Godfrey M. Hewitt

The genetic legacy of the Quaternary ice ages

Global climate has fluctuated greatly during the past three million years, leading to the recent major ice ages. An inescapable consequence for most living organisms is great changes in their distribution, which are expressed differently in boreal, temperate and tropical zones. Such range changes can be expected to have genetic consequences, and the advent of DNA technology provides most suitable markers to examine these. Several good data sets are now available, which provide tests of expectations, insights into species colonization and unexpected genetic subdivision and mixture of species. The genetic structure of human populations may be viewed in the same context. The present genetic structure of populations, species and communities has been mainly formed by Quaternary ice ages, and genetic, fossil and physical data combined can greatly help our understanding of how organisms were so affected.

Mitochondrial pseudogenes: evolution's misplaced witnesses

Nuclear copies of mitochondrial DNA (mtDNA) have contaminated PCR-based mitochondrial studies of over 64 different animal species. Since the last review of these nuclear mitochondrial pseudogenes (Numts) in animals, Numts have been found in 53 of the species studied. The recent evidence suggests that Numts are not equally abundant in all species, for example they are more common in plants than in animals, and also more numerous in humans than in Drosophila. Methods for avoiding Numts have now been tested, and several recent studies demonstrate the potential utility of Numt DNA sequences in evolutionary studies. As relics of ancient mtDNA, these pseudogenes can be used to infer ancestral states or root mitochondrial phylogenies. Where they are numerous and selectively unconstrained, Numts are ideal for the study of spontaneous mutation in nuclear genomes.

Speciation, hybrid zones and phylogeography - or seeing genes in space and time.

The origins and development of the study of speciation, hybrid zones and phylogeography are outlined using evolutionary iconography. This traces the ideas in this field from Lamarck and Darwin through to the present as represented in diagrams and figures. A ‘tree of trees’ summarizes this growth and current vitality. The new facility to use various DNA sequences from nuclear, mitochondrial and chloroplast genomes to determine genetic variation throughout a species range is examined particularly. There is great genomic subdivision across species distributions, which can be interpreted in the light of the recent demonstrations of severe palaeoclimatic oscillations. Refugia and postglacial colonization routes are proposed for several organisms across Europe. The role of geography in speciation through the Pleistocene is considered. These emerging principles and analyses are applied to data available on a variety of organisms in other regions of the world, such as the Arctic, North America and the Tropics, and including the progress of Homo sapiens through the last ice age. Some suggestions are made for future research directions.

Nuclear integrations: challenges for mitochondrial DNA markers

The combined use of mitochondrial DNA markers and polymerase chain reaction (PCR) techniques has greatly enhanced evolutionary studies. These techniques have also promoted the discovery of mitochondrial-like sequences in the nuclear genomes of many animals. While the nuclear sequences themselves are interesting, and capable of serving as valuable molecular tools, they can also confound phylogenetic and population genetic studies. Clearly, a better understanding of these phenomena and vigilance towards misleading data are needed.

Hybridization and speciation

Hybridization has many and varied impacts on the process of speciation. Hybridization may slow or reverse differentiation by allowing gene flow and recombination. It may accelerate speciation via adaptive introgression or cause near‐instantaneous speciation by allopolyploidization. It may have multiple effects at different stages and in different spatial contexts within a single speciation event. We offer a perspective on the context and evolutionary significance of hybridization during speciation, highlighting issues of current interest and debate. In secondary contact zones, it is uncertain if barriers to gene flow will be strengthened or broken down due to recombination and gene flow. Theory and empirical evidence suggest the latter is more likely, except within and around strongly selected genomic regions. Hybridization may contribute to speciation through the formation of new hybrid taxa, whereas introgression of a few loci may promote adaptive divergence and so facilitate speciation. Gene regulatory networks, epigenetic effects and the evolution of selfish genetic material in the genome suggest that the Dobzhansky–Muller model of hybrid incompatibilities requires a broader interpretation. Finally, although the incidence of reinforcement remains uncertain, this and other interactions in areas of sympatry may have knock‐on effects on speciation both within and outside regions of hybridization.

