ALBERT

Description: The genomes of related species contain valuable information on the history of the considered taxa. Great apes in particular exhibit variation of evolutionary patterns along their genomes. However, the great ape data also bring new challenges, such as the presence of incomplete lineage sorting and ancestral shared polymorphisms. Previous methods for genome-scale analysis are restricted to very few individuals or cannot disentangle the contribution of mutation rates and fixation biases. This represents a limitation both for the understanding of these forces as well as for the detection of regions affected by selection. Here, we present a new model designed to estimate mutation rates and fixation biases from genetic variation within and between species. We relax the assumption of instantaneous substitutions, modeling substitutions as mutational events followed by a gradual fixation. Hence, we straightforwardly account for shared ancestral polymorphisms and incomplete lineage sorting. We analyze genome-wide synonymous site alignments of human, chimpanzee, and two orangutan species. From each taxon, we include data from several individuals. We estimate mutation rates and GC-biased gene conversion intensity. We find that both mutation rates and biased gene conversion vary with GC content. We also find lineage-specific differences, with weaker fixation biases in orangutan species, suggesting a reduced historical effective population size. Finally, our results are consistent with directional selection acting on coding sequences in relation to exonic splicing enhancers.

Description: Genetic control of male or female gonad development displays between different groups of organisms a remarkable diversity of "master sex-determining genes" at the top of the genetic hierarchies, whereas downstream components surprisingly appear to be evolutionarily more conserved. Without much further studies, conservation of sequence has been equalized to conservation of function. We have used the medaka fish to investigate the generality of this paradigm. In medaka, the master male sex-determining gene is dmrt1bY , a highly conserved downstream regulator of sex determination in vertebrates. To understand its function in orchestrating the complex gene regulatory network, we have identified targets genes and regulated pathways of Dmrt1bY. Monitoring gene expression and interactions by transgenic fluorescent reporter fish lines, in vivo tissue-chromatin immunoprecipitation and in vitro gene regulation assays revealed concordance but also major discrepancies between mammals and medaka, notably amongst spatial, temporal expression patterns and regulations of the canonical Hedgehog and R-spondin/Wnt/Follistatin signaling pathways. Examination of Foxl2 protein distribution in the medaka ovary defined a new subpopulation of theca cells, where ovarian-type aromatase transcriptional regulation appears to be independent of Foxl2. In summary, these data show that the regulation of the downstream regulatory network of sex determination is less conserved than previously thought.

Description: Synonymous codon usage patterns are shaped by a balance between mutation, drift, and natural selection. To date, detection of translational selection in vertebrates has proven to be a challenging task, obscured by small long-term effective population sizes in larger animals and the existence of isochores in some species. The consensus is that, in such species, natural selection is either completely ineffective at overcoming mutational pressures and genetic drift or perhaps is effective but so weak that it is not detectable. The aim of this research is to understand the interplay between mutation, selection, and genetic drift in vertebrates. We observe that although variation in mutational bias is undoubtedly the dominant force influencing codon usage, translational selection acts as a weak additional factor influencing synonymous codon usage. These observations indicate that translational selection is a widespread phenomenon in vertebrates and is not limited to a few species.

Description: Class IV homeodomain leucine zipper (C4HDZ) genes are plant-specific transcription factors that, based on phenotypes in Arabidopsis thaliana , play an important role in epidermal development. In this study, we sampled all major extant lineages and their closest algal relatives for C4HDZ homologs and phylogenetic analyses result in a gene tree that mirrors land plant evolution with evidence for gene duplications in many lineages, but minimal evidence for gene losses. Our analysis suggests an ancestral C4HDZ gene originated in an algal ancestor of land plants and a single ancestral gene was present in the last common ancestor of land plants. Independent gene duplications are evident within several lineages including mosses, lycophytes, euphyllophytes, seed plants, and, most notably, angiosperms. In recently evolved angiosperm paralogs, we find evidence of pseudogenization via mutations in both coding and regulatory sequences. The increasing complexity of the C4HDZ gene family through the diversification of land plants correlates to increasing complexity in epidermal characters.

Description: Resolving difficult nodes for any part of the vertebrate tree of life often requires analyzing a large number of loci. Developing molecular markers that are workable for the groups of interest is often a bottleneck in phylogenetic research. Here, on the basis of a nested polymerase chain reaction (PCR) strategy, we present a universal toolkit including 102 nuclear protein-coding locus (NPCL) markers for vertebrate phylogenomics. The 102 NPCL markers have a broad range of evolutionary rates, which makes them useful for a wide range of time depths. The new NPCL toolkit has three important advantages compared with all previously developed NPCL sets: 1) the kit is universally applicable across vertebrates, with a PCR success rate of 94.6% in 16 widely divergent tested vertebrate species; 2) more than 90% of PCR reactions produce strong and single bands of the expected sizes that can be directly sequenced; and 3) all cleanup PCR reactions can be sequenced with only two specific universal primers. To test its actual phylogenetic utility, 30 NPCLs from this toolkit were used to address the higher level relationships of living salamanders. Of the 639 target PCR reactions performed on 19 salamanders and several outgroup species, 632 (98.9%) were successful, and 602 (94.1%) were directly sequenced. Concatenation and species-tree analyses on this 30-locus data set produced a fully resolved phylogeny and showed that Cryptobranchoidea (Cryptobranchidae + Hynobiidae) branches first within the salamander tree, followed by Sirenidae. Our experimental tests and our demonstration for a particular case show that our NPCL toolkit is a highly reliable, fast, and cost-effective approach for vertebrate phylogenomic studies and thus has the potential to accelerate the completion of many parts of the vertebrate tree of life.

Description: Genetic incompatibilities are commonly observed between hybridizing species. Although this type of isolating mechanism has received considerable attention, we have few examples describing how genetic incompatibilities evolve. We investigated the evolution of two loci involved in a classic example of a Bateson–Dobzhansky–Muller (BDM) incompatibility in Xiphophorus , a genus of freshwater fishes from northern Central America. Hybrids develop a lethal melanoma due to the interaction of two loci, an oncogene and its repressor. We cloned and sequenced the putative repressor locus in 25 Xiphophorus species and an outgroup species, and determined the status of the oncogene in those species from the literature. Using phylogenetic analyses, we find evidence that a repeat region in the proximal promoter of the repressor is coevolving with the oncogene. The data support a hypothesis that departs from the standard BDM model: it appears the alleles that cause the incompatibilities have coevolved simultaneously within lineages, rather than in allopatric or temporal isolation.

Description: Incompatibility systems in which individuals bearing identical alleles reject each other favor the maintenance of a diversity of alleles. Mushroom mating type loci ( MAT ) encode for dozens or hundreds of incompatibility alleles whose loss from the population is greatly restricted through negative frequency selection, leading to a system of alleles with highly divergent sequences. Here, we use DNA sequences of homeodomain (HD) encoding genes at the MAT locus of five closely related species of the root rot basidiomycete Heterobasidion annosum sensu lato to show that the extended coalescence time of MAT alleles greatly predates speciation in the group, contrasting loci outside of MAT that show allele divergences largely consistent with the species phylogeny with those of MAT , which show rampant trans-species polymorphism. We observe a roughly 6-fold greater genealogical depth and polymorphism of MAT compared with non- MAT that argues for the maintenance of balanced polymorphism for a minimum duration of 24 My based on a molecular-clock calibrated species phylogeny. As with other basidiomycete HD genes, balancing selection appears to be concentrated at the specificity-determining region in the N-terminus of the protein based on identification of codons under selection and the absence of recombination within the region. However, the elevated polymorphism extends into the nonspecificity determining regions as well as a neighboring non- MAT gene, the mitochondrial intermediate peptidase ( MIP ). In doing so, increased divergence should decrease recombination among alleles and as a by-product create incompatibilities in the functional domains not involved in allele recognition but in regulating sexual development.

Description: Here we present computational machinery to efficiently and accurately identify transposable element (TE) insertions in 146 next-generation sequenced inbred strains of Drosophila melanogaster . The panel of lines we use in our study is composed of strains from a pair of genetic mapping resources: the Drosophila Genetic Reference Panel (DGRP) and the Drosophila Synthetic Population Resource (DSPR). We identified 23,087 TE insertions in these lines, of which 83.3% are found in only one line. There are marked differences in the distribution of elements over the genome, with TEs found at higher densities on the X chromosome, and in regions of low recombination. We also identified many more TEs per base pair of intronic sequence and fewer TEs per base pair of exonic sequence than expected if TEs are located at random locations in the euchromatic genome. There was substantial variation in TE load across genes. For example, the paralogs derailed and derailed-2 show a significant difference in the number of TE insertions, potentially reflecting differences in the selection acting on these loci. When considering TE families, we find a very weak effect of gene family size on TE insertions per gene, indicating that as gene family size increases the number of TE insertions in a given gene within that family also increases. TEs are known to be associated with certain phenotypes, and our data will allow investigators using the DGRP and DSPR to assess the functional role of TE insertions in complex trait variation more generally. Notably, because most TEs are very rare and often private to a single line, causative TEs resulting in phenotypic differences among individuals may typically fail to replicate across mapping panels since individual elements are unlikely to segregate in both panels. Our data suggest that "burden tests" that test for the effect of TEs as a class may be more fruitful.

Description: Sex determination mechanisms are highly variable across teleost fishes and sexual development is often plastic. Nevertheless, downstream factors establishing the two sexes are presumably conserved. Here, we study sequence evolution and gene expression of core genes of sexual development in a prime model system in evolutionary biology, the East African cichlid fishes. Using the available five cichlid genomes, we test for signs of positive selection in 28 genes including duplicates from the teleost whole-genome duplication, and examine the expression of these candidate genes in three cichlid species. We then focus on a particularly striking case, the A- and B-copies of the aromatase cyp19a1 , and detect different evolutionary trajectories: cyp19a1A evolved under strong positive selection, whereas cyp19a1B remained conserved at the protein level, yet is subject to regulatory changes at its transcription start sites. Importantly, we find shifts in gene expression in both copies. Cyp19a1 is considered the most conserved ovary-factor in vertebrates, and in all teleosts investigated so far, cyp19a1A and cyp19a1B are expressed in ovaries and the brain, respectively. This is not the case in cichlids, where we find new expression patterns in two derived lineages: the A-copy gained a novel testis-function in the Ectodine lineage, whereas the B-copy is overexpressed in the testis of the speciest-richest cichlid group, the Haplochromini. This suggests that even key factors of sexual development, including the sex steroid pathway, are not conserved in fish, supporting the idea that flexibility in sexual determination and differentiation may be a driving force of speciation.

Description: Greenstreet, S. P. R., Fraser, H. M., Rogers, S. I., Trenkel, V. M., Simpson, S. D., and Pinnegar, J. K. 2012. Redundancy in metrics describing the composition, structure, and functioning of the North Sea demersal fish community. – ICES Journal of Marine Science, 69: 8–22. Broader ecosystem management objectives for North Sea demersal fish currently focus on restoring community size structure. However, most policy drivers explicitly concentrate on restoring and conserving biodiversity, and it has not yet been established that simply restoring demersal fish size composition will be sufficient to reverse declines in biodiversity and ensure a generally healthy community. If different aspects of community composition, structure, and function vary independently, then to monitor all aspects of community general health will require application of a suite of metrics. This assumes low redundancy among the metrics used in any such suite and implies that addressing biodiversity issues specifically will require explicit management objectives for particular biodiversity metrics. This issue of metric redundancy is addressed, and 15 metrics covering five main attributes of community composition, structure, and function are applied to groundfish survey data. Factor analysis suggested a new interpretation of the metric information and indicated that a minimum suite of seven metrics was necessary to ensure that all changes in the general health of the North Sea demersal fish community were monitored properly. Covariance among size-based and species-diversity metrics was low, implying that restoration of community size structure would not necessarily reverse declines in species diversity.

Description: Dankel, D. J., Aps, R., Padda, G., Röckmann, C., van der Sluijs, J. P., Wilson, D. C., and Degnbol, P. 2012. Advice under uncertainty in the marine system. – ICES Journal of Marine Science, 69: 3–7. There is some uncertainty in the fisheries science–policy interface. Although progress has been made towards more transparency and participation in fisheries science in ICES Areas, routine use of state-of-the-art quantitative and qualitative tools to address uncertainty systematically is still lacking. Fisheries science that gives advice to policy-making is plagued by uncertainties; the stakes of the policies are high and value-laden and need therefore to be treated as an example of "post-normal science" (PNS). To achieve robust governance, understanding of the characteristics and implications of the scientific uncertainties for management strategies need to come to the centre of the table. This can be achieved using state-of-the-art tools such as pedigree matrices and uncertainty matrices, as developed by PNS scholars and used in similar science–policy arenas on other complex issues. An explicit extension of the peer community within maritime systems will be required to put these new tools in place. These new competences become even more important as many countries within the ICES Area are now embarking on new policies.

Description: Browman, H.I. 2012. Quo Vadimus. – ICES Journal of Marine Science, 69: 1–2.

