Understanding Trichotomy
Are you more closely related to a sibling or a first cousin? The answer is obvious, but the question is not as silly as it sounds. If you were trying to determine the relationship of three people (two siblings and a first cousin) based on their DNA sequence, you would get contradictory information. For most informative loci, you would make the correct conclusion. However some of the time you would erroneously group the cousin and one of the siblings together because they shared genes that were absent in the other sibling. So overall, you are more closely related (genetically) to your sibling, but there are parts of your genome where your cousin is your closest relative.
Applying this scenario to a larger scale helps us to understand the trichotomy problem in determining species relationships, such as the relationship between humans, chimpanzees, and gorillas. You've probably heard that our genome sequence is very similar (94-98.5%, depending on how you measure) to chimpanzees. The problem is that both genomes are also very similar to gorilla. Depending on where you look, you might group gorillas and humans together, or gorillas and chimpanzees together. The problem is illustrated in panel A of the the following figure [1]. (Click for larger image.)
The meaning of PI-characters and RGCs is not important here. All you need to know is that, along with genes, these are characteristics in genomes that can be compared. As the figure shows, in most cases the human-chimpanzee relationship is supported, however in a minority of cases other relationships are supported.
Why is there support for alternative relationships? The answer is analogous to the sibling/cousin relationship scenario above. Humans, chimpanzees and gorillas each have their roots in a common gene pool. However, the sorting of ancestral polymorphic alleles in the diverging lineages is subject to evolutionary processes such as genetic drift. Thus purely by chance, alleles can become fixed in a way that gives a different picture than the true species-branching pattern.
The concept is illustrated in the following figure [2]:
Here, the broad lines represent what the true relationship of humans, chimpanzees, and gorillas is thought to be. The colored lines represent hypothetical gene polymorphisms. Notice that the true species relationship can differ from that inferred by looking only at the polymorphisms (esp. HG and CG, which give results as in panel A of the first figure).
1. Rokas A, Carroll SB (2006) Bushes in the Tree of Life. PLoS Biol 4(11)
2. Hobolth A, Christensen OF, Mailund T, Schierup MH (2007) Genomic Relationships and Speciation Times of Human, Chimpanzee, and Gorilla Inferred from a Coalescent Hidden Markov Model. PLoS Genet 3(2): e7