Bioinformatic strategies to provide functional clues to the unknown genes in plasmodium falciparum genome

Parasite

Volumn 17, Issue 4, 2010, Pages 273-283

  Florent, I ^a   Marechal E ^b   Gascuel, O ^c   Brehelin L ^c  

^a MUSÉUM NATIONAL D'HISTOIRE NATURELLE   (France)

^b CNRS   (France)

^c LIRMM   (France)

Author keywords

Annotations; Bioinformatics; Genome; Novel mining methods; Plasmcdium

Indexed keywords

PLASMODIUM FALCIPARUM;

EID: 78650932931     PISSN: 1252607X     EISSN: 17761042     Source Type: Journal    
DOI: 10.1051/parasite/2010174273     Document Type: Article

References (69)

1. ACHIDI E.A. ET AL. A global network for investigating the genomic epidemiology of malaria. Nature, 2008, 456, 732-737.
2. AHOUIDI A.D., BEI A.K., NEAFSEY D.E., SARR O., VOLKMAN S., MlLNER D., COX-SlNGH J., FERREIRA M.U., NDIR O., PREMJI Z., MBOUP S. & DURAISINGH M.T. Population genetic analysis of large sequence polymorphisms in Plasmodium falciparum blood-stage antigens. Infect, Genet. Evol., 2010, 10, 200-206.
3. ALTSCHUL S.F., GISH W., MILLER W., MYERS E.W. & LIPMAN D.J. Basic local alignment search tool. J. MoI. Biol., 1990, 215, 403-410.
4. ARAVIND L., IYER L.M., WELLEMS T.E. & MILLER L.H. Plasmodium biology: genomic gleanings. Cell, 2003, 115, 771-785.
5. AURRECOECHEA C., BRESTELLI J., BRUNK B.P., CARLTON J.M., DOMMER J., FISCHER S., GAJRIA B., GAO X., GINGLE A., GRANT G., HARB O.S., HEIGES M., INNAMORATO F., IODICE J., KISSINGER J.C., KRAEMER E., LI W., MILLER J.A., MORRISON H.G., NAYAK V., PENNINGTON C., PINNEY D.F., Roos D.S., Ross C., STOECKERT C.J., JR., SULLIVAN S., TREATMAN C. & WANG H. GiardiaDB and TrichDB: integrated genomic resources for the eukaryotic protist pathogens Giardia lamblia and Trichomonas vaginalis. Nucleic Acids Res., 2009a, 37, D526-530.
6. AURRECOECHEA C., BRESTELLI J., BRUNK B.P., DOMMER J., FISCHER S., GAJRIA B., GAO X., GINGLE A., GRANT G., HARB O.S., HEIGES M., INNAMORATO F., IODICE J., KISSINGER J.C., KRAEMER E., Li W., MILLER J.A., NAYAK V., PENNINGTON C., PINNEY D.F., Roos D.S., Ross C., STOECKERT C.J., JR., TREATMAN C. & WANG H. PlasmoDB: a functional genomic database for malaria parasites. Nucleic Acids Res., 2009b, 37, D539-543.
7. BASTIEN O., LESPINATS S., ROY S., METAYER K., FERTIL B., CODANI J.J. & MARECHAL E. Analysis of the compositional biases in Plasmodium falciparum genome and proteome using Arabidopsis thaliana as a reference. Gene, 2004, 336, 163-173.
8. BASTIEN O. & MARECHAL E. Evolution of biological sequences implies an extreme value distribution of type I for both global and local pairwise alignment scores. BMC Bioinformatics, 2008, 9, 332.
9. BASTIEN O., ROY S. & MARECHAL E. Construction of non-symmetric substitution matrices derived from proteomes with biased amino acid distributions. C. R. Biol., 2005, 328, 445-453.
10. BIRKHOLTZ L., VAN BRUMMELEN A.C., CLARK K., NIEMAND J., MARECHAL E., LLINAS M. & Louw A.I. Exploring functional genomics for drug target and therapeutics discovery in Plasmodia. Acta Trop., 2008, 105, 113-123.
