Avaliação 1 PEMARF

Nome: Caio Gueratto Coelho da Silva

Público-Alvo: Estudantes de graduaçaã o em Biologia

Título

Neutral theory in Taxonomy: Implications and recent debates.

Introdução

The neutral theory was one of the most impactful ideas of 20 th century (Dietrich, 1994). It was so
because the prevailing hypothesis by the time Kimura presented his findings, in 1968, was that mutations in
the genetic material would rarely occur. Scentists accepted this idea because they believed that nucleotide
mutations in DNA would be predominantly deleterious. Following this rational, populations would have
mutated DNA strings eliminated. Therefore, the mutational rate would be very slow. Researchers analyzed
proteins structure of different mammals groups (Buettner-Janusch & Hill, 1965; Kaplan, 1965) and estimated
that one aminoacid was replaced in a scale of 106-107 year, depending on the molecule. People thought that
this was also the case for genome.
What the neutral theory (Kimura, 1968) states is that, in genome level, the opposite occurs. When
Kimura calculated mutational rate of some coding genes, it was clear that it was much faster than expected by
the prevailing hypothesis. The estimated mutational rate in the genomic level per generation was 200~2000
replacements (Kimura, 1968). The conclusion was that most part of mutations occuring in the genome is
neutral. These changes are not deleterious neither advantageous. Because neutral mutations are not
deleterious, they occur without being eliminated, and can occur at high evolutionary rate (Kimura, 1991).
The publishing of the neutral theory changed scientists' view about genomic mutation and it is
possible to see its repercussion in several biologic areas. One of these areas is Taxonomy. In the following
sessions I will explain how this theory opened space to scientists apply molecular techniques to species
inference and delimitation. I will also expose one debate topic that still occur recently entangled by the
neutral theory.

Desenvolvimento

Molecular techniques applied to species inference and delimitation is one of the implications of neutral
theory for Taxonomy (Dietrich, 1994). With the arising of the concept that genome changes occur in a
constant rate, it is possible to access the time scale separating two or more lineages. This means that if one
has the the genetic distance between lineages, it is possible to infer how much time took their divergence and
in which pattern. So, genetic distance can be converted to time, once we know that mutational rates occur in a
constant pace (Ricklefs, 2006). This idea of a constant rate came from the neutral theory, as we saw in the
previous session. One example of this constancy is in mitochondrial DNA. A common calibration for mtDNA
genes in birds is 2% sequence divergence between sister lineages per million years (Lovette, 2004 apud
Ricklefs, 2006). This calibrations varies greatly in different groups of organisms.
In fact, besides using these estimations for species delimitation, taxonomists can use them to help
estimating divergence time of any lineages: species, genus, family or even more inclusive taxa. The only
difference is the gene set one uses to make estimations. Different genes have different substitution rates. As I
said in the previous paragraph, mtDNA calibration is 2% divergence. These are very fast evolving genes.
Therefore, taxonomists use some of these genes (and other fast evolving regions of genome, as microsatilites)
to estimate divergences between species or even populations. For 28S, a nuclear very conservative gene, the
divergence rates between sister species is probably 0 or a very low value. This makes the usage of this gene
imprecise to separate species. However, 28S is used to infer phylogenetic relatioship of more inclusive taxa,
such as families (Pinto-da-Rocha et al., 2014), orders (Ishiwata et al., 2011) and even huge domains (Baldauf,
2008).
The use of molecular data to create hypothesis regarding species is particularly important in cryptic or
very variable species. This happens because, in these cases, morphological data only is not informative to
define species. A good example is with the cryptic butterfly species Astraptes fulgerator (Herbert et al., 2004).
In a work analyzing the Citochrome Oxidase I, a mitochondrial gene, the authors found out that instead of only
one species, the individuals called of Astrapes fulgerator could be divided into 10 distinct clades. These clades
varied accordingly to ecological and some subtle morphological differences.
As we can see, implications of neutral theory in Taxonomy are important. These implications have
effects until nowadays' Taxonomy discussions. One very important discussion topic is about species
delimitation using molecular data only (DeSalle et al., 2005). Recently, the publishing of works showing only
the molecular data of lineages and concluding that the lineages are new species has become common (Herbert
et al., 2003a,b; 2004a,b). Traditional morphologists state that only molecular data does not say much about
whether a lineage is a species or not (Lipscomb et al, 2003). They state that because methods of species
delimitation are controversial (e.g. Sukumaran & Knowles, 2016). Moreover, divergence values in DNA
sequence can be arbitrary to define different species. Each group of species present one specific value of
divergence. Therefore, it becomes complicated to infer which is the cutoff value to state that two different
sister lineages are two different species or two populations of a single species (Lipscomb et al., 2003). On the
other hand, “molecular taxonomists” state that this problem of species delimitation is real when analysing
morphology too (Herbert et al., 2003a). They use the examples of cryptic species as a strong argument to say
that is impossible to delimit these species without use of molecular methods. This debate only occurs as an
implications of the neutral theory, as it would be impossible inferring anything regarding molecular
relationships of very closely related lineages without this unifying theory.

Conclusão

In light of the foregoing, we can consider that Neutral theory had, and still has, a great impact on Taxonomy
studies. Without it, one could not infer about molecular species delimitation and population structure. It is
due to the neutralists findings about genes evolving in a constant rate, and about differences in the
substitution rates for each gene, that we can create hypothesis about what delimits a species, which set of
genes use to build phylogenetic hypothesis for a particular taxonomic level, and whether two closely related
taxa are different species or not.

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