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A refined genome‐scale reconstruction of Chlamydomonas metabolism provides a platform for systems‐level analyses

Imam, Saheed, Schäuble, Sascha, Valenzuela, Jacob, López García de Lomana, Adrián, Carter, Warren, Price, Nathan D., Baliga, Nitin S.·2015·The Plant Journal
DOI 10.1111/tpj.13059PMC PMC4715634
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Abstract

Summary: Microalgae have reemerged as organisms of prime biotechnological interest due to their ability to synthesize a suite of valuable chemicals. To harness the capabilities of these organisms, we need a comprehensive systems‐level understanding of their metabolism, which can be fundamentally achieved through large‐scale mechanistic models of metabolism. In this study, we present a revised and significantly improved genome‐scale metabolic model for the widely‐studied microalga, Chlamydomonas reinhardtii . The model, i C re1355, represents a major advance over previous models, both in content and predictive power. i C re1355 encompasses a broad range of metabolic functions encoded across the nuclear, chloroplast and mitochondrial genomes accounting for 1355 genes (1460 transcripts), 2394 and 1133 metabolites. We found improved performance over the previous metabolic model based on comparisons of predictive accuracy across 306 phenotypes (from 81 mutants), lipid yield analysis and growth rates derived from chemostat‐grown cells (under three conditions). Measurement of macronutrient uptake revealed carbon and phosphate to be good predictors of growth rate, while nitrogen consumption appeared to be in excess. We analyzed high‐resolution time series transcriptomics data using i C re1355 to uncover dynamic pathway‐level changes that occur in response to nitrogen starvation and changes in light intensity. This approach enabled accurate prediction of growth rates, the cessation of growth and accumulation of triacylglycerols during nitrogen starvation, and the temporal response of different growth‐associated pathways to increased light intensity. Thus, i C re1355 represents an experimentally validated genome‐scale reconstruction of C. reinhardtii metabolism that should serve as a useful resource for studying the metabolic processes of this and related microalgae.

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  1. Alper H Jin YS Moxley JF Stephanopoulos G 2005 Identifying gene targets for the metabolic engineering of lycopene biosynthesis in Escherichia coli Metab Eng 7 155 164 15885614
  2. Atteia A Adrait A Brugiere S Tardif M van Lis R Deusch O Dagan T Kuhn L Gontero B Martin W Garin J Joyard J Rolland N 2009 A proteomic survey of Chlamydomonas reinhardtii mitochondria sheds new light on the metabolic plasticity of the organelle and on the nature of the alpha-proteobacterial mitochondrial ancestor Mol Biol Evol 26 1533 1548 19349646
  3. Bateman A Coin L Durbin R Finn RD Hollich V Griffiths-Jones S Khanna A Marshall M Moxon S Sonnhammer EL Studholme DJ Yeats C Eddy SR 2004 The Pfam protein families database Nucleic Acids Res 32 D138 141 14681378 PMC308855
  4. Bishop CL Purton S Nugent JH 2003 Molecular analysis of the Chlamydomonas nuclear gene encoding PsbW and demonstration that PsbW is a subunit of photosystem II, but not photosystem I Plant Mol Biol 52 285 289 12856936