11. BIRKHOLTZ L.M., BASTIEN O., WELLS G., GRANDO D., JOUBERT F., KASAM V., ZIMMERMANN M., ORTET P., JACQ N., SAIDANI N., ROY S., HOFMANN-APITIUS M., BRETON V., Louw A.I. & MARECHAL E. Integration and mining of malaria molecular, functional and pharmacological data: how far are we from a chemogenomic knowledge space? Malar. J., 2006, 5, 110.
12. BOZDECH Z., LLINAS M., PULLIAM B.L., WONG E.D., ZHU J. & DERISI J.L. The transcriptome of the intraerythrocytic developmental cycle of Plasmodium falciparum. PLoS Biol, 2003a, 1, E5.
13. Bozdech Z., ZHU J., JOACHIMIAK M.P., COHEN F.E., PULLIAM B. & DERISI J.L. Expression profiling of the schizont and trophozoite stages of Plasmodium falciparum with a long-oligonucleotide microarray. Genome Biol, 2003b, 4.
14. BREHEUN L., DUFAYARD J.F. & GASCUEL O. PlasmoDraft: a database of Plasmodium falciparum gene function predictions based on postgenomic data. BMC Bioinformatics, 2008, 9, 440.
15. BREHELIN L., FLORENT I., GASCUEL O. & MARECHAL E. Assessing functional annotation transfers with inter-species conserved coexpression: application to Plasmodium falciparum. BMC Genomics, 2010, 11, 35.
16. CARLTON J.M., ADAMS J.H., SILVA J.C., BIDWELL S.L., LORENZI H., CALER E., CRABTREE J., ANGIUOLI S.V., MERINO E.F., AMEDEO P., CHENG Q., COULSON R.M., CRABB B.S., DEL PORTILLO H.A., ESSIEN K., FELDBLYUM T. V., FERNANDEZ-BECERRA C., GILSON P.R., GUEYE A.H., Guo X., KANG'A S., KOOIJP.M. T.W., KORSINCZKY M., MEYER E.V., NENE V., PAULSEN I., WHITE O., RALPH S.A., REN Q., SARGEANT T.J., SALZBERG S.L., STOECKERT C.J., SULUVAN S.A., YAMAMOTO M.M., HOFFMAN S.L., WORTMAN J.R., GARDNER M.J., GAUNSKI M.R., BARNWELL J.W. AND FRASER-LIGGETT C.M. Comparative genomics of the neglected human malaria parasite Plasmodium vivax. Nature, 2008, 455, 757-763.
17. CARLTON J.M., ANGIUOLI S.V., SUH B.B., KOOIJ T.W., PERTEA M., SILVA J.C., ERMOLAEVA M.D., ALLEN J.E., SELENGUT J.D., KOO H.L., PETERSON J.D., POP M., KOSACK D.S., SHUMWAY M.F., BIDWELL S.L., SHALLOM S.J., VAN AKEN S.E., RIEDMULLER S.B., FELDBLYUM T.V., CHO J.K., QUACKENBUSH J., SEDEGAH M., SHOAIBI A., CUMMINGS L.M., FLORENS L., YATES J.R., RAINE J.D., SINDEN R.E., HARRIS M.A., CUNNINGHAM D.A., PREISER P.R., BERGMAN L.W., VAIDYA A.B., VAN LIN L.H., JANSE CJ., WATERS A.P., SMITH H.O., WHITE O.R., SALZBERG S.L., VENTER J.C, FRASER C.M., HOFFMAN S.L., GARDNER MJ. & CARUCCI DJ. Genome sequence and comparative analysis of the model rodent malaria parasite Plasmodium yoelii yoelii. Nature, 2002, 419, 512-519.
18. CLARK K., DHOOGRA M., LOUW A.I. & BIRKHOLTZ L.M. Transcriptional responses of Plasmodium falciparum to alpha-difluoromethylornithine-induced polyamine depletion. Biol. Chem., 2008, 389, 111-125.
19. DAHL E.L., SHOCK J.L., SHENAI B.R., GUT J., DERISI J.L. & ROSENTHAL P.J. Tetracyclines specifically target the apicoplast of the malaria parasite Plasmodium falciparum. Antimicrob. Agents Chemother., 2006, 50, 3124-3131.
20. DATE S.V. & STOECKERT C.J., JR. Computational modeling of the Plasmodium falciparum interactome reveals protein function on a genome-wide scale. Genome Res., 2006, 16, 542-549.