Description: Hüssy, K., Coad, J. O., Farrell, E. D., Clausen, L. A. W., and Clarke, M. W. 2012. Age verification of boarfish ( Capros aper ) in the Northeast Atlantic. – ICES Journal of Marine Science, 69: 34–40. The boarfish ( Capros aper ) is a pelagic species of recent interest to the fishing industry, with landings increasing by 〉500% over the past 3 years. The objective of the study was to provide a method for age determination based on whole sagittal otoliths, with the results to be used in stock assessment. Translucent zones laid down at regular intervals are identified by marginal increment analysis as seasonally recurring. Translucent zones are formed between September/October and March/April, regardless of fish age. The occurrence of the first annulus is validated by analysis of presumed daily growth increments. Subsequent annulus deposition is homogenous between individuals and allows general guidelines to be derived for interpreting the age of boarfish using their otoliths.

Description: Kim, D., Yang, E. J., Kim, K. H., Shin, C-W., Park, J., Yoo, S., and Hyun, J-H. 2012. Impact of an anticyclonic eddy on the summer nutrient and chlorophyll a distributions in the Ulleung Basin, East Sea (Japan Sea). – ICES Journal of Marine Science, 69: 23–29. The impact of the anticyclonic Ulleung Warm Eddy (UWE) on the vertical distributions of nutrient and chlorophyll a (Chl a ) concentrations in the Ulleung Basin (UB) was investigated during the contrasting summers of 2005 and 2007. The physical structure of the water column was characterized by an intrathermocline eddy (ITE) in 2005, whereas the UWE remained distant from the sampling transect in 2007. Water column structures appeared to be highly stratified, and nutrients in the surface waters were totally depleted at all stations. In 2005, an exceptionally high concentration of Chl a (5.5 mg m –3 ) was measured below the surface mixed layer in the eddy core (station D3), and values of ~2.5 mg m –3 were observed at the eddy edge (stations D2 and D4). Formation of an ITE efficiently mixed surface and deep-ocean waters, the latter supplying sufficient nutrients to generate an extremely high concentration of Chl a at the base of the subsurface layer. Overall, the results indicated that the anticyclonic UWE plays a key ecological role in supporting substantial phytoplankton biomass in the nutrient-depleted surface waters in summer and maintaining high benthic mineralization in the deep-sea sediments of the UB.

Description: Franchini, P., Sola, L., Crosetti, D., Milana, V., and Rossi, A. R. 2012. Low levels of population genetic structure in the gilthead sea bream, Sparus aurata , along the coast of Italy. – ICES Journal of Marine Science, 69: 41–50. The gilthead sea bream, Sparus aurata , is a coastal, commercially important fish. Contrasting results concerning the genetic structure of the species at different geographic scales have been reported. Here, an investigation is made into the population genetic structure of S. aurata along the coast of Italy, using samples analysed previously and material from new sampling sites (12) and using different microsatellite loci (10). One sample from the eastern Atlantic and three temporal replicates from one site were also included. The presence of a weak (overall F ST = 0.0072), but significant, genetic population subdivision was detected by F -statistics. Temporal replicates indicate genetic data consistency over time. Isolation by distance between the Atlantic and the coast of Italy is suggested by a Mantel test. The distributional pattern of genetic variance obtained by analysis of molecular variation reflects the geographic sampling areas, but is only partially congruent with the results obtained with fewer sites and loci. The dispersal of passive eggs/larvae by the main currents appears to contribute to shaping the gene flow. Given the intensity of sea bream aquaculture activities in Italy, the possibility that aquaculture may have partially contributed to the population genetic pattern detected cannot be excluded.

Description: Charrier, F., Mazel, V., Caraguel, J-M., Abdallah, Y., Le Gurun, L. L., Legault, A., and Laffaille, P. 2012. Escapement of silver-phase European eels, Anguilla anguilla , determined from fishing activities in a Mediterranean lagoon (Or, France). – ICES Journal of Marine Science, 69: 30–33. Escapement of silver eels from a Mediterranean lagoon was estimated by a capture–tagging–recapture and automated tag-reading study. The population of silver-phase eels in the lagoon was estimated to be 13.2 kg ha –1 , with an escapement rate from the commercial fishery of 76.8%.

Description: Jennings, S., and Lee, J. 2012. Defining fishing grounds with vessel monitoring system data. – ICES Journal of Marine Science, 69: 51–63. Methods for defining fishing grounds to support marine spatial planning and management are developed, applied, and compared. The methods are broadly applicable and repeatable because they use vessel monitoring system (VMS) data that are archived and increasingly accessible. For several fleets at regional and national scales, an attempt is made to assess how the choice of criteria for defining grounds influences (i) size, shape, and location, (ii) overlap among grounds, and (iii) the extent to which annual and multi-annual patterns of fishing activity describe grounds used seasonally or by individual vessels. The results show that grounds defined by excluding infrequently fished margins (areas with 〈10% of total fishing activity) are typically 50% smaller than total fished area. However, landings weight or value (LWV) per unit activity can be higher at the margins, with 10% of activity usually accounting for 10–20% of LWV. The removal of fishing activity in the margins, as a consequence of regulation or fleet behaviour, would lead to disproportionately greater reductions in interactions with other fisheries, sectors, and the environment. Accessible high-resolution information on the "anatomy" of all fishing grounds would better inform debates on the allocation and the use of marine space and the integration of fisheries and environmental management.

Description: Bentley, N., Kendrick, T. H., Starr, P. J., and Breen, P. A. 2012. Influence plots and metrics: tools for better understanding fisheries catch-per-unit-effort standardizations. – ICES Journal of Marine Science, 69: 84–88. Standardization of catch per unit effort using generalized linear models (GLMs) is a common procedure that attempts to remove the confounding effects of variables other than abundance. Simple plots and metrics are described to assist understanding the standardization effects of explanatory variables included in GLMs, illustrated with an example based on New Zealand trevally ( Caranx lutescens ) data.

Description: Aldrin, M., Mortensen, B., Storvik, G., Nedreaas, K., Aglen, A., and Aanes, S. 2012. Improving management decisions by predicting fish bycatch in the Barents Sea shrimp fishery. – ICES Journal of Marine Science, 69: 64–74. When the bycatch of juvenile fish within the Barents Sea shrimp fishery is too large, the area is closed to fishing for a certain period. Bycatch is estimated from sampled trawl hauls, for which the shrimp yield is recorded, along with the total number of various bycatch fish species. At present, bycatch estimation is based on a simple estimator, the sum of the number of fish caught within the area of interest within a small time window, divided by the corresponding shrimp yield (in weight). No historical data are used. A model-based estimation is proposed in which spatio-temporal models are constructed for the variation in both the yield of shrimp and the amount of bycatch in space and time. The main effects are described through generalized additive models, and local dependence structures are specified through correlated random effects. Model estimation includes historical and recent data. Experiments with both simulated and real data show that the model-based estimator outperforms the present simple estimator when a low or moderate number of samples (e.g. 〈20) is available, whereas the two estimators are equally good when the number of samples is high.

Description: Jiao, Y., Smith, E. P., O'Reilly, R., and Orth, D. J. 2012. Modelling non-stationary natural mortality in catch-at-age models. – ICES Journal of Marine Science, 69: 105–118. Non-stationarity in the natural mortality of many species has been discussed among research scientists, but no generally applicable models/methods have been developed where a statistical catch-at-age (SCA) model framework is used. Using the Atlantic weakfish ( Cynoscion regalis ) fishery as an example, several SCA models are developed to assess the population dynamics, then compared. Models used included (i) an SCA with constant natural mortality, (ii) an SCA with unknown natural mortality, but with a hierarchical prior distribution from a mixture of distributions based on life-history information, (iii) an SCA with age-specific unknown natural mortality, (iv) an SCA with time-varying natural mortality, following a random-walk process, and (v) an SCA with age-specific time-varying natural mortality. The last two models imply that the population dynamics are not stationary. A Bayesian approach was used to estimate parameters, and performance of the models was compared through goodness-of-fit and the retrospective patterns of the models. A simulation study was then conducted to test the uncertainty resulting from model selection, the efficiency of using the best model selected based on deviance information criterion, and whether changes in natural mortality could be detected. An SCA with time-varying natural mortality, following a random-walk process, is proposed for the example fishery here. The estimated non-stationary temporal patterns in natural mortality were linked further to climate–ocean oscillation indices, to diagnose possible mechanisms/linkages to these temporal variations in natural mortality.

Description: Juntunen, T., Vanhatalo, J., Peltonen, H., and Mäntyniemi, S. 2012. Bayesian spatial multispecies modelling to assess pelagic fish stocks from acoustic- and trawl-survey data. – ICES Journal of Marine Science, 69: 95–104. A Bayesian spatial model was constructed to estimate the abundance of multiple fish species in a pelagic environment. Acoustic- and trawl-survey data were combined with environmental data to predict the spatial distribution of (i) the acoustic backscattering of fish, (ii) the relative proportion of each species, and (iii) their mean length in the Gulf of Finland in the northeastern Baltic Sea. By combining the three spatial model layers, the spatial distribution of the biomass of each species was estimated. The model consists of a linear predictor on environmental variables and a spatial random effect given by a Gaussian process. A Bayesian approach is a natural choice for the task because it provides a theoretically justified means of summarizing the uncertainties from various model layers. In the study area, three species dominate pelagic waters: sprat ( Sprattus sprattus ), herring ( Clupea harengus ), and three-spined stickleback ( Gasterosteus aculeatus ). Results are presented for each model layer and for estimated total biomass for each species in 2 x 2 km lattices. The posterior mean and central 95% credible intervals of total biomass were sprat 45.7 kt (27.7–71.6), herring 24.6 kt (9.7–41.3), and three-spined stickleback 1.9 kt (0.9–3.2).

Description: Smith, W. K., and Solow, A. R. 2012. Missing and presumed lost: extinction in the ocean and its inference. – ICES Journal of Marine Science, 69: 89–94. The number of modern extinctions in the ocean is unknown. The actual demise of the last individual of a species is essentially unobservable, so extinction can only be inferred. Statistical methods are described for inferring extinction from sighting records, species–area considerations, and taxonomic samples collected at two different times. The methods are illustrated using a variety of real datasets, including a sighting record of the Caribbean monk seal and results from three surveys of benthic invertebrates.

Description: Delavenne, J., Metcalfe, K., Smith, R. J., Vaz, S., Martin, C. S., Dupuis, L., Coppin, F., and Carpentier, A. 2012. Systematic conservation planning in the eastern English Channel: comparing the Marxan and Zonation decision-support tools. – ICES Journal of Marine Science, 69: 75–83. The systematic conservation approach is now commonly used for the design of efficient marine protected area (MPA) networks, and identifying these priority areas often involves using specific conservation-planning software. Several such software programmes have been developed in recent years, each differing in the underlying algorithms used. Here, an investigation is made into whether the choice of software influences the location of priority areas by comparing outputs from Marxan and Zonation, two widely used conservation-planning, decision-support tools. Using biological and socio-economic data from the eastern English Channel, outputs are compared and it is shown that the two software packages identified similar sets of priority areas, although the relatively wide distribution of habitat types and species considered offered much flexibility. Moreover, the similarity increased with increasing spatial constraint, especially when using real-world cost data, suggesting that the choice of cost metric has a greater influence on conservation-planning analyses than the choice of software. However, Marxan generally produced more efficient results and Zonation produced results with greater connectivity, so the most appropriate software package will depend on the overall goals of the MPA planning process.

Description: Andersen, B. S., Ulrich, C., Eigaard, O. R., and Christensen, A-S. 2012. Short-term choice behaviour in a mixed fishery: investigating métier selection in the Danish gillnet fishery. – ICES Journal of Marine Science, 69: 131–143. The study presents a short-term effort allocation modelling approach based on a discrete choice random utility model combined with a survey questionnaire to examine the selection of métiers (a combination of fishing area and target species) in the Danish North Sea gillnet fishery. Key decision variables were identified from the survey questionnaire, and relevant proxies for the decision function were identified based on available landings and effort information. Additional variables from the survey questionnaire were further used to validate and verify the outcome of the choice model. Commercial fishers in a mixed fishery make use of a number of decision variables used previously in the literature, but also a number of decision parameters rarely explicitly accounted for, such as price, weather, and management regulation. The seasonal availability of individual target species and within-year changes in monthly catch ration were the main explanatory drivers, but gillnetters were also responsive to information on the whole fishery, fish prices, and distance travelled to fishing grounds. Heterogeneous responses were evident from geographic differences in home harbour, which underpins the need to understand alternative fishing strategies among individual gillnetters better.

Description: Calise, L., and Knutsen, T. 2012. Multifrequency target strength of northern krill ( Meganyctiphanes norvegica ) swimming horizontally. – ICES Journal of Marine Science, 69: 119–130. Multifrequency acoustic measurements on ex situ horizontally swimming krill were made in a novel experimental setting. An ensemble of northern krill ( Meganyctiphanes norvegica ) was introduced to a large enclosure (a mesocosm), and acoustic backscatter was sampled using a multifrequency (70, 120, and 200 kHz) echosounder (Simrad EK60). Two submerged lamps were placed at opposite sides of the mesocosm and switched on and off to induce the krill, by light attraction, to swim horizontally through the acoustic beams. By tracking echoes, animal displacement, swimming speed, and target strength ( TS ) by frequency were estimated. The dominant and secondary modes of the total-length distribution were 21.8 ± 3.0 and 27.8 ± 2.7 mm, respectively. Although krill orientation was assumed stable and the ping rate was high, the range and inter-ping variability of the average TS values were large, decreasing and increasing with frequency, respectively. The overall TS frequency response observed and concurrent measurements at 120 and 200 kHz confirm the theoretical expectation that the acoustic backscatter from the investigated organisms were confined to the Rayleigh and Geometric scattering regions, a finding that might both aid acoustic identification and size-group separation of in situ northern krill.

