712243a001675754261c21430668e182 pmc_31740730.pdf cbedbdf6e189be9151ccf7ca6af796b1fec2cbc4 pmc_31740730.pdf 514d04db02db12fbf10af7a37fe7649407323b9d46fa0111ae84238b62467df9 pmc_31740730.pdf Title: The molecular genealogy of sequential overlapping inversions implies both homologous chromosomes of a heterokaryotype in an inversion origin Subject: Scientific Reports, doi:10.1038/s41598-019-53582-8 Keywords: Author: Dorcas J. Orengo Creator: Springer Producer: iText® 5.3.5 ©2000-2012 1T3XT BVBA (AGPL-version) CreationDate: Sat Nov 9 04:00:28 2019 CET ModDate: Tue Nov 12 19:14:16 2019 CET Tagged: no UserProperties: no Suspects: no Form: AcroForm JavaScript: no Pages: 11 Encrypted: no Page size: 595.276 x 782.362 pts Page rot: 0 File size: 2635744 bytes Optimized: no PDF version: 1.4 name type encoding emb sub uni object ID ------------------------------------ ----------------- ---------------- --- --- --- --------- IBNBQX+Corbel-Bold CID TrueType Identity-H yes yes yes 107 0 RISBCK+Corbel-BoldItalic Type 1C WinAnsi yes yes yes 442 0 MinionPro-Regular Type 1C Custom yes no yes 67 0 MDMBDO+Corbel-Bold2 Type 1C WinAnsi yes yes yes 68 0 MinionPro-It Type 1C Custom yes no yes 70 0 CMCSVM+Corbel2 Type 1C WinAnsi yes yes yes 71 0 DBECYH+Corbel CID TrueType Identity-H yes yes yes 72 0 MyriadPro-It Type 1C WinAnsi yes no yes 73 0 EuclidSymbol Type 1C Custom yes no yes 66 0 MinionPro-Bold Type 1C WinAnsi yes no yes 69 0 HLCEXZ+TimesNewRomanPSMT Type 1C WinAnsi yes yes yes 284 0 Jhove (Rel. 1.22.1, 2019-04-17) Date: 2020-08-10 12:17:29 CEST RepresentationInformation: pmc_31740730.pdf ReportingModule: PDF-hul, Rel. 1.12.1 (2019-04-17) LastModified: 2020-07-06 04:06:31 CEST Size: 2635744 Format: PDF Version: 1.4 Status: Well-Formed, but not valid SignatureMatches: PDF-hul ID: PDF-HUL-122 ErrorMessage: edu.harvard.hul.ois.jhove.module.pdf.PdfInvalidException: Invalid indirect destination - referenced object 'SpringerNature_SciRep_53582.indd:Figure 1. Schematic representation of chromosomal arrangements Est, E1+2, E1+2+9, E1+2+9+3, and E1+2+9+12 of Drosophila subobscura. Inversions originating these arrangements are indicated by crossed lines. Breakpoint regions :51' cannot be found Offset: 2136288 ID: PDF-HUL-122 ErrorMessage: edu.harvard.hul.ois.jhove.module.pdf.PdfInvalidException: Invalid indirect destination - referenced object 'SpringerNature_SciRep_53582.indd:Figure 1. Schematic representation of chromosomal arrangements Est, E1+2, E1+2+9, E1+2+9+3, and E1+2+9+12 of Drosophila subobscura. Inversions originating these arrangements are indicated by crossed lines. Breakpoint region:57' cannot be found Offset: 2136288 ID: PDF-HUL-122 ErrorMessage: edu.harvard.hul.ois.jhove.module.pdf.PdfInvalidException: Invalid indirect destination - referenced object 'SpringerNature_SciRep_53582.indd:Figure 2. Functional annotation of breakpoint regions spanning fragment A. Annotation extracted from our previous work28,32. Breakpoint regions are named as in Fig. 1 except for the GAL region that is here presented in reverse orientatio:58' cannot be found Offset: 2136288 ID: PDF-HUL-122 ErrorMessage: edu.harvard.hul.ois.jhove.module.pdf.PdfInvalidException: Invalid indirect destination - referenced object 'SpringerNature_SciRep_53582.indd:Figure 1. Schematic representation of chromosomal arrangements Est, E1+2, E1+2+9, E1+2+9+3, and E1+2+9+12 of Drosophila subobscura. Inversions originating these arrangements are indicated by crossed lines. Breakpoint regio:59' cannot be found Offset: 2136288 ID: PDF-HUL-122 ErrorMessage: edu.harvard.hul.ois.jhove.module.pdf.PdfInvalidException: Invalid indirect destination - referenced object 'SpringerNature_SciRep_53582.indd:Figure 1. Schematic representation of chromosomal arrangements