Population structure and connectivity in Indo-Pacific deep-sea mussels of the Bathymodiolus septemdierum complex

Current pressures to mine polymetallic sulfide deposits pose threats to the animal communities found at deep-sea hydrothermal vents. Management plans aimed at preserving these unusual communities require knowledge of historical and contemporary forces that shaped the distributions and connectivity of associated species. As most vent research has focused on the eastern Pacific and mid-Atlantic ridge systems less is known about Indo-Pacific vents, where mineral extraction activities are imminent. Deep-sea mussels (Bivalvia: Mytilidae) of the genus Bathymodiolus include the morphotypic species B. septemdierum, B. brevior, B. marisindicus, and B. elongatus which are among the dominant vent taxa in western Pacific back-arc basins and the Central Indian Ridge. To assess their interpopulational relationships, we examined multilocus genotypes based on DNA sequences from four nuclear and four mitochondrial genes, and allozyme variation encoded by eleven genes. Bayesian assignment methods grouped mussels from seven widespread western Pacific localities into a single cluster, whereas the Indian Ocean mussels were clearly divergent. Thus, we designate two regional metapopulations. Notably, contemporary migration rates among all sites appeared to be low despite limited population differentiation, which highlights the necessity of obtaining realistic data on recovery times and fine-scale population structure to develop and manage conservation units effectively. Future studies using population genomic methods to address these issues in a range of species will help to inform management plans aimed at mitigating potential impacts of deep-sea mining in the Indo-Pacific region

DNA hydroxymethylation levels are altered in blood cells from Down syndrome persons enrolled in the MARK-AGE project

Down syndrome (DS) is caused by the presence of part or an entire extra copy of chromosome 21, a phenomenon that can cause a wide spectrum of clinically defined phenotypes of the disease. Most of the clinical signs of DS are typical of the ageing process including dysregulation of immune system. Beyond the causative genetic defect, DS persons display epigenetic alterations, particularly aberrant DNA methylation patterns that can contribute to the heterogeneity of the disease. In the present work we investigated the levels of 5-hydroxymethylcytosine (5hmC) and of the TET dioxygenase enzymes, which are involved in DNA demethylation processes and are often deregulated in pathological conditions as well as in ageing. Analyses were carried out on peripheral blood mononuclear cells of DS volunteers enrolled in the context of the MARK-AGE study, a large-scale cross-sectional population study with subjects representing the general population in eight European countries. We observed a decrease of 5hmC, TET1 and other components of the DNA methylation/demethylation machinery in DS subjects, indicating that aberrant DNA methylation patterns in DS, which may have consequences on the transcriptional status of immune cells, may be due to a global disturbance of methylation control in DS

Population connectivity and speciation of vent mussels from the Mid-Atlantic Ridge: An interdisciplinary approach

Knowledge about the genetic connectivity of populations and the mechanisms underlying speciation is an important requirement for biodiversity conservation. However, despite the fact that such information is generally sparse for deep sea biota, the demands for mining of seafloor mineral resources are increasing. Using mussels of the genus Bathymodiolus as a model system and integrating expertise from various research disciplines, the present study aims to elucidate patterns of geneflow between hydrothermal vent populations of the Mid-Atlantic Ridge. To characterize the level of genetic exchange on ecological time scales, we will genotype mussels with species- diagnostic SNP markers designed from high-throughput transcriptomic data. By performing hybridization experiments, we will study patterns of reproductive isolation between B. azoricus and B. childressi from shallower sites. Moreover, we will measure larval survival and behaviour of B. azoricus under a variety of environmental conditions to identify factors that influence the spreading of propagules between vents. The results from the ecophysiological experiments will be incorporated into larval dispersal studies performed with Lagrangian simulations utilizing a high-resolution model of the global ocean under realistic forcing. By comparing the outcomes of the genetic, biophysical and ecological approaches, we expect that this work will make fundamental contributions to a better understanding of population connectivity and diversification in the deep sea as well as help to manage anthropogenic threats to hydrothermal vent ecosystems

Age-dependent expression of DNMT1 and DNMT3B in PBMCs from a large European population enrolled in the MARK-AGE study

Aging is associated with alterations in the content and patterns of DNA methylation virtually throughout the entire human lifespan. Reasons for these variations are not well understood. However, several lines of evidence suggest that the epigenetic instability in aging may be traced back to the alteration of the expression of DNA methyltransferases. Here, the association of the expression of DNA methyltransferases DNMT1 and DNMT3B with age has been analysed in the context of the MARK-AGE study, a large-scale cross-sectional study of the European general population. Using peripheral blood mononuclear cells, we assessed the variation of DNMT1 and DNMT3B gene expression in more than two thousand age-stratified women and men (35-75 years) recruited across eight European countries. Significant age-related changes were detected for both transcripts. The level of DNMT1 gradually dropped with aging but this was only observed up to the age of 64 years. By contrast, the expression of DNMT3B decreased linearly with increasing age and this association was particularly evident in females. We next attempted to trace the age-related changes of both transcripts to the influence of different variables that have an impact on changes of their expression in the population, including demographics, dietary and health habits, and clinical parameters. Our results indicate that age affects the expression of DNMT1 and DNMT3B as an almost independent variable in respect of all other variables evaluated

