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Background: The shift from solitary to social behavior is one of the major evolutionary transitions. Primitively eusocial bumblebees are uniquely placed to illuminate the evolution of highly eusocial insect societies. Bumblebees are also invaluable natural and agricultural pollinators, and there is widespread concern over recent population declines in some species. High-quality

Background: The shift from solitary to social behavior is one of the major evolutionary transitions. Primitively eusocial bumblebees are uniquely placed to illuminate the evolution of highly eusocial insect societies. Bumblebees are also invaluable natural and agricultural pollinators, and there is widespread concern over recent population declines in some species. High-quality genomic data will inform key aspects of bumblebee biology, including susceptibility to implicated population viability threats.

Results: We report the high quality draft genome sequences of Bombus terrestris and Bombus impatiens, two ecologically dominant bumblebees and widely utilized study species. Comparing these new genomes to those of the highly eusocial honeybee Apis mellifera and other Hymenoptera, we identify deeply conserved similarities, as well as novelties key to the biology of these organisms. Some honeybee genome features thought to underpin advanced eusociality are also present in bumblebees, indicating an earlier evolution in the bee lineage. Xenobiotic detoxification and immune genes are similarly depauperate in bumblebees and honeybees, and multiple categories of genes linked to social organization, including development and behavior, show high conservation. Key differences identified include a bias in bumblebee chemoreception towards gustation from olfaction, and striking differences in microRNAs, potentially responsible for gene regulation underlying social and other traits.

Conclusions: These two bumblebee genomes provide a foundation for post-genomic research on these key pollinators and insect societies. Overall, gene repertoires suggest that the route to advanced eusociality in bees was mediated by many small changes in many genes and processes, and not by notable expansion or depauperation.

ContributorsSadd, Ben M. (Author) / Barribeau, Seth M. (Author) / Bloch, Guy (Author) / de Graaf, Dirk C. (Author) / Dearden, Peter (Author) / Elsik, Christine G. (Author) / Gadau, Juergen (Author) / Grimmelikhuijzen, Cornelis J. P. (Author) / Hasselmann, Martin (Author) / Lozier, Jeffrey D. (Author) / Robertson, Hugh M. (Author) / Smagghe, Guy (Author) / Stolle, Eckart (Author) / Van Vaerenbergh, Matthias (Author) / Waterhouse, Robert M. (Author) / Bornberg-Bauer, Erich (Author) / Klasberg, Steffen (Author) / Bennett, Anna K. (Author) / Camara, Francisco (Author) / Guigo, Roderic (Author) / Hoff, Katharina (Author) / Mariotti, Marco (Author) / Munoz-Torres, Monica (Author) / Murphy, Terence (Author) / Santesmasses, Didac (Author) / Amdam, Gro (Author) / Beckers, Matthew (Author) / Beye, Martin (Author) / Biewer, Matthias (Author) / Bitondi, Marcia MG (Author) / Blaxter, Mark L. (Author) / Bourke, Andrew FG (Author) / Brown, Mark JF (Author) / Buechel, Severine D. (Author) / Cameron, Rossanah (Author) / Cappelle, Kaat (Author) / Carolan, James C. (Author) / Christiaens, Olivier (Author) / Ciborowski, Kate L. (Author) / Clarke, David F. (Author) / Colgan, Thomas J. (Author) / Collins, David H. (Author) / Cridge, Andrew G. (Author) / Dalmay, Tamas (Author) / Dreier, Stephanie (Author) / du Plessis, Louis (Author) / Duncan, Elizabeth (Author) / Erler, Silvio (Author) / Evans, Jay (Author) / Falcon, Talgo (Author) / Flores, Kevin (Author) / Freitas, Flavia CP (Author) / Fuchikawa, Taro (Author) / Gempe, Tanja (Author) / Hartfelder, Klaus (Author) / Hauser, Frank (Author) / Helbing, Sophie (Author) / Humann, Fernanda (Author) / Irvine, Frano (Author) / Jermiin, Lars S (Author) / Johnson, Claire E. (Author) / Johnson, Reed M (Author) / Jones, Andrew K. (Author) / Kadowaki, Tatsuhiko (Author) / Kidner, Jonathan H. (Author) / Koch, Vasco (Author) / Kohler, Arian (Author) / Kraus, F. Bernhard (Author) / Lattorff, H. Michael G. (Author) / Leask, Megan (Author) / Lockett, Gabrielle A. (Author) / Mallon, Eamonn B. (Author) / Marco Antonio, David S. (Author) / Marxer, Monika (Author) / Meeus, Ivan (Author) / Moritz, Robin FA (Author) / Nair, Ajay (Author) / Napflin, Kathrin (Author) / Nissen, Inga (Author) / Niu, Jinzhi (Author) / Nunes, Francis MF (Author) / Oakeshott, John G. (Author) / Osborne, Amy (Author) / Otte, Marianne (Author) / Pinheiro, Daniel G. (Author) / Rossie, Nina (Author) / Rueppell, Olav (Author) / Santos, Carolina G (Author) / Schmid-Hempel, Regula (Author) / Schmitt, Bjorn D. (Author) / Schulte, Christina (Author) / Simoes, Zila LP (Author) / Soares, Michelle PM (Author) / Swevers, Luc (Author) / Winnebeck, Eva C. (Author) / Wolschin, Florian (Author) / Yu, Na (Author) / Zdobnov, Evgeny M (Author) / Aqrawi, Peshtewani K (Author) / Blakenburg, Kerstin P (Author) / Coyle, Marcus (Author) / Francisco, Liezl (Author) / Hernandez, Alvaro G. (Author) / Holder, Michael (Author) / Hudson, Matthew E. (Author) / Jackson, LaRonda (Author) / Jayaseelan, Joy (Author) / Joshi, Vandita (Author) / Kovar, Christie (Author) / Lee, Sandra L. (Author) / Mata, Robert (Author) / Mathew, Tittu (Author) / Newsham, Irene F. (Author) / Ngo, Robin (Author) / Okwuonu, Geoffrey (Author) / Pham, Christopher (Author) / Pu, Ling-Ling (Author) / Saada, Nehad (Author) / Santibanez, Jireh (Author) / Simmons, DeNard (Author) / Thornton, Rebecca (Author) / Venkat, Aarti (Author) / Walden, Kimberly KO (Author) / Wu, Yuan-Qing (Author) / Debyser, Griet (Author) / Devreese, Bart (Author) / Asher, Claire (Author) / Blommaert, Julie (Author) / Chipman, Ariel D. (Author) / Chittka, Lars (Author) / Fouks, Bertrand (Author) / Liu, Jisheng (Author) / O'Neill, Meaghan P (Author) / Sumner, Seirian (Author) / Puiu, Daniela (Author) / Qu, Jiaxin (Author) / Salzberg, Steven L (Author) / Scherer, Steven E (Author) / Muzny, Donna M. (Author) / Richards, Stephen (Author) / Robinson, Gene E (Author) / Gibbs, Richard A. (Author) / Schmid-Hempel, Paul (Author) / Worley, Kim C (Author) / College of Liberal Arts and Sciences (Contributor)
Created2015-04-24
Description

We present a phylogeographic study of at least six reproductively isolated lineages of new world harvester ants within the Pogonomyrmex barbatus and P. rugosus species group. The genetic and geographic relationships within this clade are complex: Four of the identified lineages show genetic caste determination (GCD) and are divided into

We present a phylogeographic study of at least six reproductively isolated lineages of new world harvester ants within the Pogonomyrmex barbatus and P. rugosus species group. The genetic and geographic relationships within this clade are complex: Four of the identified lineages show genetic caste determination (GCD) and are divided into two pairs. Each pair has evolved under a mutualistic system that necessitates sympatry. These paired lineages are dependent upon one another because their GCD requires interlineage matings for the production of F1 hybrid workers, and intralineage matings are required to produce queens. This GCD system maintains genetic isolation among these interdependent lineages, while simultaneously requiring co-expansion and emigration as their distributions have changed over time. It has also been demonstrated that three of these four GCD lineages have undergone historical hybridization, but the narrower sampling range of previous studies has left questions on the hybrid parentage, breadth, and age of these groups. Thus, reconstructing the phylogenetic and geographic history of this group allows us to evaluate past insights and hypotheses and to plan future inquiries in a more complete historical biogeographic context. Using mitochondrial DNA sequences sampled across most of the morphospecies’ ranges in the U.S.A. and Mexico, we conducted a detailed phylogeographic study. Remarkably, our results indicate that one of the GCD lineage pairs has experienced a dramatic range expansion, despite the genetic load and fitness costs of the GCD system. Our analyses also reveal a complex pattern of vicariance and dispersal in Pogonomyrmex harvester ants that is largely concordant with models of late Miocene, Pliocene, and Pleistocene range shifts among various arid-adapted taxa in North America.

ContributorsMott, Brendon (Author) / Gadau, Juergen (Author) / Anderson, Kirk E. (Author) / College of Liberal Arts and Sciences (Contributor)
Created2015-07-01
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Description

The electronic band structure of MoS2, MoSe2, WS2, and WSe2, crystals has been studied at various hydrostatic pressures experimentally by photoreflectance (PR) spectroscopy and theoretically within the density functional theory (DFT). In the PR spectra direct optical transitions (A and B) have been clearly observed and pressure coefficients have been

The electronic band structure of MoS2, MoSe2, WS2, and WSe2, crystals has been studied at various hydrostatic pressures experimentally by photoreflectance (PR) spectroscopy and theoretically within the density functional theory (DFT). In the PR spectra direct optical transitions (A and B) have been clearly observed and pressure coefficients have been determined for these transitions to be: αA = 2.0 ± 0.1 and αB = 3.6 ± 0.1 meV/kbar for MoS2, αA = 2.3 ± 0.1 and αB = 4.0 ± 0.1 meV/kbar for MoSe2, αA = 2.6 ± 0.1 and αB = 4.1 ± 0.1 meV/kbar for WS2, αA = 3.4 ± 0.1 and αB = 5.0 ± 0.5 meV/kbar for WSe2. It has been found that these coefficients are in an excellent agreement with theoretical predictions. In addition, a comparative study of different computational DFT approaches has been performed and analyzed. For indirect gap the pressure coefficient have been determined theoretically to be −7.9, −5.51, −6.11, and −3.79, meV/kbar for MoS2, MoSe2, WS2, and WSe2, respectively. The negative values of this coefficients imply a narrowing of the fundamental band gap with the increase in hydrostatic pressure and a semiconductor to metal transition for MoS2, MoSe2, WS2, and WSe2, crystals at around 140, 180, 190, and 240 kbar, respectively.

ContributorsDybala, F. (Author) / Polak, M. P. (Author) / Kopaczek, J. (Author) / Scharoch, P. (Author) / Wu, Kedi (Author) / Tongay, Sefaattin (Author) / Kudrawiec, R. (Author) / Ira A. Fulton Schools of Engineering (Contributor)
Created2016-05-24
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Description

Binary transition metal dichalcogenide monolayers share common properties such as a direct optical bandgap, spin-orbit splittings of hundreds of meV, light–matter interaction dominated by robust excitons and coupled spin-valley states. Here we demonstrate spin-orbit-engineering in Mo[(1-x)]WxSe2 alloy monolayers for optoelectronics and applications based on spin- and valley-control. We probe the

Binary transition metal dichalcogenide monolayers share common properties such as a direct optical bandgap, spin-orbit splittings of hundreds of meV, light–matter interaction dominated by robust excitons and coupled spin-valley states. Here we demonstrate spin-orbit-engineering in Mo[(1-x)]WxSe2 alloy monolayers for optoelectronics and applications based on spin- and valley-control. We probe the impact of the tuning of the conduction band spin-orbit spin-splitting on the bright versus dark exciton population. For MoSe2 monolayers, the photoluminescence intensity decreases as a function of temperature by an order of magnitude (4–300 K), whereas for WSe2 we measure surprisingly an order of magnitude increase. The ternary material shows a trend between these two extreme behaviors. We also show a non-linear increase of the valley polarization as a function of tungsten concentration, where 40% tungsten incorporation is sufficient to achieve valley polarization as high as in binary WSe2.

ContributorsWang, Gang (Author) / Robert, Cedric (Author) / Tuna, Aslihan (Author) / Chen, Bin (Author) / Yang, Sijie (Author) / Alamdari, Sarah (Author) / Gerber, Iann C. (Author) / Amand, Thierry (Author) / Marie, Xavier (Author) / Tongay, Sefaattin (Author) / Urbaszek, Bernhard (Author) / Ira A. Fulton Schools of Engineering (Contributor)
Created2015-12-14
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Description

We present two-dimensional Mg(OH)2 sheets and their vertical heterojunctions with CVD-MoS2 for the first time as flexible 2D insulators with anomalous lattice vibration and chemical and physical properties. New hydrothermal crystal growth technique enabled isolation of environmentally stable monolayer Mg(OH)2 sheets. Raman spectroscopy and vibrational calculations reveal that the lattice

We present two-dimensional Mg(OH)2 sheets and their vertical heterojunctions with CVD-MoS2 for the first time as flexible 2D insulators with anomalous lattice vibration and chemical and physical properties. New hydrothermal crystal growth technique enabled isolation of environmentally stable monolayer Mg(OH)2 sheets. Raman spectroscopy and vibrational calculations reveal that the lattice vibrations of Mg(OH)2 have fundamentally different signature peaks and dimensionality effects compared to other 2D material systems known to date. Sub-wavelength electron energy-loss spectroscopy measurements and theoretical calculations show that Mg(OH)2 is a 6 eV direct-gap insulator in 2D, and its optical band gap displays strong band renormalization effects from monolayer to bulk, marking the first experimental confirmation of confinement effects in 2D insulators. Interestingly, 2D-Mg(OH)2 sheets possess rather strong surface polarization (charge) effects which is in contrast to electrically neutral h-BN materials. Using 2D-Mg(OH)2 sheets together with CVD-MoS2 in the vertical stacking shows that a strong change transfer occurs from n-doped CVD-MoS2 sheets to Mg(OH)2, naturally depleting the semiconductor, pushing towards intrinsic doping limit and enhancing overall optical performance of 2D semiconductors. Results not only establish unusual confinement effects in 2D-Mg(OH)2, but also offer novel 2D-insulating material with unique physical, vibrational, and chemical properties for potential applications in flexible optoelectronics.

ContributorsTuna, Aslihan (Author) / Wu, Kedi (Author) / Sahin, Hasan (Author) / Chen, Bin (Author) / Yang, Sijie (Author) / Cai, Hui (Author) / Aoki, Toshihiro (Author) / Horzum, Seyda (Author) / Kang, Jun (Author) / Peeters, Francois M. (Author) / Tongay, Sefaattin (Author) / Ira A. Fulton Schools of Engineering (Contributor)
Created2016-02-05
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Description

Transition metal trichalcogenides form a class of layered materials with strong in-plane anisotropy. For example, titanium trisulfide (TiS3) whiskers are made out of weakly interacting TiS3 layers, where each layer is made of weakly interacting quasi-one-dimensional chains extending along the b axis. Here we establish the unusual vibrational properties of

Transition metal trichalcogenides form a class of layered materials with strong in-plane anisotropy. For example, titanium trisulfide (TiS3) whiskers are made out of weakly interacting TiS3 layers, where each layer is made of weakly interacting quasi-one-dimensional chains extending along the b axis. Here we establish the unusual vibrational properties of TiS3 both experimentally and theoretically. Unlike other two-dimensional systems, the Raman active peaks of TiS3 have only out-of-plane vibrational modes, and interestingly some of these vibrations involve unique rigid-chain vibrations and S–S molecular oscillations. High-pressure Raman studies further reveal that the AgS-S S-S molecular mode has an unconventional negative pressure dependence, whereas other peaks stiffen as anticipated. Various vibrational modes are doubly degenerate at ambient pressure, but the degeneracy is lifted at high pressures. These results establish the unusual vibrational properties of TiS3 with strong in-plane anisotropy, and may have relevance to understanding of vibrational properties in other anisotropic two-dimensional material systems.

ContributorsWu, Kedi (Author) / Torun, Engin (Author) / Sahin, Hasan (Author) / Chen, Bin (Author) / Fan, Xi (Author) / Pant, Anupum (Author) / Wright, David (Author) / Aoki, Toshihiro (Author) / Peeters, Francois M. (Author) / Soignard, Emmanuel (Author) / Tongay, Sefaattin (Author) / Ira A. Fulton Schools of Engineering (Contributor)
Created2016-09-22
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Description

A central goal of biology is to uncover the genetic basis for the origin of new phenotypes. A particularly effective approach is to examine the genomic architecture of species that have secondarily lost a phenotype with respect to their close relatives. In the eusocial Hymenoptera, queens and workers have divergent

A central goal of biology is to uncover the genetic basis for the origin of new phenotypes. A particularly effective approach is to examine the genomic architecture of species that have secondarily lost a phenotype with respect to their close relatives. In the eusocial Hymenoptera, queens and workers have divergent phenotypes that may be produced via either expression of alternative sets of caste-specific genes and pathways or differences in expression patterns of a shared set of multifunctional genes. To distinguish between these two hypotheses, we investigated how secondary loss of the worker phenotype in workerless ant social parasites impacted genome evolution across two independent origins of social parasitism in the ant genera Pogonomyrmex and Vollenhovia. We sequenced the genomes of three social parasites and their most-closely related eusocial host species and compared gene losses in social parasites with gene expression differences between host queens and workers. Virtually all annotated genes were expressed to some degree in both castes of the host, with most shifting in queen-worker bias across developmental stages. As a result, despite >1 My of divergence from the last common ancestor that had workers, the social parasites showed strikingly little evidence of gene loss, damaging mutations, or shifts in selection regime resulting from loss of the worker caste. This suggests that regulatory changes within a multifunctional genome, rather than sequence differences, have played a predominant role in the evolution of social parasitism, and perhaps also in the many gains and losses of phenotypes in the social insects.

ContributorsSmith, Chris R. (Author) / Helms Cahan, Sara (Author) / Kemena, Carsten (Author) / Brady, Sean G. (Author) / Yang, Wei (Author) / Bornberg-Bauer, Erich (Author) / Eriksson, Ti (Author) / Gadau, Juergen (Author) / Helmkampf, Martin (Author) / Gotzek, Dietrich (Author) / Okamoto Miyakawa, Misato (Author) / Suarez, Andrew V. (Author) / Mikheyev, Alexander (Author) / College of Liberal Arts and Sciences (Contributor)
Created2015-07-29
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Description

Black phosphorus attracts enormous attention as a promising layered material for electronic, optoelectronic and thermoelectric applications. Here we report large anisotropy in in-plane thermal conductivity of single-crystal black phosphorus nanoribbons along the zigzag and armchair lattice directions at variable temperatures. Thermal conductivity measurements were carried out under the condition of

Black phosphorus attracts enormous attention as a promising layered material for electronic, optoelectronic and thermoelectric applications. Here we report large anisotropy in in-plane thermal conductivity of single-crystal black phosphorus nanoribbons along the zigzag and armchair lattice directions at variable temperatures. Thermal conductivity measurements were carried out under the condition of steady-state longitudinal heat flow using suspended-pad micro-devices. We discovered increasing thermal conductivity anisotropy, up to a factor of two, with temperatures above 100 K. A size effect in thermal conductivity was also observed in which thinner nanoribbons show lower thermal conductivity. Analysed with the relaxation time approximation model using phonon dispersions obtained based on density function perturbation theory, the high anisotropy is attributed mainly to direction-dependent phonon dispersion and partially to phonon–phonon scattering. Our results revealing the intrinsic, orientation-dependent thermal conductivity of black phosphorus are useful for designing devices, as well as understanding fundamental physical properties of layered materials.