Description: Rosen, S., Engås, A., Fernö, A., and Jörgensen, T. 2012. The reactions of shoaling adult cod to a pelagic trawl: implications for commercial trawling. – ICES Journal of Marine Science, 69: 303–312. The reactions of shoaling adult Atlantic cod to a pelagic trawl were measured during fishing off the north coast of Norway. Cod remaining in the trawl track dived at rates as fast as 0.35 m s –1 following vessel passage and swam away from the vessel, in the direction of the approaching trawl, at an average rate of 0.6 m s –1 . They did not attempt to swim ahead of the trawl as documented previously, but passed into the lower half of the trawl entrance and swam slowly in the direction of trawling at a rate of 0.2–0.5 m s –1 as the trawl's greater speed through water carried them deeper into the trawl. Shoals compressed vertically once inside the trawl, suggesting that packing density increased at least fourfold. Fish remained in the lower part of the trawl as they moved through its tapered section towards the codend, with little to no clearance above the bottom panel, but significant clearance beneath the top panel. Catches were sufficient to support commercial harvest, and the behaviour observed suggests that changes in trawl design and fishing strategy might improve fuel economy and species selectivity.

Description: Shackell, N. L., Bundy, A., Nye, J. A., and Link, J. S. 2012. Common large-scale responses to climate and fishing across Northwest Atlantic ecosystems. – ICES Journal of Marine Science, 69: 151–152. Investigating whether there were common biological responses to climate and fishing across seven Northwest Atlantic ecosystems, a minimum/maximum autocorrelation factor analysis of biological indicators for each region revealed a common primary multivariate trend of a rapid change during the 1980s and early 1990s. There was a strong common pattern in the biological indicators responsible for the primary multivariate temporal trend in the five more northerly regions: an increase in the abundance of phytoplankton, an increase in biomass at mid-trophic levels, and a decline in predatory groundfish size. The common associations between patterns and drivers were fishing indices and the Atlantic Multidecadal Oscillation, but all associations weakened when co-varying drivers were held constant. The results are consistent with known long-term effects of intense fishing, such as a decline in average fish size and changes in species composition. Less fishing pressure has allowed some regions to recover to former predatory biomass levels since the late 1990s, but the bulk of the biomass consists of fewer species. However, fishing was not the only driver, and a more mechanistic understanding of how the climate affects lower trophic levels is needed to contextualize climate effects in heavily fished ecosystems.

Description: Deporte, N., Ulrich, C., Mahévas, S., Demanèche, S., and Bastardie, F. 2012. Regional métier definition: a comparative investigation of statistical methods using a workflow applied to international otter trawl fisheries in the North Sea. – ICES Journal of Marine Science, 69: 331–342. The European Common Fisheries Policy recognizes the importance of accounting for heterogeneity in fishing practices, and métier-based sampling is now at the core of the EU Data Collection Framework. The implementation of such an approach would require Member States to agree on the standard regional métier definitions and on practical rules to categorize logbook records into métiers. Several alternative approaches have been used in the past to categorize landings profiles, but no consensus has yet emerged. A generic open-source workflow is developed to test and compare a selection of methods, including principal components analysis (PCA), hierarchical agglomerative clustering (HAC), K -means, and Clustering LARge Applications (CLARA), and to provide simple allocation rules. This workflow is applied to a unique regional dataset consisting of bottom-trawl logbooks of five North Sea countries. No method proved to be infallible, but combining PCA with either CLARA or HAC performed best. For 2008, a hierarchical classification with 14 species assemblages is proposed. Discriminant analysis proved more robust than simple ordination methods for allocating a new logbook record into an existing métier. The whole approach is directly operational and could contribute to defining more objective and consistent métiers across European fisheries.

Description: Greenstreet, S. P. R., Rogers, S. I., Rice, J. C., Piet, G. J., Guirey, E. J., Fraser, H. M., and Fryer, R. J. 2012. A reassessment of trends in the North Sea Large Fish Indicator and a re-evaluation of earlier conclusions. – ICES Journal of Marine Science, 69: 343–345. Previous analysis of the Large Fish Indicator, the basis for the North Sea "fish communities" EcoQO, suggested a strong recovery between 2001 and 2008. Discovery of a mistake in this earlier analysis now suggests that this recovery is not as strong as first thought, but reanalysis of the corrected data still supports the majority of conclusions drawn in the original paper.

Description: Xie, Y., Michielsens, C. G. J., and Martens, F. J. 2012. Classification of fish and non-fish acoustic tracks using discriminant function analysis. – ICES Journal of Marine Science, 69: 313–322. Hydroacoustic data acquired for estimating fish populations contain information on both fish and non-fish targets, so sonar technicians traditionally rely on their knowledge of fish behaviour and experience with hydroacoustics to remove non-fish targets from the hydroacoustic data. This process is often labour-intensive and time-consuming, making real-time assessment of fish populations difficult. Simple solutions are not always available for all circumstances. However, the split-beam sonar data collected in the lower Fraser River, British Columbia, Canada, showed distinct signatures between actively swimming fish and non-fish objects such as drifting debris, surface bubbles, and stationary objects in the water column and off the river bottom. Acoustic tracks of fish and non-fish targets were characterized by differentiable statistical patterns that were amenable to discriminant function analysis (DFA). An application of DFA to segregate fish and non-fish targets detected by a split-beam sonar system in the lower Fraser River is presented, characteristics of user-identified fish and non-fish acoustic tracks being utilized as learning samples for the DFA. Also, a method to rank the discriminating power of individual variables is presented, providing guidance for constructing efficient and effective discriminant functions with variables that offer high discriminating power. The DFA yielded classification accuracies of 96% for fish and 91% for non-fish tracks and reduced the manual sorting time by 50–75%.

Description: Rey, F. 2012. Declining silicate concentrations in the Norwegian and Barents Seas. – ICES Journal of Marine Science, 69: 208–212. Since 1990, a decline in silicate concentrations together with increasing salinities has been observed in the Atlantic water of the Norwegian and Barents Seas. This decline in silicate has been found to be related to the relative proportion in which eastern and western source water masses from the northeastern North Atlantic enter the Norwegian Sea.

Description: Zhou, S., Milton, D. A., and Fry, G. C. 2012. Integrated risk analysis for rare marine species impacted by fishing: sustainability assessment and population trend modelling. – ICES Journal of Marine Science, 69: 271–280. Risk assessment and conservation management of rare species are challenging due to a lack of data. We developed an integrated risk assessment approach to assess human impact on population sustainability of rare species. The approach involved two components: a quantitative sustainability assessment coupled with modelling trends in relative abundance. Both components took nil catches into account through zero-inflated statistical distributions that simultaneously modelled the zero and non-zero catches separately in submodels. The sustainability assessment used detection–non-detection data for population estimation and linked sustainability to easily collected life-history traits. This component provides an assessment of population sustainability at one point in time. The trend modelling applied zero-inflated negative binomial models to temporal trends in density and dispersion of species. It provided a complement to the static sustainability assessment. We applied this integrated approach to assess the risk to 14 species of rare, protected sea snakes incidentally caught in the Australian Northern Prawn Fishery. This approach can be applicable for risk assessment of many species with limited abundance data, a large number of absences and some presence–absence information only.

Description: Frid, C. L. J., and Paramor, O. A. L. 2012. Feeding the world: what role for fisheries? – ICES Journal of Marine Science, 69: 145–150. Fisheries (wild capture and aquaculture) deliver more than 110 million tonnes of food and around 15% of the dietary protein to the 7 billion people currently living on the planet. With the global population expected to peak at 9 billion by 2050, and 〉80% of global fish stocks currently fully or overexploited (and aquaculture is at least in part dependent on capture fisheries), the contribution of fisheries looks set to decline. The challenge is therefore determining how better management, an ecosystem perspective, and more efficient utilization of fisheries waste can support fisheries products continuing to contribute significantly to "feeding the world" up to and beyond the population peak.

Description: Boström, M. K., Östman, Ö., Bergenius, M. A. J., and Lunneryd, S-G. 2012. Cormorant diet in relation to temporal changes in fish communities. – ICES Journal of Marine Science, 69: 175–183. The stomach contents of 229 great cormorants ( Phalacrocorax carbo sinensis ) shot between March and October 2009 along the Swedish east coast were analysed for differences in diet between gender, age, and breeding phase. Sticklebacks ( Gasterosteus aculeatus and Pungitius pungitius ) were the most common prey, followed by eelpout ( Zoarces viviparus ), herring ( Clupea harengus membras ), and cyprinids (Cyprinidae). Diet did not differ between age and gender, but changed over the breeding season. The different phases explained around 10% of the total variation in stomach content between cormorants, suggesting no major shift in diet over the breeding season. The diet of cormorants in 2009 was compared with the results of a study conducted in the same area in 1992. There were evident changes in the diet between 1992 and 2009, with less perch ( Perca fluviatilis ) and cyprinids and more eelpout and herring in 2009. This change in diet could partly be related to changes in the fish community. The seasonal changes in diet composition of whole stomachs were less notable than in many previous studies, but long-term changes in the fish communities may induce changes in cormorant diet. It is clearly important to use stomach contents in areas with many small fish species for a comprehensive assessment of cormorant diet.

Description: Cañás, L., Stransky, C., Schlickeisen, J., Sampedro, M. P., and Fariña, A. C. 2012. Use of the otolith shape analysis in stock identification of anglerfish ( Lophius piscatorius ) in the Northeast Atlantic. – ICES Journal of Marine Science, 69: 250–256. The geographical variability in the anglerfish Lophius piscatorius otolith shape was investigated to examine biological evidence for the current separation into Northern and Southern stocks of L. piscatorius established in the Southwest European waters (Southern Shelf area) by the International Council for the Exploration of the Sea (ICES). In all, 629 otoliths of L. piscatorius (12–90 cm total length) from commercial fishing and research surveys were collected in two consecutive years, 2008 and 2009. The otolith shape was compared between stocks for differences in univariate descriptors using non-parametric analysis of covariance, as well as elliptical Fourier shape descriptors, through a metric multidimensional scaling and a linear discriminant analysis (LDA). The differences found in the otolith shape between Northern and Southern stocks were not large enough to hold the current status of L. piscatorius regarding stock definition in the Southern Shelf area, although homogeneous mixing between both stocks is not expected as there were significant differences between three univariate descriptors and the LDA was significant. These results are in agreement with the previous L. piscatorius stock identification studies (genetics, body morphology, and migratory behaviour), which do not support the current separation of these stocks but found substructures within the area.

Description: Variation in protein sequence and gene expression each contribute to phenotypic diversity, and may be subject to similar selective pressures. Eusocial insects are particularly useful for investigating the evolutionary link between protein sequence and condition-dependent patterns of gene expression because gene expression plays a central role in determining differences between eusocial insect sexes and castes. We investigated the relationship between protein coding sequence evolution and gene expression patterns in the fire ants Solenopsis invicta, S. richteri, and their hybrids to gain greater insight into how selection jointly operates on gene expression and coding sequence. We found that genes with high expression variability within castes and sexes were frequently differentially expressed between castes and sexes, as well as between species and hybrids. These results indicate that genes showing high variation in expression in one context also tend to show high variation in expression in other contexts. Our analyses further revealed that variation in both intra- and interspecific gene expression was positively associated with rate of protein sequence evolution in Solenopsis . This suggests that selective constraints on a gene operate both at the level of protein sequence and at the level of gene expression regulation. Overall, our study provides one of the strongest demonstrations that selective constraints mediate both protein sequence evolution and gene expression variability across different biological contexts and timescales.

Description: The ability to survey polymorphism on a genomic scale has enabled genome-wide scans for the targets of natural selection. Theory that connects patterns of genetic variation to evidence of natural selection most often assumes a diallelic locus and no recurrent mutation. Although these assumptions are suitable to selection that targets single nucleotide variants, fundamentally different types of mutation generate abundant polymorphism in genomes. Moreover, recent empirical results suggest that mutationally complex, multiallelic loci including microsatellites and copy number variants are sometimes targeted by natural selection. Given their abundance, the lack of inference methods tailored to the mutational peculiarities of these types of loci represents a notable gap in our ability to interrogate genomes for signatures of natural selection. Previous theoretical investigations of mutation-selection balance at multiallelic loci include assumptions that limit their application to inference from empirical data. Focusing on microsatellites, we assess the dynamics and population-level consequences of selection targeting mutationally complex variants. We develop general models of a multiallelic fitness surface, a realistic model of microsatellite mutation, and an efficient simulation algorithm. Using these tools, we explore mutation-selection-drift equilibrium at microsatellites and investigate the mutational history and selective regime of the microsatellite that causes Friedreich’s ataxia. We characterize microsatellite selective events by their duration and cost, note similarities to sweeps from standing point variation, and conclude that it is premature to label microsatellites as ubiquitous agents of efficient adaptive change. Together, our models and simulation algorithm provide a powerful framework for statistical inference, which can be used to test the neutrality of microsatellites and other multiallelic variants.