21. DE BEER T.A., WELLS G.A., BURGER P.B., JOUBERT F., MARECHAL E., BIRKHOLTZ L. & Louw A.I. Antimalarial drug discovery: in silico structural biology and rational drug design. Infect. Disord. Drug Targets, 2009, 9, 304-318.
22. DE KONING-WARD T.F., GILSON P.R., BODDEY J.A., RUG M., SMITH B.J., PAPENFUSS A.T., SANDERS P.R., LUNDIE R.J., MAIER A.G., COWMAN A.F. & CRABB B.S. A newly discovered protein export machine in malaria parasites. Nature, 2009, 459, 945-949.
23. DECHAMPS S., MAYNADIER M., WEIN S., GANNOUN-ZAKI L., MARECHAL E. & VIAL H.J. Rodent and nonrodent malaria parasites differ in their phospholipid metabolic pathways. J. Lipid Res., 2010, 51, 81-96.
24. DOOLITTLE R.F. Biodiversity: microbial genomes multiply. Nature, 2002, 416, 697-700.
25. DURBIN R., EDDY S., KROGH A. & MITCHISON G. Biological sequence analysis: Probabilistic models of proteins and nucleic acids. Cambridge University Press, 1998, 46-152.
26. EISEN M.B., SPELLMAN P.T., BROWN P.O. & BOTSTEIN D. Cluster analysis and display of genome-wide expression patterns. Proc. Natl Acad. Sd. USA, 1998, 95, 14863-14868.
27. FLORENS L., LIU X., WANG Y., YANG S., SCHWARTZ O., PEGLAR M., CARUCCI DJ., YATES J.R., 3RD & WUB Y. Proteomics approach reveals novel proteins on the surface of malaria-infected erythrocytes. Mol. Biochem. Parasitol, 2004, 135, 1-11.
28. FLORENS L., WASHBURN M.P., RAINE J.D., ANTHONY R.M., GRAINGER M., HAYNES J.D., MOCH J.K., MUSTER N., SACCI J.B., TABB D.L., WITNEY A.A., WOLTERS D., Wu Y., GARDNER M.J., HOLDER A.A., SINDEN R.E., YATES J.R. & CARUCCI D.J. A proteomic view of the Plasmodium falciparum life cycle. Nature, 2002, 419, 520-526.
29. FLORENT I., PORCEL B.M., GUILLAUME E., DA SILVA C., ARTIGUENAVE F., MARECHAL E., BREHEUN L., GASCUEL O., CHARNEAU S., WINCKER P. & GRELLIER P. A Plasmodium falciparum FcB1schizont-EST collection providing clues to schizont specific gene structure and polymorphism. BMC Genomics, 2009, 10, 235.
30. GARDNER MJ., HALL N., FUNG E., WHITE O., BERRIMAN M., HYMAN R.W., CARLTON J.M., PAIN A., NELSON K.E., BOWMAN S., PAULSEN I.T., JAMES K., EISEN J.A., RUTHERFORD K., SALZBERG S.L., CRAIG A., KYES S., CHAN M.S., NENE V., SHALLOM S.J., SUH B., PETERSON J., ANGIUOU S., PERTEA M., ALLEN J., SELENGUT J., HAFT D., MATHER M.W., VAIDYA A.B., MARTIN D.M., FAIRLAMB A.H., FRAUNHOLZ M.J., ROOS D.S., RALPH S.A., MCFADDEN G.I., CUMMINGS L.M., SUBRAMANIAN G.M., MUNGALL C., VENTER J.C., CARUCCI D.J., HOFFMAN S.L., NEWBOLD C., DAVIS R.W., FRASER C.M. & BARRELL B. Genome sequence of the human malaria parasite Plasmodium falciparum. Nature, 2002, 419, 498-511.
31. GHOUILA A., TERRAPON N., GASCUEL O., GUERFALI F.Z., LAOUINI D., MARÉCHAL É & BREHEUN L. EuPathDomains : The divergent domain database for eukaryotic pathogènes. Infection, Genetics & Evolution, 2010 (in press).
32. GREENWOOD B. Progress in malaria control in endemic areas. TravelMed. Infect. Dis., 2008, 6, 173-176.
33. GREENWOOD B. Can malaria be eliminated? Trans. R. Soc. Trop. Med. Hyg., 2009, 103 (Suppl. 1), S2-S5.
