The carbon-nitrogen balance of the nodule and its regulation under elevated carbon dioxide concentration

BioMed Research International

Volumn 2014, Issue , 2014, Pages

  Libault, Marc ^a  

^a UNIVERSITY OF OKLAHOMA   (United States)

Author keywords

[No Author keywords available]

Indexed keywords

CARBON DIOXIDE; CARBON; NITROGEN;

BACTEROIDES; CARBON BALANCE; CARBON SEQUESTRATION; CELL COMMUNICATION; ENVIRONMENTAL CHANGE; FOOD INDUSTRY; LEGUME; MICROBIAL COMMUNITY; MIMOSA PUDICA; NITROGEN BALANCE; NITROGEN FIXATION; NODULATION; NONHUMAN; ORGANISMAL INTERACTION; PHOTOSYNTHATE; PHOTOSYNTHESIS; PLANT CELL; RALSTONIA SOLANACEARUM; REGULATORY MECHANISM; REVIEW; RHIZOBIACEAE; SINORHIZOBIUM MELILOTI; METABOLISM; MICROBIOLOGY; PHYSIOLOGY; PLANT; SYMBIOSIS;

CARBON; CARBON DIOXIDE; NITROGEN; PHOTOSYNTHESIS; PLANTS; ROOT NODULES, PLANT; SYMBIOSIS;

EID: 84902203259     PISSN: 23146133     EISSN: 23146141     Source Type: Journal    
DOI: 10.1155/2014/507946     Document Type: Review

References (78)

