Table of Contents

Biological Sciences

• Applied Biological Sciences

  • You have access

    Bioinspired sonar reflectors as guiding beacons for autonomous navigation

    Ralph Simon, Stefan Rupitsch, Markus Baumann, Huan Wu, Herbert Peremans, and Jan Steckel

    PNAS January 21, 2020 117 (3) 1367-1374; first published January 6, 2020. https://doi.org/10.1073/pnas.1909890117
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    Retroconversion of estrogens into androgens by bacteria via a cobalamin-mediated methylation

    • From the Cover

    Po-Hsiang Wang, Yi-Lung Chen, Sean Ting-Shyang Wei, Kan Wu, Tzong-Huei Lee, Tien-Yu Wu, and Yin-Ru Chiang

    PNAS January 21, 2020 117 (3) 1395-1403; first published December 17, 2019. https://doi.org/10.1073/pnas.1914380117

    See companion article:

    • Retroconversion of estrogens into androgens by bacteria via a cobalamin-mediated methylation

      - Dec 17, 2019
    • Anaerobic bacteria need their vitamin B₁₂ to digest estrogen

      - Jan 09, 2020
  • Open Access

    Bioderivatization as a concept for renewable production of chemicals that are toxic or poorly soluble in the liquid phase

    Pachara Sattayawat, Ian Sofian Yunus, and Patrik R. Jones

    PNAS January 21, 2020 117 (3) 1404-1413; first published January 8, 2020. https://doi.org/10.1073/pnas.1914069117

• Biochemistry

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    The effector mechanism of siRNA spherical nucleic acids

    Gokay Yamankurt, Robert J. Stawicki, Diana M. Posadas, Joseph Q. Nguyen, Richard W. Carthew, and Chad A. Mirkin

    PNAS January 21, 2020 117 (3) 1312-1320; first published January 3, 2020. https://doi.org/10.1073/pnas.1915907117
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    Activation by substoichiometric inhibition

    Melisa Merdanovic, Steven G. Burston, Anna Laura Schmitz, Steffen Köcher, Stefan Knapp, Tim Clausen, Markus Kaiser, Robert Huber, and Michael Ehrmann

    PNAS January 21, 2020 117 (3) 1414-1418; first published January 6, 2020. https://doi.org/10.1073/pnas.1918721117
  • Open Access

    A de novo peroxidase is also a promiscuous yet stereoselective carbene transferase

    Richard Stenner, Jack W. Steventon, Annela Seddon, and J. L. Ross Anderson

    PNAS January 21, 2020 117 (3) 1419-1428; first published January 2, 2020. https://doi.org/10.1073/pnas.1915054117
  • Open Access

    Long-range interdomain communications in eIF5B regulate GTP hydrolysis and translation initiation

    Bridget Y. Huang and Israel S. Fernández

    PNAS January 21, 2020 117 (3) 1429-1437; first published January 3, 2020. https://doi.org/10.1073/pnas.1916436117
  • Open Access

    Cryo-EM analysis of a feline coronavirus spike protein reveals a unique structure and camouflaging glycans

    Tzu-Jing Yang, Yen-Chen Chang, Tzu-Ping Ko, Piotr Draczkowski, Yu-Chun Chien, Yuan-Chih Chang, Kuen-Phon Wu, Kay-Hooi Khoo, Hui-Wen Chang, and Shang-Te Danny Hsu

    PNAS January 21, 2020 117 (3) 1438-1446; first published January 3, 2020. https://doi.org/10.1073/pnas.1908898117
  • Open Access

    Rotary catalysis of bovine mitochondrial F₁-ATPase studied by single-molecule experiments

    Ryohei Kobayashi, Hiroshi Ueno, Chun-Biu Li, and Hiroyuki Noji

    PNAS January 21, 2020 117 (3) 1447-1456; first published January 2, 2020. https://doi.org/10.1073/pnas.1909407117
  • Open Access

    Translation of the intrinsically disordered protein α-synuclein is inhibited by a small molecule targeting its structured mRNA

    Peiyuan Zhang, Hye-Jin Park, Jie Zhang, Eunsung Junn, Ryan J. Andrews, Sai Pradeep Velagapudi, Daniel Abegg, Kamalakannan Vishnu, Matthew G. Costales, Jessica L. Childs-Disney, Alexander Adibekian, Walter N. Moss, M. Maral Mouradian, and Matthew D. Disney

