Abstract
All viruses require strategies to inhibit or evade the immunity pathways of cells they infect. The viruses that infect bacteria, bacteriophages (phages), must avoid nucleic-acid targeting immune pathways such as CRISPR-Cas (clustered regularly interspaced short palindromic repeats and CRISPR-associated genes) and restriction-modification (R-M) systems to replicate efficiently1. Here, we show that jumbo phage ΦKZ, infecting Pseudomonas aeruginosa, segregates its DNA from immunity nucleases by constructing a proteinaceous nucleus-like compartment. ΦKZ resists many DNA-targeting immune systems in vivo, including two CRISPR-Cas3 subtypes, Cas9, Cas12a, and the restriction enzymes HsdRMS and EcoRI. Cas and restriction enzymes are unable to access the phage DNA throughout the infection, but engineered re-localization of EcoRI inside the compartment enables phage targeting and cell protection. Moreover, ΦKZ is sensitive to the RNA targeting CRISPR-Cas enzyme, Cas13a, likely due to phage mRNA localizing to the cytoplasm. Collectively, we propose that Pseudomonas jumbo phages evade a broad spectrum of DNA-targeting nucleases through the assembly of a protein barrier around their genome.
Highlights
Despite widespread improvement in educational attainment since 2000, gender disparity persists in 2017 in many regions
Despite continued lack of gender parity in education among the reproductive age group, vast progress towards parity has been made among the 20–24 age group
We examined a younger group aged 20–24 years. Education in this group is less directly relevant to maternal, newborn, and child health than education in the full window of reproductive age, these estimates allowed us to capture how the landscape of education has shifted over time and is more likely to pick up improvements to access and retention in education systems that have been made since 2000
Summary
We have built on previous modelling efforts that focused on the geographical distribution of average education[14] by extending our estimation to the distribution of attainment, highlighting average attainment and the proportions of individuals who completed key levels of schooling that are central to policy efforts. Administration and Economics, National Research University Higher School of Economics, Moscow, Russia. 333Department of Global Health and Population, Harvard University, Boston, MA, USA. 334Department of Economics, University of Göttingen, Göttingen, Germany. 335Foundation University Medical College, Foundation University Islamabad, Islamabad, Pakistan. 336Department of Psychiatry, University of São Paulo, São Paulo, Brazil. 337Institute of Health and Society, University of Oslo, Oslo, Norway. 338Department of Neurology, Technical University of Munich, Munich, Germany. 339School of Population Health & Environmental Sciences, King’s College London, London, UK. 340NIHR Biomedical Research Centre, Guy’s and St Thomas’ Hospital and Kings College London, London, UK. 341Department of Diabetes and Metabolic Diseases, University of Tokyo, Tokyo, Japan. 342Wolkite University, Wolkite, Ethiopia. 343Centre for Suicide Research and Prevention, University of Hong Kong, Hong Kong, China. 344Department of Social Work and Social Administration, University of Hong Kong, Hong Kong, China. 345School of Allied Health Sciences, Addis Ababa University, Addis Ababa, Ethiopia. 346Department of Psychopharmacology, National Center of Neurology and Psychiatry, Tokyo, Japan. 347Health Economics & Finance, Global Health, Jackson State University, Jackson, MS, USA. 348School of Medicine, Tsinghua University, Peking, China. 349Prevention of Cardiovascular Disease Research Center, Shahid Beheshti University of Medical Sciences, Tehran, Iran. 350Global Health Institute, Wuhan University, Wuhan, China. 351Department of Epidemiology and Biostatistics, Wuhan University, Wuhan, China. 352Department of Medicine, Monash University, Melbourne, Victoria, Australia. 353Maternal and Child Health Division, International Centre for Diarrhoeal Disease Research, Bangladesh, Dhaka, Bangladesh. 354George Warren Brown School, Washington University in St Louis, St Louis, MO, USA. 355School of Public Health, Wuhan University of Science and Technology, Wuhan, China. 356Hubei Province Key Laboratory of Occupational Hazard
Sign-in/Register to view highlights & summary Sign-in/Register to access full text options 
Talk to us
Join us for a 30 min session where you can share your feedback and ask us any queries you have
Schedule a call
