b1ab3c7b27ebd4063774d98fb655024e mbio_a2026m2v17n2pe02939.pdf 2ef1314c8e43a0b5117c107ef5aecbe759e647e3 mbio_a2026m2v17n2pe02939.pdf 7423bd7d0e0879abbf33521b86288eb0a58d07535ec2ed805bf48105291cfbe6 mbio_a2026m2v17n2pe02939.pdf Title: Nucleoside binding by a surface lipoprotein governs conjugative ICE acquisition in mycoplasmas Subject: mBio 2026.17:e02939-25 Keywords: horizontal gene transfer, integrative and conjugative element, bacterial conjugation, mycoplasma, mycoplasmopsis, nucleoside binding, lipoprotein, recipient cell Author: M'hamed Derriche, Laurent Xavier Nouvel, Calvin Fauvet, Núria Mach, Elisa Simon, Gwendoline Pot, Hortensia Robert, Alexandre Stella, Christian de la Fe, Renaud Maillard, Sergi Torres-Puig, Yonathan Arfi, Christine Citti, Eric Baranowski Creator: Antenna House XSL Formatter V7.4 MR2 Linux : 7.4.3.63807 (2024-03-22T11:49+09) Producer: Antenna House PDF Output Library 7.4.1901 CreationDate: Wed Feb 4 11:18:32 2026 CET ModDate: Wed Feb 4 11:18:32 2026 CET Custom Metadata: yes Metadata Stream: yes Tagged: yes UserProperties: no Suspects: no Form: none JavaScript: no Pages: 20 Encrypted: no Page size: 585 x 783 pts Page rot: 0 File size: 1249619 bytes Optimized: no PDF version: 1.7 name type encoding emb sub uni object ID ------------------------------------ ----------------- ---------------- --- --- --- --------- MUALFL+MyriadPro-Regular Type 1C Custom yes yes yes 16 0 MUALFL+MyriadPro-Semibold Type 1C Custom yes yes yes 19 0 MUALFL+MyriadPro-Semibold Type 1C Custom yes yes yes 81 0 MUALFL+MyriadPro-Regular Type 1C Custom yes yes yes 98 0 MUALFL+MyriadPro-It Type 1C Custom yes yes yes 110 0 MUALFL+MyriadPro-Bold Type 1C Builtin yes yes yes 152 0 MUALFL+MyriadPro-Light Type 1C Custom yes yes yes 156 0 MUALFL+MyriadPro-Light Type 1C Custom yes yes yes 159 0 Times-Roman Type 1 WinAnsi no no no 3268 0 MUALFL+MyriadPro-SemiboldIt Type 1C Builtin yes yes yes 259 0 MUALFL+STIXTwoMath-Regular Type 1C Custom yes yes yes 1796 0 MUALFL+NotoSansMath-Regular CID TrueType Identity-H yes yes yes 1852 0 MUALFL+NotoSerif CID TrueType Identity-H yes yes yes 2069 0 Jhove (Rel. 1.28.0, 2023-05-18) Date: 2026-06-16 04:07:59 CEST RepresentationInformation: mbio_a2026m2v17n2pe02939.pdf ReportingModule: PDF-hul, Rel. 1.12.4 (2023-03-16) LastModified: 2026-06-15 10:36:20 CEST Size: 1249619 Format: PDF Version: 1.7 Status: Well-Formed, but not valid SignatureMatches: PDF-hul ErrorMessage: Invalid indirect destination - referenced object 'fig-caption-kriyadocs-F1_0' cannot be found ID: PDF-HUL-149 ErrorMessage: Invalid indirect destination - referenced object 'table-caption-kriyadocs-T1_0' cannot be found ID: PDF-HUL-149 ErrorMessage: Invalid indirect destination - referenced object 'fig-caption-kriyadocs-F2_0' cannot be found ID: PDF-HUL-149 ErrorMessage: Invalid indirect destination - referenced object 'fig-caption-kriyadocs-F3_0' cannot be found ID: PDF-HUL-149 ErrorMessage: Invalid indirect destination - referenced object 'table-caption-kriyadocs-T3_0' cannot be found ID: PDF-HUL-149 ErrorMessage: Invalid indirect destination - referenced object 'table-caption-kriyadocs-T2_0' cannot be found ID: PDF-HUL-149 ErrorMessage: edu.harvard.hul.ois.jhove.module.pdf.PdfInvalidException: Invalid indirect destination - referenced object 'fig-caption-kriyadocs-F1_0' cannot be found ID: PDF-HUL-122 Offset: 1211998 ErrorMessage: edu.harvard.hul.ois.jhove.module.pdf.PdfInvalidException: Invalid indirect destination - referenced object 'fig-caption-kriyadocs-F2_0' cannot be found ID: PDF-HUL-122 Offset: 1211998 ErrorMessage: edu.harvard.hul.ois.jhove.module.pdf.PdfInvalidException: Invalid indirect destination - referenced object 'fig-caption-kriyadocs-F3_0' cannot be found ID: PDF-HUL-122 Offset: 1211998 ErrorMessage: edu.harvard.hul.ois.jhove.module.pdf.PdfInvalidException: Invalid indirect destination - referenced object 'table-caption-kriyadocs-T3_0' cannot be found ID: PDF-HUL-122 Offset: 1211998 ErrorMessage: edu.harvard.hul.ois.jhove.module.pdf.PdfInvalidException: Invalid indirect destination - referenced object 'table-caption-kriyadocs-T1_0' cannot be found ID: PDF-HUL-122 Offset: 1211998 ErrorMessage: edu.harvard.hul.ois.jhove.module.pdf.PdfInvalidException: Invalid indirect destination - referenced object 'table-caption-kriyadocs-T2_0' cannot be found ID: PDF-HUL-122 Offset: 1211998 MIMEtype: application/pdf Profile: Tagged PDF PDFMetadata: Objects: 0 FreeObjects: 1 IncrementalUpdates: 0 DocumentCatalog: ViewerPreferences: HideToolbar: false HideMenubar: false HideWindowUI: false FitWindow: false CenterWindow: false DisplayDocTitle: true NonFullScreenPageMode: UseNone Direction: L2R ViewArea: CropBox ViewClip: CropBox PrintArea: CropBox PageClip: CropBox PageLayout: SinglePage PageMode: UseNone Language: EN Outlines: Item: Title: Nucleoside binding by a surface lipoprotein governs conjugative ICE acquisition in mycoplasmas Children: Item: Title: RESULTS Children: Item: Title: M. agalactiae genome-encoded determinants influencing ICEA acquisition Item: Title: M. bovis genome-encoded determinants influencing ICEB acquisition Item: Title: Disruption of the p48 gene nearly abolishes ICEA acquisition without affecting its transfer Item: Title: Disruption of the p48 gene induces transcriptional activation of a vestigial ICE Item: Title: Lipoprotein P48 expression by recipient cells facilitates ICEA acquisition Item: Title: Sequence variations in lipoprotein P48 influence ICEA acquisition Item: Title: No detectable P48-CDS14 interaction in mating partner contact Item: Title: Lipoprotein P48 is a substrate-binding protein involved in nucleoside uptake Item: Title: The nucleoside-binding activity of lipoprotein P48 is required for efficient ICE acquisition Item: Title: ICEA acquisition relies on P48 nucleoside-binding activity but