    PNAS January 21, 2020 117 (3) 1457-1467; first published January 3, 2020. https://doi.org/10.1073/pnas.1905057117

• Biophysics and Computational Biology

  • Open Access

    Intermediate states of molecular self-assembly from liquid-cell electron microscopy

    Huan Wang, Bo Li, Ye-Jin Kim, Oh-Hoon Kwon, and Steve Granick

    PNAS January 21, 2020 117 (3) 1283-1292; first published January 7, 2020. https://doi.org/10.1073/pnas.1916065117
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    Braiding topology and the energy landscape of chromosome organization proteins

    Dana Krepel, Aram Davtyan, Nicholas P. Schafer, Peter G. Wolynes, and José N. Onuchic

    PNAS January 21, 2020 117 (3) 1468-1477; first published December 30, 2019. https://doi.org/10.1073/pnas.1917750117

    See companion article:

    • Braiding topology and the energy landscape of chromosome organization proteins

      - Dec 30, 2019
    • Insights into the energy landscapes of chromosome organization proteins from coevolutionary sequence variation and structural modeling

      - Jan 10, 2020
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    Structures of cofilin-induced structural changes reveal local and asymmetric perturbations of actin filaments

    Andrew R. Huehn, Jeffrey P. Bibeau, Anthony C. Schramm, Wenxiang Cao, Enrique M. De La Cruz, and Charles V. Sindelar

    PNAS January 21, 2020 117 (3) 1478-1484; first published January 3, 2020. https://doi.org/10.1073/pnas.1915987117

    See companion article:

    • Structures of cofilin-induced structural changes reveal local and asymmetric perturbations of actin filaments

      - Jan 03, 2020
    • Cofilin-induced structural changes in actin filaments stay local

      - Jan 31, 2020
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    Cotranslational folding allows misfolding-prone proteins to circumvent deep kinetic traps

    Amir Bitran, William M. Jacobs, Xiadi Zhai, and Eugene Shakhnovich

    PNAS January 21, 2020 117 (3) 1485-1495; first published January 7, 2020. https://doi.org/10.1073/pnas.1913207117
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    Improved protein structure prediction using predicted interresidue orientations

    Jianyi Yang, Ivan Anishchenko, Hahnbeom Park, Zhenling Peng, Sergey Ovchinnikov, and David Baker

    PNAS January 21, 2020 117 (3) 1496-1503; first published January 2, 2020. https://doi.org/10.1073/pnas.1914677117
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    Molecular mechanism of the chitinolytic peroxygenase reaction

    Bastien Bissaro, Bennett Streit, Ingvild Isaksen, Vincent G. H. Eijsink, Gregg T. Beckham, Jennifer L. DuBois, and Åsmund K. Røhr

    PNAS January 21, 2020 117 (3) 1504-1513; first published January 6, 2020. https://doi.org/10.1073/pnas.1904889117

• Cell Biology

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    A cell atlas of the adult Drosophila midgut

    Ruei-Jiun Hung, Yanhui Hu, Rory Kirchner, Yifang Liu, Chiwei Xu, Aram Comjean, Sudhir Gopal Tattikota, Fangge Li, Wei Song, Shannan Ho Sui, and Norbert Perrimon

    PNAS January 21, 2020 117 (3) 1514-1523; first published January 8, 2020. https://doi.org/10.1073/pnas.1916820117
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    Loss of TSC complex enhances gluconeogenesis via upregulation of Dlk1-Dio3 locus miRNAs

    Dritan Liko, Andrzej Rzepiela, Vanja Vukojevic, Mihaela Zavolan, and Michael N. Hall

    PNAS January 21, 2020 117 (3) 1524-1532; first published January 9, 2020. https://doi.org/10.1073/pnas.1918931117
  • Open Access

    Quantitative microscopy reveals dynamics and fate of clustered IRE1α

    Vladislav Belyy, Ngoc-Han Tran, and Peter Walter

    PNAS January 21, 2020 117 (3) 1533-1542; first published December 23, 2019. https://doi.org/10.1073/pnas.1915311117

    See companion article:

    • Quantitative microscopy reveals dynamics and fate of clustered IRE1α

      - Dec 23, 2019
    • Dynamics and clustering of IRE1α during ER stress

      - Feb 04, 2020
  • Open Access

    The clock gene Bmal1 inhibits macrophage motility, phagocytosis, and impairs defense against pneumonia