1. Phillips D. A., Efficiency of symbiotic nitrogen fixation in legumes. Annual Review of Plant Physiology 1980 31 29 49
2. Stacey G., Libault M., Brechenmacher L., Wan J., May G. D., Genetics and functional genomics of legume nodulation. Current Opinion in Plant Biology 2006 9 2 110 121 2-s2.0-33344461862 10.1016/j.pbi.2006.01.005 (Pubitemid 43289304)
3. Oldroyd G. E. D., Downie J. A., Coordinating nodule morphogenesis with rhizobial infection in legumes. Annual Review of Plant Biology 2008 59 519 546 2-s2.0-44649141367 10.1146/annurev.arplant.59.032607.092839 (Pubitemid 351813041)
4. Murray J. D., Invasion by invitation: rhizobial infection in legumes. Molecular Plant-Microbe Interactions 2011 24 6 631 639 2-s2.0-79955734637 10.1094/MPMI-08-10-0181
5. Oldroyd G. E. D., Murray J. D., Poole P. S., Downie J. A., The rules of engagement in the legume-rhizobial symbiosis. Annual Review of Genetics 2011 45 119 144 2-s2.0-80455174952 10.1146/annurev-genet-110410-132549
6. Oldroyd G. E. D., Speak, friend, and enter: signalling systems that promote beneficial symbiotic associations in plants. Nature Reviews Microbiology 2013 11 4 252 263 2-s2.0-84876379435 10.1038/nrmicro2990
7. Venkateshwaran M., Volkening J. D., Sussman M. R., Ané J.-M., Symbiosis and the social network of higher plants. Current Opinion in Plant Biology 2013 16 1 118 127 2-s2.0-84873725522 10.1016/j.pbi.2012.11.007
8. Kosslak R. M., Bohlool B. B., Suppression of nodule development of one side of a split-root system of soybeans caused by prior inoculation of the other side. Plant Physiology 1984 75 125 130
9. Delves A. C., Mathews A., Day D. A., Carter A. S., Carroll B. J., Gresshoff P. M., Regulation of the soybean-Rhizobium nodule symbiosis by shoot and root factors. Plant Physiology 1986 82 588 590
10. Caetano-Anollés G., Gresshoff P. M., Plant genetic control of nodulation. Annual Review of Microbiology 1991 45 345 382 2-s2.0-0025919587 (Pubitemid 21903796)
11. Ferguson B. J., Indrasumunar A., Hayashi S., Lin M.-H., Lin Y.-H., Reid D. E., Gresshoff P. M., Molecular analysis of legume nodule development and autoregulation. Journal of Integrative Plant Biology 2010 52 1 61 76 2-s2.0-73949119030 10.1111/j.1744-7909.2010.00899.x
12. Nishimura R., Hayashit M., Wu G.-J., Kouchi H., Imaizumi-Anrakull H., Murakami Y., Kawasaki S., Akao S., Ohmori M., Nagasawa M., Harada K., Kawaguchi M., HAR1 mediates systemic regulation of symbiotic organ development. Nature 2002 420 6914 426 429 2-s2.0-18744395527 10.1038/nature01231 (Pubitemid 35415654)
13. Lohar D. P., VandenBosch K. A., Grafting between model legumes demonstrates roles for roots and shoots in determining nodule type and host/rhizobia specificity. Journal of Experimental Botany 2005 56 416 1643 1650 2-s2.0-20044388541 10.1093/jxb/eri160 (Pubitemid 40764452)
14. Schnabel E., Journet E.-P., de Carvalho-Niebel F., Duc G., Frugoli J., The Medicago truncatula SUNN gene encodes a CLV1-like leucine-rich repeat receptor kinase that regulates nodule number and root length. Plant Molecular Biology 2005 58 6 809 822 2-s2.0-27144513326 10.1007/s11103-005-8102-y (Pubitemid 41486873)
15. Magori S., Oka-Kira E., Shibata S., Umehara Y., Kouchi H., Hase Y., Tanaka A., Sato S., Tabata S., Kawaguchi M., Too much love, a root regulator associated with the long-distance control of nodulation in Lotus japonicus. Molecular Plant-Microbe Interactions 2009 22 3 259 268 2-s2.0-58449100531 10.1094/MPMI-22-3-0259
16. Yoshida C., Funayama-Noguchi S., Kawaguchi M., Plenty, a novel hypernodulation mutant in Lotus japonicus. Plant and Cell Physiology 2010 51 9 1425 1435 2-s2.0-77956892303 10.1093/pcp/pcq115
17. Bourion V., Laguerre G., Depret G., Voisin A.-S., Salon C., Duc G., Genetic variability in nodulation and root growth affects nitrogen fixation and accumulation in pea. Annals of Botany 2007 100 3 589 598 2-s2.0-34548232752 10.1093/aob/mcm147 (Pubitemid 47327703)
