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1.  The existence of species rests on a metastable equilibrium between inbreeding and outbreeding. An essay on the close relationship between speciation, inbreeding and recessive mutations 
Biology Direct  2011;6:62.
Speciation corresponds to the progressive establishment of reproductive barriers between groups of individuals derived from an ancestral stock. Since Darwin did not believe that reproductive barriers could be selected for, he proposed that most events of speciation would occur through a process of separation and divergence, and this point of view is still shared by most evolutionary biologists today.
I do, however, contend that, if so much speciation occurs, the most likely explanation is that there must be conditions where reproductive barriers can be directly selected for. In other words, situations where it is advantageous for individuals to reproduce preferentially within a small group and reduce their breeding with the rest of the ancestral population. This leads me to propose a model whereby new species arise not by populations splitting into separate branches, but by small inbreeding groups "budding" from an ancestral stock. This would be driven by several advantages of inbreeding, and mainly by advantageous recessive phenotypes, which could only be retained in the context of inbreeding. Reproductive barriers would thus not arise as secondary consequences of divergent evolution in populations isolated from one another, but under the direct selective pressure of ancestral stocks. Many documented cases of speciation in natural populations appear to fit the model proposed, with more speciation occurring in populations with high inbreeding coefficients, and many recessive characters identified as central to the phenomenon of speciation, with these recessive mutations expected to be surrounded by patterns of limited genomic diversity.
Whilst adaptive evolution would correspond to gains of function that would, most of the time, be dominant, this type of speciation by budding would thus be driven by mutations resulting in the advantageous loss of certain functions since recessive mutations very often correspond to the inactivation of a gene. A very important further advantage of inbreeding is that it reduces the accumulation of recessive mutations in genomes. A consequence of the model proposed is that the existence of species would correspond to a metastable equilibrium between inbreeding and outbreeding, with excessive inbreeding promoting speciation, and excessive outbreeding resulting in irreversible accumulation of recessive mutations that could ultimately only lead to extinction.
Reviewer names
Eugene V. Koonin, Patrick Nosil (nominated by Dr Jerzy Jurka), Pierre Pontarotti
PMCID: PMC3275546  PMID: 22152499
speciation; inbreeding; saeptation; mutation load; extinction; evolution
2.  The orthology of HLA-E and H2-Qa1 is hidden by their concerted evolution with other MHC class I molecules 
Biology Direct  2006;1:2.
Whether MHC molecules undergo concerted evolution or not has been the subject of a long-standing debate.
By comparing sequences of eight functional homologues of HLA-E from primates and rodents with those of MHC class Ia molecules from the same eight species, we find that different portions of MHC class I molecules undergo different patterns of evolution. By focusing our analyses sequentially on these various portions, we have obtained clear evidence for concerted evolution of MHC class I molecules, suggesting the occurrence of extensive interallelic and intergenic exchanges. Intra-species homogenisation of sequences is particularly noticeable at the level of exon 4, which codes for the α3 domain, but our results suggest that homogenisation also concerns certain residues of the α1–α2 codomain that lie outside the antigen recognition site.
A model is presented in which Darwinian selective pressures due to pathogens could, at the same time, favour diversification of MHC class Ia molecules and promote concerted evolution of separate loci by spreading advantageous motifs arising by mutations in individual MHC molecules to other alleles and to other loci of the MHC region. This would also allow MHC molecules to co-evolve with the proteins with which they interact to fulfil their functions of antigen presentation and regulation of NK cell activity. One of the raisons d'être of the MHC may therefore be to favour at the same time both diversification of MHC class Ia molecules and homogenisation of the whole pool of MHC class I molecules (Ia and Ib) involved in antigen presentation.
This article was reviewed by Stephan Beck, Lutz Walter and Pierre Pontarotti.
PMCID: PMC1403747  PMID: 16542007
3.  Various hypotheses on MHC evolution suggested by the concerted evolution of CD94L and MHC class Ia molecules 
Biology Direct  2006;1:3.
In the accompanying paper by Virginie Rouillon and myself, our demonstration that homogenisation by gene conversion occurs readily among MHC class I genes was made possible because of the exceptional conservation of the CD94L locus between divergent species of separate taxa, suggesting that the molecules of this family are endowed with very important and well preserved biological functions. These results lead me to elaborate various hypotheses on several aspects of MHC evolution.
In a first part, I propose a highly hypothetical scenario of MHC evolution that could explain how modern day CD94L molecules can have so many diverse and well preserved biological functions. Next, I propose that MHC class I molecules evolve more rapidly and exuberantly than class II molecules because the former are subjected to more direct selective pressures, in particular from viruses. Third, I suggest that concerted evolution, by increasing inter-genic homogeneity would in turn favour further inter-allelic and inter-loci exchanges, hence resulting in a more evolvable MHC. As a fourth and last point, I propose that the high GC content of sequences coding for classical class I molecules could be a consequence of biased gene conversion.
Testing of these various hypotheses should occur naturally over the coming years, with the ever increasing availability of more sequences related to MHC class I genes from various organisms. Ultimately, a better understanding of how MHC molecules evolve may help to decipher where and how our adaptive immune system arose, and keeps evolving in the face of the permanent challenge of infectious organisms.
This article was reviewed by Stephan Beck, Lutz Walter and Pierre Pontarotti.
PMCID: PMC1397804  PMID: 16542034

Results 1-3 (3)