Est, E1+2, E1+2+9, E1+2+9+3, and E1+2+9+12 of Drosophila subobscura. Inversions originating these arrangements are indicated by crossed lines. Breakpoint reg:66' cannot be found Offset: 2136288 ID: PDF-HUL-122 ErrorMessage: edu.harvard.hul.ois.jhove.module.pdf.PdfInvalidException: Invalid indirect destination - referenced object 'SpringerNature_SciRep_53582.indd:Figure 2. Functional annotation of breakpoint regions spanning fragment A. Annotation extracted from our previous work28,32. Breakpoint regions are named as in Fig. 1 except for the GAL region that is here presented in reverse orientat:67' cannot be found Offset: 2136288 ID: PDF-HUL-122 ErrorMessage: edu.harvard.hul.ois.jhove.module.pdf.PdfInvalidException: Invalid indirect destination - referenced object 'SpringerNature_SciRep_53582.indd:Figure 1. Schematic representation of chromosomal arrangements Est, E1+2, E1+2+9, E1+2+9+3, and E1+2+9+12 of Drosophila subobscura. Inversions originating these arrangements are indicated by crossed lines. Breakpoint regi:63' cannot be found Offset: 2136288 ID: PDF-HUL-122 ErrorMessage: edu.harvard.hul.ois.jhove.module.pdf.PdfInvalidException: Invalid indirect destination - referenced object 'SpringerNature_SciRep_53582.indd:Figure 1. Schematic representation of chromosomal arrangements Est, E1+2, E1+2+9, E1+2+9+3, and E1+2+9+12 of Drosophila subobscura. Inversions originating these arrangements are indicated by crossed lines. Breakpoint re:71' cannot be found Offset: 2136288 ID: PDF-HUL-122 ErrorMessage: edu.harvard.hul.ois.jhove.module.pdf.PdfInvalidException: Invalid indirect destination - referenced object 'SpringerNature_SciRep_53582.indd:Figure 2. Functional annotation of breakpoint regions spanning fragment A. Annotation extracted from our previous work28,32. Breakpoint regions are named as in Fig. 1 except for the GAL region that is here presented in reverse orienta:72' cannot be found Offset: 2136288 ID: PDF-HUL-122 ErrorMessage: edu.harvard.hul.ois.jhove.module.pdf.PdfInvalidException: Invalid indirect destination - referenced object 'SpringerNature_SciRep_53582.indd:Figure 1. Schematic representation of chromosomal arrangements Est, E1+2, E1+2+9, E1+2+9+3, and E1+2+9+12 of Drosophila subobscura. Inversions originating these arrangements are indicated by crossed lines. Breakpoint r:73' cannot be found Offset: 2136288 ID: PDF-HUL-122 ErrorMessage: edu.harvard.hul.ois.jhove.module.pdf.PdfInvalidException: Invalid indirect destination - referenced object 'SpringerNature_SciRep_53582.indd: Table 1. Nucleotide polymorphism and divergence at fragment A from different breakpoint regions. aAfter excluding sites with alignment gaps. Some more sites were excluded to estimate K due to additional alignment gaps.:75' cannot be found Offset: 2136288 ID: PDF-HUL-122 ErrorMessage: edu.harvard.hul.ois.jhove.module.pdf.PdfInvalidException: Invalid indirect destination - referenced object 'SpringerNature_SciRep_53582.indd: Table 1. Nucleotide polymorphism and divergence at fragment A from different breakpoint regions. aAfter excluding sites with alignment gaps. Some more sites were excluded to estimate K due to additional alignment gaps:78' cannot be found Offset: 2136288 ID: PDF-HUL-122 ErrorMessage: edu.harvard.hul.ois.jhove.module.pdf.PdfInvalidException: Invalid indirect destination - referenced object 'SpringerNature_SciRep_53582.indd:Figure 1. Schematic representation of chromosomal arrangements Est, E1+2, E1+2+9, E1+2+9+3, and E1+2+9+12 of Drosophila subobscura. Inversions originating these arrangements are indicated by crossed lines. Breakpoint :79' cannot be found Offset: 2136288 ID: PDF-HUL-122 ErrorMessage: edu.harvard.hul.ois.jhove.module.pdf.PdfInvalidException: Invalid indirect destination - referenced object 'SpringerNature_SciRep_53582.indd: Table 2. Genetic differentiation between the different breakpoint regions. FST estimates between each pair of chromosomal regions are shown in the lower part of the matrix and the corresponding P-values obtained from:80' cannot be found Offset: 2136288 ID: PDF-HUL-122 ErrorMessage: edu.harvard.hul.ois.jhove.module.pdf.PdfInvalidException: Invalid indirect destination - referenced object 'SpringerNature_SciRep_53582.indd:Figure 1. Schematic representation of chromosomal arrangements Est, E1+2, E1+2+9, E1+2+9+3, and E1+2+9+12 of Drosophila subobscura. Inversions originating these arrangements are indicated by crossed lines. Breakpoint:81' cannot be found Offset: 2136288 ID: PDF-HUL-122 ErrorMessage: edu.harvard.hul.ois.jhove.module.pdf.PdfInvalidException: Invalid indirect destination - referenced object 'SpringerNature_SciRep_53582.indd:Figure 1. Schematic representation of chromosomal arrangements Est, E1+2, E1+2+9, E1+2+9+3, and E1+2+9+12 of Drosophila subobscura. Inversions originating these arrangements are indicated by crossed lines. Breakpoin:83' cannot be found Offset: 2136288 ID: PDF-HUL-122 ErrorMessage: edu.harvard.hul.ois.jhove.module.pdf.PdfInvalidException: Invalid indirect destination - referenced object 'SpringerNature_SciRep_53582.indd:Figure 1. Schematic representation of chromosomal arrangements Est, E1+2, E1+2+9, E1+2+9+3, and E1+2+9+12 of Drosophila subobscura. Inversions originating these arrangements are indicated by crossed lines. Breakpoi:84' cannot be found Offset: 2136288 ID: PDF-HUL-122 ErrorMessage: edu.harvard.hul.ois.jhove.module.pdf.PdfInvalidException: Invalid indirect destination - referenced object 'SpringerNature_SciRep_53582.indd:Figure 3. Neighbor-joining tree of the A fragment sequences corresponding to the breakpoint regions of different E chromosomal arrangements. Bootstrap values >70% (based on 1000 replicates) are shown on the tree. Positions with over 5% alignment:85' cannot be found Offset: 2136288 ID: PDF-HUL-122 ErrorMessage: edu.harvard.hul.ois.jhove.module.pdf.PdfInvalidException: Invalid indirect destination - referenced object 'SpringerNature_SciRep_53582.indd:Figure 1. Schematic representation of chromosomal arrangements Est, E1+2, E1+2+9, E1+2+9+3, and E1+2+9+12 of Drosophila subobscura. Inversions originating these arrangements are indicated by crossed lines. Breakpo:87' cannot be found Offset: 2136288 ID: PDF-HUL-122 ErrorMessage: edu.harvard.hul.ois.jhove.module.pdf.PdfInvalidException: Invalid indirect destination - referenced object 'SpringerNature_SciRep_53582.indd:Figure 1. Schematic representation of chromosomal arrangements Est, E1+2, E1+2+9, E1+2+9+3, and E1+2+9+12 of Drosophila subobscura. Inversions originating these arrangements are indicated by crossed lines. Breakp:91' cannot be found Offset: 2136288 ID: PDF-HUL-122 ErrorMessage: edu.harvard.hul.ois.jhove.module.pdf.PdfInvalidException: Invalid indirect destination - referenced object 'SpringerNature_SciRep_53582.indd:Figure 2. Functional annotation of breakpoint regions spanning fragment A. Annotation extracted from our previous work28,32. Breakpoint regions are named as in Fig. 1 except for the GAL region that is here presented in reverse orient:93' cannot be found Offset: 2136288 ID: PDF-HUL-122 ErrorMessage: edu.harvard.hul.ois.jhove.module.pdf.PdfInvalidException: Invalid indirect destination - referenced object 'SpringerNature_SciRep_53582.indd: 32. Orengo, D. J., Puerma, E., Papaceit, M., Segarra, C. & Aguadé, M. A molecular perspective on a complex polymorphic inversion system with cytological evidence of multiply reused breakpoints. Heredity (Edinb). 