18. Cabeza R. A., Lingner A., Liese R., Sulieman S., Senbayram M., Trankner M., Dittert K., Schulze J., The activity of nodules of the supernodulating mutant Mt sunn is not limited by photosynthesis under optimal growth conditions. International Journal of Molecular Sciences 2014 15 6031 6045
19. Burdon J. J., Gibson A. H., Searle S. D., Woods M. J., Brockwell J., Variation in the effectiveness of symbiotic associations between native rhizobia and temperate Australian Acacia within-species interactions. Journal of Applied Ecology 1999 36 3 398 408 2-s2.0-0032812868 10.1046/j.1365-2664.1999.00409.x (Pubitemid 29341765)
20. Denton M. D., Coventry D. R., Bellotti W. D., Howieson J. G., Distribution, abundance and symbiotic effectiveness of Rhizobium leguminosarum bv. trifolii from alkaline pasture soils in South Australia. Australian Journal of Experimental Agriculture 2000 40 1 25 35 2-s2.0-0034056768 10.1071/EA99035 (Pubitemid 30236553)
21. Garau G., Reeve W. G., Brau L., Deiana P., Yates R. J., James D., Tiwari R., O'Hara G. W., Howieson J. G., The symbiotic requirements of different Medicago spp. suggest the evolution of Sinorhizobium meliloti and S. medicae with hosts differentially adapted to soil pH. Plant and Soil 2005 276 1-2 263 277 2-s2.0-28344433035 10.1007/s11104-005-0374-0 (Pubitemid 41719504)
22. 2 fixation with the sequenced microsymbiont Sinorhizobium meliloti 1021. New Phytologist 2008 179 1 62 66 2-s2.0-44649165228 10.1111/j.1469-8137. 2008.02464.x (Pubitemid 351783181)
23. Amarger N., Selection of Rhizobium strains on their competitive ability for nodulation. Soil Biology and Biochemistry 1981 13 6 481 486 2-s2.0-0000521366
24. Singleton P. W., Stockinger K. R., Compensation against ineffective nodulation in soybean. Crop Science 1983 23 69 72
25. Hahn M., Studer D., Competitiveness of anif- Bradyrhizobium japonicum mutant against the wild-type strain. FEMS Microbiology Letters 1986 33 1 143 148 2-s2.0-0022644833 10.1016/0378-1097(86)90202-8 (Pubitemid 16180431)
26. Kiers E. T., Rousseau R. A., West S. A., Denlson R. F., Host sanctions and the legume- rhizobium mutualism. Nature 2003 425 6953 78 81 2-s2.0-0041821981 10.1038/nature01931 (Pubitemid 37101720)
27. Gage D. J., Analysis of infection thread development using Gfp- and DsRed-expressing Sinorhizobium meliloti. Journal of Bacteriology 2002 184 24 7042 7046 2-s2.0-0036956040 10.1128/JB.184.24.7042-7046.2002 (Pubitemid 36356137)
28. Denison R. F., Kiers E. T., Life histories of symbiotic rhizobia and mycorrhizal fungi. Current Biology 2011 21 18 R775 R785 2-s2.0-80053298333 10.1016/j.cub.2011.06.018
29. Oono R., Anderson C. G., Denison R. F., Failure to fix nitrogen by non-reproductive symbiotic rhizobia triggers host sanctions that reduce fitness of their reproductive clonemates. Proceedings of the Royal Society B: Biological Sciences 2011 278 1718 2698 2703 2-s2.0-79960761072 10.1098/rspb.2010.2193
30. West S. A., Kiers E. T., Simms E. L., Denison R. F., Sanctions and mutualism stability: why do rhizobia fix nitrogen? Proceedings of the Royal Society B: Biological Sciences 2002 269 1492 685 694 2-s2.0-41049083079 10.1098/rspb.2001.1878 (Pubitemid 34407375)
31. Denison R. F., Kiers E. T., Lifestyle alternatives for rhizobia: mutualism, parasitism, and forgoing symbiosis. FEMS Microbiology Letters 2004 237 2 187 193 2-s2.0-4143069107 10.1016/j.femsle.2004.07.013 (Pubitemid 39091952)
32. Aguilar O. M., Riva O., Peltzer E., Analysis of Rhizobium etli and of its symbiosis with wild Phaseolus vulgaris supports coevolution in centers of host diversification. Proceedings of the National Academy of Sciences of the United States of America 2004 101 37 13548 13553 2-s2.0-4544269622 10.1073/pnas. 0405321101 (Pubitemid 39238440)
33. Zanetti M. E., Blanco F. A., Beker M. P., Battaglia M., Aguilar O. M., A C subunit of the plant nuclear factor NF-Y required for rhizobial infection and nodule development affects partner selection in the common bean- Rhizobium etli symbiosis. Plant Cell 2010 22 12 4142 4157