  5. Blaby IK Blaby-Haas CE Tourasse N Hom EF Lopez D Aksoy M Grossman A Umen J Dutcher S Porter M King S Witman GB Stanke M Harris EH Goodstein D Grimwood J Schmutz J Vallon O Merchant SS Prochnik S 2014 The Chlamydomonas genome project: a decade on Trends Plant Sci 19 672 680 24950814 PMC4185214
  6. Blaby IK Glaesener AG Mettler T Fitz-Gibbon ST Gallaher SD Liu B Boyle NR Kropat J Stitt M Johnson S Benning C Pellegrini M Casero D Merchant SS 2013 Systems-level analysis of nitrogen starvation-induced modifications of carbon metabolism in a Chlamydomonas reinhardtii starchless mutant Plant Cell 25 4305 4323 24280389 PMC3875720
  7. Bolling C Fiehn O 2005 Metabolite profiling of Chlamydomonas reinhardtii under nutrient deprivation Plant Physiol 139 1995 2005 16306140 PMC1310576
  8. Boyle NR Morgan JA 2009 Flux balance analysis of primary metabolism in Chlamydomonas reinhardtii BMC Syst Biol 3 4 19128495 PMC2628641
  9. Boyle NR Page MD Liu B Blaby IK Casero D Kropat J Cokus SJ Hong-Hermesdorf A Shaw J Karpowicz SJ Gallaher SD Johnson S Benning C Pellegrini M Grossman A Merchant SS 2012 Three acyltransferases and nitrogen-responsive regulator are implicated in nitrogen starvation-induced triacylglycerol accumulation in Chlamydomonas J Biol Chem 287 15811 15825 22403401 PMC3346115
  10. Branco-Price C Kaiser KA Jang CJ Larive CK Bailey-Serres J 2008 Selective mRNA translation coordinates energetic and metabolic adjustments to cellular oxygen deprivation and reoxygenation in Arabidopsis thaliana Plant J 56 743 755 18665916
  11. Cardol P Gonzalez-Halphen D Reyes-Prieto A Baurain D Matagne RF Remacle C 2005 The mitochondrial oxidative phosphorylation proteome of Chlamydomonas reinhardtii deduced from the Genome Sequencing Project Plant Physiol 137 447 459 15710684 PMC1065347
  12. Castruita M Casero D Karpowicz SJ Kropat J Vieler A Hsieh SI Yan W Cokus S Loo JA Benning C Pellegrini M Merchant SS 2011 Systems biology approach in Chlamydomonas reveals connections between copper nutrition and multiple metabolic steps Plant Cell 23 1273 1292 21498682 PMC3101551
  13. Chang RL Ghamsari L Manichaikul A Hom EF Balaji S Fu W Shen Y Hao T Palsson BO Salehi-Ashtiani K Papin JA 2011 Metabolic network reconstruction of Chlamydomonas offers insight into light-driven algal metabolism Mol Syst Biol 7 518 21811229 PMC3202792
  14. Chisti Y 2007 Biodiesel from microalgae Biotechnol Adv 25 294 306 17350212
  15. Colijn C Brandes A Zucker J Lun DS Weiner B Farhat MR Cheng TY Moody DB Murray M Galagan JE 2009 Interpreting expression data with metabolic flux models: predicting Mycobacterium tuberculosis mycolic acid production PLoS Comput Biol 5 e1000489 19714220 PMC2726785
  16. Croft MT Warren MJ Smith AG 2006 Algae need their vitamins Eukaryot Cell 5 1175 1183 16896203 PMC1539151
  17. Davis MC Fiehn O Durnford DG 2013 Metabolic acclimation to excess light intensity in Chlamydomonas reinhardtii Plant Cell Environ 36 1391 1405 23346954
  18. Duanmu D Casero D Dent RM Gallaher S Yang W Rockwell NC Martin SS Pellegrini M Niyogi KK Merchant SS Grossman AR Lagarias JC 2013 Retrograde bilin signaling enables Chlamydomonas greening and phototrophic survival Proc Natl Acad Sci U S A 110 3621 3626 23345435 PMC3587268
  19. Emanuelsson O Brunak S von Heijne G Nielsen H 2007 Locating proteins in the cell using TargetP, SignalP and related tools Nat Protoc 2 953 971 17446895