Description: Although endogenous retroviruses are common across vertebrate genomes, the koala retrovirus (KoRV) is the only retrovirus known to be currently invading the germ line of its host. KoRV is believed to have first infected koalas in northern Australia less than two centuries ago. We examined KoRV in 28 koala museum skins collected in the late 19th and 20th centuries and deep sequenced the complete proviral envelope region from five northern Australian specimens. Strikingly, KoRV env sequences were conserved among koalas collected over the span of a century, and two functional motifs that affect viral infectivity were fixed across the museum koala specimens. We detected only 20 env polymorphisms among the koalas, likely representing derived mutations subject to purifying selection. Among northern Australian koalas, KoRV was already ubiquitous by the late 19th century, suggesting that KoRV evolved and spread among koala populations more slowly than previously believed. Given that museum and modern koalas share nearly identical KoRV sequences, it is likely that koala populations, for more than a century, have experienced increased susceptibility to diseases caused by viral pathogenesis.

Description: Protein interaction networks play central roles in biological systems, from simple metabolic pathways through complex programs permitting the development of organisms. Multicellularity could only have arisen from a careful orchestration of cellular and molecular roles and responsibilities, all properly controlled and regulated. Disease reflects a breakdown of this organismal homeostasis. To better understand the evolution of interactions whose dysfunction may be contributing factors to disease, we derived the human protein coevolution network using our MatrixMatchMaker algorithm and using the Orthologous MAtrix project (OMA) database as a source for protein orthologs from 103 eukaryotic genomes. We annotated the coevolution network using protein–protein interaction data, many functional data sources, and we explored the evolutionary rates and dates of emergence of the proteins in our data set. Strikingly, clustering based only on the topology of the coevolution network partitions it into two subnetworks, one generally representing ancient eukaryotic functions and the other functions more recently acquired during animal evolution. That latter subnetwork is enriched for proteins with roles in cell–cell communication, the control of cell division, and related multicellular functions. Further annotation using data from genetic disease databases and cancer genome sequences strongly implicates these proteins in both ciliopathies and cancer. The enrichment for such disease markers in the animal network suggests a functional link between these coevolving proteins. Genetic validation corroborates the recruitment of ancient cilia in the evolution of multicellularity.

Description: Here, we sequenced the 5,419,609 bp circular genome of an Enterobacter aerogenes clinical isolate that killed a patient and was resistant to almost all current antibiotics (except gentamicin) commonly used to treat Enterobacterial infections, including colistin. Genomic and phylogenetic analyses explain the discrepancies of this bacterium and show that its core genome originates from another genus, Klebsiella. Atypical characteristics of this bacterium (i.e., motility, presence of ornithine decarboxylase, and lack of urease activity) are attributed to genomic mosaicism, by acquisition of additional genes, such as the complete 60,582 bp flagellar assembly operon acquired "en bloc" from the genus Serratia. The genealogic tree of the 162,202 bp multidrug-resistant conjugative plasmid shows that it is a chimera of transposons and integrative conjugative elements from various bacterial origins, resembling a rhizome. Moreover, we demonstrate biologically that a G53S mutation in the pmrA gene results in colistin resistance. E. aerogenes has a large RNA population comprising 8 rRNA operons and 87 cognate tRNAs that have the ability to translate transferred genes that use different codons, as exemplified by the significantly different codon usage between genes from the core genome and the "mobilome." On the basis of our findings, the evolution of this bacterium to become a "killer bug" with new genomic repertoires was from three criteria that are "opportunity, power, and usage" to indicate a sympatric lifestyle: "opportunity" to meet other bacteria and exchange foreign sequences since this bacteria was similar to sympatric bacteria; "power" to integrate these foreign sequences such as the acquisition of several mobile genetic elements (plasmids, integrative conjugative element, prophages, transposons, flagellar assembly system, etc.) found in his genome; and "usage" to have the ability to translate these sequences including those from rare codons to serve as a translator of foreign languages.

Description: Diatoms are the most species-rich group of microalgae, and their contribution to marine primary production is important on a global scale. Diatoms can form dense blooms through rapid asexual reproduction; mutations acquired and propagated during blooms likely provide the genetic, and thus phenotypic, variability upon which natural selection may act. Positive selection was tested using genome and transcriptome-wide pair-wise comparisons of homologs in three genera of diatoms ( Pseudo-nitzschia, Ditylum, and Thalassiosira ) that represent decreasing phylogenetic distances. The signal of positive selection was greatest between two strains of Thalassiosira pseudonana . Further testing among seven strains of T. pseudonana yielded 809 candidate genes of positive selection, which are 7% of the protein-coding genes. Orphan genes and genes encoding protein-binding domains and transcriptional regulators were enriched within the set of positively selected genes relative to the genome as a whole. Positively selected genes were linked to the potential selective pressures of nutrient limitation and sea surface temperature based on analysis of gene expression profiles and identification of positively selected genes in subsets of strains from locations with similar environmental conditions. The identification of positively selected genes presents an opportunity to test new hypotheses in natural populations and the laboratory that integrate selected genotypes in T. pseudonana with their associated phenotypes and selective forces.

Description: Antisense transcription, or transcription on the opposite strand of the same genomic locus as another transcript, has been observed in many organisms, including yeast. Several antisense transcripts are known to be conserved across various species of yeast, and a few antisense transcripts are associated with functional regulation of the sense transcript. We detect antisense transcription from approximately 90% of protein-coding genes, and antisense transcription is generally associated with histone modifications indicative of a transcriptionally active state. The pattern of genome-wide antisense transcription in two species of budding yeast, Saccharomyces cerevisiae and S. paradoxus , is widely evolutionarily conserved: Antisense transcripts exhibit conserved expression levels and localization with respect to gene annotations. Regions of genes exhibiting conserved antisense transcription also show less sequence divergence than regions of genes without antisense transcription. These findings provide further support that widespread antisense transcription is functional in yeast, and expand the catalog of putative functional antisense transcripts to include nonpolyadenylated transcripts. Because antisense transcripts are less divergent in sequence than expected, they likely contain sequences important to their function.

Description: Reconstruction of the past is an important task of evolutionary biology. It takes place at different points in a hierarchy of molecular variation, including genes, individuals, populations, and species. Statistical inference about population histories has recently received considerable attention, following the development of computational tools to provide tractable approaches to this very challenging problem. Here, we introduce a likelihood-based approach which generalizes a recently developed model for random fluctuations in allele frequencies based on an approximation to the neutral Wright–Fisher diffusion. Our new framework approximates the infinite alleles Wright–Fisher model and uses an implementation with an adaptive Markov chain Monte Carlo algorithm. The method is especially well suited to data sets harboring large population samples and relatively few loci for which other likelihood-based models are currently computationally intractable. Using our model, we reconstruct the global population history of a major human pathogen, Streptococcus pneumoniae . The results illustrate the potential to reach important biological insights to an evolutionary process by a population genetics approach, which can appropriately accommodate very large population samples.

Description: Humans’ ability for rapid dispersal and adaptation has allowed us to colonize diverse geographic and climatic regions of the planet, creating a complex evolutionary history. This complexity can be understood, at least partially, by modeling the underlying demographic parameters in the evolutionary process. In this study, we analyze a model of human evolution in which population size, gene flow (GF), and time are varied. Specifically, we simulate mitochondrial DNA for 42 demographic scenarios, represented by 42 parameter combinations, to describe the initial dispersal of modern humans out of Africa. The analyses include three values for colonization size (CS; 1%, 10%, and 30% of the African population), seven values for rate of GF (10 –6 –0.5), and two values for time of colonization (50,000 and 100,000 years ago). We then estimate summary statistics for the simulated data sets to calculate the percent of explained variation by each parameter and to identify which parameter combinations generate distinct differences in genetic variation, that is, which demographic scenarios can be distinguished from each other. On the basis of these results, we make recommendations about which summary statistics to use according to the parameter of interest. Our results show that CS, GF, and their interaction have the largest effect on genetic variation under our model of human evolution. Comparison with empirical data suggests that 1% of the existing African mitochondrial genetic variation left and colonized the rest of the world (i.e., CS = 1%) and bidirectional GF continued at a level of ~10 individuals per generation (i.e., GF = 10 –3 ) after the initial colonization. Our study serves as a model to bridge the gap between the use of simulations for theoretical population genetics and empirical data analysis such as approximate Bayesian computation approaches and is, thus, applicable to the study of molecular evolution in any organism.

Description: In recent years, ancient DNA has increasingly been used for estimating molecular timescales, particularly in studies of substitution rates and demographic histories. Molecular clocks can be calibrated using temporal information from ancient DNA sequences. This information comes from the ages of the ancient samples, which can be estimated by radiocarbon dating the source material or by dating the layers in which the material was deposited. Both methods involve sources of uncertainty. The performance of Bayesian phylogenetic inference depends on the information content of the data set, which includes variation in the DNA sequences and the structure of the sample ages. Various sources of estimation error can reduce our ability to estimate rates and timescales accurately and precisely. We investigated the impact of sample-dating uncertainties on the estimation of evolutionary timescale parameters using the software BEAST. Our analyses involved 11 published data sets and focused on estimates of substitution rate and root age. We show that, provided that samples have been accurately dated and have a broad temporal span, it might be unnecessary to account for sample-dating uncertainty in Bayesian phylogenetic analyses of ancient DNA. We also investigated the sample size and temporal span of the ancient DNA sequences needed to estimate phylogenetic timescales reliably. Our results show that the range of sample ages plays a crucial role in determining the quality of the results but that accurate and precise phylogenetic estimates of timescales can be made even with only a few ancient sequences. These findings have important practical consequences for studies of molecular rates, timescales, and population dynamics.

Description: Genetic exchange by conjugation is responsible for the spread of resistance, virulence, and social traits among prokaryotes. Recent works unraveled the functioning of the underlying type IV secretion systems (T4SS) and its distribution and recruitment for other biological processes (exaptation), notably pathogenesis. We analyzed the phylogeny of key conjugation proteins to infer the evolutionary history of conjugation and T4SS. We show that single-stranded DNA (ssDNA) and double-stranded DNA (dsDNA) conjugation, while both based on a key AAA + ATPase, diverged before the last common ancestor of bacteria. The two key ATPases of ssDNA conjugation are monophyletic, having diverged at an early stage from dsDNA translocases. Our data suggest that ssDNA conjugation arose first in diderm bacteria, possibly Proteobacteria, and then spread to other bacterial phyla, including bacterial monoderms and Archaea. Identifiable T4SS fall within the eight monophyletic groups, determined by both taxonomy and structure of the cell envelope. Transfer to monoderms might have occurred only once, but followed diverse adaptive paths. Remarkably, some Firmicutes developed a new conjugation system based on an atypical relaxase and an ATPase derived from a dsDNA translocase. The observed evolutionary rates and patterns of presence/absence of specific T4SS proteins show that conjugation systems are often and independently exapted for other functions. This work brings a natural basis for the classification of all kinds of conjugative systems, thus tackling a problem that is growing as fast as genomic databases. Our analysis provides the first global picture of the evolution of conjugation and shows how a self-transferrable complex multiprotein system has adapted to different taxa and often been recruited by the host. As conjugation systems became specific to certain clades and cell envelopes, they may have biased the rate and direction of gene transfer by conjugation within prokaryotes.

Description: Hydrothermal vents are typically located in midocean ridges and back-arc basins and are usually generated by the movement of tectonic plates. Life thrives in these environments despite the extreme conditions. In addition to chemoautotrophic bacteria, decapod crustaceans are dominant in many of the hydrothermal vents discovered to date. Contrary to the hypothesis that these species are remnants of relic fauna, increasing evidence supports the notion that hydrothermal vent decapods have diversified in more recent times with previous research attributing the origin of alvinocarid shrimps to the Miocene. This study investigated seven representative decapod species from four hydrothermal vents throughout the Western Pacific and Indian Oceans. A partitioned mix-model phylogenomic analysis of mitochondrial DNA produced a consistent phylogenetic topology of these vent-endemic species. Additionally, molecular dating analysis calibrated using multiple fossils suggested that both bythograeid crabs and alvinocarid shrimps originated in the late Mesozoic and early Cenozoic. Although of limited sampling, our estimates support the extinction/repopulation hypothesis, which postulates recent diversification times for most hydrothermal vent species due to their mass extinction by global deep-water anoxic/dysoxic events during the Late Cretaceous and Early Tertiary. The continental-derived property of the West Pacific province is compatible with the possibility that vent decapods diversified from ancestors from shallow-water regions such as cold seeps. Our results move us a step closer toward understanding the evolutionary origin of hydrothermal vent species and their distribution in the Western Pacific–Indian Ocean Region.