    Gareth B. Kitchen, Peter S. Cunningham, Toryn M. Poolman, Mudassar Iqbal, Robert Maidstone, Matthew Baxter, James Bagnall, Nicola Begley, Ben Saer, Tracy Hussell, Laura C. Matthews, David H. Dockrell, Hannah J. Durrington, Julie E. Gibbs, John F. Blaikley, Andrew S. Loudon, and David W. Ray

    PNAS January 21, 2020 117 (3) 1543-1551; first published January 3, 2020. https://doi.org/10.1073/pnas.1915932117

• Developmental Biology

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    Global shape of Toll activation is determined by wntD enhancer properties

    Neta Rahimi, Shari Carmon, Inna Averbukh, Farzaneh Khajouei, Saurabh Sinha, Eyal D. Schejter, Naama Barkai, and Ben-Zion Shilo

    PNAS January 21, 2020 117 (3) 1552-1558; first published January 3, 2020. https://doi.org/10.1073/pnas.1918268117

• Ecology

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    Geographically divergent evolutionary and ecological legacies shape mammal biodiversity in the global tropics and subtropics

    John Rowan, Lydia Beaudrot, Janet Franklin, Kaye E. Reed, Irene E. Smail, Andrew Zamora, and Jason M. Kamilar

    PNAS January 21, 2020 117 (3) 1559-1565; first published December 16, 2019. https://doi.org/10.1073/pnas.1910489116

    See companion article:

    • Geographically divergent evolutionary and ecological legacies shape mammal biodiversity in the global tropics and subtropics

      - Dec 16, 2019
    • Present-day drivers do not explain biodiversity patterns in mammals

      - Jan 03, 2020
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    Infrared optical and thermal properties of microstructures in butterfly wings

    Anirudh Krishna, Xiao Nie, Andrew D. Warren, Jorge E. Llorente-Bousquets, Adriana D. Briscoe, and Jaeho Lee

    PNAS January 21, 2020 117 (3) 1566-1572; first published January 9, 2020. https://doi.org/10.1073/pnas.1906356117
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    Land-use history impacts functional diversity across multiple trophic groups

    Gaëtane Le Provost, Isabelle Badenhausser, Yoann Le Bagousse-Pinguet, Yann Clough, Laura Henckel, Cyrille Violle, Vincent Bretagnolle, Marilyn Roncoroni, Peter Manning, and Nicolas Gross

    PNAS January 21, 2020 117 (3) 1573-1579; first published January 6, 2020. https://doi.org/10.1073/pnas.1910023117
  • Open Access

    Scaling the risk landscape drives optimal life-history strategies and the evolution of grazing

    • From the Cover

    Uttam Bhat, Christopher P. Kempes, and Justin D. Yeakel

    PNAS January 21, 2020 117 (3) 1580-1586; first published December 17, 2019. https://doi.org/10.1073/pnas.1907998117

    See companion article:

    • Scaling the risk landscape drives optimal life-history strategies and the evolution of grazing

      - Dec 17, 2019
    • Life-history models reconstruct mammalian evolution

      - Jan 08, 2020
  • Open Access

    Recovery of critically endangered Nassau grouper (Epinephelus striatus) in the Cayman Islands following targeted conservation actions

    • From the Cover

    Lynn Waterhouse, Scott A. Heppell, Christy V. Pattengill-Semmens, Croy McCoy, Phillippe Bush, Bradley C. Johnson, and Brice X. Semmens

    PNAS January 21, 2020 117 (3) 1587-1595; first published January 6, 2020. https://doi.org/10.1073/pnas.1917132117

    See companion article:

    • Recovery of critically endangered Nassau grouper (Epinephelus striatus) in the Cayman Islands following targeted conservation actions

      - Jan 06, 2020
    • Island of hope for the threatened Nassau grouper

      - Jan 22, 2020

• Environmental Sciences

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    High productivity in hybrid-poplar plantations without isoprene emission to the atmosphere

    Russell K. Monson, Barbro Winkler, Todd N. Rosenstiel, Katja Block, Juliane Merl-Pham, Steven H. Strauss, Kori Ault, Jason Maxfield, David J. P. Moore, Nicole A. Trahan, Amberly A. Neice, Ian Shiach, Greg A. Barron-Gafford, Peter Ibsen, Joel T. McCorkel, Jörg Bernhardt, and Joerg-Peter Schnitzler