not on nucleoside uptake Item: Title: DISCUSSION Children: Item: Title: Recipient determinants of ICE acquisition Item: Title: P48 as a nucleoside-binding lipoprotein Item: Title: One protein, two possible mechanisms Item: Title: Nucleoside metabolism and ICE dissemination Item: Title: A target for the control of HGT in mycoplasmas Item: Title: Conclusion Item: Title: MATERIALS AND METHODS Children: Item: Title: Mycoplasmas, cell lines, and culture conditions Item: Title: Random transposon mutagenesis and mapping of insertion sites Item: Title: High-throughput mating assays Item: Title: Cell culture mating assays Item: Title: DNA constructs for protein expression in mycoplasmas Item: Title: Western blotting Item: Title: 5-FU inhibition and nucleoside competition assays Item: Title: Co-IP assays Item: Title: Proteomic analysis, data-independent acquisition, and data processing Item: Title: RNA isolation and RNA-seq Filters: FilterPipeline: FlateDecode FilterPipeline: DCTDecode Images: Image: NisoImageMetadata: FormatName: image/png ImageWidth: 32 ImageHeight: 51 ColorSpace: RGB BitsPerSample: 8 BitsPerSampleUnit: integer Filter: FlateDecode Image: NisoImageMetadata: FormatName: image/jpg CompressionScheme: JPEG ImageWidth: 557 ImageHeight: 136 ColorSpace: RGB BitsPerSample: 8 BitsPerSampleUnit: integer Image: NisoImageMetadata: FormatName: image/jpg CompressionScheme: JPEG ImageWidth: 2069 ImageHeight: 1811 ColorSpace: black is zero BitsPerSample: 8 BitsPerSampleUnit: integer Image: NisoImageMetadata: FormatName: image/jpg CompressionScheme: JPEG ImageWidth: 1702 ImageHeight: 1946 ColorSpace: black is zero BitsPerSample: 8 BitsPerSampleUnit: integer Image: NisoImageMetadata: FormatName: image/jpg CompressionScheme: JPEG ImageWidth: 2137 ImageHeight: 1727 ColorSpace: RGB BitsPerSample: 8 BitsPerSampleUnit: integer Image: NisoImageMetadata: FormatName: image/jpg CompressionScheme: JPEG ImageWidth: 33 ImageHeight: 32 ColorSpace: RGB BitsPerSample: 8 BitsPerSampleUnit: integer Fonts: Type0: Font: BaseFont: MUALFL+NotoSerif Encoding: Identity-H ToUnicode: true Font: BaseFont: MUALFL+NotoSansMath-Regular Encoding: Identity-H ToUnicode: true Type1: Font: BaseFont: MUALFL+MyriadPro-Regular FontSubset: true FirstChar: 32 LastChar: 158 FontDescriptor: FontName: MUALFL+MyriadPro-Regular Flags: Nonsymbolic FontBBox: -157, -250, 1126, 952 FontFile3: true EncodingDictionary: Differences: true ToUnicode: true Font: BaseFont: MUALFL+MyriadPro-Semibold FontSubset: true FirstChar: 44 LastChar: 133 FontDescriptor: FontName: MUALFL+MyriadPro-Semibold Flags: Nonsymbolic FontBBox: -161, -250, 1198, 973 FontFile3: true EncodingDictionary: Differences: true ToUnicode: true Font: BaseFont: MUALFL+MyriadPro-Regular FontSubset: true FirstChar: 40 LastChar: 133 FontDescriptor: FontName: MUALFL+MyriadPro-Regular Flags: Nonsymbolic FontBBox: -157, -250, 1126, 952 FontFile3: true EncodingDictionary: Differences: true ToUnicode: true Font: BaseFont: MUALFL+MyriadPro-Semibold FontSubset: true FirstChar: 32 LastChar: 129 FontDescriptor: FontName: MUALFL+MyriadPro-Semibold Flags: Nonsymbolic FontBBox: -161, -250, 1198, 973 FontFile3: true EncodingDictionary: Differences: true ToUnicode: true Font: BaseFont: MUALFL+MyriadPro-SemiboldIt FontSubset: true FirstChar: 32 LastChar: 118 FontDescriptor: FontName: MUALFL+MyriadPro-SemiboldIt Flags: Nonsymbolic, Italic FontBBox: -193, -250, 1164, 972 FontFile3: true ToUnicode: true Font: BaseFont: Times-Roman Encoding: WinAnsiEncoding Font: BaseFont: MUALFL+STIXTwoMath-Regular FontSubset: true FirstChar: 128 LastChar: 128 FontDescriptor: FontName: MUALFL+STIXTwoMath-Regular Flags: Serif, Nonsymbolic FontBBox: -978, -1641, 3072, 2627 FontFile3: true EncodingDictionary: Differences: true ToUnicode: true Font: BaseFont: MUALFL+MyriadPro-Bold FontSubset: true FirstChar: 48 LastChar: 57 FontDescriptor: FontName: MUALFL+MyriadPro-Bold Flags: Nonsymbolic FontBBox: -163, -250, 1256, 989 FontFile3: true ToUnicode: true Font: BaseFont: MUALFL+MyriadPro-Light FontSubset: true FirstChar: 32 LastChar: 129 FontDescriptor: FontName: MUALFL+MyriadPro-Light Flags: Nonsymbolic FontBBox: -153, -250, 1046, 935 FontFile3: true EncodingDictionary: Differences: true ToUnicode: true Font: BaseFont: MUALFL+MyriadPro-It FontSubset: true FirstChar: 32 LastChar: 128 FontDescriptor: FontName: MUALFL+MyriadPro-It Flags: Nonsymbolic, Italic FontBBox: -185, -250, 1090, 953 FontFile3: true EncodingDictionary: Differences: true ToUnicode: true Font: BaseFont: MUALFL+MyriadPro-Light FontSubset: true FirstChar: 99 LastChar: 128 FontDescriptor: FontName: MUALFL+MyriadPro-Light Flags: Nonsymbolic FontBBox: -153, -250, 1046, 935 FontFile3: true EncodingDictionary: Differences: true ToUnicode: true CIDFontType2: Font: BaseFont: MUALFL+NotoSansMath-Regular CIDSystemInfo: Registry: Adobe Registry: Identity Supplement: 0 FontDescriptor: FontName: MUALFL+NotoSansMath-Regular Flags: Symbolic FontBBox: -354, -420, 1695, 1069 FontFile2: true Font: BaseFont: MUALFL+NotoSerif CIDSystemInfo: Registry: Adobe Registry: Identity Supplement: 0 FontDescriptor: FontName: MUALFL+NotoSerif Flags: Serif, Symbolic FontBBox: -558, -250, 1361, 1048 FontFile2: true XMP: application/pdfNucleoside binding by a surface lipoprotein governs conjugative ICE acquisition in mycoplasmasmBio02939-25M'hamed DerricheLaurent Xavier NouvelCalvin FauvetNúria MachElisa SimonGwendoline PotHortensia RobertAlexandre StellaChristian de la FeRenaud MaillardSergi Torres-PuigYonathan ArfiChristine CittiEric BaranowskiM'hamed DerricheLaurent Xavier NouvelCalvin FauvetNúria MachElisa SimonGwendoline PotHortensia RobertAlexandre StellaChristian de la FeRenaud MaillardSergi Torres-PuigYonathan ArfiChristine CittiEric BaranowskiAmerican Society for MicrobiologyCopyright © 2025 Derriche et al.This is an