Description: Bachiller, E., Cotano, U., Boyra, G., and Irigoien, X. 2013. Spatial distribution of the stomach weights of juvenile anchovy ( Engraulis encrasicolus L.) in the Bay of Biscay. – ICES Journal of Marine Science, 70: 362–378. Previous studies have shown that the survival of larval and juvenile anchovy off the continental shelf in the Bay of Biscay is not significantly different from that observed on the shelf, even though the food concentration is significantly higher on the shelf. In this paper we investigate the causes for the lack of relationship between food and survival for anchovy juvenile through analyses of feeding activity versus zooplankton distribution (in the transition from summer to fall from 2003 to 2010). The spatial distribution of the stomach weights of juvenile anchovy in relation to body size on and off the shelf revealed higher stomach fullness in areas off the shelf, where less zooplankton biomass was available. This result indicates that the food concentration is not always the main factor determining ingestion in fish. A situation of comparatively lesser prey abundance may provide better feeding conditions when combined with lower predation risk and lower light attenuation. In addition, the relatively heavier stomachs found in anchovies caught in years of higher recruitment indices suggest a large stomach content may be a symptom of good biological condition that may favour the winter survival of a larger fraction of the juvenile stock.

Description: Van Hoof, L. 2013. Design or pragmatic evolution: applying ITQs in EU fisheries management. – ICES Journal of Marine Science, 70: 462–470. Among the proposals for the 2012 revision of the EU Common Fisheries Policy, a strong case is made for the introduction of a system of rights-based management. The EU perceives individual fishing concessions as an important instrument for capacity management. We will use the introduction of individual tradable quotas in the management of the Dutch North Sea beam trawl fisheries as a case for exploring the effect of the introduction of such an instrument. The effect will be assessed in terms of reduction of fishing capacity in the Dutch beam trawl fleet and its economic and social impact. These Dutch experiences will be translated to the current debate on the reform of the EU Common Fisheries Policy. Especially, we will focus on the issues of "relative stability", the concentration of rights, and the effects on the small-scale fisheries sector. Some of the negative effects associated with individual tradable rights can be addressed through design. However, trying to maintain stability and counter perceived negative impacts on fishing communities will modify the effect of introducing individual fishing concessions.

Description: Spitz, J., Chouvelon, T., Cardinaud, M., Kostecki, C., and Lorance, P. 2013. Prey preferences of adult sea bass Dicentrarchus labrax in the northeastern Atlantic: implications for bycatch of common dolphin Delphinus delphis – ICES Journal of Marine Science, 70: 452–461. In the northeastern Atlantic, adult sea bass ( Dicentrarchus labrax ) is one of largest fish living on the shelf, and this species has important commercial value. However, pelagic trawl fisheries that target sea bass have negative operational interactions with common dolphins ( Delphinus delphis ). Our goal was to determine the diet of adult sea bass in the Bay of Biscay from stomach-content and stable-isotope analyses, and explore the dietary overlap between sea bass and common dolphins. We found that sea bass primarily target small pelagic fish, most notably mackerel ( Scomber scombrus ), scads ( Trachurus spp.), anchovy ( Engraulis encrasicolus ), and sardine ( Sardina pilchardus ). These four species also dominated the diets of common dolphins. This overlap in feeding preferences could increase the risk of dolphins being caught by trawl fisheries while feeding among sea bass, and may be an underlying mechanism to explain the high rate of common dolphin bycatch observed in the pelagic trawl fishery for sea bass in the Bay of Biscay. Understanding the foraging ecology and trophic interactions of predator species is an essential step for identifying and resolving management issues in the northeastern Atlantic and other marine ecosystems.

Description: Lassen, H., Pedersen, S. A., Frost, H., and Hoff. A. 2013. Fishery management advice with ecosystem considerations. – ICES Journal of Marine Science, 70: 471–479. The European Union Marine Strategy Framework Directive (MSFD) includes four descriptors of Good Environmental Status (GES) which are affected by fishing activity. These descriptors are: biodiversity, fish stocks, foodweb, and seabed integrity. This paper shows how these descriptors can be related to variables within an ecological model and how an ecological model can be used to analyse whether the fishing pressure that is estimated based on bioeconomic criteria is within general sustainable limits. The paper presents an example of such an analysis of the Eastern Baltic cod fishery using two models: a bioeconomic model and an ecological model. The models are calibrated based on historic data. The mapping between the descriptors specified by MSFD and variables available for analysis in the models is incomplete, e.g. genetics and spatial structures are not included in the models. The models can be used strategically, providing a qualitative understanding of the anticipated relative changes.

Description: Finley, C. and Oreskes, N. 2013. Maximum sustained yield: a policy disguised as science. – ICES Journal of Marine Science, 70: 245–250. Overfishing is most commonly explained as an example of the tragedy of the commons, where individuals are unable to control their activities, leading to the destruction of the resource they are dependent on. The historical record suggests otherwise. Between1949 and 1958, the US State Department used fisheries science, and especially the concept of maximum sustained yield (MSY) as a political tool to achieve its foreign policy objectives. During the Cold War, the Department thought that if countries were allowed to restrict fishing in their waters, it might lead to restrictions on passage of military vessels. While there has been much criticism of MSY and its failure to conserve fish stocks, there has been little attention paid to the political context in which MSY was adopted.

Description: The sharing of the Northeast Atlantic mackerel ( Scomber scombrus ) stock is analysed as a game between four parties: the European Union, Norway, the Faroe Islands, and Iceland. Consideration is given to how the outcome depends on the nature of the stock's migrations. Two types of migrations are considered: (i) density-dependent, where the mackerel migrates into the Icelandic economic zone only if it exceeds 3.5 million t, and (ii) stochastic migrations, where the said migrations are stochastic. It is determined that the Faroe Islands would never accept a cooperative solution wherein they can only fish with the globally optimal fishing mortality within their own zone. This is also true for Iceland when the migrations into her zone are stochastic, but not if they are density-dependent. In the latter case, the other players have incentives to retaliate to Icelandic overfishing by fishing harder, which greatly reduces the number of years when mackerel are available in the Icelandic zone. It is assumed that the objective is maximization of the catch volume over a time-horizon of 50 years.

Description: Harris, P. T., Bridge, T. C. L., Beaman, R. J., Webster, J. M., Nichol, S. L., and Brooke, B. P. 2013. Submerged banks in the Great Barrier Reef, Australia, greatly increase available coral reef habitat. – ICES Journal of Marine Science, 70: 284–293. Anthropogenic global ocean warming is predicted to cause bleaching of many near-sea-surface (NSS) coral reefs, placing increased importance on deeper reef habitats to maintain coral reef biodiversity and ecosystem function. However, the location and spatial extent of many deep reef habitats is poorly known. The question arises: how common are deep reef habitats in comparison with NSS reefs? We used a dataset from the Great Barrier Reef (GBR) to show that only about 39% of available seabed on submerged banks is capped by NSS coral reefs (16 110 km 2 ); the other 61% of bank area (25 600 km 2 ) is submerged at a mean depth of around 27 m and represents potential deep reef habitat that is spatially distributed along the GBR continental shelf in the same latitudinal distribution as NSS reefs. Out of 25 600 km 2 of submerged bank area, predictive habitat modelling indicates that more than half (around 14 000 km 2 ) is suitable habitat for coral communities.

Description: Dutertre, M., Hamon D., Chevalier C., and Ehrhold, A. 2013. The use of the relationships between environmental factors and benthic macrofaunal distribution in the establishment of a baseline for coastal management – ICES Journal of Marine Science, 70: 294–308. Relationships between benthic macrofauna and natural abiotic factors were studied along the coastal fringe of South Brittany, situated north of the Gulf of Biscay on the French Atlantic continental shelf. Within the framework of the REBENT network, sediment characteristics, depth, and macrofaunal abundance were determined for 95 stations spread over five subtidal sectors, using a combination of seabed acoustic remote sensing systems and grab sampling. The physico-chemical properties of the water column and the hydrodynamic conditions were generated by validated three-dimensional environmental models which take into account variations over shorter temporal scales. Multivariate analyses ranked 16 natural abiotic variables according to the significance of their influence on the macrofauna. Together these variables explained 51% of spatial variation in the macrofauna, with morpho-sedimentological and hydrological factors contributing 22% and 26%, respectively. The outputs from validated three-dimensional environmental models appear to be useful interpretational tools for benthic ecology studies, especially in estuarine and coastal ecosystems with high environmental variability due to regular freshwater inputs. Ten major species assemblages were identified using biological and physical characteristics. The results provide important baseline knowledge for future ecosystem and resource management.

Description: Prigge, E., Marohn, L., Oeberst R., and Hanel, R. 2013. Model prediction vs. reality—testing the predictions of a European eel ( Anguilla anguilla ) stock dynamics model against the in situ observation of silver eel escapement in compliance with the European eel regulation – ICES Journal of Marine Science, 70: 309–318. A direct monitoring of European silver eel ( Anguilla anguilla , L) escapement from rivers and estuaries has been proven to be challenging, and a Europe-wide documentation of escaping silver eel numbers therefore hardly seems realistic. To reinforce management decisions, policy-makers are thus widely reliant on the accuracy of escapement models. A 3-year programme of silver eel escapement monitoring was undertaken to compile model input data and revise an eel population model (German Eel Model II; GEM II) already used in the decision-making process of management authorities. By compiling necessary input data and analysing vital system-specific population characteristics, it was possible to compare the documented silver eel escapement with the modelled potential silver eel escapement. Resulting model predictions were close to actually monitored escapement numbers, which were distinctly lower than reference escapement values for the same freshwater system given in the implementation report of the German Eel Management Plans. Applying different commercial and recreational catch scenarios revealed the sensitivity of the model. The results show the potential of the GEM II and highlight the importance of high-quality input data to use model predictions as the basis for management measures.

Description: Sumaila, U. R. 2013. How to make progress in disciplining overfishing subsidies. ICES Journal of Marine Science, 70: 251–258. The World Trade Organization (WTO) has been working for more than seven years now to discipline overfishing subsidies, as mandated by the global community, without success. I argue that this failure is partly because WTO negotiators aim for an all-inclusive deal, i.e. negotiations are conducted as a "single undertaking", whereby results must be achieved in all areas. Negotiators are required to broker an all-inclusive deal for all maritime WTO member countries and for all fisheries, whether domestic or international; small or large scale; developing or developed country fisheries. It is argued here that this commitment to a "single undertaking" does not align the incentive to remove subsidies with national interests, and therefore needs to be changed by splitting the world's fisheries into domestic and international fisheries. In this way, the battle for eliminating overfishing subsidies for some stocks would shift to home countries, and for others this would still rest with the international community. This split, it is argued, would align the incentives and improve the chances of eliminating overfishing subsidies.

Description: Morgan, M. J., Garabana, D., Rideout, R. M., Román, E., Pérez-Rodriguez, A. and Saborido-Rey, F. 2013. Changes in distribution of Greenland halibut in a varying environment. – ICES Journal of Marine Science, 70: 352–361. Fish are expected to respond to changing oceanographic temperature by altering their distribution. Off the coast of Newfoundland, Canada, there have been major changes in oceanographic temperature over the last several decades, with both record cold and record warm years being observed. Greenland halibut is an important flatfish species in the area, and is distributed in deep waters over a very wide geographic range. Thus, it might be buffered from temperature change in the overall area by reduced temperature variation at depth, and the diverse temperature conditions over its wide range. We examined intrapopulation variation in temperature and depth distribution, and the biological changes in relation to changes in available temperature. On the Flemish Cap, variation in available temperature was limited, and changes in depth were related to changing age composition and the differential depth distribution with age/size. In other areas there was a larger decline in available temperature, and associated with this, Greenland halibut moved to deeper waters and occupied warmer temperatures than they had previously. Concurrently, growth declined and condition increased. This study shows that shifts in distribution may not result in maintenance of homogeneous environmental conditions, and that resulting biological changes will be difficult to predict.

Description: Hufnagl, M., Huebert, K. B., and Temming, A. 2013. How does seasonal variability in growth, recruitment, and mortality affect the performance of length-based mortality and asymptotic length estimates in aquatic resources? – ICES Journal of Marine Science, 70: 329–341. We tested the sensitivity of eight methods for estimating total mortality from size frequencies (modified Wetherall; Powell; Beverton and Holt; Jones and van Zalinge; Hoenig; Ssentongo and Larkin; seasonal and non-seasonal Length Converted Catch Curve) to violations of basic assumptions, such as seasonal growth, mortality, recruitment and variable asymptotic length L or growth parameter K . For each method, bias was estimated by simulating length frequency distributions with different combinations of known L , Z and K values, calculating ( Z / K ) and L estimates, and comparing the true input with the estimated output values. Input mortality was generally underestimated by all methods and in 27% of all simulations no method provided estimates within ± 1. Spring recruitment especially negatively influenced the mortality estimate. A decision tree was developed that provides general guidance in selecting appropriate methods despite violated assumptions, but species-specific case studies are recommended. An example of a species-specific study is provided for the brown shrimp, Crangon crangon. Despite inherent limitations for all methods, the results illustrate that estimates of and Z for brown shrimp can be improved substantially by selecting suitable methods and correcting for observed bias.