34. HALL. N., KARRAS M., RAINE J.D., CARLTON J.M., KOOIJ T.W., BERRIMAN M., FLORENS L., JANSSEN C.S., PAIN A., CHRISTOPHIDES G.K., JAMES K., RUTHERFORD K., HARRIS B., HARRIS D., CHURCHER C., QUAIL M.A., ORMOND D., DOGGETT J., TRUEMAN H.E., MENDOZA J., BIDWELL S.L., RAJANDREAM M.A., CARUCCI D.J., YATES J.R., 3RD, KAFATOS F.C., JANSE C.J., BARRELL B., TURNER C.M., WATERS A.P. & SINDEN R.E. A comprehensive survey of the Plasmodium life cycle by genomic, transcriptomic, and proteomic analyses. Science, 2005,307, 82-86.
35. Hu G., CABRERA A., KONO M., MOK S., CHAAL B.K., HAASE S., ENGELBERG K., CHEEMADAN S., SPIELMANN T., PREISER P.R., GILBERGER T.W. & BozDECH Z. Transcriptional profiling of growth perturbations of the human malaria parasite Plasmodium falciparum. Nat. Biotechnol, 2010, 28, 91-98.
36. JEFFARES D.C., PAIN A., BERRY A., COX A.V., STALKER J., INGLE C.E., THOMAS A., QUAIL M.A., SIEBENTHALL K., UHLEMANN A.C., KYES S., KRISHNA S., NEWBOLD C., DERMITZAKIS E.T. & BERRIMAN M. Genome variation and evolution of the malaria parasite Plasmodium falciparum. Nat. Genet., 2007, 39, 120-125.
37. JOUBERT F., HARRISON C.M., KOEGELENBERG R.J., ODENDAAL C.J. & DE BEER T.A. Discovery: an interactive resource for the rational selection and comparison of putative drug target proteins in malaria. Malar.J., 2009, 8, 178.
38. KHAN S.M., FRANKE-FAYARD B., MAIR G.R., LASONDER E., JANSE C.J., MANN M. & WATERS A.P. Proteome analysis of separated male and female gametocytes reveals novel sex-specific Plasmodium biology. Cell, 2005, 121, 675-687.
39. Koou T.W., CARLTON J.M., BIDWELL S.L., HALL N., RAMESAR J., JANSE C.J. & WATERS A.P. A Plasmodium Whole-Genome Synteny Map: lndels and Synteny Breakpoints as Foci for Species-Specific Genes. PLoS Pathog, 2005, 1, e44.
40. LACOUNT D.J., IGNAU M., CHETTIER R., PHANSALKAR A., BELL R., HESSELBERTH J.R., ScHOENFELD L.W., OTA I., SAHASRABUDHE S., KURSCHNER C., FIELDS S. & HUGHES R.E. A protein interaction network of the malaria parasite Plasmodium falciparum. Nature, 2005, 438, 103-107.
41. LAL K., PRIETO J.H., BROMLEY E., SANDERSON S.J., YATES J.R., 3RD, WASTUNG J.M., TOMLEY F.M. & SINDEN R.E. Characterisation of Plasmodium invasive organelles; an ookinete microneme proteome. Proteomics, 2009, 9, 1142-1151.
42. LAMARQUE M., TASTET C., PONCET J., DÉMETTRE E., JOUIN P., VIAL H.J. & DUBREMETZ J.F. Food vacuole proteome of the malarial parasite Plasmodium falciparum. Proteomics, 2008, 2, 1361-1374.
43. LASONDER E., ISHIHAMA Y., ANDERSEN J.S., VERMUNT A.M., PAIN A., SAUERWEIN R.W., ELING W.M., HALL. N., WATERS A.P., STUNNENBERG H.G. & MANN M. Analysis of the Plasmodium falciparum proteome by high-accuracy mass spectrometry. Nature, 2002, 419, 537-542.
44. LASONDER E., JANSE C.J., VAN GEMERT G.J., MAIR G.R., VERMUNT A.M., DOURADINHA B.G., VAN NOORT V., HUYNEN M.A., LUTY A.J., KROEZE H., KHAN S.M., SAUERWEIN R.W., WATERS A.P., MANN M. & STUNNENBERG H.G. Proteomic profiling of Plasmodium sporozoite maturation identifies new proteins essential for parasite development and infectivity. PLoS Pathog, 2008, 4, e1000195.