open-access article distributed under the terms of the Creative Commons Attribution 4.0 International license.20252023Integrative and conjugative elements (ICEs) are major mediators of horizontal gene transfer in bacteria. However, the role of recipient cells in their acquisition has received little attention. Using the ruminant pathogens Mycoplasma agalactiae and Mycoplasma bovis as minimal models, we combined genome-wide transposon mutagenesis with high-throughput mating assays to identify recipient factors required for ICE acquisition. The surface lipoprotein P48 emerged as the primary determinant of ICE uptake in both species. Structural and functional analyses revealed that P48 is the substrate-binding component of an ABC transporter with nucleoside-binding capacity. A single-point mutation that abolished nucleoside binding drastically reduced ICE acquisition, demonstrating that P48-mediated nucleoside recognition is essential for conjugative transfer. However, ICE uptake did not require nucleoside transport, as inactivation of the transporter permease blocked nucleoside analog toxicity but not ICE invasion. Loss of P48 also triggered transcriptional activation of vestigial ICE genes, suggesting that surface recognition affects the intracellular state of the recipient. Remarkably, ICE transfer from recipient-derived donors was unaffected by P48 loss, underscoring its acquisition-specific role. Together, these results reveal a previously unrecognized, surface-exposed recipient factor critical for efficient ICE transfer in mycoplasmas and identify nucleotide binding as a central function in conjugation. By demonstrating that recipient-encoded functions can directly control ICE dissemination, this work challenges the donor-centric paradigm of bacterial conjugation and suggests new strategies to restrict horizontal gene flow in pathogenic and synthetic mycoplasmas.Integrative and conjugative elements (ICEs) are mobile DNA elements that drive bacterial conjugation, a major process by which bacteria exchange genes. Although conjugation has been studied for decades, the focus has been almost exclusively on donor cells and the ICE itself, leaving the role of recipient cells largely overlooked. Using the wall-less ruminant pathogens Mycoplasma agalactiae and Mycoplasma bovis as minimal models, we discovered that a single recipient lipoprotein is required for efficient ICE uptake. Our data show that nucleoside recognition by P48, but not transport, is critical for conjugation, revealing an unexpected mechanistic link between nutrient sensing and gene acquisition. These findings shift the paradigm of conjugation from a donor-driven process to one jointly determined by donor and recipient functions. By identifying a recipient-encoded determinant of ICE transfer, this work opens new avenues to control horizontal gene flow in both pathogenic and engineered bacteria.20250923202512082150-7511uuid:956e75b0-7f53-44da-9d6a-942b04016d12uuid:55e19d6b-c9ff-4bc8-805f-d3eb19ceb73a Antenna House PDF Output Library 7.4.1901 Antenna House XSL Formatter V7.4 MR2 Linux : 7.4.3.63807 (2024-03-22T11:49+09) 2026-02-04T10:18:32+00:00 2026-02-04T10:18:32+00:00 2026-02-04T10:18:32+00:00 8457fdd4-bbca-4417-8024-e64de4bc486f 1 Pages: Page: Sequence: 1 Annotations: Annotation: Subtype: Link Contents: CrossMarkLogoLink Rect: 505, 701, 544, 742 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: 10.1128/mbio.02939-25 Rect: 462, 23, 534, 32 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 391, 47, 525, 65 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: 1 Rect: 116, 579, 121, 590 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2485 Annotation: Subtype: Link Contents: 1 Rect: 216, 579, 221, 590 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2485 Annotation: Subtype: Link Contents: 1 Rect: 279, 579, 284, 590 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2485 Annotation: Subtype: Link Contents: 1 Rect: 334, 579, 339, 590 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2485 Annotation: Subtype: Link Contents: 1 Rect: 390, 579, 395, 590 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2485 Annotation: Subtype: Link Contents: 1 Rect: 463, 579, 468, 590 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2485 Annotation: Subtype: Link Contents: 1 Rect: 541, 579, 545, 590 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2485 Annotation: Subtype: Link Contents: 2 Rect: 106, 567, 111, 578 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2485 Annotation: Subtype: Link Contents: 3 Rect: 179, 567, 184, 578 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2485 Annotation: Subtype: Link Contents: 1 Rect: 249, 567, 254, 578 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2485 Annotation: Subtype: Link Contents: 4 Rect: 322, 567, 327, 578 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2485 Annotation: Subtype: Link Contents: 5 Rect: 382, 567, 387, 578 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2485 Annotation: Subtype: Link Contents: 1 Rect: 442, 567, 447, 578 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2485 Annotation: Subtype: Link Contents: 1 Rect: 508, 567, 512, 578 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2485 Annotation: Subtype: Link Contents: 17 Rect: 220, 543, 231, 554 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2485 Annotation: Subtype: Link Contents: See the funding table on p. 17 Rect: 391, 125, 477, 134 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2485 Page: Sequence: 2 Annotations: Annotation: Subtype: Link Contents: 10.1128/mbio.02939-25 Rect: 462, 23, 534, 32 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: 1 Rect: 79, 663, 85, 674 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents:  6 Rect: 83, 663, 94, 674 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents: 7 Rect: 260, 639, 266, 650 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents:  9 Rect: 264, 639, 275, 650 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents: 10 Rect: 131, 603, 142, 614 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents: 7, Rect: 216, 567, 228, 578 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents: 9 Rect: 226, 567, 233, 578 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents: 7, Rect: 136, 531, 148, 542 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents: 11 Rect: 146, 531, 157, 542 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents: 12, Rect: 295, 519, 310, 530 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents: 13 Rect: 308, 519, 319, 530 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents: 14 Rect: 175, 495, 186, 506 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents:  17 Rect: 184, 495, 200, 506 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Annotation: Subtype: Link Contents: 15 Rect: 43, 459, 54, 470 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents: 14 Rect: 256, 447, 268, 458 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents:  17 Rect: 266, 447, 281, 458 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Annotation: Subtype: Link Contents: 14, Rect: 108, 411, 123, 422 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents: 18 Rect: 121, 411, 132, 422 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Annotation: Subtype: Link Contents: 1, Rect: 268, 387, 280, 398 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents: 4, Rect: 278, 387, 290, 398 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents: 19 Rect: 288, 387, 300, 398 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Annotation: Subtype: Link Contents:  21 Rect: 298, 387, 313, 398 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Annotation: Subtype: Link Contents: 22 Rect: 75, 351, 86, 362 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Annotation: Subtype: Link Contents: 23 Rect: 192, 327, 203, 338 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Annotation: Subtype: Link Contents: 1, Rect: 198, 303, 210, 314 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents: 24 Rect: 208, 303, 219, 314 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Annotation: Subtype: Link Contents: 25 Rect: 122, 279, 133, 290 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Annotation: Subtype: Link Contents: 26 Rect: 328, 267, 339, 278 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Annotation: Subtype: Link Contents: 27 Rect: 43, 207, 54, 218 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Annotation: Subtype: Link Contents: 28, Rect: 155, 195, 171, 206 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Annotation: Subtype: Link Contents: 29 Rect: 169, 195, 180, 206 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Page: Sequence: 3 Annotations: Annotation: Subtype: Link Contents: ExternalLinking Rect: 197, 74, 234, 83 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 236, 74, 272, 83 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 286, 74, 322, 83 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: 10.1128/mbio.02939-25 Rect: 462, 23, 534, 32 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: Table 1 Rect: 99, 579, 128, 590 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Annotation: Subtype: Link Contents: Table 2 Rect: 342, 567, 371, 578 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Annotation: Subtype: Link Contents: Table 2 Rect: 233, 543, 264, 554 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Annotation: Subtype: Link Contents: Fig. 1A Rect: 200, 447, 229, 458 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Annotation: Subtype: Link Contents: Table 2 Rect: 231, 447, 261, 458 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Annotation: Subtype: Link Contents: Fig. 1B Rect: 283, 411, 310, 422 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Annotation: Subtype: Link Contents: a Rect: 61, 336, 65, 346 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 252 Annotation: Subtype: Link Contents: 14 Rect: 292, 299, 302, 309 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents: 23 Rect: 273, 287, 283, 297 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Annotation: Subtype: Link Contents: 23 Rect: 360, 275, 371, 285 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Annotation: Subtype: Link Contents: 14 Rect: 183, 227, 193, 237 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents: 23 Rect: 215, 131, 225, 141 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Annotation: Subtype: Link Contents: 23 Rect: 182, 95, 192, 105 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Annotation: Subtype: Link Contents: 17, Rect: 300, 65, 313, 74 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Annotation: Subtype: Link Contents: 18 Rect: 311, 65, 320, 74 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Annotation: Subtype: Link Contents: 14 Rect: 270, 57, 279, 66 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents: 30 Rect: 57, 40, 66, 49 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Page: Sequence: 4 Annotations: Annotation: Subtype: Link Contents: ExternalLinking Rect: 324, 446, 361, 455 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 319, 404, 411, 413 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: 10.1128/mbio.02939-25 Rect: 462, 23, 534, 32 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: 31 Rect: 344, 317, 356, 328 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Annotation: Subtype: Link Contents: 18 Rect: 299, 305, 310, 316 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Annotation: Subtype: Link Contents: Table 1 Rect: 100, 245, 130, 256 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Annotation: Subtype: Link Contents: 23 Rect: 165, 221, 176, 232 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Annotation: Subtype: Link Contents: Table 3 Rect: 117, 173, 147, 184 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Annotation: Subtype: Link Contents: Fig. 1B Rect: 226, 149, 253, 160 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Annotation: Subtype: Link Contents: Fig. S1 Rect: 324, 137, 353, 148 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents: Fig. S2 Rect: 197, 89, 226, 100 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents: Table 3 Rect: 232, 89, 264, 100 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Annotation: Subtype: Link Contents: Fig. 1B Rect: 325, 65, 353, 76 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Annotation: Subtype: Link Contents: Table 3 Rect: 43, 41, 74, 52 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Annotation: Subtype: Link Contents: Fig. 1B Rect: 267, 41, 296, 52 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Annotation: Subtype: Link Contents: a Rect: 66, 684, 70, 694 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 486 Annotation: Subtype: Link Contents: b Rect: 142, 684, 146, 694 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 486 Annotation: Subtype: Link Contents: c Rect: 171, 684, 175, 694 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 486 Annotation: Subtype: Link Contents: d Rect: 257, 684, 262, 694 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 486 Annotation: Subtype: Link Contents: e Rect: 333, 684, 337, 694 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 486 Annotation: Subtype: Link Contents: f Rect: 439, 684, 442, 694 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 486 Annotation: Subtype: Link Contents: Fig. 1A Rect: 159, 412, 180, 421 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Page: Sequence: 5 Annotations: Annotation: Subtype: Link Contents: ExternalLinking Rect: 233, 75, 280, 86 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: 10.1128/mbio.02939-25 Rect: 462, 23, 534, 32 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: Fig. S2 Rect: 314, 99, 340, 110 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents: Table 3 Rect: 342, 99, 371, 110 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Annotation: Subtype: Link Contents: Table 1 Rect: 489, 237, 515, 247 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Annotation: Subtype: Link Contents: Table 1 Rect: 81, 141, 105, 151 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Page: Sequence: 6 Annotations: Annotation: Subtype: Link Contents: ExternalLinking Rect: 43, 339, 90, 350 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 327, 458, 363, 467 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 319, 416, 411, 425 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: 10.1128/mbio.02939-25 Rect: 462, 23, 534, 32 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: Fig. S3 Rect: 242, 101, 270, 112 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents: 32 Rect: 333, 77, 345, 88 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Annotation: Subtype: Link Contents: 1 Rect: 41, 41, 376, 64 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Annotation: Subtype: Link Contents: a Rect: 63, 685, 67, 694 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 1193 Annotation: Subtype: Link Contents: b Rect: 145, 685, 149, 694 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 1193 Annotation: Subtype: Link Contents: c Rect: 168, 685, 172, 694 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 1193 Annotation: Subtype: Link Contents: d Rect: 262, 685, 266, 694 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 1193 Annotation: Subtype: Link Contents: e Rect: 342, 685, 346, 694 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 1193 Annotation: Subtype: Link Contents: f Rect: 443, 685, 446, 694 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 1193 Annotation: Subtype: Link Contents: Table 2 Rect: 273, 466, 296, 475 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Annotation: Subtype: Link Contents: Fig. S2 Rect: 158, 424, 179, 433 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Page: Sequence: 7 Annotations: Annotation: Subtype: Link Contents: 10.1128/mbio.02939-25 Rect: 462, 23, 534, 32 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: Table 1 Rect: 81, 585, 110, 596 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Annotation: Subtype: Link Contents: Fig. S4 Rect: 98, 549, 124, 560 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents: Table S1 Rect: 128, 513, 161, 524 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents: Table S1 Rect: 65, 501, 100, 512 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents: Fig. S5 Rect: 43, 489, 69, 500 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents: Table S1 Rect: 84, 429, 119, 440 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents: 33, Rect: 184, 405, 199, 416 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Annotation: Subtype: Link Contents: 34 Rect: 197, 405, 208, 416 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Annotation: Subtype: Link Contents: Table 1 Rect: 341, 279, 371, 290 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Annotation: Subtype: Link Contents: Fig. 1C Rect: 179, 231, 209, 242 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Annotation: Subtype: Link Contents: Fig. S6 Rect: 307, 219, 335, 230 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents: Table 2 Rect: 238, 195, 269, 206 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Annotation: Subtype: Link Contents: Fig. 1C and D Rect: 43, 99, 95, 110 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Annotation: Subtype: Link Contents: Fig. S4 Rect: 232, 87, 258, 98 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Page: Sequence: 8 Annotations: Annotation: Subtype: Link Contents: ExternalLinking Rect: 214, 161, 256, 172 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: 10.1128/mbio.02939-25 Rect: 462, 23, 534, 32 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: Fig. S1 Rect: 150, 657, 177, 668 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents: Fig. 1E Rect: 130, 633, 157, 644 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Annotation: Subtype: Link Contents: 14 Rect: 254, 463, 265, 474 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents: Table 1 Rect: 156, 403, 188, 414 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Annotation: Subtype: Link Contents: Fig. 2A Rect: 345, 391, 374, 402 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Annotation: Subtype: Link Contents: Table S2 Rect: 41, 379, 75, 390 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents: 14 Rect: 249, 319, 260, 330 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents: Fig. 2B Rect: 151, 295, 179, 306 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Annotation: Subtype: Link Contents: Table 1 Rect: 212, 283, 242, 294 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Annotation: Subtype: Link Contents: Fig. 2C Rect: 290, 271, 318, 282 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Annotation: Subtype: Link Contents: Fig. 2B Rect: 182, 259, 208, 270 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Annotation: Subtype: Link Contents: Fig. 2D Rect: 343, 259, 371, 270 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Annotation: Subtype: Link Contents: 35 Rect: 339, 149, 351, 160 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Annotation: Subtype: Link Contents: Fig. 1B Rect: 43, 113, 71, 124 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Annotation: Subtype: Link Contents: Fig. 3A Rect: 115, 89, 144, 100 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Annotation: Subtype: Link Contents: Fig. S7 Rect: 147, 89, 175, 100 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents: Fig. 3A Rect: 320, 65, 347, 76 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Page: Sequence: 9 Annotations: Annotation: Subtype: Link Contents: 10.1128/mbio.02939-25 Rect: 462, 23, 534, 32 LastModified: D:20260204101832+00'00' Flags: Print Page: Sequence: 10 Annotations: Annotation: Subtype: Link Contents: ExternalLinking Rect: 105, 271, 121, 280 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 266, 271, 304, 280 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: 10.1128/mbio.02939-25 Rect: 462, 23, 534, 32 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: Fig. 3B Rect: 223, 144, 249, 155 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Annotation: Subtype: Link Contents: Fig. 3C Rect: 240, 120, 267, 131 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Page: Sequence: 11 Annotations: Annotation: Subtype: Link Contents: 10.1128/mbio.02939-25 Rect: 462, 23, 534, 32 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: Fig. 3A Rect: 59, 621, 86, 632 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Annotation: Subtype: Link Contents: Fig. 1C Rect: 341, 609, 371, 620 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Annotation: Subtype: Link Contents: 3B Rect: 41, 573, 376, 596 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Annotation: Subtype: Link Contents: Fig. 1D Rect: 277, 549, 305, 560 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Annotation: Subtype: Link Contents: Fig. 1B Rect: 166, 435, 195, 446 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Annotation: Subtype: Link Contents: Fig. 1B Rect: 81, 411, 108, 422 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Annotation: Subtype: Link Contents: Table 2 Rect: 109, 411, 138, 422 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Annotation: Subtype: Link Contents: 3B Rect: 41, 375, 376, 398 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Annotation: Subtype: Link Contents: Fig. 1C Rect: 183, 363, 210, 374 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Annotation: Subtype: Link Contents: Fig. 1A Rect: 227, 339, 255, 350 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Annotation: Subtype: Link Contents: Fig. S2 Rect: 259, 291, 286, 302 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents: Table 3 Rect: 288, 291, 317, 302 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Annotation: Subtype: Link Contents: 1 Rect: 155, 165, 162, 176 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents: 21, Rect: 348, 153, 364, 164 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Annotation: Subtype: Link Contents: 28, Rect: 362, 153, 376, 164 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Annotation: Subtype: Link Contents: 29 Rect: 41, 141, 52, 152 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Page: Sequence: 12 Annotations: Annotation: Subtype: Link Contents: 10.1128/mbio.02939-25 Rect: 462, 23, 534, 32 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: 33, Rect: 346, 621, 362, 632 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Annotation: Subtype: Link Contents: 34 Rect: 360, 621, 371, 632 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Annotation: Subtype: Link Contents: 36 Rect: 272, 431, 283, 442 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Annotation: Subtype: Link Contents:  38 Rect: 281, 431, 297, 442 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Annotation: Subtype: Link Contents: 39 Rect: 116, 395, 127, 406 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Annotation: Subtype: Link Contents: 22, Rect: 332, 287, 348, 298 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Annotation: Subtype: Link Contents: 40, Rect: 346, 287, 362, 298 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Annotation: Subtype: Link Contents: 41 Rect: 360, 287, 371, 298 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Annotation: Subtype: Link Contents: 42 Rect: 140, 239, 151, 250 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Annotation: Subtype: Link Contents: 14 Rect: 43, 113, 54, 124 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents: 43 Rect: 328, 89, 339, 100 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Annotation: Subtype: Link Contents:  47 Rect: 337, 89, 353, 100 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Page: Sequence: 13 Annotations: Annotation: Subtype: Link Contents: 10.1128/mbio.02939-25 Rect: 462, 23, 534, 32 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: 48 Rect: 105, 675, 116, 686 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Annotation: Subtype: Link Contents:  52 Rect: 114, 675, 130, 686 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 3555 Annotation: Subtype: Link Contents: 26 Rect: 360, 639, 371, 650 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Annotation: Subtype: Link Contents: 10 Rect: 79, 461, 90, 472 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents: 53 Rect: 223, 437, 235, 448 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 3555 Annotation: Subtype: Link Contents:  56 Rect: 233, 437, 248, 448 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 3555 Annotation: Subtype: Link Contents: 23 Rect: 360, 425, 371, 436 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Annotation: Subtype: Link Contents: 57 Rect: 104, 299, 116, 310 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 3555 Annotation: Subtype: Link Contents:  59 Rect: 114, 299, 129, 310 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 3555 Annotation: Subtype: Link Contents: Table 1 Rect: 111, 41, 143, 52 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: null Annotation: Subtype: Link Contents: 60 Rect: 362, 41, 373, 52 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 3555 Page: Sequence: 14 Annotations: Annotation: Subtype: Link Contents: 10.1128/mbio.02939-25 Rect: 462, 23, 534, 32 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: 61 Rect: 235, 555, 246, 566 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 3555 Annotation: Subtype: Link Contents: 34 Rect: 360, 513, 371, 524 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Annotation: Subtype: Link Contents: 23 Rect: 217, 489, 228, 500 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Annotation: Subtype: Link Contents: 62 Rect: 137, 477, 149, 488 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 3555 Annotation: Subtype: Link Contents: 13 Rect: 43, 417, 54, 428 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents: 63 Rect: 273, 369, 285, 380 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 3555 Annotation: Subtype: Link Contents: Table S3 Rect: 97, 357, 130, 368 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents: Table S3 Rect: 226, 345, 259, 356 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents: 23 Rect: 82, 141, 93, 152 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Page: Sequence: 15 Annotations: Annotation: Subtype: Link Contents: 10.1128/mbio.02939-25 Rect: 462, 23, 534, 32 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: 34 Rect: 159, 669, 170, 680 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Annotation: Subtype: Link Contents: 34 Rect: 274, 657, 285, 668 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Annotation: Subtype: Link Contents: Table S3 Rect: 280, 645, 316, 656 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents: Table S3 Rect: 299, 621, 333, 632 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents: Table S3 Rect: 226, 537, 261, 548 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents: Table S3 Rect: 267, 489, 301, 500 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents: 34 Rect: 279, 453, 290, 464 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2958 Annotation: Subtype: Link Contents: 64 Rect: 83, 377, 94, 388 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 3555 Annotation: Subtype: Link Contents: 64 Rect: 317, 341, 328, 352 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 3555 Annotation: Subtype: Link Contents: 14 Rect: 312, 137, 324, 148 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 2665 Annotation: Subtype: Link Contents: 64 Rect: 330, 65, 341, 76 LastModified: D:20260204101832+00'00' Flags: Print ActionDest: 3555 Page: Sequence: 16 Annotations: Annotation: Subtype: Link Contents: ExternalLinking Rect: 199, 537, 246, 548 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 250, 537, 296, 548 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 43, 323, 90, 334 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: 10.1128/mbio.02939-25 Rect: 462, 23, 534, 32 LastModified: D:20260204101832+00'00' Flags: Print Page: Sequence: 17 Annotations: Annotation: Subtype: Link Contents: http://orcid.org/0000-0003-3789-4418 Rect: 135, 446, 280, 457 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: http://orcid.org/0000-0002-8001-6314 Rect: 95, 433, 240, 444 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: http://orcid.org/0000-0002-6064-7899 Rect: 103, 420, 248, 431 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: http://orcid.org/0000-0003-0660-1854 Rect: 111, 407, 256, 418 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 41, 351, 170, 362 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 41, 221, 200, 252 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: 10.1128/mbio.02939-25 Rect: 462, 23, 534, 32 LastModified: D:20260204101832+00'00' Flags: Print Page: Sequence: 18 Annotations: Annotation: Subtype: Link Contents: ExternalLinking Rect: 312, 537, 357, 548 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 256, 