  20. Fan J Andre C Xu C 2011 A chloroplast pathway for the de novo biosynthesis of triacylglycerol in Chlamydomonas reinhardtii FEBS Lett 585 1985 1991 21575636
  21. Feely RA Sabine CL Lee K Berelson W Kleypas J Fabry VJ Millero FJ 2004 Impact of anthropogenic CO2 on the CaCO3 system in the oceans Science 305 362 366 15256664
  22. Guo Y Sheng Q Li J Ye F Samuels DC Shyr Y 2013 Large scale comparison of gene expression levels by microarrays and RNAseq using TCGA data PLoS One 8 e71462 23977046 PMC3748065
  23. Harris EH 2001 Chlamydomonas as a model organism Annu Rev Plant Physiol Plant Mol Biol 52 363 406 11337403
  24. Hemschemeier A Duner M Casero D Merchant SS Winkler M Happe T 2013 Hypoxic survival requires a 2-on-2 hemoglobin in a process involving nitric oxide Proc Natl Acad Sci U S A 110 10854 10859 23754374 PMC3696821
  25. Henry CS Zinner JF Cohoon MP Stevens RL 2009 iBsu1103: a new genome-scale metabolic model of Bacillus subtilis based on SEED annotations Genome Biol 10 R69 19555510 PMC2718503
  26. Honisch B Ridgwell A Schmidt DN Thomas E Gibbs SJ Sluijs A Zeebe R Kump L Martindale RC Greene SE Kiessling W Ries J Zachos JC Royer DL Barker S Marchitto TM Jr Moyer R Pelejero C Ziveri P Foster GL Williams B 2012 The geological record of ocean acidification Science 335 1058 1063 22383840
  27. Iwai M Ikeda K Shimojima M Ohta H 2014 Enhancement of extraplastidic oil synthesis in Chlamydomonas reinhardtii using a type-2 diacylglycerol acyltransferase with a phosphorus starvation-inducible promoter Plant Biotechnol J 12 808 819 24909748 PMC4160818
  28. Jones PC Jiang W Fillingame RH 1998 Arrangement of the multicopy H+-translocating subunit c in the membrane sector of the Escherichia coli F1F0 ATP synthase J Biol Chem 273 17178 17185 9642286
  29. Kanehisa M Goto S Kawashima S Nakaya A 2002 The KEGG databases at GenomeNet Nucleic Acids Res 30 42 46 11752249 PMC99091
  30. Kanehisa M Goto S Sato Y Kawashima M Furumichi M Tanabe M 2014 Data, information, knowledge and principle: back to metabolism in KEGG Nucleic Acids Res 42 D199 205 24214961 PMC3965122
  31. Kellis M Wold B Snyder MP Bernstein BE Kundaje A Marinov GK Ward LD Birney E Crawford GE Dekker J Dunham I Elnitski LL Farnham PJ Feingold EA Gerstein M Giddings MC Gilbert DM Gingeras TR Green ED Guigo R Hubbard T Kent J Lieb JD Myers RM Pazin MJ Ren B Stamatoyannopoulos JA Weng Z White KP Hardison RC 2014 Defining functional DNA elements in the human genome Proc Natl Acad Sci U S A 111 6131 6138 24753594 PMC4035993
  32. Kim J Reed JL 2012 RELATCH: relative optimality in metabolic networks explains robust metabolic and regulatory responses to perturbations Genome Biol 13 R78 23013597 PMC3506949
  33. Kleessen S Irgang S Klie S Giavalisco P Nikoloski Z 2015 Integration of transcriptomics and metabolomics data specifies the metabolic response of Chlamydomonas to rapamycin treatment Plant J 81 822 835 25600836
  34. Kliphuis AM Klok AJ Martens DE Lamers PP Janssen M Wijffels RH 2011 Metabolic modeling of Chlamydomonas reinhardtii: energy requirements for photoautotrophic growth and maintenance J Appl Phycol 24 253 266 22427720 PMC3289792
  35. Krasensky J Jonak C 2012 Drought, salt, and temperature stress-induced metabolic rearrangements and regulatory networks J Exp Bot 63 1593 1608 22291134 PMC4359903