45. LE ROCH K.G., JOHNSON J.R., AHIBOH H., CHUNG D.W., PRUDHOMME J., PLOUFFE D., HENSON K., ZHOU Y., WITOLA W., YATES J.R., MAMOUN C.B., WINZELER E.A. & VIAL H. A systematic approach to understand the mechanism of action of the bisthiazollum compound T4 on the human malaria parasite, Plasmodium falciparum. BMC Genomics, 2008, 9, 513.
46. LE ROCH K.G., ZHOU Y., BLAIR P.L., GRAINGER M., MOCH J.K., HAYNES J.D., DE LA VEGA P., HOLDER A.A., BATALOV S., CARUCCI D.J. & WINZELER E.A. Discovery of gene function by expression profiling of the malaria parasite life cycle. Science, 2003, 301, 1503-1508.
47. LIEW K.J., Hu G., BOZDECH Z. & PETER P.R. Defining species specific genome differences in malaria parasites. BMC Genomics, 2010, 11, 128.
48. LIPMAN D.J. & PEARSON W.R. Rapid and sensitive protein similarity searches. Science, 1985, 227, 1435-1441.
49. LLINAS M., BOZDECH Z., WONG E.D., ADAI A.T. & DERISI J.L. Comparative whole genome transcrlptome analysis of three Plasmodium falciparum strains. Nucleic Acids Res., 2006, 34, 1166-1173.
50. LLINAS M. & DEL PORTILLO H.A. Mining the malaria transcriptome. Trends Parasitol, 2005, 21, 350-352.
51. MAIR G.R., BRAKS J.A., GARVER L.S., WIEGANTJ.C., HALL N., DIRKS R.W., KHAN S.M., DIMOPOULOS G., JANSE C.J. & WATERS A.P. Regulation of sexual development of Plasmodium by translational repression. Science, 2006, 313, 667-669.
52. NATALANG O., BISCHOFF E., DEPLAINE G., PROUX C., DILUES M.A., SISMEIRO O., GUIGON G., BONNEFOY S., PATARAPOTIKUL J., MERCEREAU-PUIJALON O., COPPEE J.Y. & DAVID P.H. Dynamic RNA profiling in Plasmodium falciparum synchronized blood stages exposed to lethal doses of artesunate. BMC Genomics, 2008, 9, 388.
53. NYALWIDHE J. & LINGELBACH K. Proteases and chaperones are the most abundant proteins in the parasitophorous vacuole of Plasmodium falciparum-infected erythrocytes. Proteomics, 2006, 6, 1563-1573.
54. PAIN A., BOHME U., BERRY A.E., MUNGALL K., FINN R.D., JACKSON A.P., MOURIER T., MISTRY J., PASINI E.M., ASLETT M.A., BALA-SUBRAMMANIAM S., BORGWARDT K., BROOKS K., CARRET C., CARVER T.J., CHEREVACH I., CHILUNGWORTH T., CLARK T.G., GAUNSKI M.R., HALL N., HARPER D., HARRIS D., HAUSER H., IVENS A., JANSSEN C.S., KEANE T., LARKE N., LAPP S., MARTI M., MOULE S., MEYER I.M., ORMOND D., PETERS N., SANDERS M., SANDERS S., SARGEANT T.J., SIMMONDS M., SMITH F., SQUARES R., THURSTON S., TIVEY A.R., WALKER D., WHITE B., ZUIDERWUK E., CHURCHER C., QUAIL M.A., COWMAN A.F., TURNER C.M., RAJANDREAM M.A., KOCKEN C.H., THOMAS A.W., NEWBOLD C.I., BARRELL B.G. & BERRIMAN M. The genome of the simian and human malaria parasite Plasmodium knowlesi. Nature, 2008, 455, 799-803.
55. RADFAR A., DIEZ A. & BAUTISTA J.M. Chloroquine mediates specific proteome oxidative damage across the erythrocytic cycle of resistant Plasmodium falciparum. Free Radic. Biol. Med., 2008, 44, 2034-2042.
56. SAIDANI N., GRANDO D., VALADIE H., BASTIEN O. & MARECHAL E. Potential and limits of in silico target discovery - Case study of the search for new antimalarial chemotherapeutic targets. Infect. Genet. Evol., 2009, 9, 359-367.
