Encyclopedia of Evolution.pdf - Online Reading Center
Encyclopedia of Evolution.pdf - Online Reading Center
Encyclopedia of Evolution.pdf - Online Reading Center
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cladistics<br />
The above examples used easily visible characteristics to<br />
classify organisms into groups. The characteristics need not<br />
be visible. In fact, many cladistic analyses use DNA sequence<br />
information (see bioinformatics; DNA [evidence for<br />
evolution]). Suppose the scientist has segments <strong>of</strong> DNA<br />
from three species. If the segment <strong>of</strong> DNA is 100 nucleotides<br />
in length, this allows 100 characters to be used to test<br />
each cladogram. Nearly every phylogenetic study includes at<br />
least one analysis that uses DNA nucleotides as traits. It is<br />
no easier to use DNA than to use visible characters in cladistic<br />
analysis. The reason is that one cannot compare just any<br />
old piece <strong>of</strong> DNA <strong>of</strong> one species with any old piece <strong>of</strong> DNA<br />
in another, any more than one can make comparisons <strong>of</strong> just<br />
any visible trait. Just as visible traits must be homologous,<br />
so the DNA segments must be homologous. The investigator<br />
must compare the same gene in all <strong>of</strong> the species in the analysis.<br />
Further complications arise when one considers that some<br />
mutations exchange one nucleotide for another, while other<br />
mutations add or delete nucleotides, or reverse segments, or<br />
even move segments to different places. Usually, researchers<br />
use certain well-studied DNA sequences rather than trying to<br />
figure a new one out for themselves. In comparisons among<br />
plants, matK (the gene for an enzyme that acts upon the<br />
amino acid tyrosine) and rbcL (the gene for the large subunit<br />
<strong>of</strong> the enzyme rubisco) are frequently used.<br />
Some analyses use noncoding DNA for phylogenetic<br />
analysis. Researchers may use transposable elements (see<br />
horizontal gene transfer) because homoplasy is very<br />
unlikely to occur with them. It is very unlikely that the same<br />
transposable element would evolve twice: It would neither<br />
insert itself in the same location in the chromosomes <strong>of</strong> two<br />
different species, nor transfer to a new location without gaining<br />
or losing a few nucleotides in the process. Other analyses<br />
use short or long interspersed sequences in the noncoding<br />
DNA. The figure on page 73 uses the presence or absence <strong>of</strong><br />
20 segments <strong>of</strong> noncoding DNA to construct a cladogram <strong>of</strong><br />
mammals. Cows and deer are clustered together, because they<br />
share all <strong>of</strong> the segments for which information is available.<br />
Whales and hippos are clustered together, because they share<br />
15 <strong>of</strong> the 17 segments for which information is available.<br />
Whales and cows are not clustered closely together, because<br />
they share only nine <strong>of</strong> the 17 segments for which information<br />
is available. Camels are not closely related to cows, as<br />
they have none <strong>of</strong> the 11 DNA segments that cows possess.<br />
Choosing the best cladogram<br />
A computer can generate many possible cladograms for any<br />
set <strong>of</strong> species that are being compared. Which <strong>of</strong> these cladograms<br />
is the correct one? The correct one is the one that most<br />
closely matches what actually happened during evolutionary<br />
history. Nobody can know exactly what happened during<br />
evolutionary history; cladists must make an assumption about<br />
which cladogram is best. They <strong>of</strong>ten make the assumption<br />
<strong>of</strong> parsimony (see scientific method), that is, they choose<br />
the simplest explanation. To do this, they choose the cladogram<br />
that requires the fewest number <strong>of</strong> character changes.<br />
Consider this simplified example in the classification <strong>of</strong> dogs,<br />
cats, and lions. Since each group has two branches, one could<br />
classify them in three ways:<br />
1. [Dogs] [Cats, Lions]<br />
2. [Dogs, Cats] [Lions]<br />
3. [Dogs, Lions] [Cats]<br />
Cats and lions both have retractable claws; dogs do not.<br />
Cladograms 2 and 3 require that retractable claws should<br />
have evolved twice, separately in cats and lions. Therefore<br />
cladogram 1 is the most parsimonious. It would not be safe<br />
to draw a conclusion based upon just one characteristic (in<br />
this case, retractable claws). Cladograms based on any <strong>of</strong> a<br />
number <strong>of</strong> other characteristics would lead to the same conclusion:<br />
Cats and lions form the feline group, while dogs are<br />
part <strong>of</strong> a separate (canine) group. From this one would conclude<br />
that cats and lions shared a common ancestor that lived<br />
much more recently than the common ancestor <strong>of</strong> canines<br />
and felines.<br />
It is easy enough to compare the three cladograms that<br />
are possible with three species. However, as more species are<br />
compared, the analysis becomes exponentially more difficult.<br />
When comparing even a slightly larger number <strong>of</strong> species, the<br />
calculations become so complex that they can only be performed<br />
by a computer. As the number <strong>of</strong> species in the analysis<br />
increases, even a computer cannot sort through all <strong>of</strong> the<br />
possibilities. For 10 species, there are more than two million<br />
possible cladograms; for 50 species, there are 3 × 10 76 possible<br />
cladograms. To provide some idea <strong>of</strong> how large a number<br />
this is, consider that the universe has existed for less than<br />
10 18 seconds! The computer must randomly generate a subset<br />
<strong>of</strong> several hundred possible cladograms and make comparisons<br />
among them.<br />
Even when a cladistic analysis compares only a few hundred<br />
<strong>of</strong> the possible cladograms, there may be a large number<br />
<strong>of</strong> almost equally parsimonious ones. The computer must<br />
then generate a consensus tree that preserves the patterns that<br />
the best cladograms share in common with one another.<br />
Applying cladistics to classification<br />
A cladistic approach to classification would group organisms<br />
together on the basis <strong>of</strong> the branch points in the cladogram.<br />
A clade (whether it corresponds to a genus, a family,<br />
an order, a class, a phylum, or a kingdom; see Linnaean<br />
system) must consist <strong>of</strong> all species that share a common<br />
ancestor, and only those species. Such a clade is called monophyletic<br />
(mono- means one). Cladists will not accept a genus,<br />
family, order, class, phylum, or kingdom that includes only<br />
some <strong>of</strong> the species that have evolved from a common ancestor.<br />
Such a group is called paraphyletic (para- means beside).<br />
Cladists most vigorously reject taxonomic groups that both<br />
omit some species that share a common ancestor and include<br />
some species that do not, a group referred to as polyphyletic<br />
(poly- means many).<br />
A cladistic approach can lead to a different system <strong>of</strong><br />
classification than what is traditionally recognized, as in these<br />
examples: