Effect of social structure and introduction history on genetic diversity and differentiation.

Invasive species are a global threat to biodiversity, and understanding their history and biology is a major goal of invasion biology. Population-genetic approaches allow insights into these features, as population structure is shaped by factors such as invasion history (number, origin and age of introductions) and life-history traits (e.g., mating system, dispersal capability). We compared the relative importance of these factors by investigating two closely related ants, Tetramorium immigrans and Tetramorium tsushimae, that differ in their social structure and invasion history in North America. We used mitochondrial DNA sequences and microsatellite alleles to estimate the source and number of introduction events of the two species, and compared genetic structure among native and introduced populations. Genetic diversity of both species was strongly reduced in introduced populations, which also differed genetically from native populations. Genetic differentiation between ranges and the reduction in microsatellite diversity were more severe in the more recently introduced and supercolonial T. tsushimae. However, the loss of mitochondrial haplotype diversity was more pronounced in T. immigrans, which has single-queen colonies and was introduced earlier. Tetramorium immigrans was introduced at least twice from Western Europe to North America and once independently to South America. Its monogyny might have limited genetic diversity per introduction, but new mutations and successive introductions over a long time may have added to the gene pool in the introduced range. Polygyny in T. tsushimae probably facilitated the simultaneous introduction of several queens from a Japanese population to St. Louis, USA. In addition to identifying introduction pathways, our results reveal how social structure can influence the population-genetic consequences of founder events.

Intraspecific support for the polygyny-vs.-polyandry hypothesis in the bulldog ant Myrmecia brevinoda.

The number of queens per colony and the number of matings per queen are the most important determinants of the genetic structure of ant colonies, and understanding their interrelationship is essential to the study of social evolution. The polygyny-vs.-polyandry hypothesis argues that polygyny and polyandry should be negatively associated because both can result in increased intracolonial genetic variability and have costs. However, evidence for this long-debated hypothesis has been lacking at the intraspecific level. Here, we investigated the colony genetic structure in the Australian bulldog ant Myrmecia brevinoda. The numbers of queens per colony varied from 1 to 6. Nestmate queens within polygynous colonies were on average related (r(qq) = 0.171 ± 0.019), but the overall relatedness between queens and their mates was indistinguishable from zero (r(qm) = 0.037 ± 0.030). Queens were inferred to mate with 1-10 males. A lack of genetic isolation by distance among nests indicated the prevalence of independent colony foundation. In accordance with the polygyny-vs.-polyandry hypothesis, the number of queens per colony was significantly negatively associated with the estimated number of matings (Spearman rank correlation R = -0.490, P = 0.028). This study thus provides the rare intraspecific evidence for the polygyny-vs.-polyandry hypothesis. We suggest that the high costs of multiple matings and the strong effect of multiple mating on intracolonial genetic diversity may be essential to the negative association between polygyny and polyandry and that any attempt to empirically test this hypothesis should place emphasis upon these two key underlying aspects.

A role for parasites in stabilising the fig-pollinator mutualism.

Mutualisms are interspecific interactions in which both players benefit. Explaining their maintenance is problematic, because cheaters should outcompete cooperative conspecifics, leading to mutualism instability. Monoecious figs (Ficus) are pollinated by host-specific wasps (Agaonidae), whose larvae gall ovules in their "fruits" (syconia). Female pollinating wasps oviposit directly into Ficus ovules from inside the receptive syconium. Across Ficus species, there is a widely documented segregation of pollinator galls in inner ovules and seeds in outer ovules. This pattern suggests that wasps avoid, or are prevented from ovipositing into, outer ovules, and this results in mutualism stability. However, the mechanisms preventing wasps from exploiting outer ovules remain unknown. We report that in Ficus rubiginosa, offspring in outer ovules are vulnerable to attack by parasitic wasps that oviposit from outside the syconium. Parasitism risk decreases towards the centre of the syconium, where inner ovules provide enemy-free space for pollinator offspring. We suggest that the resulting gradient in offspring viability is likely to contribute to selection on pollinators to avoid outer ovules, and by forcing wasps to focus on a subset of ovules, reduces their galling rates. This previously unidentified mechanism may therefore contribute to mutualism persistence independent of additional factors that invoke plant defences against pollinator oviposition, or physiological constraints on pollinators that prevent oviposition in all available ovules.

Specificity and transmission mosaic of ant nest-wall fungi.

Mutualism, whereby species interact to their mutual benefit, is extraordinary in a competitive world. To recognize general patterns of origin and maintenance from the plethora of mutualistic associations proves a persisting challenge. The simplest situation is believed to be that of a single mutualist specific to a single host, vertically transmitted from one host generation to the next. We characterized ascomycete fungal associates cultured for nest architecture by the ant subgenera Dendrolasius and Chthonolasius. The ants probably manage their fungal mutualists by protecting them against fungal competitors. The ant subgenera display different ant-to-fungus specificity patterns, one-to-two and many-to-one, and we infer vertical transmission, in the latter case overlaid by horizontal transmission. Possible evolutionary trajectories include a reversal from fungiculture by other Lasius subgenera and inheritance of fungi through life cycle interactions of the ant subgenera. The mosaic indicates how specificity patterns can be shaped by an interplay between host life-cycles and transmission adaptations.

Integrative taxonomy: a multisource approach to exploring biodiversity.

Good alpha taxonomy is central to biology. On the basis of a survey of arthropod studies that used multiple disciplines for species delimitation, we evaluated the performance of single disciplines. All included disciplines had a considerable failure rate. Rigor in species delimitation can thus be increased when several disciplines chosen for complementarity are used. We present a flexible procedure and stopping rule for integrative taxonomy that uses the information from different disciplines separately. Disagreement among disciplines over the number and demarcation of species is resolved by elucidating and invoking evolutionary explanations for disagreement. With the identification of further promising study organisms and of new questions for in-depth analysis, evolutionary biology should profit from integrative taxonomy. An important rationale is clarity in researcher bias in the decision-making process. The success of integrative taxonomy will further increase through methodological progress, taxonomic training of evolutionary biologists, and balanced resource allocation.

Abandoning aggression but maintaining self-nonself discrimination as a first stage in ant supercolony formation.

An ant supercolony is a very large entity with very many queens. Although normal colonies of small extent and few queens remain distinct, a supercolony is integrated harmoniously over a large area [1, 2]. The lack of aggression is advantageous: Aggression is costly, involving direct and indirect losses and recognition errors [3, 4]. Indeed, supercolonial ants are among the ecologically most successful organisms [5-7]. But how supercolonies arise remains mysterious [1, 2, 8]. Suggestions include that reduced within-colony relatedness or reduced self-nonself discrimination would foster supercolony formation [1, 2, 5, 7, 9-12]. However, one risks confusing correlation and causality in deducing the evolution from distinct colonies to supercolonies when observing established supercolonies. It might help to follow up observations of another lack of aggression, that between single-queened colonies in some ant species. We show that the single-queened Lasius austriacus lacks aggression between colonies and occasionally integrates workers across colonies but maintains high within-colony relatedness and self-nonself discrimination. Provided that the ecological framework permits, reduced aggression might prove adaptive for any ant colony irrespective of within-colony relatedness. Abandoning aggression while maintaining discrimination might be a first stage in supercolony formation. This adds to the emphasis of ecology as central to the evolution of cooperation in general [13].

Weaver ants Oecophylla smaragdina encounter nasty neighbors rather than dear enemies.

The evolution of territorial behavior requires that the benefits of territoriality outweigh the costs. The costs are primarily those of territorial defense against encroaching neighbors or against floaters seeking to establish their own territory. One way to reduce the cost of defense might be to restrict serious conflict to encounters with those posing the greatest threat. Studies of many animals have found that less aggression is shown toward neighbors than toward, strangers, a phenomenon known as the "dear enemy" effect. However, the opposite can also be true, namely, that more aggression is shown toward neighbors than strangers: the "nasty neighbor" effect. This may be particularly true of group-living species that defend a resource-based territory. Here we show that (1) colonies of the weaver ant Oecophylla smaragdina were able to recognize a greater proportion of workers from neighboring colonies as non-colony members; and (2) when recognized as non-colony members, more aggression was exhibited toward neighbors than non-neighbors. We present for the first time evidence that differential levels of aggression involve both a perceptual and behavioral component. On the other hand, we found no evidence that weaver ant workers were better able to recognize workers from previously unknown colonies or responded more aggressively to them, even after a 10-day period of contact. This contrasts with other species in which rapid learning of the identity of new potential enemies has been demonstrated. We suggest that such a response is unnecessary for weaver ants, as encounters with intruders from non-neighboring colonies are probably rare and of little consequence. This study adds to the growing body of evidence that the nasty neighbor effect may be much more common than the dear enemy effect among group-living species.

Genetic caste determination in termites: out of the shade but not from Mars.

Several ant species are known with genetic effects on caste determination but, for termites, the role of environment has been assumed to be omnipotent. Now Hayashi et al. report that commitment to the nymph and worker pathways in Reticulitermes speratus follows a simple model involving two alleles at a sex-linked locus. The spread of this system of genetic caste determination seems best explained by selection at the colony level. This remarkable system may be widely applicable throughout termites, although it cannot be universal, and may provide a window into causal aspects of the molecular biology of caste determination.

Ground dwelling ants as surrogates for establishing conservation priorities in the Australian wet tropics.

This study aims to identify a set of areas with high biodiversity value over a small spatial scale within the Australian Wet Tropics. We identified sites of high biodiversity value across an altitudinal gradient of ground dwelling ant communities using three measures of biodiversity. The three measures considered were estimated species richness, complementarity between sites and evolutionary history. The latter measure was derived using the systematic nomenclature of the ants to infer a surrogate phylogeny. The goal of conservation assessments could then be achieved by choosing the most diverse site combinations. This approach was found to be valuable for identifying the most diverse site combinations across an altitudinal gradient that could ensure the preservation of terrestrial ground dwelling invertebrates in the Australian Wet Tropics.

Meta-population structure in a coral reef fish demonstrated by genetic data on patterns of migration, extinction and re-colonisation.

BACKGROUND: Management strategies for coral reefs are dependant on information about the spatial population structure and connectivity of reef organisms. Genetic tools can reveal important information about population structure, however, this information is lacking for many reef species. We used a mitochondrial molecular marker to examine the population genetic structure and the potential for meta-population dynamics in a direct developing coral reef fish using 283 individuals from 15 reefs on the Great Barrier Reef, Australia. We employed a hierarchical sampling design to test genetic models of population structure at multiple geographical scales including among regions, among shelf position and reefs within regions. Predictions from island, isolation-by-distance and meta-population models, including the potential for asymmetric migration, local extinction and patterns of re-colonisation were examined. RESULTS: Acanthochromis polyacanthus displayed strong genetic structure among regions (PhiST = 0.81, P < 0.0001) that supported an equilibrium isolation-by-distance model (r = 0.77, P = 0.001). Significant structuring across the continental shelf was only evident in the northern region (PhiST = 0.31, P < 0.001) and no evidence of isolation-by-distance was found within any region. Pairwise PhiST values indicated overall strong but variable genetic structure (mean PhiST among reefs within regions = 0.28, 0.38, 0.41), and asymmetric migration rates among reefs with low genetic structure. Genetic differentiation among younger reefs was greater than among older reefs supporting a meta-population propagule-pool colonisation model. Variation in genetic diversities, demographic expansion and population growth estimates indicated more frequent genetic bottlenecks/founder effects and subsequent population expansion in the central and southern regions compared to the northern one. CONCLUSION: Our findings provide genetic evidence for meta-population dynamics in a direct developing coral reef fish and we reject the equilibrium island and isolation-by distance models at local spatial scales. Instead, strong non-equilibrium genetic structure appears to be generated by genetic bottlenecks/founder effects associated with population reductions/extinctions and asymmetric migration/(re)-colonisation of such populations. These meta-population dynamics varied across the geographical range examined with edge populations exhibiting lower genetic diversities and higher rates of population expansion than more central populations. Therefore, coral reef species may experience local population reductions/extinctions that promote overall meta-population genetic differentiation.

A DNA and morphology based phylogenetic framework of the ant genus Lasius with hypotheses for the evolution of social parasitism and fungiculture.

BACKGROUND: Ants of the genus Lasius are ecologically important and an important system for evolutionary research. Progress in evolutionary research has been hindered by the lack of a well-founded phylogeny of the subgenera, with three previous attempts disagreeing. Here we employed two mitochondrial genes (cytochrome c oxidase subunit I, 16S ribosomal RNA), comprising 1,265 bp, together with 64 morphological characters, to recover the phylogeny of Lasius by Bayesian and Maximum Parsimony inference after exploration of potential causes of phylogenetic distortion. We use the resulting framework to infer evolutionary pathways for social parasitism and fungiculture. RESULTS: We recovered two well supported major lineages. One includes Acanthomyops, Austrolasius, Chthonolasius, and Lasius pallitarsis, which we confirm to represent a seventh subgenus, the other clade contains Dendrolasius, and Lasius sensu stricto. The subgenus Cautolasius, displaying neither social parasitism nor fungiculture, probably belongs to the second clade, but its phylogenetic position is not resolved at the cutoff values of node support we apply. Possible causes for previous problems with reconstructing the Lasius phylogeny include use of other reconstruction techniques, possibly more prone to instabilities in some instances, and the inclusion of phylogenetically distorting characters. CONCLUSION: By establishing an updated phylogenetic framework, our study provides the basis for a later formal taxonomic revision of subgenera and for studying the evolution of various ecologically and sociobiologically relevant traits of Lasius, although there is need for future studies to include nuclear genes and additional samples from the Nearctic. Both social parasitism and fungiculture evolved twice in Lasius, once in each major lineage, which opens up new opportunities for comparative analyses. The repeated evolution of social parasitism has been established for other groups of ants, though not for temporary social parasitism as found in Lasius. For fungiculture, the independent emergence twice in a monophyletic group marks a novel scenario in ants. We present alternative hypotheses for the evolution of both traits, with one of each involving loss of the trait. Though less likely for both traits than later evolution without reversal, we consider reversal as sufficiently plausible to merit independent testing.

From transcriptome to biological function: environmental stress in an ectothermic vertebrate, the coral reef fish Pomacentrus moluccensis.

BACKGROUND: Our understanding of the importance of transcriptional regulation for biological function is continuously improving. We still know, however, comparatively little about how environmentally induced stress affects gene expression in vertebrates, and the consistency of transcriptional stress responses to different types of environmental stress. In this study, we used a multi-stressor approach to identify components of a common stress response as well as components unique to different types of environmental stress. We exposed individuals of the coral reef fish Pomacentrus moluccensis to hypoxic, hyposmotic, cold and heat shock and measured the responses of approximately 16,000 genes in liver. We also compared winter and summer responses to heat shock to examine the capacity for such responses to vary with acclimation to different ambient temperatures. RESULTS: We identified a series of gene functions that were involved in all stress responses examined here, suggesting some common effects of stress on biological function. These common responses were achieved by the regulation of largely independent sets of genes; the responses of individual genes varied greatly across different stress types. In response to heat exposure over five days, a total of 324 gene loci were differentially expressed. Many heat-responsive genes had functions associated with protein turnover, metabolism, and the response to oxidative stress. We were also able to identify groups of co-regulated genes, the genes within which shared similar functions. CONCLUSION: This is the first environmental genomic study to measure gene regulation in response to different environmental stressors in a natural population of a warm-adapted ectothermic vertebrate. We have shown that different types of environmental stress induce expression changes in genes with similar gene functions, but that the responses of individual genes vary between stress types. The functions of heat-responsive genes suggest that prolonged heat exposure leads to oxidative stress and protein damage, a challenge of the immune system, and the re-allocation of energy sources. This study hence offers insight into the effects of environmental stress on biological function and sheds light on the expected sensitivity of coral reef fishes to elevated temperatures in the future.

The evolution of worker caste diversity in social insects.

Morphological diversification of workers is predicted to improve the division of labor within social insect colonies, yet many species have monomorphic workers. Individual-level selection on the reproductive capacities of workers may counter colony-level selection for diversification, and life-history differences between species (timing of caste determination, colony size, genetic variation available) may mediate the strength of this selection. We tested this through phylogenetically independent contrast analyses on a new data set for 35 ant species. Evidence was found that early divergence of queen-worker developmental pathways may facilitate the evolution of worker diversity because queen-worker dimorphism was strongly positively associated with diversity. By contrast, risks for colonies that invest in specialized workers and colony size effects on costs of worker reproduction seem unlikely to strongly affect the evolution of worker diversity because there was no significant association between colony size and diversity when controlling statistically for queen-worker dimorphism. Finally, worker diversity was greater in species with multiple lineages per colony, and it was negatively associated with relatedness in monogynous species. This could be due to high intracolonial genetic variance favoring the expression and evolution of great worker diversity or to diversity evolving more easily when there is selection for repression of worker reproduction (worker policing).

Population and colony genetic structure of the primitive termite Mastotermes darwiniensis.

The termite Mastotermes darwiniensis is the sole extant member of its family and occupies the basal position in the phylogeny of the eusocial order Isoptera. In this study, we investigated the micro- and macrogeographic genetic structure of M. darwiniensis in its native range in Australia. A total of 1591 workers were sampled from 136 infested trees in 24 locales. Each locale was separated by 2-350 km, and these locales were found within two broader geographic regions approximately 1500 km apart. The multilocus genotypes of all termites were assayed at six polymorphic microsatellite loci. The genetic data indicated that colonies typically fed on multiple trees within locales and extended over linear distances of up to 320 m. Single colonies were frequently headed by multiple reproductives. Workers were highly related (r = 0.40) and substantially inbred (f = 0.10). Thus, M. darwiniensis colonies are characterized by the input of alleles from multiple reproductives, which sometimes engage in consanguineous matings. Our analyses of population genetic structure above the level of the colony indicated that locales and regions were significantly differentiated (theta(locale) = 0.50, theta(region) = 0.37). Moreover, locales showed a pattern of genetic isolation by distance within regions. Thus, M. darwiniensis populations display restricted gene flow over moderate geographic distances. We suggest that the genetic patterns displayed by M. darwiniensis result primarily from selective pressures acting to maintain high relatedness among colonymates while allowing colonies to grow rapidly and dominate local habitats.

Quantitative Analysis of Directionality in Mammalian Karyotype Evolution.

The ancestral mammalian karyotype had been hypothesized to have had 2n ≈ 80 (the "fusion hypothesis"), 2n = 6-14 (the "fission hypothesis") or a diploid number close to the present mode (the "modal hypothesis"). The fusion hypothesis has long been the dominant paradigm in the study of karyotype evolution, but recent evidence favors the fission hypothesis, and our analysis also strongly supports fission as the predominant rearrangement compared to fusion. To formalize our analysis, we first define $$\overline A$$ chromosomes as a group containing both acrocentrics and telocentrics, and $$\overline M$$ chromosomes as all the rest. Given this dichotomy, we then divide pericentric inversions into three types according to whether they convert $$\overline A$$ chromosomes to $$\overline M$$ , $$\overline M$$ to $$\overline A$$ or make no interchange. Only the first two types, denoted p.i. $$(\overline {AM})$$ and p.i. $$(\overline {MA})$$ , are important for the analysis. If fusion predominates, the direction of karyotype evolution is determined by the overall joint action of centric fusions and p.i. $$(\overline {MA})$$ (the "fusion cycle"), whereas, if fission predominates, the chief rearrangements are fission and p.i. $$(\overline {AM})$$ (the "fission cycle"). The necessary predominance of p.i. $$(\overline {MA})$$ over p.i. $$(\overline {AM})$$ in the fusion cycle makes it extremely unlikely on a priori grounds that this cycle has been very important in mammalian karyotype evolution, because a probabilistic analysis shows that p.i. $$(\overline {AM})$$ should occur many times more often than the reverse rearrangement, especially when the chromosome number is low. Chromosomes will therefore seldom be available for fusion. On the other hand, given duplication of both heterochromatic and centromeric material, there are no obstacles to the operation of the fission cycle. The modal hypothesis is also implausible on a priori grounds because of the many fusions required for most groups. We next define the karyograph as a graph on which karyotypes are plotted in terms of chromosome number (2n) versus arm number (2AN). We determine 2AN by counting one for each $$\overline A$$ and two for each $$\overline M$$ chromosome. Robertsonian changes each alter 2n by one while leaving 2AN constant while p.i. $$(\overline AM)$$ and p.i. $$(\overline MA)$$ do the reverse. An extensive karyograph analysis of the known mammalian karyotypes shows that there is little correlation between karyotypic and morphological level under any of the three hypotheses, and that there is a strong tendency for linear patterns to emerge when families are plotted separately. This linearity consists of either vertical or horizontal lines on the karyograph, or some combination of the two. Such linearity would be unexpected if the fusion cycle dominated mammalian karyotype evolution, but is readily understandable under the fission cycle as resulting from the development of synchrony between large sections of the genome. This synchrony can develop readily under the fission cycle in that fission produces two $$\overline A$$ chromosomes constrained to evolve by inversion for some time. The resulting $$\overline M$$ chromosomes would later become available for fission following duplication of centromeric material and give rise to four $$\overline A$$ chromosomes, again constrained to evolve by inversion for a while also. The frequently observed linearity of family karyograph distributions, and the above argument concerning the development of synchrony, suggests that mammalian karyotypes tend to follow an upwardly zig-zag course (with occasional "back eddies" by centric fusion) during evolution when plotted on the karyograph.

The evolution of parasites from their hosts: intra- and interspecific parasitism and Emery's rule.

In some taxa of Hymenoptera, fungi, red algae and mistletoe, parasites and their hosts are either sibling species or at least closely related (Emery's rule). Three evolutionary mechanisms have been proposed for this phenomenon: (i) intraspecific parasitism is followed by sympatric speciation; (ii) allopatric speciation is followed by secondary sympatry and the subsequent parasitism of one sibling species by the other; and (iii) allopatric speciation of a species with intraspecific parasitism is followed by secondary sympatry, in which one species becomes an obligate parasite of the other. Mechanisms (i) and (ii) are problematic, while mechanism (iii) has not, to our knowledge, been analysed quantitatively. In this paper, we develop a model for single- and two-species evolutionary stable strategies (ESSs) to examine the basis for Emery's rule and to determine whether mechanism (iii) is consistent with ESS reasoning. In secondary sympatry after allopatric speciation, the system's evolution depends on the relative abundances of the two sibling species and on the proportional damage wrought by parasites of each species on non-parasitic members of the other. Depending on these interspecific effects, either the rarer or the commoner species may become the parasite and the levels of within-species parasitism need not determine which evolves to obligate parasitism.

Infection with the trypanosome Crithidia bombi and expression of immune-related genes in the bumblebee Bombus terrestris.

Social bees and other insects are frequently parasitized by a large range of different microorganisms. Among these is Crithidia bombi (Kinetoplastida: Trypanosomatidae), a common gut parasite of bumblebees, Bombus spp. (Insecta: Apidae). Bumblebees are important pollinators in commercial and natural environments. There are clear detrimental effects of C. bombi infections on the fitness of bumblebees. However, little has been known about how the bee's immune system responds to infections with trypanosome parasites. Here, we study the immune response of Bombus terrestris on infection by C. bombi. We measured the expression of four immune-related genes (Hemomucin, MyD88, Relish, and TEP7) using RT-qPCR in adult B. terrestris workers that were either healthy or infected with the trypanosome parasite C. bombi. The potential recognition gene Hemomucin was significantly upregulated in the infected bees. Further, there was substantial and significant variation in all four genes among different bumblebee colonies irrespective of infection status.

Animal performance and stress: responses and tolerance limits at different levels of biological organisation.

Recent advances in molecular biology and the use of DNA microarrays for gene expression profiling are providing new insights into the animal stress response, particularly the effects of stress on gene regulation. However, interpretation of the complex transcriptional changes that occur during stress still poses many challenges because the relationship between changes at the transcriptional level and other levels of biological organisation is not well understood. To confront these challenges, a conceptual model linking physiological and transcriptional responses to stress would be helpful. Here, we provide the basis for one such model by synthesising data from organismal, endocrine, cellular, molecular, and genomic studies. We show using available examples from ectothermic vertebrates that reduced oxygen levels and oxidative stress are common to many stress conditions and that the responses to different types of stress, such as environmental, handling and confinement stress, often converge at the challenge of dealing with oxygen imbalance and oxidative stress. As a result, a common set of stress responses exists that is largely independent of the type of stressor applied. These common responses include the repair of DNA and protein damage, cell cycle arrest or apoptosis, changes in cellular metabolism that reflect the transition from a state of cellular growth to one of cellular repair, the release of stress hormones, changes in mitochondrial densities and properties, changes in oxygen transport capacities and changes in cardio-respiratory function. Changes at the transcriptional level recapitulate these common responses, with many stress-responsive genes functioning in cell cycle control, regulation of transcription, protein turnover, metabolism, and cellular repair. These common transcriptional responses to stress appear coordinated by only a limited number of stress-inducible and redox-sensitive transcription factors and signal transduction pathways, such as the immediate early genes c-fos and c-jun, the transcription factors NFkappaB and HIF-1alpha, and the JNK and p38 kinase signalling pathways. As an example of environmental stress responses, we present temperature response curves at organismal, cellular and molecular levels. Acclimation and physiological adjustments that can shift the threshold temperatures for the onset of these responses are discussed and include, for example, adjustments of the oxygen delivery system, the heat shock response, cellular repair system, and transcriptome. Ultimately, however, an organism's ability to cope with environmental change is largely determined by its ability to maintain aerobic scope and to prevent loss in performance. These systemic constraints can determine an organism's long-term survival well before cellular and molecular functions are disturbed. The conceptual model we propose here discusses some of the crosslinks between responses at different levels of biological organisation and the central role of oxygen balance and oxidative stress in eliciting these responses with the aim to help the interpretation of environmental genomic data in the context of organismal function and performance.

Phylogenetic analysis of honey bee behavioral evolution.

DNA sequences from three mitochondrial (rrnL, cox2, nad2) and one nuclear gene (itpr) from all 9 known honey bee species (Apis), a 10th possible species, Apis dorsata binghami, and three outgroup species (Bombus terrestris, Melipona bicolor and Trigona fimbriata) were used to infer Apis phylogenetic relationships using Bayesian analysis. The dwarf honey bees were confirmed as basal, and the giant and cavity-nesting species to be monophyletic. All nodes were strongly supported except that grouping Apis cerana with A. nigrocincta. Two thousand post-burnin trees from the phylogenetic analysis were used in a Bayesian comparative analysis to explore the evolution of dance type, nest structure, comb structure and dance sound within Apis. The ancestral honey bee species was inferred with high support to have nested in the open, and to have more likely than not had a silent vertical waggle dance and a single comb. The common ancestor of the giant and cavity-dwelling bees is strongly inferred to have had a buzzing vertical directional dance. All pairwise combinations of characters showed strong association, but the multiple comparisons problem reduces the ability to infer associations between states between characters. Nevertheless, a buzzing dance is significantly associated with cavity-nesting, several vertical combs, and dancing vertically, a horizontal dance is significantly associated with a nest with a single comb wrapped around the support, and open nesting with a single pendant comb and a silent waggle dance.