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▪ Abstract  Trans-species polymorphism (TSP) is the occurrence of similar alleles in related species. Excluding instances in which the similarity arose by convergent evolution, TSP is generated by the passage of alleles from ancestral to descendant species. Closely related, recently diverged species, such as those of the Lake Victoria cichlid flock, may share neutral alleles, but long-lasting TSPs occur only in genetic systems evolving under balancing selection. Two such systems have been studied extensively, the major histocompatibility complex (Mhc) of jawed vertebrates and the self-incompatibility (SI) system of flowering plants. Allelic lineages that diverged many millions of years ago and passed through numerous speciation events have been described in both systems. The lineages may differ at up to 50% of their coding sites, both synonymous and nonsynonymous. The differences arise by the process of incorporation of mutations, which is different from the process of fixation. TSP, on the one hand, complicates phylogenetic analysis, but on the other, it is a useful tool for the study of speciation.

Although osteocytes are the most abundant cells in bone, little is known about their function, and no specific marker protein for osteocytes has been described. Dentin matrix protein 1 (DMP1) is an acidic phosphoprotein expressed in tooth organ and bone. Our previous work showed that in the chicken, which is not capable of forming tooth, DMP1 messenger RNA (mRNA) is highly expressed in bone by Northern blot analysis. To clarify the significance of DMP1 expression in bone, the expression of DMP1 mRNA and its protein was examined in the chicken and rat. In the chicken, DMP1 mRNA was detected only in bone tissues and was localized in osteocytes and preosteocytes but not in osteoblasts. Similarly, in the rat, DMP1 mRNA was predominantly expressed in osteocytes and preosteocytes in bone matrix but not in osteoblasts located at the bone surface. Antiserum was raised against the peptide from rat DMP1, and the localization of DMP1 was examined by immunohistochemistry. In the development of bone, DMP1 was first detected in newly formed bone matrix after osteoblastic cells had been embedded within it. After the appearance of typical osteocytes, DMP1 was localized in the pericellular bone matrix of osteocytes, including their processes. These data show that DMP1 is a bone matrix protein specifically expressed in osteocytes and preosteocytes and suggest that DMP1 plays a role in bone homeostasis because of its high calcium ion-binding capacity.

The acronym Mhc, major histocompatibility complex, is customarily not allied with topics in evolutionary biology. Here, however, we attempt to demonstrate that the Mhc has much to offer to this discipline and intimate that evolutionary biologists who ignore its contributions miss out on a chance of applying a new approach to vexing questions. One aspect of the Mhc in particular affords a fresh look at the population processes that transform one species into another: the trans-species polymorphism, the passage of allelic lineages from ancestral to descendant species. We provide examples of using the Mhc polymorphism in estimating the size of the founding population of new species, and of analyzing the long-term population demographies of phylogenetic lineages. We then extend the concept of trans-species polymorphism to other genes, even those not evolving under balancing selection, and argue that the phenomenon is widespread between closely related species. On the example of the cichlid fishes of Lake Victoria, we demonstrate how the concept changes the interpretation of this so-called "species flock." We contend that the conclusions reached regarding the cichlid fishes apply also to other examples of adaptive radiation, for example that of Darwin's finches, and so provide new insights into the nature of speciation in general.

Darwin's finches comprise a group of passerine birds first collected by Charles Darwin during his visit to the Galápagos Archipelago. The group, a textbook example of adaptive radiation (the diversification of a founding population into an array of species differentially adapted to diverse environmental niches), encompasses 14 currently recognized species, of which 13 live on the Galápagos Islands and one on the Cocos Island in the Pacific Ocean. Although Darwin's finches have been studied extensively by morphologists, ecologists, and ethologists, their phylogenetic relationships remain uncertain. Here, sequences of two mtDNA segments, the cytochrome b and the control region, have been used to infer the evolutionary history of the group. The data reveal the Darwin's finches to be a monophyletic group with the warbler finch being the species closest to the founding stock, followed by the vegetarian finch, and then by two sister groups, the ground and the tree finches. The Cocos finch is related to the tree finches of the Galápagos Islands. The traditional classification of ground finches into six species and tree finches into five species is not ref lected in the molecular data. In these two groups, ancestral polymorphisms have not, as yet, been sorted out among the cross-hybridizing species.