513, 303, 524 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 100, 501, 153, 512 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 167, 501, 219, 512 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 311, 501, 364, 512 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 41, 489, 93, 500 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 189, 489, 242, 500 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 255, 489, 307, 500 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 169, 435, 196, 446 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 59, 263, 287, 282 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 59, 227, 287, 246 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 59, 200, 287, 219 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 151, 173, 268, 183 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 59, 137, 287, 156 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 59, 101, 287, 120 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 100, 74, 223, 84 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 59, 47, 287, 66 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 361, 263, 501, 273 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 318, 236, 546, 255 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 349, 200, 473, 210 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 395, 164, 527, 174 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 318, 119, 546, 138 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 377, 74, 501, 84 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: 10.1128/mbio.02939-25 Rect: 462, 23, 534, 32 LastModified: D:20260204101832+00'00' Flags: Print Page: Sequence: 19 Annotations: Annotation: Subtype: Link Contents: ExternalLinking Rect: 137, 692, 249, 702 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 59, 656, 287, 675 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 59, 611, 287, 630 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 59, 566, 287, 585 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 59, 539, 287, 558 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 59, 503, 287, 522 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 59, 467, 287, 486 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 79, 422, 220, 432 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 102, 377, 224, 387 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 117, 341, 249, 351 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 83, 305, 217, 315 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 59, 269, 287, 288 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 59, 242, 287, 261 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 151, 215, 263, 225 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 104, 170, 227, 180 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 59, 125, 287, 144 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 59, 89, 287, 108 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 59, 44, 287, 63 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 318, 674, 546, 693 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 318, 638, 546, 657 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 318, 593, 546, 612 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 318, 557, 546, 576 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 318, 512, 546, 531 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 318, 476, 546, 495 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 318, 386, 546, 405 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 318, 341, 546, 360 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 357, 296, 497, 306 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 318, 269, 546, 288 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 318, 233, 546, 252 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 357, 197, 511, 207 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 361, 161, 506, 171 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 318, 125, 546, 144 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 328, 98, 449, 108 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 318, 62, 546, 81 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: 10.1128/mbio.02939-25 Rect: 462, 23, 534, 32 LastModified: D:20260204101832+00'00' Flags: Print Page: Sequence: 20 Annotations: Annotation: Subtype: Link Contents: ExternalLinking Rect: 59, 692, 287, 711 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 59, 665, 287, 684 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 59, 629, 287, 648 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 143, 584, 287, 594 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 118, 548, 250, 558 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 59, 503, 287, 522 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 126, 467, 266, 477 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 108, 422, 238, 432 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 318, 656, 546, 675 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 353, 620, 483, 630 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 318, 539, 546, 558 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 318, 494, 546, 513 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 364, 467, 479, 477 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: ExternalLinking Rect: 318, 431, 546, 450 LastModified: D:20260204101832+00'00' Flags: Print Annotation: Subtype: Link Contents: 10.1128/mbio.02939-25 Rect: 462, 23, 534, 32 LastModified: D:20260204101832+00'00' Flags: Print Checksum: 6f91c0cc Type: CRC32 Checksum: b1ab3c7b27ebd4063774d98fb655024e Type: MD5 Checksum: 2ef1314c8e43a0b5117c107ef5aecbe759e647e3 Type: SHA-1 Checksum: 7423bd7d0e0879abbf33521b86288eb0a58d07535ec2ed805bf48105291cfbe6 Type: SHA-256