    PNAS January 21, 2020 117 (3) 1596-1605; first published January 6, 2020. https://doi.org/10.1073/pnas.1912327117

• Evolution

  • Open Access

    Biophysical principles of choanoflagellate self-organization

    Ben T. Larson, Teresa Ruiz-Herrero, Stacey Lee, Sanjay Kumar, L. Mahadevan, and Nicole King

    PNAS January 21, 2020 117 (3) 1303-1311; first published January 2, 2020. https://doi.org/10.1073/pnas.1909447117
  • Open Access

    A compartment size-dependent selective threshold limits mutation accumulation in hierarchical tissues

    Dániel Grajzel, Imre Derényi, and Gergely J. Szöllősi

    PNAS January 21, 2020 117 (3) 1606-1611; first published January 6, 2020. https://doi.org/10.1073/pnas.1913104117
  • Open Access

    Fin ray patterns at the fin-to-limb transition

    Thomas A. Stewart, Justin B. Lemberg, Natalia K. Taft, Ihna Yoo, Edward B. Daeschler, and Neil H. Shubin

    PNAS January 21, 2020 117 (3) 1612-1620; first published December 30, 2019. https://doi.org/10.1073/pnas.1915983117

• Genetics

  • Open Access

    Cystic fibrosis carriers are at increased risk for a wide range of cystic fibrosis-related conditions

    Aaron C. Miller, Alejandro P. Comellas, Douglas B. Hornick, David A. Stoltz, Joseph E. Cavanaugh, Alicia K. Gerke, Michael J. Welsh, Joseph Zabner, and Philip M. Polgreen

    PNAS January 21, 2020 117 (3) 1621-1627; first published December 27, 2019. https://doi.org/10.1073/pnas.1914912117

    See companion article:

    • Cystic fibrosis carriers are at increased risk for a wide range of cystic fibrosis-related conditions

      - Dec 27, 2019
    • Cystic fibrosis heterozygosity: Carrier state or haploinsufficiency?

      - Jan 28, 2020
  • Open Access

    Large-scale contractions of Friedreich’s ataxia GAA repeats in yeast occur during DNA replication due to their triplex-forming ability

    Alexandra N. Khristich, Jillian F. Armenia, Robert M. Matera, Anna A. Kolchinski, and Sergei M. Mirkin

    PNAS January 21, 2020 117 (3) 1628-1637; first published January 7, 2020. https://doi.org/10.1073/pnas.1913416117
  • Open Access

    Presynaptic MAST kinase controls opposing postsynaptic responses to convey stimulus valence in Caenorhabditis elegans

    Shunji Nakano, Muneki Ikeda, Yuki Tsukada, Xianfeng Fei, Takamasa Suzuki, Yusuke Niino, Rhea Ahluwalia, Ayana Sano, Rumi Kondo, Kunio Ihara, Atsushi Miyawaki, Koichi Hashimoto, Tetsuya Higashiyama, and Ikue Mori

    PNAS January 21, 2020 117 (3) 1638-1647; first published January 7, 2020. https://doi.org/10.1073/pnas.1909240117

• Immunology and Inflammation

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    Structural insight into T cell coinhibition by PD-1H (VISTA)

    Benjamin T. Slater, Xue Han, Lieping Chen, and Yong Xiong

    PNAS January 21, 2020 117 (3) 1648-1657; first published January 9, 2020. https://doi.org/10.1073/pnas.1908711117

• Medical Sciences

  • Open Access

    Identification and characterization of extrachromosomal circular DNA in maternal plasma

    Sarah T. K. Sin, Peiyong Jiang, Jiaen Deng, Lu Ji, Suk Hang Cheng, Anindya Dutta, Tak Y. Leung, K. C. Allen Chan, Rossa W. K. Chiu, and Y. M. Dennis Lo

    PNAS January 21, 2020 117 (3) 1658-1665; first published January 3, 2020. https://doi.org/10.1073/pnas.1914949117
  • Open Access

    High-dose ascorbic acid synergizes with anti-PD1 in a lymphoma mouse model

    Rebecca A. Luchtel, Tushar Bhagat, Kith Pradhan, William R. Jacobs Jr, Mark Levine, Amit Verma, and Niraj Shenoy