34. Petroni K., Kumimoto R. W., Gnesutta N., Calvenzani V., Fornari M., Tonelli C., Holt B. F. Jr., Mantovani R., The promiscuous life of plant NUCLEAR FACTOR Y transcription factors. Plant Cell 2012 24 12 4777 4792 2-s2.0-84873036600 10.1105/tpc.112.105734
35. Benedito V. A., Torres-Jerez I., Murray J. D., Andriankaja A., Allen S., Kakar K., Wandrey M., Verdier J., Zuber H., Ott T., Moreau S., Niebel A., Frickey T., Weiller G., He J., Dai X., Zhao P. X., Tang Y., Udvardi M. K., A gene expression atlas of the model legume Medicago truncatula. Plant Journal 2008 55 3 504 513 2-s2.0-47849128399 10.1111/j.1365-313X.2008.03519.x (Pubitemid 352037081)
36. He J., Benedito V. A., Wang M., Murray J. D., Zhao P. X., Tang Y., Udvardi M. K., The Medicago truncatula gene expression atlas web server. BMC Bioinformatics 2009 10, article 441 2-s2.0-75149193517 10.1186/1471-2105-10-441
37. Libault M., Farmer A., Joshi T., Takahashi K., Langley R. J., Franklin L. D., He J., Xu D., May G., Stacey G., An integrated transcriptome atlas of the crop model Glycine max, and its use in comparative analyses in plants. Plant Journal 2010 63 1 86 99 2-s2.0-77954249340 10.1111/j.1365-313X.2010.04222.x
38. Severin A. J., Woody J. L., Bolon Y.-T., Joseph B., Diers B. W., Farmer A. D., Muehlbauer G. J., Nelson R. T., Grant D., Specht J. E., Graham M. A., Cannon S. B., May G. D., Vance C. P., Shoemaker R. C., RNA-Seq Atlas of Glycine max: a guide to the soybean transcriptome. BMC Plant Biology 2010 10, article 160 2-s2.0-77955214742 10.1186/1471-2229-10-160
39. Joshi T., Patil K., Fitzpatrick M. R., Franklin L. D., Yao Q., Cook J. R., Wang Z., Libault M., Brechenmacher L., Valliyodan B., Wu X., Cheng J., Stacey G., Nguyen H. T., Xu D., Soybean Knowledge Base (SoyKB): a web resource for soybean translational genomics. BMC Genomics 2012 13 supplement 1, article S15 2-s2.0-84862247547 10.1186/1471-2164-13-S1-S15
40. Hackenberg D., Keetman U., Grimm B., Homologous NF-YC2 subunit from arabidopsis and tobacco is activated by photooxidative stress and induces flowering. International Journal of Molecular Sciences 2012 13 3 3458 3477 2-s2.0-84858953527 10.3390/ijms13033458
41. Roth L. E., Stacey G., Bacterium release into host cells of nitrogen-fixing soybean nodules: the symbiosome membrane comes from three sources. European Journal of Cell Biology 1989 49 1 13 23 2-s2.0-0024676720
42. Lodwig E. M., Leonard M., Marroqui S., Wheeler T. R., Findlay K., Downie J. A., Poole P. S., Role of polyhydroxybutyrate and glycogen as carbon storage compounds in pea and bean bacteroids. Molecular Plant-Microbe Interactions 2005 18 1 67 74 2-s2.0-11144312983 10.1094/MPMI-18-0067 (Pubitemid 40033931)
43. Oono R., Schmitt I., Sprent J. I., Denison R. F., Multiple evolutionary origins of legume traits leading to extreme rhizobial differentiation. New Phytologist 2010 187 2 508 520 2-s2.0-77954149703 10.1111/j.1469-8137.2010. 03261.x
44. Van de Velde W., Zehirov G., Szatmari A., Debreczeny M., Ishihara H., Kevei Z., Farkas A., Mikulass K., Nagy A., Tiricz H., Satiat-Jeunemaître B., Alunni B., Bourge M., Kucho K.-I., Abe M., Kereszt A., Maroti G., Uchiumi T., Kondorosi E., Mergaert P., Plant peptides govern terminal differentiation of bacteria in symbiosis. Science 2010 327 5969 1122 1126 2-s2.0-77649220369 10.1126/science.1184057
45. Saalbach G., Erik P., Wienkoop S., Characterisation by proteomics of peribacteroid space and peribacteroid membrane preparations from pea (Pisum sativum) symbiosomes. Proteomics 2002 2 325 337 (Pubitemid 34269986)
46. Hohnjec N., Lenz F., Fehlberg V., Vieweg M. F., Baier M. C., Hause B., Küster H., The signal peptide of the Medicago truncatula modular nodulin MtNOD25 operates as an address label for the specific targeting of proteins to nitrogen-fixing symbiosomes. Molecular Plant-Microbe Interactions 2009 22 1 63 72 2-s2.0-58249102865 10.1094/MPMI-22-1-0063
