THE LOCI OF EVOLUTION: HOW PREDICTABLE IS GENETIC EVOLUTION?

Is genetic evolution predictable? Evolutionary developmental biologists have argued that, at least for morphological traits, the answer is a resounding yes. Most mutations causing morphological variation are expected to reside in the cis‐regulatory, rather than the coding, regions of developmental genes. This “cis‐regulatory hypothesis” has recently come under attack. In this review, we first describe and critique the arguments that have been proposed in support of the cis‐regulatory hypothesis. We then test the empirical support for the cis‐regulatory hypothesis with a comprehensive survey of mutations responsible for phenotypic evolution in multicellular organisms. Cis‐regulatory mutations currently represent approximately 22% of 331 identified genetic changes although the number of cis‐regulatory changes published annually is rapidly increasing. Above the species level, cis‐regulatory mutations altering morphology are more common than coding changes. Also, above the species level cis‐regulatory mutations predominate for genes not involved in terminal differentiation. These patterns imply that the simple question “Do coding or cis‐regulatory mutations cause more phenotypic evolution?” hides more interesting phenomena. Evolution in different kinds of populations and over different durations may result in selection of different kinds of mutations. Predicting the genetic basis of evolution requires a comprehensive synthesis of molecular developmental biology and population genetics.     

HOW COMMON IS HOMOPLOID HYBRID SPECIATION?

Hybridization has long been considered a process that prevents divergence between species. In contrast to this historical view, an increasing number of empirical studies claim to show evidence for hybrid speciation without a ploidy change. However, the importance of hybridization as a route to speciation is poorly understood, and many claims have been made with insufficient evidence that hybridization played a role in the speciation process. We propose criteria to determine the strength of evidence for homoploid hybrid speciation. Based on an evaluation of the literature using this framework, we conclude that although hybridization appears to be common, evidence for an important role of hybridization in homoploid speciation is more circumscribed.     

ESTIMATION OF TOTAL BODY WATER IN HARBOR SEALS: HOW USEFUL IS BIOELECTRICAL IMPEDANCE ANALYSIS?

We evaluated bioelectrical impedance analysis (BIA) as a means of rapidly and inexpensively estimating total body water (TBW) of harbor seals ( Phoca vitulina ). Deuterium oxide dilution was used to estimate TBW in 17 adult females and 16 of their pups between birth and late lactation. Isotope dilution was also used to determine TBW in 12 adult males early and 10 of these males late in the breeding season. At the same time, resistance ( Rs ) and reactance ( Xc ) measurements were taken using a tetrapolar, impedance plethysmograph (Model 101 A, RJL Systems). Seals were sedated with diazepam prior to taking BIA measurements. Within-day duplicate Rs measurements on pups and adults, taken 2-240 min apart, differed by an average of 3.0%± 1.4% ( n = 42, CV = 102%). Movement of the seal during BIA measurements caused variability in both Rs and Xc values. BIA measurements were generally poor predictors of TBW. Rs was significantly correlated with TBW in pups only ( Rs = 0.93, P = 0.001, n = 11). Bioelectrical conductor volume (length2/ Rs ) was significantly correlated with TBW only in adult females ( Rs = 0.63, P = 0.02, n = 14). We conclude that BIA is not a reliable method of estimating TBW in wild harbor seals.     

THE CLASSIFICATION OF PROBOSCIDEA: HOW MANY CLADISTIC CLASSIFICATIONS?

Abstract— Hennig conceived a method to build a "phylogenetic system", with the stipulation that a "properly drawn phylogenetic tree must be directly translatable into the language of phylogenetic systematics". Consequently, this system could be the general reference system of biology. A review of the classificatory technical improvements, conventions and rules which have been proposed for the past twenty years together with their application to the classification of the Proboscidea, leads to the conclusion that more than one formal system can be built upon one given cladogram. As words are used more frequently for communication than diagrams, schemes or graphs, the "general reference system of biology1' remains somewhere in Utopia. The "phylogenetic system" is rather more synonymous with a cladogram than with a written classification.     

UNREQUESTED TERMINATION OF LIFE: IS IT PERMISSIBLE?

Conclusion: One can appreciate and understand why doctors take action to terminate life without being requested by the patient who is suffering unbearably and pointlessly. But in my opinion such action is not permissible from a moral point of view. In the first place there is a danger that interests other than those of the patient will play a decisive role. Furthermore, it is not permissible because the interest of society as a whole is at stake. The safety of all future patients would no longer be guaranteed. It is also not allowed because the medical profession must constantly be on its guard so that it does not become caught up in the situations described above. In nearly every case it should be possible for doctor and patient to discuss the termination of life explicitly and at an early stage. The ultimate reason why termination of life without request of the patient is not permissible is that in principle it will never be possible to draw a clear dividing line between unrequested and unwanted termination of life. However, it is conceivable that in certain cases the conflict of loyalties will be such that unrequested termination of life is justified. In the rare extreme case of an inescapable conflict of equally stringent duties, it may just be necessary to do something which in essence is impermissible.     

HOW MANY SPECIES OF ALGAE ARE THERE?