114, 610–618 (2015). :95' cannot be found Offset: 2136288 ID: PDF-HUL-122 ErrorMessage: edu.harvard.hul.ois.jhove.module.pdf.PdfInvalidException: Invalid indirect destination - referenced object 'SpringerNature_SciRep_53582.indd:Figure 2. Functional annotation of breakpoint regions spanning fragment A. Annotation extracted from our previous work28,32. Breakpoint regions are named as in Fig. 1 except for the GAL region that is here presented in reverse orien:96' cannot be found Offset: 2136288 ID: PDF-HUL-122 ErrorMessage: edu.harvard.hul.ois.jhove.module.pdf.PdfInvalidException: Invalid indirect destination - referenced object 'SpringerNature_SciRep_53582.indd:Figure 2. Functional annotation of breakpoint regions spanning fragment A. Annotation extracted from our previous work28,32. Breakpoint regions are named as in Fig. 1 except for the GAL region that is here presented in reverse orie:98' cannot be found Offset: 2136288 ID: PDF-HUL-122 ErrorMessage: edu.harvard.hul.ois.jhove.module.pdf.PdfInvalidException: Invalid indirect destination - referenced object 'SpringerNature_SciRep_53582.indd:Figure 2. Functional annotation of breakpoint regions spanning fragment A. Annotation extracted from our previous work28,32. Breakpoint regions are named as in Fig. 1 except for the GAL region that is here presented in reverse ori:100' cannot be found Offset: 2136288 ID: PDF-HUL-122 ErrorMessage: edu.harvard.hul.ois.jhove.module.pdf.PdfInvalidException: Invalid indirect destination - referenced object 'SpringerNature_SciRep_53582.indd:Figure 4. Schematic representation of the NHEJ-4-chromosome model for the origin of inversion E9. The sequential steps of how arrangement E1+2+9 could have originated from an Est/E1+2 heterokaryotypic individual through inversion E9:105' cannot be found Offset: 2136288 ID: PDF-HUL-122 ErrorMessage: edu.harvard.hul.ois.jhove.module.pdf.PdfInvalidException: Invalid indirect destination - referenced object 'SpringerNature_SciRep_53582.indd:Figure 5. Schematic representation of the NHEJ-3-chromosome model for the origin of inversion E9. The sequential steps of how arrangement E1+2+9 could have originated from an Est/E1+2 heterokaryotypic individual through inversion E9:107' cannot be found Offset: 2136288 ID: PDF-HUL-122 ErrorMessage: edu.harvard.hul.ois.jhove.module.pdf.PdfInvalidException: Invalid indirect destination - referenced object 'SpringerNature_SciRep_53582.indd:Figure 6. Schematic representation of the BIR-NHEJ-chromosome model for the origin of inversion E9. The sequential steps of how arrangement E1+2+9 could have originated from an Est/E1+2 heterokaryotypic individual through inversion E:111' cannot be found Offset: 2136288 ID: PDF-HUL-122 ErrorMessage: edu.harvard.hul.ois.jhove.module.pdf.PdfInvalidException: Invalid indirect destination - referenced object 'SpringerNature_SciRep_53582.indd:Figure 4. Schematic representation of the NHEJ-4-chromosome model for the origin of inversion E9. The sequential steps of how arrangement E1+2+9 could have originated from an Est/E1+2 heterokaryotypic individual through inversion E:114' cannot be found Offset: 2136288 ID: PDF-HUL-122 ErrorMessage: edu.harvard.hul.ois.jhove.module.pdf.PdfInvalidException: Invalid indirect destination - referenced object 'SpringerNature_SciRep_53582.indd:Figure 2. Functional annotation of breakpoint regions spanning fragment A. Annotation extracted from our previous work28,32. Breakpoint regions are named as in Fig. 1 except for the GAL region that is here presented in reverse or:116' cannot be found Offset: 2136288 MIMEtype: application/pdf PDFMetadata: Objects: 479 FreeObjects: 1 IncrementalUpdates: 0 DocumentCatalog: ViewerPreferences: HideToolbar: false HideMenubar: false HideWindowUI: false FitWindow: false CenterWindow: false DisplayDocTitle: false NonFullScreenPageMode: UseNone Direction: L2R ViewArea: CropBox ViewClip: CropBox PrintArea: CropBox PageClip: CropBox PageLayout: SinglePage PageMode: UseThumbs Language: EN Outlines: Item: Title: The molecular genealogy of sequential overlapping inversions implies both homologous chromosomes of a heterokaryotype in an ... Children: Item: Title: Results and Discussion Children: Item: Title: Nucleotide variation at a fragment flanking multiple inversion breakpoints. Item: Title: Genetic differentiation between the A fragments of the different breakpoint regions. Item: Title: Inversion E9 originated in an inversion heterokaryotype. Item: Title: New models to explain the origin of inversion E9. Item: Title: Materials and Methods Item: Title: Acknowledgements Item: Title: Figure 1 Schematic representation of chromosomal arrangements Est, E1+2, E1+2+9, E1+2+9+3, and E1+2+9+12 of Drosophila subobscura. Item: Title: Figure 2 Functional annotation of breakpoint regions spanning fragment A. Item: Title: Figure 3 Neighbor-joining tree of the A fragment sequences corresponding to the breakpoint regions of different E chromosomal arrangements. Item: Title: Figure 4 Schematic representation of the NHEJ-4-chromosome model for the origin of inversion E9. Item: Title: Figure 5 Schematic representation of the NHEJ-3-chromosome model for the origin of inversion E9. Item: Title: Figure 6 Schematic representation of the BIR-NHEJ-chromosome model for the origin of inversion E9. Item: Title: Table 1 Nucleotide polymorphism and divergence at fragment A from different breakpoint regions. Item: Title: Table 2 Genetic differentiation between the different breakpoint regions. Info: Title: The molecular genealogy of sequential overlapping inversions implies both homologous chromosomes of a heterokaryotype in an inversion origin Author: Dorcas J. Orengo Subject: Scientific Reports, doi:10.1038/s41598-019-53582-8 Keywords: Creator: Springer Producer: iText® 5.3.5 ©2000-2012 1T3XT BVBA (AGPL-version) CreationDate: Sat Nov 09 04:30:28 CET 2019 ModDate: Tue Nov 12 19:14:16 CET 2019 ID: 0xfd328a1e4dd72c15cdead7e3d9a1d535, 0x62720f28bb5e06e92cbc4e537dffa713 Filters: FilterPipeline: FlateDecode FilterPipeline: DCTDecode Images: Image: NisoImageMetadata: FormatName: image/jpg CompressionScheme: JPEG ImageWidth: 1900 ImageHeight: 1003 BitsPerSample: 8 BitsPerSampleUnit: integer Intent: RelativeColorimetric Name: X Image: NisoImageMetadata: FormatName: image/png ImageWidth: 1350 ImageHeight: 942 BitsPerSample: 8 BitsPerSampleUnit: integer Filter: FlateDecode Intent: RelativeColorimetric Name: X Image: NisoImageMetadata: FormatName: image/png ImageWidth: 1281 ImageHeight: 2300 BitsPerSample: 8 BitsPerSampleUnit: integer Filter: FlateDecode Intent: RelativeColorimetric Name: X Image: NisoImageMetadata: FormatName: image/jpg CompressionScheme: JPEG ImageWidth: 1900 ImageHeight: 1069 BitsPerSample: 8 BitsPerSampleUnit: integer Intent: RelativeColorimetric Name: X Image: NisoImageMetadata: FormatName: image/png ImageWidth: 1900 ImageHeight: 1108 BitsPerSample: 8 BitsPerSampleUnit: integer Filter: FlateDecode Intent: RelativeColorimetric Name: X Image: NisoImageMetadata: FormatName: image/png ImageWidth: 1900 ImageHeight: 1043 BitsPerSample: 8 BitsPerSampleUnit: integer Filter: FlateDecode Intent: RelativeColorimetric Name: X Image: NisoImageMetadata: FormatName: image/png ImageWidth: 1100 ImageHeight: 90 ColorSpace: black is zero BitsPerSample: 1 BitsPerSampleUnit: integer Filter: FlateDecode Intent: RelativeColorimetric Name: X Fonts: Type0: Font: BaseFont: DBECYH+Corbel Encoding: Identity-H ToUnicode: true Font: BaseFont: IBNBQX+Corbel-Bold Encoding: Identity-H ToUnicode: true Type1: Font: BaseFont: EuclidSymbol FirstChar: 