergence of duplicated genes?

2. How does an entirely new function originate after gene duplication?

3. What roles does gene duplication play in the establishment of complex gene expression networks and protein–protein inter- action networks, which are key characteristics of biological systems?

Source: J. Zhang 2003. Evolution by gene duplication: an update. Trends in Ecology & Evolution 18: p. 292.

Source: J. Zhang2003. Evolution by gene duplication: an update. Trends in Ecology & Evolution 18: p. 292.

Source: J. Zhang2003. Evolution by gene duplication: an update. Trends in Ecology & Evolution 18: p. 292.

Page 28: 1 Chapter 11 Evolution by Gene Duplication Chau-Ti Ting ctting@ntu.edu.tw Unless noted, the course materials are licensed under Creative Commons Attribution-NonCommercial-ShareAlike.

Outstanding questions about the evolution of duplicate genes (II)

4. How does genetic buffering function?

5. How important is gene duplication to the origin of species-specific features and speciation?

Source: J. Zhang2003. Evolution by gene duplication: an update. Trends in Ecology & Evolution 18: p. 292.

Source: J. Zhang2003. Evolution by gene duplication: an update. Trends in Ecology & Evolution 18: p. 292.

Page 29: 1 Chapter 11 Evolution by Gene Duplication Chau-Ti Ting ctting@ntu.edu.tw Unless noted, the course materials are licensed under Creative Commons Attribution-NonCommercial-ShareAlike.

Copyright DeclarationWork Licensing Author/Source Page

National Taiwan University Chau-Ti Ting

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W. H. Freeman and P. Benjamin2005. Genetics: A Conceptual Approach, 2nd ed., p. 239. W. H. Freeman and Company. New York, USA.http://www.nature.com/scitable/topicpage/dna-deletion-and-duplication-and-the-associated-331It is used subject to the fair use doctrine of:•Taiwan Copyright Act Articles 52 & 65•Scitable by Nature Education Terms of Use•Nature Terms and Conditions•The "Code of Best Practices in Fair Use for OpenCourseWare 2009 (http://www.centerforsocialmedia.org/sites/default/files/10-305-OCW-Oct29.pdf)" by A Committee of Practitioners of OpenCourseWare in the U.S. The contents are based on Section 107 of the 1976 U.S. Copyright Act

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C. B. Bridges1936. The Bar “gene” a duplication. Science 83: p. 210.http://www.sciencemag.org/content/83/2148/210.extractIt is used subject to the fair use doctrine of:•Taiwan Copyright Act Articles 52 & 65•Science/AAAS Copyright Statement•The "Code of Best Practices in Fair Use for OpenCourseWare 2009 (http://www.centerforsocialmedia.org/sites/default/files/10-305-OCW-Oct29.pdf)" by A Committee of Practitioners of OpenCourseWare in the U.S. The contents are based on Section 107 of the 1976 U.S. Copyright Act

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Wikipedia Lee M. Silverhttp://en.wikipedia.org/wiki/File:Unequalcrossingover.gif2012/06/28 visited

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National Taiwan University Chau-Ti TingP7

National Taiwan University Chau-Ti TingP8

National Taiwan University Chau-Ti TingP9

National Taiwan University Chau-Ti TingP10

“a necessary concomitant to increased complexity of organization.”

Roy J. Britten and Eric H. Davidson1969. Gene Regulation for Higher Cells: A Theory. Science 25: p. 349. http://www.sciencemag.org/content/165/3891/349.extract?sid=43f1093b-ba99-4437-b8ea-b728b4aa468fIt is used subject to the fair use doctrine of:•Taiwan Copyright Act Articles 52 & 65It is used subject to the fair use doctrine of:•Taiwan Copyright Act Articles 52 & 65•The "Code of Best Practices in Fair Use for OpenCourseWare 2009 (http://www.centerforsocialmedia.org/sites/default/files/10-305-OCW-Oct29.pdf)" by A Committee of Practitioners of OpenCourseWare in the U.S. The contents are based on Section 107 of the 1976 U.S. Copyright Act

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Wikipedia M. A. Clarkhttp://en.wikipedia.org/wiki/File:Susumu_Ohno.png2012/06/28 visited

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Work Licensing Author/Source

J. Zhang2003. Evolution by gene duplication: an update. Trends in Ecology & Evolution 18: p. 292.http://www.sciencedirect.com/science/article/pii/S0169534703000338It is used subject to the fair use doctrine of:•Taiwan Copyright Act Articles 52 & 65•Cell Press Terms and Conditions•Elsevier Website Terms and Conditions•The "Code of Best Practices in Fair Use for OpenCourseWare 2009 (http://www.centerforsocialmedia.org/sites/default/files/10-305-OCW-Oct29.pdf)" by A Committee of Practitioners of OpenCourseWare in the U.S. The contents are based on Section 107 of the 1976 U.S. Copyright Act