Description: Brunel, T., and Piet, G. J. 2013. Is age structure a relevant criterion for the health of fish stocks? – ICES Journal of Marine Science, 70: 270–283. The age and size structure of exploited fish stocks is one of the criteria for Good Environmental Status of commercial fish. However, two underlying assumptions to this criterion remain to be tested: first, that a well-balanced age structure is indeed indicative of a "healthier" stock, and second, that managers can exert a control on the age structure, independently from the regulation of stock abundance. This study investigates these questions using simulations from a population model in which recruitment is based on egg production, which is more sensitive to age structure variations than the traditionally used spawning stock biomass (SSB) and that was parameterized to represent the population dynamics of North Sea cod, plaice, and herring. Our results show that (i) the age structure is highly dependent on the selection pattern, as well as on the level of fishing mortality; (ii) the selection pattern determines the ability of fish stocks to withstand, and recover from, external perturbation; and (iii) the selection pattern determines the output of the fishery providing the management option to balance stable but relatively low yields vs strongly fluctuating high yields. Therefore, we propose to make the selection pattern, for which clear management targets can be set, a policy goal instead of the age structure that is currently in place.

Description: Bréchon, A. L., Coombs S. H., Sims D. W., and Griffiths A. M. 2013. Development of a rapid genetic technique for the identification of clupeid larvae in the Western English Channel and investigation of mislabelling in processed fish products – ICES Journal of Marine Science, 70: 399–407. A novel genetic technique, involving real-time polymerase chain reaction (PCR) and high-resolution melt (HRM) analysis was developed, which successfully identified four species of clupeid larvae from the Western English Channel and revealed the presence of herring ( Clupea harengus ). This was unexpected in a context of global warming as herring are at the southern limit of their range off southwest England, where the larvae are generally very rare. The application of the method in cases of degraded DNA, such as in processed fishery products, was also assessed, revealing a low level of misidentification or equivocal identity (8.3%). Eight-year-old formalin-preserved samples were also analysed successfully. This highlights the potentially broad applications for HRM-based approaches to species identification in marine science.

Description: Quetglas, A., Ordines, F., Hidalgo, M., Monserrat, S., Ruiz, S., Amores, Á., Moranta, J., and Massutí, E. 2013. Synchronous combined effects of fishing and climate within a demersal community. – ICES Journal of Marine Science, 70: 319–328. Accumulating evidence shows that fishing exploitation and environmental variables can synergistically affect the population dynamics of exploited populations. Here, we document an interaction between fishing impact and climate variability that triggered a synchronic response in the population fluctuations of six exploited species in the Mediterranean from 1965–2008. Throughout this period, the fishing activity experienced a sharp increase in fishing effort, which caused all stocks to shift from an early period of underexploitation to a later period of overexploitation. This change altered the population resilience of the stocks and brought about an increase in the sensitivity of its dynamics to climate variability. Landings increased exponentially when underexploited but displayed an oscillatory behaviour once overexploited. Climatic indices, related to the Mediterranean mesoscale hydrography and large-scale north Atlantic climatic variability, seemed to affect the species with broader age structure and longer lifespan, while the global-scale El Niño Southern Oscillation index (ENSO) positively influenced the population abundances of species with a narrow age structure and short lifespan. The species affected by ENSO preferentially inhabit the continental shelf, suggesting that Mediterranean shelf ecosystems are sensitive to the hydroclimatic variability linked to global climate.

Description: Papetti, C., Di Franco, A., Zane, L., Guidetti, P., De Simone, V., Spizzotin, M., Zorica, B., Cikes Kec, V. and Mazzoldi, C. 2013. Single population and common natal origin for Adriatic Scomber scombrus stocks: evidence from an integrated approach – ICES Journal of Marine Science, 70: 387–398. In order to implement proper fishery management strategies aimed at avoiding stock declines, information about connectivity among stocks and populations is critically required. In this perspective, the present study investigated population structure of the Atlantic mackerel Scomber scombrus in the northern–central Adriatic Sea by integrating multiple approaches (analysis of fisheries data, population genetics, and otolith chemistry). Monthly data of fishery landings indicate a latitudinal trend along the western Adriatic coast, with Atlantic mackerel disappearing from the northern waters in winter, corresponding to the reproductive season. Population genetic analyses by genotyping of eight microsatellites clearly point to the presence of a single panmictic population in the northern–central Adriatic Sea. Otolith cores of samples from the northern–central Adriatic were chemically homogeneous, suggesting a common spawning ground. These results strongly suggest that Atlantic mackerel perform an autumn–winter migration in the northern–central Adriatic Sea, from the northern to the central sector, to reach a single spawning ground, and that a single population is present in this area. Considering that S. scombrus has shown a marked decline in the last 40 years in the Adriatic, this study highlights a potential high vulnerability to collapse by overfishing for the Atlantic mackerel stocks in this geographic area.

Description: Uusitalo, L., Fleming-Lehtinen, V., Hällfors, H., Jaanus A., Hällfors, S., and London, L. 2013. A novel approach for estimating phytoplankton biodiversity – ICES Journal of Marine Science, 70: 408–417. Maintaining biodiversity is one of the main priorities in environmental protection. The biodiversity of phytoplankton, the key primary producers in the marine ecosystem, is, however, often very difficult to estimate, since the phytoplankton assemblage includes a vast number of taxa, many of which occur in such small quantities that they may not be recorded in routine sampling. Moreover, many taxa cannot be identified to species level by routine methods such as light microscopy of preserved samples, even by a skilled taxonomist. This means that, in general, we cannot assume to have a complete list of species in the ecosystem at any given point in time. In this paper, we present an approach for evaluating phytoplankton biodiversity in spite of this challenge. Since eutrophication, which increases phytoplankton biomass, has been identified as the most important factor causing degradation of the Baltic Sea ecosystem, the proposed approach was evaluated against total phytoplankton biomass. When phytoplankton biomass was low, both low and high biodiversity values were observed, and, as the phytoplankton biomass increased, the high biodiversity values disappeared. These results were consistent both using data based on individual samples and using yearly sampling station averages.

Description: Frie, A. K., Hammill, M. O., Hauksson, E., Lind, Y., Lockyer, C., Stenman, O., and Svetocheva, O. 2013. Error patterns in age estimation and tooth readability assignment of grey seals ( Halichoerus grypus ): results from a transatlantic, image-based, blind-reading study using known-age animals – ICES Journal of Marine Science, 70: 418–430. We analysed error patterns in a first interlaboratory grey seal ( Halichoerus grypus ) age-reading experiment. The experiment involved ten readers, who estimated age using images of cementum growth layers from teeth of 68 known-age seals (0–22 years). The percentages of correct estimates ranged from 32.4% to 60.3% among readers, and 89.3% of all errors were by ±1–2 years. Six readers showed increasing underageing with increasing seal age. An elevated risk of underestimation by 1 year occurred in teeth collected 0–5 months after breeding and was attributed to more frequent absence of a distinct growth layer for the new year and lack of information on months between the last birthday and the date of sample collection (plusmonths). For plusmonths 6–11, positive bias was predominant, suggesting that overestimation is the more common error when plusmonth information is available. Readers assigned readability scores to the tooth sections, and 79.1% of all ageing errors occurred in sections of low or intermediate readability. Excluding these sections would, however, also exclude 43.0% of all correct estimates. Neither levels of age estimation error nor predictive values of readability assignments were associated with reader experience levels. Analyses of image markings identified common errors in delineations of annual increment layers.

Description: Domínguez-Petit, R., Ouellet, P., and Lambert, Y. 2013. Reproductive strategy, egg characteristics and embryonic development of Greenland halibut ( Reinhardtius hippoglossoides ). – ICES Journal of Marine Science, 70: 342–351. Despite the commercial importance of Greenland halibut (GH), important gaps exist in our knowledge of the reproductive and early life stage biology for this species. The present study examined through laboratory experiments the spawning strategy, realized fecundity, egg characteristics, biochemical composition, and embryonic development of GH. The results confirmed the hypothesis that GH is a single-batch spawner producing large eggs, resulting in low realized fecundity. Embryonic development and hatching time are highly dependent on incubation temperature; 50% hatching occurred after 46, 30, and 24 days at 2, 4, and 6°C, respectively. Few changes in the biochemical composition of the eggs are observed during embryonic development. Newly hatched larvae are not well developed, having a large yolk sac, no pigmentation and incomplete development of the jaws. Egg specific density confirmed the mesopelagic distribution of the eggs at sea. However, important buoyancy changes occurring in the last 3–4 days before hatching indicate that larvae hatch higher in the water column. These results are important for understanding advection and dispersion processes of GH eggs and larvae and the connectivity between spawning grounds and nursery areas.

Description: Johnsen, E., and Harbitz, A. 2013. Small-scale spatial structuring of burrowed sandeels and the catching properties of the dredge – ICES Journal of Marine Science, 70: 379–386. The lesser sandeel ( Ammodytes marinus ) remains burrowed for long periods in the seabed in sandy areas, and its habitat preference is reflected in a large-scale (10 km) patchy distribution in the North Sea. The small-scale spatial structure of sandeels in the seabed is less known. In this study, it was found that sandeel catches in grabs and dredges fit gamma distributions with a common shape parameter on both a 10 m and a 100 m scale. These spatial models were used to examine the catching properties of the dredge, which, in the new ICES sandeel assessment, is used to estimate recruitment and maturity curves. Analyses of the empirical data and simulations of the dredge and grab catch rates show very low dredge catchability (~5%), but no length-dependent catchability in the dredge was found. However, analyses of the catch rates of parallel dredge hauls indicated increasing dredge catchability with sandeel density. The observed small-scale patchiness and the low and seemingly density-dependent dredge catchability result in biased abundance estimates with low precision. Therefore, we recommend a joint international effort to develop a more catch-efficient sampling dredge, where the effect of density on catchability is further examined.

Description: Rousi, H., Laine, A. O., Peltonen, H., Kangas, P., Andersin, A-B., Rissanen, J., Sandberg-Kilpi, E., and Bonsdorff, E. 2013. Long-term changes in coastal zoobenthos in the northern Baltic Sea: the role of abiotic environmental factors – ICES Journal of Marine Science, 70: 440–451. We investigated site-specific changes in a coastal zoobenthic community during 1964–2007 in the northern Baltic Sea. Multivariate analysis indicated that the community structure had changed. The amphipods Monoporeia affinis and Pontoporeia femorata decreased drastically in the late 1970s–early 1980s, and by the early 1990s P. femorata vanished and M. affinis abundance was low. The decline of M . affinis and P . femorata was followed by an increase of the bivalve Macoma balthica and the arrival of the polychaete genus Marenzelleria in the 1990s. Trend analyses showed a rising trend for near-bottom temperature from the late 1960s. There was a significant decline in salinity during the early 1980s, which stabilized during the early 1990s. A negative trend was observed for oxygen concentration during the entire study period. There were interannual variations in the phosphorus and nitrogen concentrations, and a step-like increase in phosphorus concentration at the beginning of the 21st century. The correlations of temperature, salinity, oxygen, total phosphorus and nitrogen with zoobenthic communities were examined using Constrained Correspondence Analysis. Temperature was the most highly correlated explanatory variable for the benthic species. The study highlights the importance of long-term data sets in assessing the state and ecological processes of zoobenthic systems.

Description: O'Driscoll, R. L., Oeffner, J., and Dunford, A. J. 2013. In situ target strength estimates of optically verified southern blue whiting ( Micromesistius australis ) – ICES Journal of Marine Science, 70: 431–439. Estimates of the acoustic target strength (TS) of southern blue whiting ( Micromesistius australis ) at 38 kHz were obtained using an autonomous acoustic–optical system (AOS) mounted on a demersal trawl. Data were collected from aggregations of spawning adult [mean fork length (FL) 34.4 cm] and immature (mean FL 24.6 cm) southern blue whiting south of New Zealand. Mean TS was estimated from 162 tracks containing 695 echoes from targets identified from video as southern blue whiting. The mean TS was –37.9 dB with a 95% confidence interval ( CI ) of –39.7 to –36.6 dB for 21 immature fish and –34.6 dB (95% CI –35.4 to –34.0 dB) for 141 adults. A logarithmic fit through the mean TS values produced a TS–fork length (FL) relationship from optically verified targets of TS = 22.06 log 10 FL – 68.54. This new relationship gives TS values within 1 dB of those estimated using the relationship recently adopted by ICES for blue whiting ( Micromesistius poutassou ) of TS = 20 log 10 TL – 65.2 (where TL is total length) obtained from in situ measurements, but higher values than those estimated from the previous relationship for southern blue whiting of TS = 38 log 10 FL – 97, which was based on swimbladder modelling.

Description: De Robertis, A., Wilson, C. D., Furnish, S. R., and Dahl, P. H. 2013. Underwater radiated noise measurements of a noise-reduced fisheries research vessel. – ICES Journal of Marine Science, 70: 480–484. Vessel-radiated noise is traditionally measured at naval acoustic ranges, but lower-cost options are desirable for routine monitoring of research vessels. Measurements of a noise-reduced research vessel made at a naval noise range are compared to those made using an experimental mooring equipped with commercially available instrumentation. The measurements from the mooring were precise and within 2.5 dB of those from the noise range at third-octave bands 〈500 Hz. At higher frequencies, direct comparisons were precluded by an intermittent shaft-related noise present only during the mooring measurements, but previously observed at the navy range. The agreement between the two methods suggests that simplified, field-deployable hydrophone systems can be used to accurately characterize the noise signatures of research vessels.

