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. 2024 Feb 6;19(1):6.
doi: 10.1186/s13015-023-00242-2.

Predicting horizontal gene transfers with perfect transfer networks

Affiliations

Predicting horizontal gene transfers with perfect transfer networks

Alitzel López Sánchez et al. Algorithms Mol Biol. .

Abstract

Background: Horizontal gene transfer inference approaches are usually based on gene sequences: parametric methods search for patterns that deviate from a particular genomic signature, while phylogenetic methods use sequences to reconstruct the gene and species trees. However, it is well-known that sequences have difficulty identifying ancient transfers since mutations have enough time to erase all evidence of such events. In this work, we ask whether character-based methods can predict gene transfers. Their advantage over sequences is that homologous genes can have low DNA similarity, but still have retained enough important common motifs that allow them to have common character traits, for instance the same functional or expression profile. A phylogeny that has two separate clades that acquired the same character independently might indicate the presence of a transfer even in the absence of sequence similarity.

Our contributions: We introduce perfect transfer networks, which are phylogenetic networks that can explain the character diversity of a set of taxa under the assumption that characters have unique births, and that once a character is gained it is rarely lost. Examples of such traits include transposable elements, biochemical markers and emergence of organelles, just to name a few. We study the differences between our model and two similar models: perfect phylogenetic networks and ancestral recombination networks. Our goals are to initiate a study on the structural and algorithmic properties of perfect transfer networks. We then show that in polynomial time, one can decide whether a given network is a valid explanation for a set of taxa, and show how, for a given tree, one can add transfer edges to it so that it explains a set of taxa. We finally provide lower and upper bounds on the number of transfers required to explain a set of taxa, in the worst case.

Keywords: Character-based; Gene-expression; Horizontal gene transfer; Indirect phylogenetic methods; Perfect phylogenies; Tree-based networks.

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Conflict of interest statement

The authors declare that they have no competing interests.

Figures

Fig. 1
Fig. 1
1 A species tree T with a set of taxa S on characters C={a,b,c}. 2 A PTN with T as base tree that explains S. Red arrows represent transfer edges. 3 Another PTN with T as base tree that also explains S. Note that this PTN requires two less transfers. 4 A tree-based network G with S-map σ for which no labeling can explain S
Fig. 2
Fig. 2
a A PPN network with one character c with two states {0,1}. b A tree displayed by G that admits a labeling such that every state forms a connected component. Note that this tree can be obtained by the removal of the green edge and suppression of the resulting subdivision node. The interpretation of this scenario is that state 0 was transferred using the birectional edge. On the other hand, a solution for PTNs would forbid the transfer of state absence, and would also disregard the removal of the green edge, in which case no C-labeling is possible
Fig. 3
Fig. 3
(1) Left: a network G with 5 taxa at the leaves. The fat red edge is a transfer edge. Two taxa have no character, two have all characters of C, and one has only the odd numbered taxa (we assume odd d in the figure). Right: a C-labeling that explains G (the reticulation receives the odd characters from the transfer edge). This network has no explanation with d-crossovers. (2) Left: a network G with 5 taxa on character set C={c1,c2}. Right: a binary C-labeling with single crossovers that explains G. This network is not a PTN
Algorithm 1
Algorithm 1
Check if a given tree-based network G explains S.
Algorithm 2
Algorithm 2
Place an edge between all the first-appearance trees.
Fig. 4
Fig. 4
An example of a solution that will be given by our greedy algorithm (1) The given instance T with C={a,b,c}. Triangles represent subtrees and colors represent the characters that can be found on them (2) One possible solution output by Algorithm 2. The green arrow joins the first two subtrees that contain {a,b}, the yellow arrows joins the second subtree with the subtree that contains only {a} and the final arrow connects two subtrees that contain only {c}
Fig. 5
Fig. 5
The greedy Algorithm 2 can add more transfers than the minimum. 1 A network G whose leaves are on two characters a and b. 2 One possible solution output by Algorithm 2. The left clade is chosen to originate a and a transfer is greedily added above all the a’s on the right side. Then, the left clade is also chosen to originate b, but for that two more separate transfers are added. 3 A solution that is somewhat less natural, but with one less transfer
Fig. 6
Fig. 6
An illustration of T with k=3 and |S|=8. Each color corresponds to a distinct character. Each node labeled as 1 is a first-appearance of the character associated with that color. Notice that each level has its corresponding character
Fig. 7
Fig. 7
An illustration of f(x)=z for a marked node px that has a closest marked descendant py. Here, the characters of x must be a subset of the characters of y, as otherwise x would be a first appearance. We can then argue that a character of y\x has its first appearance between px and py

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