Algae have been estimated to include anything from 30,000 to more than 1 million species. An attempt is made here to arrive at a more accurate estimate using species numbers in phyla and classes included in the on‐line taxonomic database AlgaeBase ( http://www.algaebase.org ). Despite uncertainties regarding what organisms should be included as algae and what a species is in the context of the various algal phyla and classes, a conservative approach results in an estimate of 72,500 algal species, names for 44,000 of which have probably been published, and 33,248 names have been processed by AlgaeBase to date (June 2012). Some published estimates of diatom numbers are of over 200,000 species, which would result in four to five diatom species for every other algal species. Concern is expressed at the decline and potential extinction of taxonomists worldwide capable of improving and completing the necessary systematic studies.     

HOW REPEATABLE IS ADAPTIVE EVOLUTION? THE ROLE OF GEOGRAPHICAL ORIGIN AND FOUNDER EFFECTS IN LABORATORY ADAPTATION

The importance of contingency versus predictability in evolution has been a long-standing issue, particularly the interaction between genetic background, founder effects, and selection. Here we address experimentally the effects of genetic background and founder events on the repeatability of laboratory adaptation in Drosophila subobscura populations for several functional traits. We found disparate starting points for adaptation among laboratory populations derived from independently sampled wild populations for all traits. With respect to the subsequent evolutionary rate during laboratory adaptation, starvation resistance varied considerably among foundations such that the outcome of laboratory evolution is rather unpredictable for this particular trait, even in direction. In contrast, the laboratory evolution of traits closely related to fitness was less contingent on the circumstances of foundation. These findings suggest that the initial laboratory evolution of weakly selected characters may be unpredictable, even when the key adaptations under evolutionary domestication are predictable with respect to their trajectories.     

SELECTION ON MOTHERS AND OFFSPRING: WHOSE PHENOTYPE IS IT AND DOES IT MATTER?

Reproductive and early life-history traits can be considered aspects of either offspring or maternal phenotype, and their evolution will therefore depend on selection operating through offspring and maternal components of fitness. Furthermore, selection at these levels may be antagonistic, with optimal offspring and maternal fitness occurring at different phenotypic values. We examined selection regimes on the correlated traits of birth weight, birth date, and litter size in Soay sheep (Ovis aries) using data from a long-term study of a free-living population on the archipelago of St. Kilda, Scotland. We tested the hypothesis that selective constraints on the evolution of the multivariate phenotype arise through antagonistic selection, either acting at offspring and maternal levels, or on correlated aspects of phenotype. All three traits were found to be under selection through variance in short-term and lifetime measures of fitness. Analysis of lifetime fitness revealed strong positive directional selection on birth weight and weaker selection for increased birth date at both levels. However, there was also evidence for stabilizing selection on these traits at the maternal level, with reduced fitness at high phenotypic values indicating lower phenotypic optima for mothers than for offspring. Additionally, antagonistic selection was found on litter size. From the offspring's point of view it is better to be born a singleton, whereas maternal fitness increases with average litter size. The decreased fitness of twins is caused by their reduced birth weight; therefore, this antagonistic selection likely results from trade-offs between litter size and birth weight that have different optimal resolutions with respect to offspring and maternal fitness. Our results highlight how selection regimes may vary depending on the assignment of reproductive and early life-history traits to either offspring or maternal phenotype.     

HOW IS THE RATE OF CLIMATIC‐NICHE EVOLUTION RELATED TO CLIMATIC‐NICHE BREADTH?

The rate of climatic‐niche evolution is important to many research areas in ecology, evolution, and conservation biology, including responses of species to global climate change, spread of invasive species, speciation, biogeography, and patterns of species richness. Previous studies have implied that clades with higher rates of climatic‐niche evolution among species should have species with narrower niche breadths, but there is also evidence suggesting the opposite pattern. However, the relationships between rate and breadth have not been explicitly analyzed. Here, we examine the relationships between the rate of climatic‐niche evolution and climatic‐niche breadth using phylogenetic and climatic data for 250 species in the salamander family Plethodontidae, a group showing considerable variation in both rates of climatic‐niche evolution and climatic‐niche breadths. Contrary to some expectations, we find no general relationship between climatic‐niche breadth and the rate of climatic‐niche evolution. Climatic‐niche breadths for some ecologically important climatic variables considered separately (temperature seasonality and annual precipitation) do show significant relationships with the rate of climatic‐niche evolution, but rates are faster in clades in which species have broader (not narrower) niche breadths. In summary, our results show that narrower niche breadths are not necessarily associated with faster rates of niche evolution.     

MATHEMATICS OF KIN‐ AND GROUP‐SELECTION: FORMALLY EQUIVALENT?

Evolutionary game theory is a general mathematical framework that describes the evolution of social traits. This framework forms the basis of many multilevel selection models and is also frequently used to model evolutionary dynamics on networks. Kin selection, which was initially restricted to describe social interactions between relatives, has also led to a broader mathematical approach, inclusive fitness, that can not only describe social evolution among relatives, but also in group structured populations or on social networks. It turns out that the underlying mathematics of game theory is fundamentally different from the approach of inclusive fitness. Thus, both approaches—evolutionary game theory and inclusive fitness—can be helpful to understand the evolution of social traits in group structured or spatially extended populations.