Description: Proteins in the superfamily of voltage-gated ion channels mediate behavior across the tree of life. These proteins regulate the movement of ions across cell membranes by opening and closing a central pore that controls ion flow. The best-known members of this superfamily are the voltage-gated potassium, calcium (Ca v ), and sodium (Na v ) channels, which underlie impulse conduction in nerve and muscle. Not all members of this family are opened by changes in voltage, however. NALCN (NA + leak channel nonselective) channels, which encode a voltage-insensitive "sodium leak" channel, have garnered a growing interest. This study examines the phylogenetic relationship among Na v /Ca v voltage-gated and voltage-insensitive channels in the eukaryotic group Opisthokonta, which includes animals, fungi, and their unicellular relatives. We show that NALCN channels diverged from voltage-gated channels before the divergence of fungi and animals and that the closest relatives of NALCN channels are fungal calcium channels, which they functionally resemble.

Description: Newer parts of sex chromosomes, neo-sex chromosomes, offer unique possibilities for studying gene degeneration and sequence evolution in response to loss of recombination and population size decrease. We have recently described a neo-sex chromosome system in Sylvioidea passerines that has resulted from a fusion between the first half (10 Mb) of chromosome 4a and the ancestral sex chromosomes. In this study, we report the results of molecular analyses of neo-Z and neo-W gametologs and intronic parts of neo-Z and autosomal genes on the second half of chromosome 4a in three species within different Sylvioidea lineages (Acrocephalidea, Timaliidae, and Alaudidae). In line with hypotheses of neo-sex chromosome evolution, we observe 1) lower genetic diversity of neo-Z genes compared with autosomal genes, 2) moderate synonymous and weak nonsynonymous sequence divergence between neo-Z and neo-W gametologs, and 3) lower GC content on neo-W than neo-Z gametologs. Phylogenetic reconstruction of eight neo-Z and neo-W gametologs suggests that recombination continued after the split of Alaudidae from the rest of the Sylvioidea lineages (i.e., after ~42.2 Ma) and with some exceptions also after the split of Acrocephalidea and Timaliidae (i.e., after ~39.4 Ma). The Sylvioidea neo-sex chromosome shares classical evolutionary features with the ancestral sex chromosomes but, as expected from its more recent origin, shows weaker divergence between gametologs.

Description: In bacteria, physiological change may be effected by a single gene acquisition, producing ecological differentiation without genetic isolation. Natural selection acting on such differences can reduce the frequency of genotypes that arise from recombination at these loci. However, gene acquisition can only account for recombination interference in the fraction of the genome that is tightly linked to the integration site. To identify additional loci that contribute to adaptive differences, we examined orthologous genes in species of Enterobacteriaceae to identify significant differences in the degree of codon selection. Significance was assessed using the Adaptive Codon Enrichment metric, which accounts for the variation in codon usage bias that is expected to arise from mutation and drift; large differences in codon usage bias were identified in more genes than would be expected to arise from stochastic processes alone. Genes in the same operon showed parallel differences in codon usage bias, suggesting that changes in the overall levels of gene expression led to changes in the degree of adaptive codon usage. Most significant differences between orthologous operons were found among those involved with specific environmental adaptations, whereas "housekeeping" genes rarely showed significant changes. When considered together, the loci experiencing significant changes in codon selection outnumber potentially adaptive gene acquisition events. The identity of genes under strong codon selection seems to be influenced by the habitat from which the bacteria were isolated. We propose a two-stage model for how adaptation to different selective regimes can drive bacterial speciation. Initially, gene acquisitions catalyze rapid ecological differentiation, which modifies the utilization of genes, thereby changing the strength of codon selection on them. Alleles develop fitness variation by substitution, producing recombination interference at these loci in addition to those flanking acquired genes, allowing sequences to diverge across the entire genome and establishing genetic isolation (i.e., protection from frequent homologous recombination).

Description: Gene expression levels correlate with multiple aspects of gene sequence and gene structure in phylogenetically diverse taxa, suggesting an important role of gene expression levels in the evolution of protein-coding genes. Here we present results of a genome-wide study of the influence of gene expression on synonymous codon usage, amino acid composition, and gene structure in the red flour beetle, Tribolium castaneum . Consistent with the action of translational selection, we find that synonymous codon usage bias increases with gene expression. However, the correspondence between tRNA gene copy number and optimal codons is weak. At the amino acid level, translational selection is suggested by the positive correlation between tRNA gene numbers and amino acid usage, which is stronger for highly expressed genes. In addition, there is a clear trend for increased use of metabolically cheaper, less complex amino acids as gene expression increases. tRNA gene numbers also correlate negatively with amino acid size/complexity (S/C) score indicating the coupling between translational selection and selection to minimize the use of large/complex amino acids. Interestingly, the analysis of 10 additional genomes suggests that the correlation between tRNA gene numbers and amino acid S/C score is widespread and might be explained by selection against negative consequences of protein misfolding. At the level of gene structure, three major trends are detected: 1) complete coding region length increases across low and intermediate expression levels but decreases in highly expressed genes; 2) the average intron size shows the opposite trend, first decreasing with expression, followed by a slight increase in highly expressed genes; and 3) intron density remains nearly constant across all expression levels. These changes in gene architecture are only in partial agreement with selection favoring reduced cost of biosynthesis.

Description: New protein-coding genes can originate either through modification of existing genes or de novo. Recently, the importance of de novo origination has been recognized in eukaryotes, although eukaryotic genes originated de novo are relatively rare and difficult to identify. In contrast, viruses contain many de novo genes , namely those in which an existing gene has been "overprinted" by a new open reading frame, a process that generates a new protein-coding gene overlapping the ancestral gene. We analyzed the evolution of 12 experimentally validated viral genes that originated de novo and estimated their relative ages. We found that young de novo genes have a different codon usage from the rest of the genome. They evolve rapidly and are under positive or weak purifying selection. Thus, young de novo genes might have strain-specific functions, or no function, and would be difficult to detect using current genome annotation methods that rely on the sequence signature of purifying selection. In contrast to young de novo genes, older de novo genes have a codon usage that is similar to the rest of the genome. They evolve slowly and are under stronger purifying selection. Some of the oldest de novo genes evolve under stronger selection pressure than the ancestral gene they overlap, suggesting an evolutionary tug of war between the ancestral and the de novo gene.

Description: Here, we present a study of the molecular evolution of the pheromone receptor genes ( pre-1 and pre-2 ) in Neurospora taxa with different mating systems. We focus on comparisons between heterothallic and homothallic taxa, reproducing sexually by outcrossing and by intrahaploid selfing, respectively. Our general aim was to use a phylogenetic framework to investigate whether the evolutionary trajectory of the pheromone and receptor genes in Neurospora differs between heterothallic and homothallic taxa, and among the homothallic lineages/clades previously indicated to represent independent switches from heterothallism to homothallism in the evolutionary history of the genus. We complemented molecular evolution analyses with an expression study of the pre genes and their upstream regulators, the mating-type ( mat ) genes, in homothallic taxa. Our analyses suggest that the pheromone receptor gene pre-1 is functionally conserved in both heterothallic and homothallic taxa. Moreover, we found evidence of positive selection for a small fraction of codons in the cytoplasmic signal-transducing C-terminal region of the protein PRE-1. Distribution of positively selected codons differs between heterothallic and homothallic groups, suggesting functional divergence associated with mating system. The gene pre-2 was shown to evolve under high selective constraints, with no strong evidence for positive selection. Although our data suggest that both pre-1 and pre-2 are overall functional in homothallic taxa, individual taxa display frame-shift mutations causing premature stop codons, which might indicate loss of function. Transcriptional patterns of pre and mat genes in six homothallic taxa, selected to represent six different switches from heterothallism to homothallism, do not support a universal pattern of regulation of these genes during reproductive tissue development. Taken together, our analyses suggest that the pheromone receptor genes pre-1 and pre-2 are in general functional in homothallic Neurospora taxa, in contrast with the situation for the mat genes that are generally degenerate in these taxa.

Description: Comparative genome biology has unveiled the polyploid origin of all angiosperms and the role of recurrent polyploidization in the amplification of gene families and the structuring of genomes. Which species share certain ancient polyploidy events, and which do not, is ill defined because of the limited number of sequenced genomes and transcriptomes and their uneven phylogenetic distribution. Previously, it has been suggested that most, but probably not all, of the eudicots have shared an ancient hexaploidy event, referred to as the gamma triplication. In this study, detailed phylogenies of subfamilies of MADS-box genes suggest that the gamma triplication has occurred before the divergence of Gunnerales but after the divergence of Buxales and Trochodendrales. Large-scale phylogenetic and K S - based approaches on the inflorescence transcriptomes of Gunnera manicata (Gunnerales) and Pachysandra terminalis (Buxales) provide further support for this placement, enabling us to position the gamma triplication in the stem lineage of the core eudicots. This triplication likely initiated the functional diversification of key regulators of reproductive development in the core eudicots, comprising 75% of flowering plants. Although it is possible that the gamma event triggered early core eudicot diversification, our dating estimates suggest that the event occurred early in the stem lineage, well before the rapid speciation of the earliest core eudicot lineages. The evolutionary significance of this paleopolyploidy event may thus rather lie in establishing a species lineage that was resilient to extinction, but with the genomic potential for later diversification. We consider that the traits generated from this potential characterize extant core eudicots both chemically and morphologically.

Description: Genome reduction in obligately intracellular bacteria is one of the most well-established patterns in the field of molecular evolution. In the extreme, many sap-feeding insects harbor nutritional symbionts with genomes that are so reduced that it is not clear how they perform basic cellular functions. For example, the primary symbiont of psyllids ( Carsonella ) maintains one of the smallest and most AT-rich bacterial genomes ever identified and has surprisingly lost many genes that are thought to be essential for its role in provisioning its host with amino acids. However, our understanding of this extreme case of genome reduction is limited, as genomic data for Carsonella are available from only a single host species, and little is known about the functional role of "secondary" bacterial symbionts in psyllids. To address these limitations, we analyzed complete Carsonella genomes from pairs of congeneric hosts in three divergent genera within the Psyllidae ( Ctenarytaina , Heteropsylla , and Pachypsylla ) as well as complete secondary symbiont genomes from two of these host species ( Ctenarytaina eucalypti and Heteropsylla cubana ). Although the Carsonella genomes are generally conserved in size, structure, and GC content and exhibit genome-wide signatures of purifying selection, we found that gene loss has remained active since the divergence of the host species and had a particularly large impact on the amino acid biosynthesis pathways that define the symbiotic role of Carsonella . In some cases, the presence of additional bacterial symbionts may compensate for gene loss in Carsonella , as functional gene content indicates a high degree of metabolic complementarity between co-occurring symbionts. The genomes of the secondary symbionts also show signatures of long-term evolution as vertically transmitted, intracellular bacteria, including more extensive genome reduction than typically observed in facultative symbionts. Therefore, a history of co-evolution with secondary bacterial symbionts can partially explain the ongoing genome reduction in Carsonella . However, the absence of these secondary symbionts in other host lineages indicates that the relationships are dynamic and that other mechanisms, such as changes in host diet or functional coordination with the host genome, must also be at play.

Description: Gene duplication is a major driver of organismal adaptation and evolution and plays an important role in multiple human diseases. Whole-genome analyses have shown similar and high rates of gene duplication across a variety of eukaryotic species. Most of these studies, however, did not address the possible impact of interlocus gene conversion (IGC) on the evolution of gene duplicates. Because IGC homogenizes pairs of duplicates, widespread conversion would cause gene duplication events that happened long ago to appear more recent, resulting in artificially high estimates of duplication rates. Although the majority of genome-wide studies (including in the budding yeast Saccharomyces cerevisiae [Scer]) point to levels of IGC between paralogs ranging from 2% to 18%, Gao and Innan (Gao LZ, Innan H. 2004. Very low gene duplication rate in the yeast genome. Science 306:1367–1370.) found that gene conversion in yeast affected 〉80% of paralog pairs. If conversion rates really are this high, it would imply that the rate of gene duplication in eukaryotes is much lower than previously reported. In this work, we apply four different methodologies—including one approach that closely mirrors Gao and Innan’s method—to estimate the level of IGC in Scer. Our analyses point to a maximum conversion level of 13% between paralogs in this species, in close agreement with most estimates of IGC in eukaryotes. We also show that the exceedingly high levels of conversion found previously derive from application of an accurate method to an inappropriate data set. In conclusion, our work provides the most striking evidence to date supporting the reduced incidence of IGC among Scer paralogs and sets up a framework for future analyses in other eukaryotes.