47. Meckfessel M. H., Blancaflor E. B., Plunkett M., Dong Q., Dickstein R., Multiple domains in mtenod8 protein including the signal peptide target it to the symbiosome. Plant Physiology 2012 159 1 299 310 2-s2.0-84860591451 10.1104/pp.111.191403
48. Mergaert P., Nikovics K., Kelemen Z., Maunoury N., Vaubert D., Kondorosi A., Kondorosi E., A novel family in Medicago truncatula consisting of more than 300 nodule-specific genes coding for small, secreted polypeptides with conserved cysteine motifs. Plant Physiology 2003 132 1 161 173 2-s2.0-0038663127 10.1104/pp.102.018192 (Pubitemid 36590408)
49. Mergaert P., Uchiumi T., Alunni B., Evanno G., Cheron A., Catrice O., Mausset A.-E., Barloy-Hubler F., Galibert F., Kondorosi A., Kondorosi E., Eukaryotic control on bacterial cell cycle and differentiation in the Rhizobium -legume symbiosis. Proceedings of the National Academy of Sciences of the United States of America 2006 103 13 5230 5235 2-s2.0-33645510060 10.1073/pnas. 0600912103
50. Alunni B., Kevei Z., Redondo-Nieto M., Kondorosi A., Mergaert P., Kondorosi E., Genomic organization and evolutionary insights on GRP and NCR genes, two large nodule-specific gene families in Medicago truncatula. Molecular Plant-Microbe Interactions 2007 20 9 1138 1148 2-s2.0-34548017391 10.1094/MPMI-20-9-1138 (Pubitemid 47281780)
51. Kereszt A., Mergaert P., Kondorosi E., Bacteroid development in legume nodules: evolution of mutual benefit or of sacrificial victims? Molecular Plant-Microbe Interactions 2011 24 11 1300 1309 2-s2.0-80054741756 10.1094/MPMI-06-11-0152
52. Wang D., Griffitts J., Starker C., Fedorova E., Limpens E., Ivanov S., Bisseling T., Long S., A nodule-specific protein secretory pathway required for nitrogen-fixing symbiosis. Science 2010 327 5969 1126 1129 2-s2.0-77649212137 10.1126/science.1184096
53. Cabeza R., Koester B., Liese R., Lingner A., Baumgarten V., Dirks J., Salinas-Riester G., Pommerenke C., Dittert K., Schulze J., An RNA sequencing transcriptome analysis reveals novel insights into molecular aspects of the nitrate impact on the nodule activity of Medicago truncatula. Plant Physiology 2014 164 400 411
54. Haag A. F., Baloban M., Sani M., Kerscher B., Pierre O., Farkas A., Longhi R., Boncompagni E., Hérouart D., Dall'Angelo S., Kondorosi E., Zanda M., Mergaert P., Ferguson G. P., Protection of Sinorhizobium against host cysteine-rich antimicrobial peptides is critical for symbiosis. PLoS Biology 2011 9 10 2-s2.0-80055041906 10.1371/journal.pbio.1001169 e1001169
55. Marlow V. L., Haag A. F., Kobayashi H., Fletcher V., Scocchi M., Walker G. C., Ferguson G. P., Essential role for the BacA protein in the uptake of a truncated eukaryotic peptide in Sinorhizobium meliloti. Journal of Bacteriology 2009 191 5 1519 1527 2-s2.0-62649148439 10.1128/JB.01661-08
56. Karunakaran R., Haag A. F., East A. K., Ramachandran V. K., Prell J., James E. K., Scocchi M., Ferguson G. P., Poole P. S., BacA is essential for bacteroid development in nodules of galegoid, but not phaseoloid, legumes. Journal of Bacteriology 2010 192 11 2920 2928 2-s2.0-77952558034 10.1128/JB.00020-10
57. Udvardi M. K., Day D. A., Metabolite transport across symbiotic membranes of legume nodules. Annual Review of Plant Biology 1997 48 493 523 2-s2.0-0345950222
58. Benedito V. A., Li H., Dai X., Wandrey M., He J., Kaundal R., Torres-Jerez I., Karen Gomez S., Harrison M. J., Tang Y., Zhao P. X., Udvardi M. K., Genomic inventory and transcriptional analysis of Medicago truncatula transporters. Plant Physiology 2010 152 3 1716 1730 2-s2.0-77949519092 10.1104/pp.109.148684
59. Masalkar P., Wallace I. S., Hwang J. H., Roberts D. M., Interaction of cytosolic glutamine synthetase of soybean root nodules with the C-terminal domain of the symbiosome membrane nodulin 26 aquaglyceroporin. Journal of Biological Chemistry 2010 285 31 23880 23888 2-s2.0-77954898936 10.1074/jbc.M110.135657