Combined genotyping, microbial diversity and metabolite profiling studies on farmed Mytilus spp. from Kiel Fjord

The blue mussel Mytilus is a popular food source with high economical value. Species of the M. edulis complex (M. edulis, M. galloprovincialis and M. trossulus) hybridise whenever their geographic ranges overlap posing difficulties to species discrimination, which is important for blue mussel aquaculture. The aim of this study was to determine the genetic structure of farmed blue mussels in Kiel Fjord. Microbial and metabolic profile patterns were studied to investigate a possible dependency on the genotype of the bivalves. Genotyping confirmed the complex genetic structure of the Baltic Sea hybrid zone and revealed an unexpected dominance of M. trossulus alleles being in contrast to the predominance of M. edulis alleles described for wild Baltic blue mussels. Culture-dependent and -independent microbial community analyses indicated the presence of a diverse Mytilus-associated microbiota, while an LC-MS/MS-based metabolome study identified 76 major compounds dominated by pigments, alkaloids and polyketides in the whole tissue extracts. Analysis of mussel microbiota and metabolome did not indicate genotypic dependence, but demonstrated high intraspecific variability of farmed mussel individuals. We hypothesise that individual differences in microbial and metabolite patterns may be caused by high individual plasticity and might be enhanced by e.g. nutritional condition, age and gender

Population genetics and morphometric variation of blue mussels in the western Baltic Sea

The investigation of hybridizing taxa can provide intriguing insights into the process of speciation and the adaptive potential of populations to environmental change. Due to its gradual nature in terms of ecology and genetics, the European blue mussel hybrid zone between North Sea Mytilus edulis and Baltic M. trossulus is an ideal study system to analyse the significance of inter-specific hybridization in evolution. Although much research has already focussed on the genetic and phenotypic structures in the Baltic Proper, less is known about the situation in the western transition area. By means of a multi-locus genotypic assessment and multivariate morphometric analyses I have examined the genetic and morphological constitutions of several Baltic Mytilus populations. Paying special attention to the poorly resolved outer parts, I find that most blue mussels in the western Baltic Sea and Skagerrak are higher generation backcross hybrids (99 % in the Kiel Fjord; 81 % in Tjärnö). While my results suggest that gene variants of M. edulis still prevail over that of M. trossulus in these mytilids, they also demonstrate that a minority of individuals in the Kiel Fjord are introgressed by M. galloprovincialis alleles. Although the origin of these alien polymorphisms cannot be told from my data, I postulate that their movement into the Baltic gene pool is most likely human-mediated (e.g. ship traffic). Considering the increasing records of anthropogenic introductions of Mytilus species to non-native habitats and the limited power of the applied molecular markers to discriminate M. galloprovincialis, it is to expect that more cryptic invasions are detected in future, technically advanced investigations. My analysis of M. edulis allele frequency changes from the North Sea to the Baltic Proper extends and supports an earlier work by Stuckas et al. (2009), providing evidence for discordant patterns of gene flow across the hybrid zone. These discrepancies probably arise from the concerted action of direct selection, genetic hitchhiking, stochastic evolutionary forces and shifts in the geographical position of the secondary contact area. Different to this previous study and in line with the observations by Kijewski et al. (2006), I find that introgression into the central Baltic is restricted at least for some M. edulis maternal haplotypes (D-loop), indicative of cytonuclear incompatibilities between inner and outer Baltic mussels. While differential environmental pressures might contribute to the maintenance of semi-permeable genetic barriers between M. edulis-like and M. trossulus-like hybrid populations, they can also account for the observed dissimilarities in shell morphology and phenotypic diversity. In contrast to individuals of the Baltic Proper, which express a Abstract | 7 population-specific, M. trossulus-like morphotype, mussels of the transition zone build a morphological continuum between parental forms. Following the reasoning by Gardner (1996) I propose that the strong environmental variability in the western Baltic selects for multiple phenotypes, while the constant and extreme conditions (e.g. low salinities, absence of predators) in the eastern Baltic favour only a single shell morphotype. It remains to be shown whether these opposing patterns are consequences of environmentally-induced discrepancies in genetic variability at fixed causal loci (level of heterozygosity, selection for multiple alleles) and/or differences in phenotypic plasticity

Analysis of the machinery and intermediates of the 5hmC-mediated DNA demethylation pathway in aging on samples from the MARKAGE Study