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J. Zhang2003. Evolution by gene duplication: an update. Trends in Ecology & Evolution 18: p. 293.http://www.sciencedirect.com/science/article/pii/S0169534703000338It is used subject to the fair use doctrine of:•Taiwan Copyright Act Articles 52 & 65•Cell Press Terms and Conditions•Elsevier Website Terms and Conditions•The "Code of Best Practices in Fair Use for OpenCourseWare 2009 (http://www.centerforsocialmedia.org/sites/default/files/10-305-OCW-Oct29.pdf)" by A Committee of Practitioners of OpenCourseWare in the U.S. The contents are based on Section 107 of the 1976 U.S. Copyright Act

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A. Force, M. Lynch, F. B. Pickett, A. Amores, Y. Yan, and J. Postlethwait1999. Preservation of duplicate genes by complementary, degenerative mutations. Genetics 151: p. 1534.http://www.genetics.org/content/151/4/1531.full.pdfIt is used subject to the fair use doctrine of:•Taiwan Copyright Act Articles 52 & 65•The "Code of Best Practices in Fair Use for OpenCourseWare 2009 (http://www.centerforsocialmedia.org/sites/default/files/10-305-OCW-Oct29.pdf)" by A Committee of Practitioners of OpenCourseWare in the U.S. The contents are based on Section 107 of the 1976 U.S. Copyright Act

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J. Zhang2003. Evolution by gene duplication: an update. Trends in Ecology & Evolution 18: p. 294.http://www.sciencedirect.com/science/article/pii/S0169534703000338It is used subject to the fair use doctrine of:•Taiwan Copyright Act Articles 52 & 65•Cell Press Terms and Conditions•Elsevier Website Terms and Conditions•The "Code of Best Practices in Fair Use for OpenCourseWare 2009 (http://www.centerforsocialmedia.org/sites/default/files/10-305-OCW-Oct29.pdf)" by A Committee of Practitioners of OpenCourseWare in the U.S. The contents are based on Section 107 of the 1976 U.S. Copyright Act

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M. Long, E. Betran, K. Thornton, and W. Wang2003. The origin of new genes: glimpses from the young and old. Nature Reviews Genetics 4: p. 866.http://www.nature.com/nrg/journal/v4/n11/pdf/nrg1204.pdfIt is used subject to the fair use doctrine of:•Taiwan Copyright Act Articles 52 & 65•Nature Terms and Conditions•The "Code of Best Practices in Fair Use for OpenCourseWare 2009 (http://www.centerforsocialmedia.org/sites/default/files/10-305-OCW-Oct29.pdf)" by A Committee of Practitioners of OpenCourseWare in the U.S. The contents are based on Section 107 of the 1976 U.S. Copyright Act

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M. Long, E. Betran, K. Thornton, and W. Wang2003. The origin of new genes: glimpses from the young and old. Nature Reviews Genetics 4: p. 867.http://www.nature.com/nrg/journal/v4/n11/pdf/nrg1204.pdfIt is used subject to the fair use doctrine of:•Taiwan Copyright Act Articles 52 & 65•Nature Terms and Conditions•The "Code of Best Practices in Fair Use for OpenCourseWare 2009 (http://www.centerforsocialmedia.org/sites/default/files/10-305-OCW-Oct29.pdf)" by A Committee of Practitioners of OpenCourseWare in the U.S. The contents are based on Section 107 of the 1976 U.S. Copyright Act

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Work Licensing Author/Source

M. Long, E. Betran, K. Thornton, and W. Wang2003. The origin of new genes: glimpses from the young and old. Nature Reviews Genetics 4: p. 869.http://www.nature.com/nrg/journal/v4/n11/pdf/nrg1204.pdfIt is used subject to the fair use doctrine of:•Taiwan Copyright Act Articles 52 & 65•Nature Terms and Conditions•The "Code of Best Practices in Fair Use for OpenCourseWare 2009 (http://www.centerforsocialmedia.org/sites/default/files/10-305-OCW-Oct29.pdf)" by A Committee of Practitioners of OpenCourseWare in the U.S. The contents are based on Section 107 of the 1976 U.S. Copyright Act

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M. Long, E. Betran, K. Thornton, and W. Wang2003. The origin of new genes: glimpses from the young and old. Nature Reviews Genetics 4: p. 872.http://www.nature.com/nrg/journal/v4/n11/pdf/nrg1204.pdfIt is used subject to the fair use doctrine of:•Taiwan Copyright Act Articles 52 & 65•Nature Terms and Conditions•The "Code of Best Practices in Fair Use for OpenCourseWare 2009 (http://www.centerforsocialmedia.org/sites/default/files/10-305-OCW-Oct29.pdf)" by A Committee of Practitioners of OpenCourseWare in the U.S. The contents are based on Section 107 of the 1976 U.S. Copyright Act

