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Molecular clocks

Molecular clocks. Molecular clock?. The molecular clock hypothesis was put forward by Zuckerkandl and Pauling in 1962. They noted that rates of amino acid replacements in animal hemoglobins were proportional to time of divergence—as judged from the fossil record. Molecular clocks?.

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Molecular clocks

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  1. Molecular clocks

  2. Molecular clock? • The molecular clock hypothesis was put forward by Zuckerkandl and Pauling in 1962. • They noted that rates of amino acid replacements in animal hemoglobins were proportional to time of divergence—as judged from the fossil record.

  3. Molecular clocks? • Zuckerkandl and Pauling, therefore, proposed that for any given protein, the rate of molecular evolution is approximately constant over time in all lineages.

  4. The molecular clock hypothesis If proteins evolve at constant rates, then the number of substitutions between two sequences may be used to estimate divergence times. This is analogous to the dating of geological times by radioactive decay.

  5. Example: The rate of nonsynonymous substitution for a-globin is 0.56  10–9 nonsynonymous substitutions per nonsynonymous site per year. Rat and human a-globins differ by 0.093 nonsynonymous substitutions per nonsynonymous site. If the universal molecular-clock hypothesis is correct, then human and rat diverged from a common ancestor 0.093/2  0.56  10 –9 = 83 million years ago.

  6. Pro Con Allan C. Wilson Morris Goodman

  7. The “sacrament” of the straight line

  8. Q: How to draw a straight line? A1: Have no more than two observation points.

  9. Q: How to draw a straight line? A2: With more than two observation points, use a very thick line.

  10. Q: How to draw a straight line? A3: With more than two observation points, deny the accuracy of the measurements on one or both axes.

  11. Relative Rate Tests

  12. Sarich & Wilson’s Test

  13. KAB = KOA + KOB KAC = KOA + KOC KBC = KOB + KOC

  14. KOA = (KAC + KAB –KBC)/2 KOB = (KAB + KBC –KAC)/2 KOC = (KAC + KBC –KAB)/2

  15. If the molecular clock hypothesis is correct, then KAC – KBC = 0

  16. Not significantly different from 0

  17. No such difference is seen at nonsynonymous sites, indicating that mutational differences, rather than selectional differences, are involved.

  18. The results of the relative rate test depend on knowledge of true tree.

  19. Tests involving duplicated genes

  20. If A1 evolves at the same rate as A2, and B1 evolves at the same rate as B2, then

  21. A = adult; E = embryonic; F = fetal

  22. Relative rate tests have shown that there is no universal molecular clock.However, sufficiently accurate local clocks may exist.

  23. slow fast

  24. Mutation rate per site per year versus genome size (Gago S, Elena SF, Flores R, Sanjuán R. Extremely high mutation rate of a hammerhead viroid. 2009. Science 323:1308.)

  25. The ranking of organisms started with the Aristotelian Scala Naturae…

  26. … and was used by Linnaeus in his Systema Naturae. Primates (humans and monkeys) Secundates (mammals) Tertiates (all others)

  27. In the literature one often encounters the adjective “primitive” attached to the name of an organism. For example, sponges are defined as “primitive.”

  28. Humans, on the other hand, are always referred to as “advanced.”

  29. Advanced Primitive

  30. Causes of variation in substitution rates among evolutionary lineages The factors most commonly invoked to explain the differences in the rate of substitution among lineages are: (1) replication-dependent factors, i.e., mutation. (2) replication-independent factors, i.e., selection.

  31. Generation Time

  32. Rates of evolution tend to correlate with generation times.

  33. Metabolic rate = amounts of O2 consumed per weight unit per time unit.

  34. metabolic-rate effect mice whales sharks newts

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