57. SAM-YELLOWE T.Y., BANKS T.L., FUJIOKA H., DRAZBA J.A. & YADAV S.P. Plasmodium yoelii: novel rhoptry proteins identified within the body of merozoite rhoptries in rodent Plasmodium malaria. Exp. Parasitol, 2008, 120, 113-117.
58. SAM-YELLOWE T.Y., FLORENS L., WANG T., RAINE J.D., CARUCCI D.J., SINDEN R. & YATES J.R., 3RD Proteome analysis of rhoptryenrlched fractions isolated from Plasmodium merozoites. J. Proteome Res., 2004, 3, 995-1001.
59. SANDERS P.R., CANTIN G.T., GREENBAUM D.C., GILSON P.R., NEBL T., MORITZ R.L., YATES J.R., 3RD, HODDER A.N. & CRABB B.S. Identification of protein complexes in detergent-resistant membranes of Plasmodium falciparum schizonts. Mol. Biochem. Parasitol., 2007, 154, 148-157.
60. SHOCK J.L., FISCHER K.F. & DERISI J.L. Whole-genome analysis of mRNA decay in Plasmodium falciparum reveals a global lengthening of mRNA half-life during the intra-erythrocytlc development cycle. Genome Biol, 2007, 8, R134.
61. SPELLMAN P.T., SHERLOCK G., ZHANG M.Q., IYER V.R., ANDERS K., EISEN M.B., BROWN P.O., BOTSTEIN D. & FUTCHER B. Comprehensive identification of cell cycle-regulated genes of the yeast Saccharomyces cerevisiae by microarray hybridization. Mol. Biol. Cell, 1998, 9, 3273-3297.
62. STOECKERT C.J., J.R., FISCHER S., KISSINGER J.C., HEIGES M., AURRECOECHEA C., GAJRIA B. & Roos D.S. PlasmoDB v5: new looks, new genomes. Trends Parasitol, 2006, 22, 543-546.
63. TARUN A.S., PENG X., DUMPIT R.F., OGATA Y., SILVA-RIVERA H., CAMARGO N., DALY T.M., BERGMAN L.W. & KAPPE S.H. A combined transcriptome and proteome survey of malaria parasite liver stages. Proc. Natl Acad. Scl. USA, 2008, 105, 305-310.
64. TERRAPON N., GASCUEL O., MARECHAL E. & BREHEUN L. Detection of new protein domains using co-occurrence: application to Plasmodium falciparum. Bioinformatics, 2009, 25, 3077-3083.
65. TUTEJA R. Malaria - an overview. FEBS J., 2007, 274, 4670-4679.
66. VOLKMAN S.K., SABETI P.C., DECAPRIO D., NEAFSEY D.E., SCHAFFNER S.F., MILNER D.A., JR., DAILY J.P., SARR O., NDIAYE D., NDIR O., MBOUP S., DURAISINGH M.T., LUKENS A., DERR A., STANGETHOMANN N., WAGGONER S., ONOFRIO R., ZIAUGRA L., MAUCEU E., GNERRE S., JAFFE D.B., ZAINOUN J., WIEGAND R.C., BIRREN B.W., HARTL D.L., GALAGAN J.E., UNDER E.S. & WIRTH D.F. A genome-wide map of diversity in Plasmodium falciparum. Nat. Genet., 2007, 39, 113-119.
67. WINZELER E.A. Malaria research in the post-genomic era. Nature, 2008, 455, 751-756.
68. YOUNG J.A., FIVELMAN Q.L., BLAIR P.L., DE LA VEGA P., LE ROCH K.G., ZHOU Y., CARUCCI D.J., BAKER D.A. & WINZELER E.A. The Plasmodium falciparum sexual development transcriptome: a microarray analysis using ontology-based pattern identification. Mol. Biochem. Parasitol, 2005, 143, 67-79.
69. ZHOU Y., RAMACHANDRAN V., KUMAR K.A., WESTENBERGER S., REFOUR P., ZHOU B., LI F., YOUNG J.A., CHEN K., PLOUFFE D., HENSON K., NUSSENZWEIG V., CARLTON J., VINETZ J.M., DURAISINGH M.T. & WINZELER E.A. Evidence-based annotation of the malaria parasite's genome using comparative expression profiling. PLoS One, 2008,3, e1570.