Molecular Clock Problems

Wessen-Simulating-human-origin-evo.pdf

Simulating Human Origins and Evolution by Ken Wessen (2005) pp 9-11:

As is apparent from the above discussion of the work by Cann et al. (1987), molecular methods rely on knowledge of the mutation rate of DNA across time and between species. The molecular clock hypothesis is a consequence of the neutral theory of evolution (Kimura, 1968) and implies an approximately constant rate of mutation, so long as the DNA sequence retains its original function. If this is the case, then the degree of difference between sequences being compared is simply proportional to the time since the sequences diverged. By incorporating fossil evidence, the clock can be calibrated, and thus divergence times can be attached to a molecular phylogeny.  

In fact, particular DNA sequences and proteins can mutate at vastly different rates at different times and in different lineages, and although there may be some local validity of the molecular clock hypothesis, in general there is global failure (Avise, 2000; Gibbons, 1998; Ruvolo, 1996; Strauss, 1999; Wills, 1995). The fast-mutating microsatellite loci, i.e. short repetitive sections of DNA that lie between genes, have been used to construct an alternative method for timing lineages that does not rely on external calibration of the rate of molecular evolution (Goldstein et al., 1995). However, because of mutational saturation, nuclear microsatellites are only useful for timing relatively recent events. In particular, the deepest split in the human phylogeny can be recovered with such a method, but saturation will occur in less time than the five million years or more back to the human–chimpanzee common ancestor (Jorde et al., 1998).

This situation also affects substantially the common ancestor calculations described above. For example, Wills (1995) includes a variable mutation rate across mtDNA sites and obtains a range of 436 000 to 800 000 years ago for the mitochondrial common ancestor, depending on the date used for the human–chimpanzee common ancestor.

 In general, the molecular data seem to support the replacement hypothesis, but when all the aforementioned caveats are considered, it remains far from conclusive. The dates vary widely, depending on the method and assumptions employed. Furthermore, a recent African origin has difficulty with the observed continuity of regional morphological traits, especially outside of Europe, whereas the multiregional hypothesis has difficulty with the amount of gene flow required for its support, as well as with a number of aspects of the molecular data. Perhaps the only thing that is truly clear is that population size, breeding patterns, local geographic events, migrations and reproductive barriers present a severe challenge when it comes to interpreting these results (Lahr and Foley, 1998). So long as positions at both extremes in this debate consider themselves equally well supported by the same data, be it fossil or molecular, substantial further study into the basis of all these methods is obviously of great importance.