Gradual changes in the DNA methylation landscape occur throughout aging virtually in all human tissues. A widespread reduction of 5-methylcytosine (5mC), associated with highly reproducible site-specific hypermethylation, characterizes the genome in aging. Therefore, an equilibrium seems to exist between general and directional deregulating events concerning DNA methylation controllers, which may underpin the age-related epigenetic changes. In this context, 5mC-hydroxylases (TET enzymes) are new potential players. In fact, TETs catalyze the stepwise oxidation of 5mC to 5-hydroxymethylcytosine (5hmC), 5-formylcytosine (5fC) and 5-carboxylcytosine (5caC), driving the DNA demethylation process based on thymine DNA glycosylase (TDG)-mediated DNA repair pathway. The present paper reports the expression of DNA hydroxymethylation components, the levels of 5hmC and of its derivatives in peripheral blood mononuclear cells of age-stratified donors recruited in several European countries in the context of the EU Project ‘MARK-AGE’. The results provide evidence for an age-related decline of TET1, TET3 and TDG gene expression along with a decrease of 5hmC and an accumulation of 5caC. These associations were independent of confounding variables, including recruitment center, gender and leukocyte composition. The observed impairment of 5hmC-mediated DNA demethylation pathway in blood cells may lead to aberrant transcriptional programs in the elderly

Competing Ultrafast Energy Relaxation Pathways in Photoexcited Graphene

For most optoelectronic applications of graphene a thorough understanding of the processes that govern energy relaxation of photoexcited carriers is essential. The ultrafast energy relaxation in graphene occurs through two competing pathways: carrier-carrier scattering -- creating an elevated carrier temperature -- and optical phonon emission. At present, it is not clear what determines the dominating relaxation pathway. Here we reach a unifying picture of the ultrafast energy relaxation by investigating the terahertz photoconductivity, while varying the Fermi energy, photon energy, and fluence over a wide range. We find that sufficiently low fluence ($\lesssim$ 4 $\mu$J/cm$^2$) in conjunction with sufficiently high Fermi energy ($\gtrsim$ 0.1 eV) gives rise to energy relaxation that is dominated by carrier-carrier scattering, which leads to efficient carrier heating. Upon increasing the fluence or decreasing the Fermi energy, the carrier heating efficiency decreases, presumably due to energy relaxation that becomes increasingly dominated by phonon emission. Carrier heating through carrier-carrier scattering accounts for the negative photoconductivity for doped graphene observed at terahertz frequencies. We present a simple model that reproduces the data for a wide range of Fermi levels and excitation energies, and allows us to qualitatively assess how the branching ratio between the two distinct relaxation pathways depends on excitation fluence and Fermi energy.Comment: Nano Letters 201

Generation of photovoltage in graphene on a femtosecond time scale through efficient carrier heating

Graphene is a promising material for ultrafast and broadband photodetection. Earlier studies addressed the general operation of graphene-based photo-thermoelectric devices, and the switching speed, which is limited by the charge carrier cooling time, on the order of picoseconds. However, the generation of the photovoltage could occur at a much faster time scale, as it is associated with the carrier heating time. Here, we measure the photovoltage generation time and find it to be faster than 50 femtoseconds. As a proof-of-principle application of this ultrafast photodetector, we use graphene to directly measure, electrically, the pulse duration of a sub-50 femtosecond laser pulse. The observation that carrier heating is ultrafast suggests that energy from absorbed photons can be efficiently transferred to carrier heat. To study this, we examine the spectral response and find a constant spectral responsivity between 500 and 1500 nm. This is consistent with efficient electron heating. These results are promising for ultrafast femtosecond and broadband photodetector applications.Comment: 6 pages, 4 figure

Tuning ultrafast electron thermalization pathways in a van der Waals heterostructure

Ultrafast electron thermalization - the process leading to Auger recombination, carrier multiplication via impact ionization and hot carrier luminescence - occurs when optically excited electrons in a material undergo rapid electron-electron scattering to redistribute excess energy and reach electronic thermal equilibrium. Due to extremely short time and length scales, the measurement and manipulation of electron thermalization in nanoscale devices remains challenging even with the most advanced ultrafast laser techniques. Here, we overcome this challenge by leveraging the atomic thinness of two-dimensional van der Waals (vdW) materials in order to introduce a highly tunable electron transfer pathway that directly competes with electron thermalization. We realize this scheme in a graphene-boron nitride-graphene (G-BN-G) vdW heterostructure, through which optically excited carriers are transported from one graphene layer to the other. By applying an interlayer bias voltage or varying the excitation photon energy, interlayer carrier transport can be controlled to occur faster or slower than the intralayer scattering events, thus effectively tuning the electron thermalization pathways in graphene. Our findings, which demonstrate a novel means to probe and directly modulate electron energy transport in nanoscale materials, represent an important step toward designing and implementing novel optoelectronic and energy-harvesting devices with tailored microscopic properties.Comment: Accepted to Nature Physic