Description: Cyclin B3 evolution has the unique peculiarity of an abrupt 3-fold increase of the protein size in the mammalian lineage due to the extension of a single exon. We have analyzed the evolution of the gene to define the modalities of this event and the possible consequences on the function of the protein. Database searches can trace the appearance of the gene to the origin of metazoans. Most introns were already present in early metazoans, and the intron–exon structure as well as the protein size were fairly conserved in invertebrates and nonmammalian vertebrates. Although intron gains are considered as rare events, we identified two cases, one at the prochordate–chordate transition and one in murids, resulting from different mechanisms. At the emergence of mammals, the gene was relocated from chromosome 6 of platypus to the X chromosome in marsupials, but the exon extension occurred only in placental mammals. A repetitive structure of 18 amino acids, of uncertain origin, is detectable in the 3,000-nt mammalian exon-encoded sequence, suggesting an extension by multiple internal duplications, some of which are still detectable in the primate lineage. Structure prediction programs suggest that the repetitive structure has no associated three-dimensional structure but rather a tendency for disorder. Splice variant isoforms were detected in several mammalian species but without conserved pattern, notably excluding the constant coexistence of premammalian-like transcripts, without the extension. The yeast two-hybrid method revealed that, in human, the extension allowed new interactions with ten unrelated proteins, most of them with specific three-dimensional structures involved in protein–protein interactions, and some highly expressed in testis, as is cyclin B3. The interactions with activator of cAMP-responsive element modulator in testis (ACT), germ cell-less homolog 1, and chromosome 1 open reading frame 14 remain to be verified in vivo since they may not be expressed at the same stages of spermatogenesis as cyclin B3.

Description: The X chromosome has a large effect on hybrid dysfunction, particularly on hybrid male sterility. Although the evidence for this so-called large-X effect is clear, its molecular causes are not yet fully understood. One possibility is that, under certain conditions, evolution proceeds faster in X-linked than in autosomal loci (i.e., faster-X effect) due to both natural selection and their hemizygosity in males, an effect that is expected to be greatest in genes with male-biased expression. Here, I study genome-wide variation in transcript abundance between Drosophila yakuba and D. santomea , within these species and in their hybrid males to evaluate both the faster-X and large-X effects at the level of expression. I find that in X-linked male-biased genes (MBGs) expression evolves faster than in their autosomal counterparts, an effect that is accompanied by a unique reduction in expression polymorphism. This suggests that Darwinian selection is driving expression differences between species, likely enhanced by the hemizygosity of the X chromosome in males. Despite the recent split of the two sister species under study, abundant changes in both cis - and trans -regulatory elements underlie expression divergence in the majority of the genes analyzed, with significant differences in allelic ratios of transcript abundance between the two reciprocal F 1 hybrid males. Cis–trans coevolution at molecular level, evolved shortly after populations become isolated, may therefore contribute to explain the breakdown of the regulation of gene expression in hybrid males. Additionally, the X chromosome plays a large role in this hybrid male misexpression, which affects not only MBG but also, to a lesser degree, nonsex-biased genes. Interestingly, hybrid male misexpression is concentrated mostly in autosomal genes, likely facilitated by the rapid evolution of sex-linked trans -acting factors. I suggest that the faster evolution of X-linked MBGs, at both protein and expression levels, contributes to explain the large effect of the X chromosome on hybrid male sterility, likely mediating widespread autosomal misexpression through the preferential recognition of cis -regulatory elements by conspecific trans -acting factors (i.e., cis–trans conspecific recognition).

Description: The sex chromosomes of the tropical crop papaya ( Carica papaya ) are evolutionarily young and consequently allow for the examination of evolutionary mechanisms that drive early sex chromosome divergence. We conducted a molecular population genetic analysis of four X/Y gene pairs from a collection of 45 wild papaya accessions. These population genetic analyses reveal striking differences in the patterns of polymorphism between the X and Y chromosomes that distinguish them from other sex chromosome systems. In most sex chromosome systems, the Y chromosome displays significantly reduced polymorphism levels, whereas the X chromosome maintains a level of polymorphism that is comparable to autosomal loci. However, the four papaya sex-linked loci that we examined display diversity patterns that are opposite this trend: the papaya X alleles exhibit significantly reduced polymorphism levels, whereas the papaya Y alleles maintain greater than expected levels of diversity. Our analyses suggest that selective sweeps in the regions of the X have contributed to this pattern while also revealing geographically restricted haplogroups on the Y. We discuss the possible role sexual selection and/or genomic conflict have played in shaping the contrasting patterns of polymorphism found for the papaya X and Y chromosomes.

Description: The HLA region shows diversity concerning the number and content of DRB genes present per haplotype. Similar observations are made for the equivalent regions in other primate species. To elucidate the evolutionary history of the various HLA-DRB genes, a large panel of intron sequences obtained from humans, chimpanzees, rhesus macaques, and common marmosets has been subjected to phylogenetic analyses. Special attention was paid to the presence and absence of particular transposable elements and/or to their segments. The sharing of different parts of the same long interspersed nuclear element-2 (LINE2, L2) and various Alu insertions by the species studied demonstrates that one precursor gene must have been duplicated several times before the Old World monkey (OWM) and hominid (HOM) divergence. At least four ancestral DRB gene families appear to have been present before the radiation of OWM and HOM, and one of these even predates the speciation of Old and New World primates. Two of these families represent the pseudogenes DRB6/DRB2 and DRB7 , which have been locked in the genomes of various primate species over long evolutionary time spans. Furthermore, all phylogenies of different intron segments show consistently that, apart from the pseudogenes, only DRB5 genes are shared by OWM and HOM, and they demonstrate the common history of certain DRB genes/lineages of humans and chimpanzees. In contrast, the evolutionary history of some other DRB loci is difficult to decipher, thus illustrating the complex history of the evolution of DRB genes due to a combination of mutations and recombination-like events. The selected approach allowed us to shed light on the ancestral DRB gene pool in primates and on the evolutionary relationship of the various HLA-DRB genes.

Description: Chemosensory genes are frequently the target of positive selection and are often present in large gene families, but little is known about heterogeneity of selection in these cases and its relation to function. Here, we use the vomeronasal-1 receptor ( V1R ) repertoire of mouse lemurs ( Microcebus spp.) as a model system to study patterns of selection of chemosensory genes at several different levels. Mouse lemurs are small nocturnal strepsirrhine primates and have a large (~200 loci) repertoire of V1R loci that are likely important for intraspecific pheromonal communication and interspecific interactions, for example, recognition of predator cues. We investigated signals and patterns of positive selection among the 105 identified full length V1R loci in the gray mouse lemur and within 7 V1R loci amplified across multiple mouse lemur species. Phylogenetic reconstructions of published sequences revealed at least nine monophyletic clusters of V1R s in gray mouse lemurs that have diversified since the split between lemurs and lorisoid primates. A large majority of clusters evolved under significant positive selection. Similar results were found in V1R s of closely related greater galagos. Comparison with function of related V1R clusters in mice suggested a potential relationship between receptor function and strength of selection. Interestingly, most codons identified as being under positive selection are located in the extracellular domains of the receptors and hence likely indicate the position of residues involved in ligand binding. Positive selection was also detected within five V1R loci (=71% of analyzed loci) sequenced from 6 to 10 mouse lemur species, indicating ongoing selection within the genus, which may be related to sexual selection and, potentially, speciation processes. Variation in strength of positive selection on V1R s showed no simple relationship to cluster size. The diversity of V1R loci in mouse lemurs reflects their adaptive evolution and is most likely related to the fundamental relevance of olfactory communication and predator recognition in these primates. Overall, adaptive evolution is the predominant mode of evolution of V1R loci at all levels, and the substantial heterogeneity in the strength of selection may be related to receptor function.

Description: The large number of sexually transmitted diseases and ocular trachoma cases that are caused globally each year by Chlamydia trachomatis has made this organism a World Health Organization priority for vaccine development. However, there is no gene transfer system for Chlamydia to help identify potential vaccine targets. To accelerate discoveries toward this goal, here we analyzed the broadest diversity of C. trachomatis genomes to date, including 25 geographically dispersed clinical and seven reference strains representing 14 of the 19 known serotypes. Strikingly, all 32 genomes were found to have evidence of DNA acquisition by homologous recombination in their history. Four distinct clades were identified, which correspond to all C. trachomatis disease phenotypes: lymphogranuloma venereum (LGV; Clade 1); noninvasive urogenital infections (Clade 2); ocular trachoma (Clade 3); and protocolitis (Clade 4; also includes some noninvasive urogenital infections). Although the ancestral relationship between clades varied, most strains acted as donor and recipient of recombination with no evidence for barriers to genetic exchange. The niche-specific LGV and trachoma clades have undergone less recombination, although the opportunity for mixing with strains from other clades that infect the rectal and ocular mucosa, respectively, is evident. Furthermore, there are numerous occasions for gene conversion events through sequential infections at the same anatomic sites. The size of recombinant segments is relatively small (~357 bp) compared with in vitro experiments of various C. trachomatis strains but is consistent with in vitro estimates for other bacterial species including Escherichia coli and Helicobacter pylori . Selection has also played a crucial role during the diversification of the organism. Clade 2 had the lowest nonsynonymous to synonymous ratio (d N /d S ) but the highest effect of recombination, which is consistent with the widespread occurrence of synonymous substitutions in recombined genomic segments. The trachoma Clade 3 had the highest d N /d S estimates, which may be caused by an increased effect of genetic drift from niche specialization and a reduced effective population size. The degree of drift, selection, and recombination in C. trachomatis suggests that the challenge will remain to identify genomic regions that are stable and cross protective for the development of an efficacious vaccine.

Description: We surveyed genetic variation in alr2 , an allodeterminant of the colonial hydroid Hydractinia symbiolongicarpus . We generated cDNA from a sample of 239 Hydractinia colonies collected at Lighthouse Point, Connecticut, and identified 473 alr2 alleles, 198 of which were unique. Rarefaction analysis suggested that the sample was near saturation. Most alleles were rare, with 86% occurring at frequencies of 1% or less. Alleles were highly variable, diverging on average by 18% of the amino acids in a predicted extracellular domain of the molecule. Analysis of 152 full-length alleles confirmed the existence of two structural types, defined by exons 4–8 of the gene. Several residues of the predicted immunoglobulin superfamily-like domains display signatures of positive selection. We also identified 77 unique alr2 pseudogene sequences from 85 colonies. Twenty-seven of these sequences matched expressed alr2 sequences from other colonies. This observation is consistent with pseudogenes contributing to alr2 diversification through sequence donation. A more limited collection of animals was made from a distant, relict population of H. symbiolongicarpus . Sixty percent of the unique sequences identified in this sample were found to match sequences from the Lighthouse Point population. The large number of alr2 alleles, their degree of divergence, the predominance of rare alleles in the population, their persistence over broad spatial and temporal scales, and the signatures of positive selection in multiple residues of the putative recognition domain paint a consistent picture of negative-frequency-dependent selection operating in this system. The genetic diversity observed at alr2 is comparable to that of the most highly polymorphic genetic systems known to date.

Description: The mechanosensory lateral line, found only in fishes and amphibians, is an important sense organ associated with aquatic life. Lateral line patterns differ among teleost, the most diverse vertebrate taxa, hypothetically in response to selective pressures from different aquatic habitats. In this article, we conduct evolutionary genomic analyses of 34 genes associated with lateral line system development in teleosts to elucidate the significance of contrasting evolutionary rates and changes in the protein coding sequences. We find that duplicated copies of these genes are preferentially retained in the teleost genomes and that episodic events of positive selection have occurred in 22 of the 30 postduplication branches. In general, teleost genes evolved at a faster rate relative to their tetrapod counterparts, and the mutation rates of 26 of the 34 genes differed among teleosts and tetrapods. We conclude that following whole genome duplication, evolutionary rates and episodic events of positive selection on the lateral line system development genes might have been one of the factors favoring the subsequent adaptive radiation of teleosts into diverse habitats. These results provide the foundation for further detailed explorations into lateral line system genes and the evolution of diverse phenotypes and adaptations.

Description: In any comparative studies striving to understand the similarities and differences of the living organisms at the molecular genetic level, the crucial first step is to establish the homology (orthology and paralogy) of genes between different organisms. Determination of the homology of genes becomes complicated when the genes have undergone a rapid divergence in sequence or when the involved genes are members of a gene family that has experienced a differential gain or loss of its constituents in different taxonomic groups. Organisms with duplicated genomes such as teleost fishes might have been especially prone to these problems because the functional redundancies provided by the duplicate copies of genes would have allowed a rapid divergence or loss of genes during evolution. In this study, we will demonstrate that much of the ambiguities in the determination of the homology between fish and tetrapod genes resulting from the problems like these can be eliminated by complementing the sequence-based phylogenies with nonsequence information, such as the exon–intron structure of a gene or the composition of a gene’s genomic neighbors. We will use the Tbx6/16 subfamily genes of zebrafish ( tbx6 , tbx16 , tbx24 , and mga genes), which have been well known for the ambiguity of their evolutionary relationships to the Tbx6/16 subfamily genes of tetrapods, as an illustrative example. We will show that, despite the similarity of sequence and expression to the tetrapod Tbx6 genes, zebrafish tbx6 gene is actually a novel T-box gene more closely related to the tetrapod Tbx16 genes, whereas the zebrafish tbx24 gene, hitherto considered to be a novel gene due to the high level of sequence divergence, is actually an ortholog of tetrapod Tbx6 genes. We will also show that, after their initial appearance by the multiplication of a common ancestral gene at the beginning of vertebrate evolution, the Tbx6/16 subfamily of vertebrate T-box genes might have experienced differential losses of member genes in different vertebrate groups and gradual pooling of member gene’s functions in surviving members, which might have prevented the revelation of the true identity of member genes by way of the comparison of sequence and function.