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WO 2017/214476 Al O (12) INTERNATIONAL APPLICATION PUBLISHED UNDER THE PATENT COOPERATION TREATY (PCT) (19) World Intellectual Property Organization International Bureau (10) International Publication Number (43) International Publication Date W O 2017/214476 A l 14 December 2017 (14.12.2017) W ! P O PCT (51) International Patent Classification: (81) Designated States (unless otherwise indicated, for every C12N 15/87 (2006.01) C12R 1/01 (2006.01) kind of national protection available): AE, AG, AL, AM, A01K 67/033 {2006.01) AO, AT, AU, AZ, BA, BB, BG, BH, BN, BR, BW, BY, BZ, CA, CH, CL, CN, CO, CR, CU, CZ, DE, DJ, DK, DM, DO, (21) International Application Number: DZ, EC, EE, EG, ES, FI, GB, GD, GE, GH, GM, GT, HN, PCT/US20 17/036693 HR, HU, ID, IL, IN, IR, IS, JO, JP, KE, KG, KH, KN, KP, (22) International Filing Date: KR, KW, KZ, LA, LC, LK, LR, LS, LU, LY, MA, MD, ME, 09 June 2017 (09.06.2017) MG, MK, MN, MW, MX, MY, MZ, NA, NG, NI, NO, NZ, OM, PA, PE, PG, PH, PL, PT, QA, RO, RS, RU, RW, SA, (25) Filing Language: English SC, SD, SE, SG, SK, SL, SM, ST, SV, SY,TH, TJ, TM, TN, (26) Publication Langi English TR, TT, TZ, UA, UG, US, UZ, VC, VN, ZA, ZM, ZW. (30) Priority Data: (84) Designated States (unless otherwise indicated, for every 62/347,818 09 June 2016 (09.06.2016) US kind of regional protection available): ARIPO (BW, GH, GM, KE, LR, LS, MW, MZ, NA, RW, SD, SL, ST, SZ, TZ, (71) Applicants: VANDERBILT UNIVERSITY [US/US]; UG, ZM, ZW), Eurasian (AM, AZ, BY, KG, KZ, RU, TJ, 2201 West End Avenue, 305 Kirkland Hall, Nashville, Ten TM), European (AL, AT, BE, BG, CH, CY, CZ, DE, DK, nessee 37235 (US). YALE UNIVERSITY [US/US]; 433 EE, ES, FI, FR, GB, GR, HR, HU, IE, IS, IT, LT, LU, LV, Temple Street, First Floor, New Haven, Connecticut 065 11 MC, MK, MT, NL, NO, PL, PT, RO, RS, SE, SI, SK, SM, (US). TR), OAPI (BF, BJ, CF, CG, CI, CM, GA, GN, GQ, GW, (72) Inventors: METCALF, Jason; 11981 Trinkle Rd., Dex KM, ML, MR, NE, SN, TD, TG). ter, MI 48130 (US). BORDENSTEIN, Seth R.; 528 Sandpiper Circle, Nashville, Tennessee 37221 (US). LEP¬ Declarations under Rule 4.17: AGE, Daniel; 10422 N. Rock Creek Rd., Centertown, — of inventorship (Rule 4.1 7(iv)) MO 65023 (US). BORDENSTEIN, Sarah; 528 San d Published: piper Circle, Nashville, Tennessee 37221 (US). HOCHS- — with international search report (Art. 21(3)) TRASSER, Mark; 72 Spring Glen Terrace, Hamden, Con — before the expiration of the time limit for amending the necticut 065 17 (US). BECKMANN, John F.; 79 Kens claims and to be republished in the event of receipt of ington Street #1, New Haven, Connecticut 065 11 (US). amendments (Rule 48.2(h)) RONAU, Judith; 65 Beacon Street, Hamden, Connecticut — with sequence listing part of description (Rule 5.2(a)) 065 14 (US). (74) Agent: PRATHER, Donald M . et al; 999 Peachtree Street NE, Suite 1300, Atlanta, 30309 (US). (54) Title: CYTOPLASMIC INCOMPATIBILITY FACTORS AND METHODS FOR CONTROLLING ARTHROPODS c n=220 ::! n= 13 l D063 1, '«0632 9 n=178 « 1 C WD0631, 0 32 n= 0 n=57 i Q i % of embryos FIGURE 4 Q o (57) Abstract: The disclosure relates to genetically modified bacteria, genetically modified arthropods, and methods for controlling o and/or reducing arthropod populations. CYTOPLASMIC INCOMPATIBILITY FACTORS AND METHODS FOR CONTROLLING ARTHROPODS CROSS REFERENCE TO RELATEDAPPLICATIONS This application claims the benefit of U .S. Provisional Patent Application Serial No. 62/347,81 8 filed June 9, 201 6, which is expressly incorporated herein by reference. STATEMENT REGARDING FEDERALLY SPONSORED RESEARCH This invention was made with government support under 2014-67012-22268 awarded by USDA and under GM 053756, HD086833, GM007347, AI08 1322, CA068485, DK020593, DK058404, DK059637, and EY008126 awarded by National Institutes of Health and under 15 1398 and 1456778 awarded by the Nation Science Foundation. The government has certain rights in the invention. FIELD The disclosure relates to genetically modified bacteria, genetically modified arthropods, and methods for controlling and/or reducing arthropod populations. BACKGROUND The genus Wolbachia is an archetype of maternally inherited intracellular bacteria that infect the germline of millions of invertebrate species worldwide and parasitically alter arthropod sex ratios and reproductive strategies to increase the proportion of infected females (the transmitting sex) in the population. The most common of these reproductive manipulations is cytoplasmic incompatibility (CI), typically expressed as embryonic lethality in crosses between infected males and uninfected females. This lethality is completely rescued by females infected with the same or a similar Wolbachia strain. Cytoplasmic incompatibility (CI) has important applications in disease vector control, and is currently being used in field trials to drive the spread of Dengue-resistant mosquitoes in wild populations through the release of Wolbachia-infectedfemales, and as a biological control mechanism to depress mosquito populations by releasing Wolbachia-infected males incompatible with wild females. Despite more than 40 years of research, the genes by which Wolbachia cause CI remain unknown. SUMMARY Disclosed herein are genetically modified bacteria and genetically modified arthropods useful for controlling and/or reducing populations of arthropods (for example, insects). For the first time, the inventors have identified the genes that encode the cytoplasmic incompatibility factors capable of reproducing the phenomena of cytoplasmic incompatibility. These genes are used to genetically modify bacteria and/or arthropods in order to produce sterile male arthropods and/or to replace a population of target arthropods. n one aspect, provided herein is a genetically modified arthropod, said arthropod comprising: a bacterial operon encoding a cytoplasmic incompatibility factor or a variant thereof: and a promoter operably linked to the bacterial operon: wherein the expression of the cytoplasmic incompatibility factor in a male arthropod causes a reduction in viable offspring in comparison to a male arthropod lacking the cytoplasmic incompatibillty factor. n another aspect, provided herein is a method for controlling a population of target arthropods, comprising: providing a bacterial operon encoding a cytoplasmic incompatibility factor or a variant thereof, and a promoter operably linked to the bacterial operon; transforming a population of male arthropods with the bacterial operon: and releasing the male arthropods amongst a population of target arthropods, wherein the release of the male arthropods reduces the population of target arthropods. in one aspect, provided herein is a genetically modified bacterium comprising: a bacterial operon encoding a cytoplasmic incompatibility factor or a variant thereof; and a promoter operably linked to the bacterial operon; wherein the bacterial operon occurs at a non-naturally occurring genomic location in the bacterium. In another aspect, provided herein is an arthropod infected with a bacterium, wherein the bacterium comprises: a bacterial operon encoding a cytoplasmic incompatibility factor or a variant thereof; and a promoter operably linked to the bacterial operon; wherem the bacterial operon occurs at a non-naturally occurring genomic location in the bacterium. n an additional aspect, provided herein is a method for controlling a population of target arthropods, comprising: providing a genetically modified bacterium comprising: a bacterial operon encoding a cytoplasmic incompatibility factor or a variant thereof, and a promoter operably linked to the bacterial operon; infecting a population of replacement arthropods with the genetically modified bacterium; and releasing the replacement arthropods amongst a population of target arthropods, wherein the release of the replacement arthropods reduces the population of target arthropods. BRIEF DESCRIPTION OF THE DRAWINGS The accompanying figures, which are incorporated in and constitute a part of this specification, illustrate several aspects described below. FIG. 1. Cytoplasmic incompatibility (CI) gene candidate selection and evolution (a) Venn diagram illustrating unique and shared gene sets from four Cl-inducing Wolbachia strains. The number of gene families in common between strains is indicated for each combination (b) Venn diagram illustrating the number of unique wMel genes matching each criteria combination. Bayesian phylogenies of WD0631 (c) and WD0632 (e) and their homologs are shown based on a 256-aa alignment of WD0631 reciprocal BLASTp hits and a 462-aa alignment of WD0632 reciprocal BLASTp hits. When multiple similar copies of the same operon exist in the same strain, only one copy is shown. Consensus support values are shown at the nodes. Both trees are based on the JTT+G model of evolution and are unrooted (d) CI patterns correlate with WD0631/WD0632 operon homology. Ri rescues wMel and both share a similar operon (*). The inability of wMel to rescue Ri correlates with an operon type (†) that is present in Ri but absent in wMel. Likewise, bidirectional incompatibility of all other crosses correlates to divergent operons. This diagram was adapted from Bossan et. aP . (f) Protein architecture of WD0631/WD0632 homologs is conserved for each clade and is classified according to the WD0632-like domain: Type I features Peptidase_C48; Type II lacks an annotated functional domain; and Type III features DUF1703. TM stands for transmembrane domain. For (c) and (e), the WO-prefix indicates a specific phage WO haplotype and the w-prefix refers to a "WO-like island," a small subset of conserved phage genes, within that specific Wolbachia strain. FIG. 2. Expression of CI effector candidates decrease as males age. (a-f) Expression of each gene in one-day-old and seven-day-old wMel-infected D . melanogaster testes, as determined by quantitative RT-PCR, is shown relative to groEL. Error bars indicate standard deviation. * = P<0.05, ** = P .01 by Mann-Whitney U test. FIG. 3. Dual expression of WD063 1 and WD0632 induces CI. Hatch rate assays are shown with either single-gene transgenic (a,c), or dual WD063 1/WD0632 transgenic (b,c) D .
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