0 LastChar: 255 FontDescriptor: FontName: EuclidSymbol Flags: Symbolic FontBBox: -166, -253, 1090, 950 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true ToUnicode: true Font: BaseFont: MinionPro-Regular FirstChar: 0 LastChar: 255 FontDescriptor: FontName: MinionPro-Regular Flags: Serif, Nonsymbolic FontBBox: -290, -360, 1684, 989 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true ToUnicode: true Font: BaseFont: MDMBDO+Corbel-Bold2 FontSubset: true FirstChar: 0 LastChar: 255 FontDescriptor: FontName: MDMBDO+Corbel-Bold2 Flags: Nonsymbolic FontBBox: -556, -239, 1155, 952 FontFile3: true Encoding: WinAnsiEncoding ToUnicode: true Font: BaseFont: MinionPro-Bold FirstChar: 0 LastChar: 255 FontDescriptor: FontName: MinionPro-Bold Flags: Serif, Nonsymbolic FontBBox: -319, -360, 1684, 1024 FontFile3: true Encoding: WinAnsiEncoding ToUnicode: true Font: BaseFont: MinionPro-It FirstChar: 0 LastChar: 255 FontDescriptor: FontName: MinionPro-It Flags: Serif, Nonsymbolic, Italic FontBBox: -201, -360, 1684, 1002 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true ToUnicode: true Font: BaseFont: CMCSVM+Corbel2 FontSubset: true FirstChar: 0 LastChar: 255 FontDescriptor: FontName: CMCSVM+Corbel2 Flags: Nonsymbolic FontBBox: -553, -239, 1110, 952 FontFile3: true Encoding: WinAnsiEncoding ToUnicode: true Font: BaseFont: MyriadPro-It FirstChar: 0 LastChar: 255 FontDescriptor: FontName: MyriadPro-It Flags: Nonsymbolic, Italic FontBBox: -185, -250, 1090, 953 FontFile3: true Encoding: WinAnsiEncoding ToUnicode: true Font: BaseFont: RISBCK+Corbel-BoldItalic FontSubset: true FirstChar: 0 LastChar: 255 FontDescriptor: FontName: RISBCK+Corbel-BoldItalic Flags: Nonsymbolic, Italic FontBBox: -494, -239, 1194, 952 FontFile3: true Encoding: WinAnsiEncoding ToUnicode: true Font: BaseFont: HLCEXZ+TimesNewRomanPSMT FontSubset: true FirstChar: 32 LastChar: 121 FontDescriptor: FontName: HLCEXZ+TimesNewRomanPSMT Flags: Serif, Nonsymbolic FontBBox: -568, -307, 2000, 1007 FontFile3: true Encoding: WinAnsiEncoding ToUnicode: true CIDFontType2: Font: BaseFont: DBECYH+Corbel CIDSystemInfo: Registry: Adobe Registry: Identity Supplement: 0 FontDescriptor: FontName: DBECYH+Corbel Flags: Symbolic FontBBox: -553, -239, 1110, 952 FontFile2: true Font: BaseFont: IBNBQX+Corbel-Bold CIDSystemInfo: Registry: Adobe Registry: Identity Supplement: 0 FontDescriptor: FontName: IBNBQX+Corbel-Bold Flags: Symbolic FontBBox: -556, -239, 1155, 952 FontFile2: true XMP: application/pdf doi:10.1038/s41598-019-53582-8 Springer US Scientific Reports, doi:10.1038/s41598-019-53582-8 The molecular genealogy of sequential overlapping inversions implies both homologous chromosomes of a heterokaryotype in an inversion origin Dorcas J. Orengo Eva Puerma Unai Cereijo Montserrat Aguadamp#x000E9; 10.1038/s41598-019-53582-8 http://dx.doi.org/10.1038/s41598-019-53582-8 journal Scientific Reports © 2019, The Author(s) 2045-2322 10.1038/s41598-019-53582-8 Springer 2019-11-12T19:14:16+01:00 2019-11-09T08:30:28+05:30 2019-11-12T19:14:16+01:00 True iText® 5.3.5 ©2000-2012 1T3XT BVBA (AGPL-version) VoR uuid:15864410-07de-4d26-97e6-19cf035e05f2 uuid:84c7bb1a-4826-4a73-a967-160fafb7fc53 default 1 converted uuid:bb466da8-6be8-45b7-bcb4-9bba1cbafd98 converted to PDF/A-2b pdfToolbox 2019-11-09T08:31:17+05:30 2 B http://ns.adobe.com/pdf/1.3/ pdf Adobe PDF Schema internal A name object indicating whether the document has been modified to include trapping information Trapped Text http://ns.adobe.com/xap/1.0/mm/ xmpMM XMP Media Management Schema internal UUID based identifier for specific incarnation of a document InstanceID URI internal The common identifier for all versions and renditions of a document. 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