The combinatorics of tandem duplication

L. Penso-Dolfin, T. Wu, C. Greenman

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Tandem duplication is an evolutionary process whereby a segment of DNA is replicated and proximally inserted. The different configurations that can arise from this process give rise to some interesting combinatorial questions. Firstly, we introduce an algebraic formalism to represent this process as a word producing automaton. The number of words arising from n tandem duplications can then be recursively derived. Secondly, each single word accounts for multiple evolutions. With the aid of a bi-coloured 2d- tree, a Hasse diagram corresponding to a partially ordered set is constructed, from which we can count the number of evolutions corresponding to a given word. Thirdly, we implement some subtree prune and graft operations on this structure to show that the total number of possible evolutions arising from n tandem duplications is $\prod_{k=1}^n(4^k - (2k + 1))$. The space of structures arising from tandem duplication thus grows at a super-exponential rate with leading order term $\mathcal{O}(4^{\frac{1}{2}n^2})$.
Original languageEnglish
Pages (from-to)1–22
Number of pages22
JournalDiscrete Applied Mathematics
Early online date6 Jun 2015
Publication statusPublished - 30 Oct 2015


  • Combinatorics
  • Tandem duplication
  • Posets
  • Rearrangements
  • Evolution

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