
        
        Branch Swapping as a heuristic search method
    
       
        Heuristic methods
      are exploratory approaches
      that attempt an approximate solution to a particular computational
      problem that may be difficult or impossible to solve exactly.
      
          Heuristic methods are often applied to the
      problem of finding the shortest evolutionary tree for a large
      number of taxa. One heuristic search method called Sub-tree Pruning & Re-grafting (SPR) takes a "provisional tree" of
      suspected minimum length, "prunes" one branch and
      experimentally adds it to each of the inter-nodes of the remaining
      tree. In the example, the 1+2
        branch (sub-tree) is pruned and re-grafted to the
      7 branch. The method
      explores variations on a known tree that is suspected to be close
      to the minimum length: this is computationally more efficient than
      examining large numbers of alternative trees of unknown length. 
      
          A variant is Nearest-Neighbor Interchange (NNI), which takes three
      related branch tips and exchanges the outside one for one of the
      two inside ones. In this example, in the first network 2 would
      be interchanged with 1 & 3, or 5 & 6 with respect to 7. 
    
            A third method is Tree Bisection and Re-connection
        (TBR), in which the tree
        is cut as nearly in half as possible and re-grafted on each of
        the remaining branches or inter-nodes. In this example, the
        first network would be cut between 3 & 4, and the 1234
        tree re-connected to each branch in 567. 
        
          TBR is the most "costly"
      in terms of computational time, but is most likely to find a
      shorter alternative solution, if one exists. NNI is the
      least costly of the three, but is also the least likely to find a
      shorter tree. SPR is intermediate on both criteria. One
      strategy is to use a large number of TBR searches to
      identify an optimal or near-optimal tree, and NNI to
      assess bootstrap trees where finding the optimal tree in
      every case is not so essential.
    
    
    Figure modified from
        Krane & Raymer 2004; All text material © 2021 by Steven M.
          Carr