Reticulate evolution

Reticulate evolution, or network evolution, describes the

origination of a lineage through the partial merging of two
ancestor lineages, leading to relationships better described by
a phylogenetic network than a bifurcating tree.[1] Reticulate
patterns can be found in the phylogenetic reconstructions of
biodiversity lineages obtained by comparing the characteristics
of organisms.[2] Reticulation processes can potentially be
convergent and divergent at the same time.[3] Reticulate
Phylogenetic network depicting reticulate
evolution indicates the lack of independence between two
evolution: Lineage B results from a
evolutionary lineages.[1] Reticulation affects survival, fitness horizontal transfer between its two
and speciation rates of species.[2] The adjective reticulate ancestors A and C (blue, dotted lines).
stems from the Latin words reticulatus “having a net-like
pattern” and reticulum “little net”, according to the Etymology
Dictionary (http://etymonline.com/index.php).

Reticulate evolution can happen between lineages separated only for a short time, for example through
hybrid speciation in a species complex. Nevertheless, it also takes place over larger evolutionary distances,
as exemplified by the presence of organelles of bacterial origin in eukaryotic cells.[2]

Reticulation occurs at various levels:[4] at a chromosomal level, meiotic recombination causes evolution to
be reticulate; at a species level, reticulation arises through hybrid speciation and horizontal gene transfer;
and at a population level, sexual recombination causes reticulation.[1]

Contents
Underlying mechanisms and processes
Symbiosis
Symbiogenesis
Lateral gene transfer
Hybridization
Infectious heredity
Models
Applications
Examples
References
External links
Further reading

Underlying mechanisms and processes

Since the nineteenth century, scientists from different disciplines have studied how reticulate evolution
occurs. Researchers have increasingly succeeded in identifying these mechanisms and processes. It has been
found to be driven by symbiosis, symbiogenesis (endosymbiosis), lateral gene transfer, hybridization and
infectious heredity.[2]

Symbiosis

Symbiosis is a close and long-term biological interaction between two different biological organisms.[5]
Often, both of the organisms involved develop new features upon the interaction with the other organism.
This may lead to the development of new, distinct organisms.[6][7] The alterations in genetic material upon
symbiosis can occur via germline transmission or lateral transmission.[2][8][9] Therefore, the interaction
between different organisms can drive evolution of one or both organisms.[5]

Symbiogenesis

Symbiogenesis (endosymbiosis) is a special form of symbiosis whereby an organism lives inside another,
different organism. Symbiogenesis is thought to be very important in the origin and evolution of eukaryotes.
Eukaryotic organelles, such as mitochondria, have been theorized to have been originated from cell-invaded
bacteria living inside another cell.[10][11]

Lateral gene transfer

Lateral gene transfer, or horizontal gene transfer, is the movement of genetic material between unicellular
and/or multicellular organisms without a parent-offspring relationship. The horizontal transfer of genes
results in new genes, which could give new functions to the recipient and thus could drive evolution.[12]

Hybridization

In the neo-Darwinian paradigm, one of the assumed definition of a species is that of Mayr’s, which defines
species based upon sexual compatibility.[13] Mayr’s definition therefore suggests that individuals that can
produce fertile offspring must belong to the same species. However, in hybridization, two organisms
produce offspring while being distinct species.[2] During hybridization the characteristics of these two
different species are combined yielding a new organism, called a hybrid, thus driving evolution.[14]

Infectious heredity

Infectious agents, such as viruses, can infect the cells of host organisms. Viruses infect cells of other
organisms in order to enable their own reproduction. Hereto, many viruses can insert copies of their genetic
material into the host genome, potentially altering the phenotype of the host cell.[15][16][17] When these
viruses insert their genetic material in the genome of germ line cells, the modified host genome will be
passed onto the offspring, yielding genetically differentiated organisms. Therefore, infectious heredity plays
an important role in evolution,[2] for example in the formation of the female placenta.[18][19]

Models
Reticulate evolution has played a key role in the evolution of some organisms such as bacteria and flowering
plants.[20][21] However, most methods for studying cladistics have been based on a model of strictly
branching cladogeny, without assessing the importance of reticulate evolution.[22] Reticulation at
chromosomal, genomic and species levels fails to be modelled by a bifurcating tree.[1]

According to Ford Doolittle, an evolutionary and molecular biologist: “Molecular phylogeneticists will have
failed to find the “true tree,” not because their methods are inadequate or because they have chosen the
wrong genes, but because the history of life cannot properly be represented as a tree”.[23]

Reticulate evolution refers to evolutionary processes which cannot be successfully represented using a
classical phylogenetic tree model,[24] as it gives rise to rapid evolutionary change with horizontal crossings
and mergings often preceding a pattern of vertical descent with modification.[25] Reconstructing
phylogenetic relationships under reticulate evolution requires adapted analytical methods.[26] Reticulate
evolution dynamics contradict the neo-Darwininan theory, compiled in the Modern Synthesis, by which the
evolution of life occurs through natural selection and is displayed with a bifurcating or ramificating pattern.
Frequent hybridisation between species in natural populations challenges the assumption that species have
evolved from a common ancestor by simple branching, in which branches are genetically isolated.[26][27]
The study of reticulate evolution is said to have been largely excluded from the modern synthesis.[4] The
urgent need for new models which take reticulate evolution into account has been stressed by many
evolutionary biologists, such as Nathalie Gontier who has stated "reticulate evolution today is a vernacular
concept for evolutionary change induced by mechanisms and processes of symbiosis, symbiogenesis, lateral
gene transfer, hybridization, or divergence with gene flow, and infectious heredity". She calls for an
extended evolutionary synthesis that integrates these mechanisms and processes of evolution.[25]

Applications
Reticulate evolution has been extensively applied to plant hybridization in agriculture and gardening. The
first commercial hybrids appeared in the early 1920s.[28] Since then, many protoplast fusion experiments
have been carried out, some of which were aimed at improvement of crop species.[29] Wild types possessing
desirable agronomic traits are selected and fused in order to yield novel, improved species. The newly
generated plant will be improved for traits such as better yield, greater uniformity, improved color, and
disease resistance.[30]

Examples
Reticulate evolution is regarded as a process that has shaped the histories of many organisms.[31] There is
evidence of reticulation events in flowering plants, as the variation patterns between angiosperm families
strongly suggests there has been widespread hybridisation.[32] Grant[20] states that phylogenetic networks,
instead of phylogenetic trees, arise in all major groups of higher plants. Stable speciation events due to
hybridisation between angiosperm species supports the occurrence of reticulate evolution and highlights the
key role of reticulation in the evolution of plants.[33]

Genetic transfer can occur across wide taxonomic levels in microorganisms and become stably integrated
into the new microbial populations,[34][35] as has been observed through protein sequencing.[36] Reticulation
in bacteria usually only involves the transfer of only a few genes or parts of these.[22] Reticulate evolution
driven by lateral gene transfer has also been observed in marine life.[37] Lateral genetic transfer of photo-
response genes between planktonic bacteria and Archaea has been evidenced in some groups, showing an
associated increase in environmental adaptability in organisms inhabiting photic zones.[38]
Moreover, in the well-studied Darwin finches signs of reticulate evolution can be observed. Peter and
Rosemary Grant, who carried out extensive research on the evolutionary processes of the Geospiza genus,
found that hybridization occurs between some species of Darwin finches, yielding hybrid forms. This event
could explain the origin of intermediate species.[39] Jonathan Weiner[40] commented on the observations of
the Grants, suggesting the existence of reticulate evolution: "To the Grants, the whole tree of life now looks
different from a year ago. The set of young twigs and shoots they study seems to be growing together in
some seasons, apart in others. The same forces that created these lines are moving them toward fusion and
then back toward fission."; and "The Grants are looking at a pattern that was once dismissed as
insignificant in the tree of life. The pattern is known as reticulate evolution, from the Latin reticulum,
diminutive for net. The finches' lines are not so much lines or branches at all. They are more like twiggy
thickets, full of little networks and delicate webbings."

References
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External links
UA Geosciences, University of Arizona: Evolutionary Glossary (http://www.geo.arizona.edu/Ant
evs/ecol438/evolglos.html)
Hybridization and Reticulation: Trees, Networks, and Simulations (http://www.faculty.biol.ttu.ed
u/Strauss/Phylogenetics/LectureNotes/08_HybridReticul_Larsen.pdf)

Further reading
Arnold, M.L. (2008). Reticulate Evolution and Humans : Origins and Ecology. Oxford University
Press. ISBN 978-0-19-953958-1
Gontier, N. (2015). Reticulate Evolution Everywhere. In Reticulate Evolution: Symbiogenesis,
Lateral Gene Transfer, Hybridization and Infectious Heredity. Springer. ISBN 978-3-319-16344-
4
Linder, R.C.; Moret, B.M.E.; Nakhleh, L; Warnow, T. (2003). Network (Reticulate) Evolution:
Biology, Models and Algorithms (http://www.cs.utexas.edu/~phylo/resources/pdf/papers/psb04.
pdf). The University of Texas.
Linder, C. R., Rieseberg, L. H. (2004) Reconstructing patterns of reticulate evolution in plants
(http://www.amjbot.org/content/91/10/1700.full.pdf+html). American Journal of Botany,
91:1700-1708

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