ContributorsLee, Sangwook (Author) / Yang, Fan (Author) / Suh, Joonki (Author) / Yang, Sijie (Author) / Lee, Yeonbae (Author) / Li, Guo (Author) / Choe, Hwan Sung (Author) / Tuna, Aslihan (Author) / Chen, Yabin (Author) / Ko, Changhyun (Author) / Park, Joonsuk (Author) / Liu, Kai (Author) / Li, Jingbo (Author) / Hippalgaonkar, Kedar (Author) / Urban, Jeffrey J. (Author) / Tongay, Sefaattin (Author) / Wu, Junqiao (Author) / Ira A. Fulton Schools of Engineering (Contributor)
Created2015-10-16
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Description

Adaptation requires genetic variation, but founder populations are generally genetically depleted. Here we sequence two populations of an inbred ant that diverge in phenotype to determine how variability is generated. Cardiocondyla obscurior has the smallest of the sequenced ant genomes and its structure suggests a fundamental role of transposable elements

Adaptation requires genetic variation, but founder populations are generally genetically depleted. Here we sequence two populations of an inbred ant that diverge in phenotype to determine how variability is generated. Cardiocondyla obscurior has the smallest of the sequenced ant genomes and its structure suggests a fundamental role of transposable elements (TEs) in adaptive evolution. Accumulations of TEs (TE islands) comprising 7.18% of the genome evolve faster than other regions with regard to single-nucleotide variants, gene/exon duplications and deletions and gene homology. A non-random distribution of gene families, larvae/adult specific gene expression and signs of differential methylation in TE islands indicate intragenomic differences in regulation, evolutionary rates and coalescent effective population size. Our study reveals a tripartite interplay between TEs, life history and adaptation in an invasive species.

ContributorsSchrader, Lukas (Author) / Kim, Jay W. (Author) / Ence, Daniel (Author) / Zimin, Aleksey (Author) / Klein, Antonia (Author) / Wyschetzki, Katharina (Author) / Weichselgartner, Tobias (Author) / Kemena, Carsten (Author) / Stoekl, Johannes (Author) / Schultner, Eva (Author) / Wurm, Yannick (Author) / Smith, Christopher D. (Author) / Yandell, Mark (Author) / Heinze, Juergen (Author) / Gadau, Juergen (Author) / Oettler, Jan (Author) / College of Liberal Arts and Sciences (Contributor)
Created2014-12-01
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Description

Plasmonic and metamaterial based nano/micro-structured materials enable spectrally selective resonant absorption, where the resonant bandwidth and absorption intensity can be engineered by controlling the size and geometry of nanostructures. Here, we demonstrate a simple, lithography-free approach for obtaining a resonant and dynamically tunable broadband absorber based on vanadium dioxide (VO2)

Plasmonic and metamaterial based nano/micro-structured materials enable spectrally selective resonant absorption, where the resonant bandwidth and absorption intensity can be engineered by controlling the size and geometry of nanostructures. Here, we demonstrate a simple, lithography-free approach for obtaining a resonant and dynamically tunable broadband absorber based on vanadium dioxide (VO2) phase transition. Using planar layered thin film structures, where top layer is chosen to be an ultrathin (20 nm) VO2 film, we demonstrate broadband IR light absorption tuning (from similar to 90% to similar to 30% in measured absorption) over the entire mid-wavelength infrared spectrum. Our numerical and experimental results indicate that the bandwidth of the absorption bands can be controlled by changing the dielectric spacer layer thickness. Broadband tunable absorbers can find applications in absorption filters, thermal emitters, thermophotovoltaics, and sensing.

ContributorsKocer, Hasan (Author) / Butun, Serkan (Author) / Palacios, Edgar (Author) / Liu, Zizhuo (Author) / Tongay, Sefaattin (Author) / Fu, Deyi (Author) / Wang, Kevin (Author) / Wu, Junqiao (Author) / Aydin, Koray (Author) / Ira A. Fulton Schools of Engineering (Contributor)
Created2015-08-21