  36. Lecler R Vigeolas H Emonds-Alt B Cardol P Remacle C 2012 Characterization of an internal type-II NADH dehydrogenase from Chlamydomonas reinhardtii mitochondria Curr Genet 58 205 216 22814755
  37. Lewis NS Nocera DG 2006 Powering the planet: chemical challenges in solar energy utilization Proc Natl Acad Sci U S A 103 15729 15735 17043226 PMC1635072
  38. Lin H Kwan AL Dutcher SK 2010 Synthesizing and salvaging NAD: lessons learned from Chlamydomonas reinhardtii PLoS Genet 6 e1001105 20838591 PMC2936527
  39. Mahadevan R Schilling CH 2003 The effects of alternate optimal solutions in constraint-based genome-scale metabolic models Metab Eng 5 264 276 14642354
  40. Marshall WF 2008 Basal bodies platforms for building cilia Curr Top Dev Biol 85 1 22 19147000
  41. McCloskey D Palsson BO Feist AM 2013 Basic and applied uses of genome-scale metabolic network reconstructions of Escherichia coli Mol Syst Biol 9 661 23632383 PMC3658273
  42. McGlade C Ekins P 2015 The geographical distribution of fossil fuels unused when limiting global warming to 2 degrees C Nature 517 187 190 25567285
  43. Merchant SS Prochnik SE Vallon O Harris EH Karpowicz SJ Witman GB Terry A Salamov A Fritz-Laylin LK Marechal-Drouard L Marshall WF Qu LH Nelson DR Sanderfoot AA Spalding MH Kapitonov VV Ren Q Ferris P Lindquist E Shapiro H Lucas SM Grimwood J Schmutz J Cardol P Cerutti H Chanfreau G Chen CL Cognat V Croft MT Dent R Dutcher S Fernandez E Fukuzawa H Gonzalez-Ballester D Gonzalez-Halphen D Hallmann A Hanikenne M Hippler M Inwood W Jabbari K Kalanon M Kuras R Lefebvre PA Lemaire SD Lobanov AV Lohr M Manuell A Meier I Mets L Mittag M Mittelmeier T Moroney JV Moseley J Napoli C Nedelcu AM Niyogi K Novoselov SV Paulsen IT Pazour G Purton S Ral JP Riano-Pachon DM Riekhof W Rymarquis L Schroda M Stern D Umen J Willows R Wilson N Zimmer SL Allmer J Balk J Bisova K Chen CJ Elias M Gendler K Hauser C Lamb MR Ledford H Long JC Minagawa J Page MD Pan J Pootakham W Roje S Rose A Stahlberg E Terauchi AM Yang P Ball S Bowler C Dieckmann CL Gladyshev VN Green P Jorgensen R Mayfield S Mueller-Roeber B Rajamani S Sayre RT Brokstein P Dubchak I Goodstein D Hornick L Huang YW Jhaveri J Luo Y Martinez D Ngau WC Otillar B Poliakov A Porter A Szajkowski L Werner G Zhou K Grigoriev IV Rokhsar DS Grossman AR 2007 The Chlamydomonas genome reveals the evolution of key animal and plant functions Science 318 245 250 17932292 PMC2875087
  44. Mettler T Muhlhaus T Hemme D Schottler MA Rupprecht J Idoine A Veyel D Pal SK Yaneva-Roder L Winck FV Sommer F Vosloh D Seiwert B Erban A Burgos A Arvidsson S Schonfelder S Arnold A Gunther M Krause U Lohse M Kopka J Nikoloski Z Mueller-Roeber B Willmitzer L Bock R Schroda M Stitt M 2014 Systems analysis of the response of photosynthesis, metabolism, and growth to an increase in irradiance in the photosynthetic model organism Chlamydomonas reinhardtii Plant Cell 26 2310 2350 24894045 PMC4114937
  45. Miller R Wu G Deshpande RR Vieler A Gartner K Li X Moellering ER Zauner S Cornish AJ Liu B Bullard B Sears BB Kuo MH Hegg EL Shachar-Hill Y Shiu SH Benning C 2010 Changes in transcript abundance in Chlamydomonas reinhardtii following nitrogen deprivation predict diversion of metabolism Plant Physiol 154 1737 1752 20935180 PMC2996024
  46. Milne CB Kim PJ Eddy JA Price ND 2009 Accomplishments in genome-scale in silico modeling for industrial and medical biotechnology Biotechnol J 4 1653 1670 19946878 PMC3160784