Spatial patterns of genetic variation generated by different forms of dispersal during range expansion

We examined the impact of three forms of dispersal, stepping-stone, normal and leptokurtic, on spatial genetic structure of expanding populations using computer simulations. When dispersal beyond neighbouring demes is allowed, rare long-distance migration leads to the establishment of pocket populations in advance of the main invasion front and results in spatial clustering of genotypes which persists for hundreds of generations. Patchiness is more pronounced when dispersal is leptokurtic as is the case in many animal and plant species. These results are of particular interest because population genetic parameters such as gene flow and effective population size are commonly estimated using gene frequency divergence information assuming equilibrium conditions and island models. We show how the three forms of dispersal during colonization bring about contrasting population genetic structures and how this affects estimates of gene flow. The implications for experimental studies of the spatial dimension of population genetic structure are discussed.

Hybrid Zones - Natural Laboratories for Evolutionary Studies

Hybrid zones are currently of interest because they offer experimental material for studies of the characters and processes involved in divergence and speciation. Parapatric subspecies and races may differ for a whole range of genes and characters, from short DNA sequences through enzymes and chromosomes to morphology and behaviour. An understanding of the origins, evolution, dynamics and fates of these hybrid zone patchworks requires information from a multitude of lines of enwuiry, including taxonomy, genetics, molecular biology, ethology, ecology, biogeography and palaeoclimatology.

Nuclear DNA analyses in genetic studies of populations: practice, problems and prospects.

Population‐genetic studies have been remarkably productive and successful in the last decade following the invention of PCR technology and the introduction of mitochondrial and microsatellite DNA markers. While mitochondrial DNA has proven powerful for genealogical and evolutionary studies of animal populations, and microsatellite sequences are the most revealing DNA markers available so far for inferring population structure and dynamics, they both have important and unavoidable limitations. To obtain a fuller picture of the history and evolutionary potential of populations, genealogical data from nuclear loci are essential, and the inclusion of other nuclear markers, i.e. single copy nuclear polymorphic (scnp) sequences, is clearly needed. Four major uncertainties for nuclear DNA analyses of populations have been facing us, i.e. the availability of scnp markers for carrying out such analysis, technical laboratory hurdles for resolving haplotypes, difficulty in data analysis because of recombination, low divergence levels and intraspecific multifurcation evolution, and the utility of scnp markers for addressing population‐genetic questions. In this review, we discuss the availability of highly polymorphic single copy DNA in the nuclear genome, describe patterns and rate of evolution of nuclear sequences, summarize past empirical and theoretical efforts to recover and analyse data from scnp markers, and examine the difficulties, challenges and opportunities faced in such studies. We show that although challenges still exist, the above‐mentioned obstacles are now being removed. Recent advances in technology and increases in statistical power provide the prospect of nuclear DNA analyses becoming routine practice, allowing allele‐discriminating characterization of scnp loci and microsatellite loci. This certainly will increase our ability to address more complex questions, and thereby the sophistication of genetic analyses of populations.

Insect mitochondrial control region: A review of its structure, evolution and usefulness in evolutionary studies

Abstract The control region is the only major non-coding region in the mitochondrial genome of insects. It is heavily biased to A+T nucleotides and seems to evolve under a strong directional mutation pressure. Among insects, this region is variable in both size and nucleotide sequence and may contain tandem repetition which is often associated with heteroplasmy. Tandem repetition appears to undergo concerted evolution and copy number variation indicates a high mutation rate. In contrast, the nucleotide substitution rate in this region is likely to be much reduced due to high A+T content and directional mutation pressure. Insect mitochondrial control regions are not necessarily the most variable region in the genome in terms of nucleotide substitution, and may not evolve faster than single-copy nuclear non-coding sequences. These observations have implications for the use of this region as a genetic marker in evolutionary studies. Contrary to earlier expectation, this region may have limited usefulness for both inter- and intra-specific analyses, depending on the structure and evolutionary patterns of a particular sequence. As some structural elements have been observed to be highly conserved between phylogenetically very distant insect taxa, it is of great interest to study the molecular evolution of this region in the entire class, Insecta.

Colonization and diversification: towards a phylogeographic synthesis for the Canary Islands.

Recently, the Canary Islands have become a focus for studies of the colonization and the diversification of different organisms. Some authors have considered Canarian endemisms as relicts of Tertiary origin, but new molecular data suggest a general pattern of continental dispersion followed by in situ speciation. Recent phylogeographic studies are revealing variants of the simple stepping-stone colonization model that seems to hold for many Hawaiian groups. Many factors can generate deviations from such a pattern: the stochastic nature of colonization, competitive exclusion, phylogenetic constraints on adaptive evolution and extinction. An understanding of island colonization and diversification can best be developed from an ecosystem level synthesis as more data for the Canarian archipelago come to hand.