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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 development of non-volatile logic through direct coupling of spontaneous ferroelectric polarization with semiconductor charge carriers is nontrivial, with many issues, including epitaxial ferroelectric growth, demonstration of ferroelectric switching and measurable semiconductor modulation. Here we report a true ferroelectric field effect—carrier density modulation in an underlying Ge(001) substrate by switching

The development of non-volatile logic through direct coupling of spontaneous ferroelectric polarization with semiconductor charge carriers is nontrivial, with many issues, including epitaxial ferroelectric growth, demonstration of ferroelectric switching and measurable semiconductor modulation. Here we report a true ferroelectric field effect—carrier density modulation in an underlying Ge(001) substrate by switching of the ferroelectric polarization in epitaxial c-axis-oriented BaTiO3 grown by molecular beam epitaxy. Using the density functional theory, we demonstrate that switching of BaTiO3 polarization results in a large electric potential change in Ge. Aberration-corrected electron microscopy confirms BaTiO3 tetragonality and the absence of any low-permittivity interlayer at the interface with Ge. The non-volatile, switchable nature of the single-domain out-of-plane ferroelectric polarization of BaTiO3 is confirmed using piezoelectric force microscopy. The effect of the polarization switching on the conductivity of the underlying Ge is measured using microwave impedance microscopy, clearly demonstrating a ferroelectric field effect.

ContributorsPonath, Patrick (Author) / Fredrickson, Kurt (Author) / Posadas, Agham B. (Author) / Ren, Yuan (Author) / Wu, Xiaoyu (Author) / Vasudevan, Rama K. (Author) / Okatan, M. Baris (Author) / Jesse, S. (Author) / Aoki, Toshihiro (Author) / McCartney, Martha (Author) / Smith, David (Author) / Kalinin, Sergei V. (Author) / Lai, Keji (Author) / Demkov, Alexander A. (Author) / Ira A. Fulton Schools of Engineering (Contributor)
Created2015-01-01
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Description

In this paper, we report on the highly conductive layer formed at the crystalline γ-alumina/SrTiO3 interface, which is attributed to oxygen vacancies. We describe the structure of thin γ-alumina layers deposited by molecular beam epitaxy on SrTiO3 (001) at growth temperatures in the range of 400–800 °C, as determined by reflection-high-energy

In this paper, we report on the highly conductive layer formed at the crystalline γ-alumina/SrTiO3 interface, which is attributed to oxygen vacancies. We describe the structure of thin γ-alumina layers deposited by molecular beam epitaxy on SrTiO3 (001) at growth temperatures in the range of 400–800 °C, as determined by reflection-high-energy electron diffraction, x-ray diffraction, and high-resolution electron microscopy. In situ x-ray photoelectron spectroscopy was used to confirm the presence of the oxygen-deficient layer. Electrical characterization indicates sheet carrier densities of ∼1013 cm−2 at room temperature for the sample deposited at 700 °C, with a maximum electron Hall mobility of 3100 cm2V-1s-1 at 3.2 K and room temperature mobility of 22 cm2V-1s-1. Annealing in oxygen is found to reduce the carrier density and turn a conductive sample into an insulator.

ContributorsKormondy, Kristy J. (Author) / Posadas, Agham B. (Author) / Ngo, Thong Q. (Author) / Lu, Sirong (Author) / Goble, Nicholas (Author) / Jordan-Sweet, Jean (Author) / Gao, Xuan P. A. (Author) / Smith, David (Author) / McCartney, Martha (Author) / Ekerdt, John G. (Author) / Demkov, Alexander A. (Author) / College of Liberal Arts and Sciences (Contributor)
Created2015-03-07
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Description

This paper reports the molecular beam epitaxial growth and characterization of high-reflectivity and broad-bandwidth distributed Bragg reflectors (DBRs) made of ZnTe/GaSb quarter-wavelength (lambda/4) layers for optoelectronic applications in the midwave infrared spectral range (2-5 mu m). A series of ZnTe/GaSb DBRs has been successfully grown on GaSb (001) substrates using

This paper reports the molecular beam epitaxial growth and characterization of high-reflectivity and broad-bandwidth distributed Bragg reflectors (DBRs) made of ZnTe/GaSb quarter-wavelength (lambda/4) layers for optoelectronic applications in the midwave infrared spectral range (2-5 mu m). A series of ZnTe/GaSb DBRs has been successfully grown on GaSb (001) substrates using molecular beam epitaxy (MBE). During the MBE growth, a temperature ramp was applied to the initial growth of GaSb layers on ZnTe to protect the ZnTe underneath from damage due to thermal evaporation. Post-growth characterization using high-resolution x-ray diffraction, atomic force microscopy, and transmission electron microscopy reveals smooth surface morphology, low defect density, and coherent interfaces. Reflectance spectroscopy results show that a DBR sample of seven lambda/4 pairs has a peak reflectance as high as 99.0% centered at 2.56 mu m with a bandwidth of 517 nm.

ContributorsFan, Jin (Author) / Liu, Xinyu (Author) / Ouyang, Lu (Author) / Pimpinella, Richard E. (Author) / Dobrowolska, Margaret (Author) / Furdyna, Jacek K. (Author) / Smith, David (Author) / Zhang, Yong-Hang (Author) / Ira A. Fulton Schools of Engineering (Contributor)
Created2013-10-28
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Description

We report a study of the magnetic domain structure of nanocrystalline thin films of nickel-zinc ferrite. The ferrite films were synthesized using aqueous spin-spray coating at low temperature (∼90 °C) and showed high complex permeability in the GHz range. Electron microscopy and microanalysis revealed that the films consisted of columnar grains

We report a study of the magnetic domain structure of nanocrystalline thin films of nickel-zinc ferrite. The ferrite films were synthesized using aqueous spin-spray coating at low temperature (∼90 °C) and showed high complex permeability in the GHz range. Electron microscopy and microanalysis revealed that the films consisted of columnar grains with uniform chemical composition. Off-axis electron holography combined with magnetic force microscopy indicated a multi-grain domain structure with in-plane magnetization. The correlation between the magnetic domain morphology and crystal structure is briefly discussed.

ContributorsZhang, D. (Author) / Ray, N. M. (Author) / Petuskey, William (Author) / Smith, David (Author) / McCartney, Martha (Author) / Ira A. Fulton Schools of Engineering (Contributor)
Created2014-08-28
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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