    PNAS January 21, 2020 117 (3) 1666-1677; first published January 7, 2020. https://doi.org/10.1073/pnas.1908158117
  • Open Access

    Expansion, in vivo–ex vivo cycling, and genetic manipulation of primary human hepatocytes

    Eleftherios Michailidis, Koen Vercauteren, Liliana Mancio-Silva, Linda Andrus, Cyprien Jahan, Inna Ricardo-Lax, Chenhui Zou, Mohammad Kabbani, Paul Park, Corrine Quirk, Christina Pyrgaki, Brandon Razooky, Lieven Verhoye, Irene Zoluthkin, Wei-Yu Lu, Stuart J. Forbes, Luis Chiriboga, Neil D. Theise, Roland W. Herzog, Hiroshi Suemizu, William M. Schneider, Amir Shlomai, Philip Meuleman, Sangeeta N. Bhatia, Charles M. Rice, and Ype P. de Jong

    PNAS January 21, 2020 117 (3) 1678-1688; first published January 8, 2020. https://doi.org/10.1073/pnas.1919035117

• Microbiology

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    Designing cyclic competence-stimulating peptide (CSP) analogs with pan-group quorum-sensing inhibition activity in Streptococcus pneumoniae

    Yifang Yang, Jingjun Lin, Anthony Harrington, Gabriel Cornilescu, Gee W. Lau, and Yftah Tal-Gan

    PNAS January 21, 2020 117 (3) 1689-1699; first published January 8, 2020. https://doi.org/10.1073/pnas.1915812117
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    Bile acids and ceramide overcome the entry restriction for GII.3 human norovirus replication in human intestinal enteroids

    Kosuke Murakami, Victoria R. Tenge, Umesh C. Karandikar, Shih-Ching Lin, Sasirekha Ramani, Khalil Ettayebi, Sue E. Crawford, Xi-Lei Zeng, Frederick H. Neill, B. Vijayalakshmi Ayyar, Kazuhiko Katayama, David Y. Graham, Erhard Bieberich, Robert L. Atmar, and Mary K. Estes

    PNAS January 21, 2020 117 (3) 1700-1710; first published January 2, 2020. https://doi.org/10.1073/pnas.1910138117
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    Mitochondrial dysfunctions trigger the calcium signaling-dependent fungal multidrug resistance

    Yeqi Li, Yuanwei Zhang, Chi Zhang, Hongchen Wang, Xiaolei Wei, Peiying Chen, and Ling Lu

    PNAS January 21, 2020 117 (3) 1711-1721; first published December 6, 2019. https://doi.org/10.1073/pnas.1911560116

    See companion article:

    • Linking calcium signaling and mitochondrial function in fungal drug resistance

      - Jan 03, 2020
    • Mitochondrial dysfunctions trigger the calcium signaling-dependent fungal multidrug resistance

      - Dec 06, 2019
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    A promiscuous inflammasome sparks replication of a common tumor virus

    Eric M. Burton, Raphaela Goldbach-Mansky, and Sumita Bhaduri-McIntosh

    PNAS January 21, 2020 117 (3) 1722-1730; first published January 9, 2020. https://doi.org/10.1073/pnas.1919133117
  • Open Access

    Robust hepatitis E virus infection and transcriptional response in human hepatocytes

    Daniel Todt, Martina Friesland, Nora Moeller, Dimas Praditya, Volker Kinast, Yannick Brüggemann, Leonard Knegendorf, Thomas Burkard, Joerg Steinmann, Rani Burm, Lieven Verhoye, Avista Wahid, Toni Luise Meister, Michael Engelmann, Vanessa M. Pfankuche, Christina Puff, Florian W. R. Vondran, Wolfgang Baumgärtner, Philip Meuleman, Patrick Behrendt, and Eike Steinmann

    PNAS January 21, 2020 117 (3) 1731-1741; first published January 2, 2020. https://doi.org/10.1073/pnas.1912307117

• Neuroscience

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    Microglial IRF5-IRF4 regulatory axis regulates neuroinflammation after cerebral ischemia and impacts stroke outcomes

    Abdullah Al Mamun, Anjali Chauhan, Shaohua Qi, Conelius Ngwa, Yan Xu, Romana Sharmeen, Amy L. Hazen, Jun Li, Jaroslaw A. Aronowski, Louise D. McCullough, and Fudong Liu