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H. Innan and F. Kondrashov2010. The evolution of gene duplications: classifying and distinguishing between models. Nature Review Genetics 11: p. 100.http://www.nature.com/nrg/journal/v11/n2/pdf/nrg2689.pdfIt is used subject to the fair use doctrine of:•Taiwan Copyright Act Articles 52 & 65•Nature Terms and Conditions•The "Code of Best Practices in Fair Use for OpenCourseWare 2009 (http://www.centerforsocialmedia.org/sites/default/files/10-305-OCW-Oct29.pdf)" by A Committee of Practitioners of OpenCourseWare in the U.S. The contents are based on Section 107 of the 1976 U.S. Copyright Act

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What is the relative importance of positive darwinian selection and relaxation of purifying selection in functional divergence of duplicated genes?

J. Zhang2003. Evolution by gene duplication: an update. Trends in Ecology & Evolution 18: p. 292-298.http://www.sciencedirect.com/science/article/pii/S0169534703000338It is used subject to the fair use doctrine of:•Taiwan Copyright Act Articles 52 & 65•Cell Press Terms and Conditions•Elsevier Website Terms and Conditions•The "Code of Best Practices in Fair Use for OpenCourseWare 2009 (http://www.centerforsocialmedia.org/sites/default/files/10-305-OCW-Oct29.pdf)" by A Committee of Practitioners of OpenCourseWare in the U.S. The contents are based on Section 107 of the 1976 U.S. Copyright Act

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How does an entirely new function originate after gene duplication?

J. Zhang2003. Evolution by gene duplication: an update. Trends in Ecology & Evolution 18: p. 292-298.http://www.sciencedirect.com/science/article/pii/S0169534703000338It is used subject to the fair use doctrine of:•Taiwan Copyright Act Articles 52 & 65•Cell Press Terms and Conditions•Elsevier Website Terms and Conditions•The "Code of Best Practices in Fair Use for OpenCourseWare 2009 (http://www.centerforsocialmedia.org/sites/default/files/10-305-OCW-Oct29.pdf)" by A Committee of Practitioners of OpenCourseWare in the U.S. The contents are based on Section 107 of the 1976 U.S. Copyright Act

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What roles does gene duplication play in the establishment of complex gene expression networks and protein–protein inter- action networks, which are key characteristics of biological systems?

J. Zhang2003. Evolution by gene duplication: an update. Trends in Ecology & Evolution 18: p. 292-298.http://www.sciencedirect.com/science/article/pii/S0169534703000338It is used subject to the fair use doctrine of:•Taiwan Copyright Act Articles 52 & 65•Cell Press Terms and Conditions•Elsevier Website Terms and Conditions•The "Code of Best Practices in Fair Use for OpenCourseWare 2009 (http://www.centerforsocialmedia.org/sites/default/files/10-305-OCW-Oct29.pdf)" by A Committee of Practitioners of OpenCourseWare in the U.S. The contents are based on Section 107 of the 1976 U.S. Copyright Act

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Work Licensing Author/Source

How does genetic buffering function?

J. Zhang2003. Evolution by gene duplication: an update. Trends in Ecology & Evolution 18: p. 292-298.http://www.sciencedirect.com/science/article/pii/S0169534703000338It is used subject to the fair use doctrine of:•Taiwan Copyright Act Articles 52 & 65•Cell Press Terms and Conditions•Elsevier Website Terms and Conditions•The "Code of Best Practices in Fair Use for OpenCourseWare 2009 (http://www.centerforsocialmedia.org/sites/default/files/10-305-OCW-Oct29.pdf)" by A Committee of Practitioners of OpenCourseWare in the U.S. The contents are based on Section 107 of the 1976 U.S. Copyright Act

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How important is gene duplication to the origin of species-specific features and speciation?

J. Zhang2003. Evolution by gene duplication: an update. Trends in Ecology & Evolution 18: p. 292-298.http://www.sciencedirect.com/science/article/pii/S0169534703000338It is used subject to the fair use doctrine of:•Taiwan Copyright Act Articles 52 & 65•Cell Press Terms and Conditions•Elsevier Website Terms and Conditions•The "Code of Best Practices in Fair Use for OpenCourseWare 2009 (http://www.centerforsocialmedia.org/sites/default/files/10-305-OCW-Oct29.pdf)" by A Committee of Practitioners of OpenCourseWare in the U.S. The contents are based on Section 107 of the 1976 U.S. Copyright Act

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