60. 2 fixation and recycling in the model legume Lotus japonicus. Journal of Experimental Botany 2011 62 8 2959 2971 2-s2.0-79957440146 10.1093/jxb/err009
61. Udvardi M., Poole P. S., Transport and metabolism in legume-rhizobia symbioses. Annual Review of Plant Biology 2013 64 781 805 2-s2.0-84877676217 10.1146/annurev-arplant-050312-120235
62. Trainer M. A., Charles T. C., The role of PHB metabolism in the symbiosis of rhizobia with legumes. Applied Microbiology and Biotechnology 2006 71 4 377 386 2-s2.0-33745960944 10.1007/s00253-006-0354-1 (Pubitemid 44060938)
63. Libault M., Farmer A., Brechenmacher L., Drnevich J., Langley R. J., Bilgin D. D., Radwan O., Neece D. J., Clough S. J., May G. D., Stacey G., Complete transcriptome of the soybean root hair cell, a single-cell model, and its alteration in response to Bradyrhizobium japonicum infection. Plant Physiology 2010 152 2 541 552 2-s2.0-75949111391 10.1104/pp.109.148379
64. Libault M., Govindarajulu M., Berg R. H., Ong Y. T., Puricelli K., Taylor C. G., Xu D., Stacey G., A Dual-targeted soybean protein is involved in Bradyrhizobium japonicum infection of soybean root hair and cortical cells. Molecular Plant-Microbe Interactions 2011 24 9 1051 1060 2-s2.0-80051578486 10.1094/MPMI-12-10-0281
65. Newcomb E. H., Tandon S. R., Kowal R. R., Ultrastructural specialization for ureide production in uninfected cells of soybean root nodules. Protoplasma 1985 125 1-2 1 12 2-s2.0-0003110470 10.1007/BF01297345
66. Bergersen F. J., Peoples M. B., Turner G. L., A role for poly- β -hydroxybutyrate in bacteroids of soybean root nodules. Proceedings of the Royal Society B: Biological Sciences 1991 245 1312 59 64 2-s2.0-0025744187
67. Cermola M., Fedorova E., Taté R., Riccio A., Favre R., Patriarca E. J., Nodule invasion and symbiosome differentiation during Rhizobium etli-Phaseolus vulgaris symbiosis. Molecular Plant-Microbe Interactions 2000 13 7 733 741 2-s2.0-0034097053 (Pubitemid 30388081)
68. Wang C., Saldanha M., Sheng X., Shelswell K. J., Walsh K. T., Sobral B. W. S., Charles T. C., Roles of poly-3-hydroxybutyrate (PHB) and glycogen in symbiosis of Sinorhizobium meliloti with Medicago sp. Microbiology 2007 153 2 388 398 2-s2.0-33847222547 10.1099/mic.0.29214-0
69. Valls M., Genin S., Boucher C., Integrated regulation of the type III secretion system and other virulence determinants in Ralstonia solanacearum. PLoS Pathogens 2006 2 8, article e82 2-s2.0-39049195400 10.1371/journal.ppat. 0020082 (Pubitemid 44299888)
70. Marchetti M., Capela D., Glew M., Cruveiller S., Chane-Woon-Ming B., Gris C., Timmers T., Poinsot V., Gilbert L. B., Heeb P., Médigue C., Batut J., Masson-Boivin C., Experimental evolution of a plant pathogen into a legume symbiont. PLoS Biology 2010 8 1 2-s2.0-75749144287 10.1371/journal.pbio.1000280 e1000280
71. IPCC, Climate Change 2014: Mitigation of Climate Change 2014
72. Díaz S., Grime J. P., Harris J., McPherson E., Evidence of a feedback mechanism limiting plant response to elevated carbon dioxide. Nature 1993 364 6438 616 617 2-s2.0-0027293277 (Pubitemid 23274715)
73. 2 Plant and Soil 1994 165 1 67 74 2-s2.0-51249164340 10.1007/BF00009963
74. 2 fixation: a link between the carbon and nitrogen cycles in grassland ecosystems. Plant and Soil 1996 187 2 321 332 2-s2.0-0030301442
75. 2 in a grassland ecosystem. Plant Physiology 1996 112 2 575 583 2-s2.0-0030466076
76. Hungate B. A., Dukes J. S., Shaw M. R., Luo Y., Field C. B., Atmospheric science. Nitrogen and climate change. Science 2003 302 5650 1512 1513 2-s2.0-0344827224 10.1126/science.1091390 (Pubitemid 37475680)
77. Rogers A., Ainsworth E. A., Leakey A. D. B., Will elevated carbon dioxide concentration amplify the benefits of nitrogen fixation in legumes? Plant Physiology 2009 151 3 1009 1016 2-s2.0-70349785705 10.1104/pp.109.144113
78. 2 concentration. BioScience 2004 54 8 731 739 2-s2.0-4043154067 (Pubitemid 39073972)