    PNAS January 21, 2020 117 (3) 1742-1752; first published December 31, 2019. https://doi.org/10.1073/pnas.1914742117
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    Impaired endothelium-mediated cerebrovascular reactivity promotes anxiety and respiration disorders in mice

    Jan Wenzel, Cathrin E. Hansen, Carla Bettoni, Miriam A. Vogt, Beate Lembrich, Rentsenkhand Natsagdorj, Gianna Huber, Josefine Brands, Kjestine Schmidt, Julian C. Assmann, Ines Stölting, Kathrin Saar, Jan Sedlacik, Jens Fiehler, Peter Ludewig, Michael Wegmann, Nina Feller, Marius Richter, Helge Müller-Fielitz, Thomas Walther, Gabriele M. König, Evi Kostenis, Walter Raasch, Norbert Hübner, Peter Gass, Stefan Offermanns, Cor de Wit, Carsten A. Wagner, and Markus Schwaninger

    PNAS January 21, 2020 117 (3) 1753-1761; first published January 2, 2020. https://doi.org/10.1073/pnas.1907467117
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    NK cells clear α-synuclein and the depletion of NK cells exacerbates synuclein pathology in a mouse model of α-synucleinopathy

    Rachael H. Earls, Kelly B. Menees, Jaegwon Chung, Claire-Anne Gutekunst, Hyun Joon Lee, Manuel G. Hazim, Balázs Rada, Levi B. Wood, and Jae-Kyung Lee

    PNAS January 21, 2020 117 (3) 1762-1771; first published January 3, 2020. https://doi.org/10.1073/pnas.1909110117

• Physiology

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    Internal fluid pressure influences muscle contractile force

    David A. Sleboda and Thomas J. Roberts

    PNAS January 21, 2020 117 (3) 1772-1778; first published December 26, 2019. https://doi.org/10.1073/pnas.1914433117

    See companion article:

    • Internal fluid pressure influences muscle contractile force

      - Dec 26, 2019
    • Muscle force is modulated by internal pressure

      - Jan 08, 2020
  • Open Access

    Specialized stellate cells offer a privileged route for rapid water flux in Drosophila renal tubule

    Pablo Cabrero, Selim Terhzaz, Anthony J. Dornan, Saurav Ghimire, Heather L. Holmes, Daniel R. Turin, Michael F. Romero, Shireen A. Davies, and Julian A. T. Dow

    PNAS January 21, 2020 117 (3) 1779-1787; first published January 6, 2020. https://doi.org/10.1073/pnas.1915943117
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    Cryo-EM structures of a lipid-sensitive pentameric ligand-gated ion channel embedded in a phosphatidylcholine-only bilayer

    Pramod Kumar, Yuhang Wang, Zhening Zhang, Zhiyu Zhao, Gisela D. Cymes, Emad Tajkhorshid, and Claudio Grosman

    PNAS January 21, 2020 117 (3) 1788-1798; first published January 7, 2020. https://doi.org/10.1073/pnas.1906823117

• Plant Biology

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    The maize heterotrimeric G protein β subunit controls shoot meristem development and immune responses

    Qingyu Wu, Fang Xu, Lei Liu, Si Nian Char, Yezhang Ding, Byoung Il Je, Eric Schmelz, Bing Yang, and David Jackson

    PNAS January 21, 2020 117 (3) 1799-1805; first published December 18, 2019. https://doi.org/10.1073/pnas.1917577116

    See companion article:

    • The maize heterotrimeric G protein β subunit controls shoot meristem development and immune responses

      - Dec 18, 2019
    • Shoot meristem maintenance and immune response signaling converge at the G protein β subunit

      - Jan 14, 2020
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    The rhizobial autotransporter determines the symbiotic nitrogen fixation activity of Lotus japonicus in a host-specific manner

    Yoshikazu Shimoda, Yuki Nishigaya, Hiroko Yamaya-Ito, Noritoshi Inagaki, Yosuke Umehara, Hideki Hirakawa, Shusei Sato, Toshimasa Yamazaki, and Makoto Hayashi

    PNAS January 21, 2020 117 (3) 1806-1815; first published January 3, 2020. https://doi.org/10.1073/pnas.1913349117