  47. Mittler R 2002 Oxidative stress, antioxidants and stress tolerance Trends Plant Sci 7 405 410 12234732
  48. Moulin M Nguyen GT Scaife MA Smith AG Fitzpatrick TB 2013 Analysis of Chlamydomonas thiamin metabolism in vivo reveals riboswitch plasticity Proc Natl Acad Sci U S A 110 14622 14627 23959877 PMC3767531
  49. Msanne J Xu D Konda AR Casas-Mollano JA Awada T Cahoon EB Cerutti H 2012 Metabolic and gene expression changes triggered by nitrogen deprivation in the photoautotrophically grown microalgae Chlamydomonas reinhardtii and Coccomyxa sp. C-169 Phytochemistry 75 50 59 22226037
  50. Muller-Moule P Havaux M Niyogi KK 2003 Zeaxanthin deficiency enhances the high light sensitivity of an ascorbate-deficient mutant of Arabidopsis Plant Physiol 133 748 760 12972657 PMC219049
  51. Oberhardt MA Palsson BO Papin JA 2009 Applications of genome-scale metabolic reconstructions Mol Syst Biol 5 320 19888215 PMC2795471
  52. Oshlack A Robinson MD Young MD 2010 From RNA-seq reads to differential expression results Genome Biol 11 220 21176179 PMC3046478
  53. Palsson B 2000 The challenges of in silico biology Nat Biotechnol 18 1147 1150 11062431
  54. Park JH Lee KH Kim TY Lee SY 2007 Metabolic engineering of Escherichia coli for the production of L-valine based on transcriptome analysis and in silico gene knockout simulation Proc Natl Acad Sci U S A 104 7797 7802 17463081 PMC1857225
  55. Park JJ Wang H Gargouri M Deshpande RR Skepper JN Holguin FO Juergens MT Shachar-Hill Y Hicks LM Gang DR 2015 The response of Chlamydomonas reinhardtii to nitrogen deprivation: a systems biology analysis Plant J 81 611 624 25515814
  56. Pazour GJ Agrin N Leszyk J Witman GB 2005 Proteomic analysis of a eukaryotic cilium J Cell Biol 170 103 113 15998802 PMC2171396
  57. Perez-Perez ME Couso I Crespo JL 2012 Carotenoid deficiency triggers autophagy in the model green alga Chlamydomonas reinhardtii Autophagy 8 376 388 22302003
  58. Petti AA Crutchfield CA Rabinowitz JD Botstein D 2011 Survival of starving yeast is correlated with oxidative stress response and nonrespiratory mitochondrial function Proc Natl Acad Sci U S A 108 E1089 1098 21734149 PMC3215077
  59. Pogoryelov D Reichen C Klyszejko AL Brunisholz R Muller DJ Dimroth P Meier T 2007 The oligomeric state of c rings from cyanobacterial F-ATP synthases varies from 13 to 15 J Bacteriol 189 5895 5902 17545285 PMC1952053
  60. Recht L Topfer N Batushansky A Sikron N Gibon Y Fait A Nikoloski Z Boussiba S Zarka A 2014 Metabolite profiling and integrative modeling reveal metabolic constraints for carbon partitioning under nitrogen starvation in the green algae Haematococcus pluvialis J Biol Chem 289 30387 30403 25183014 PMC4215223
  61. Reed JL 2012 Shrinking the metabolic solution space using experimental datasets PLoS Comput Biol 8 e1002662 22956899 PMC3431291
  62. Reijnders MJ van Heck RG Lam CM Scaife MA dos Santos VA Smith AG Schaap PJ 2014 Green genes: bioinformatics and systems-biology innovations drive algal biotechnology Trends Biotechnol 32 617 626 25457388
  63. Rochaix JD 2002 Chlamydomonas, a model system for studying the assembly and dynamics of photosynthetic complexes FEBS Lett 529 34 38 12354609
  64. Sanz-Luque E Ocana-Calahorro F Llamas A Galvan A Fernandez E 2013 Nitric oxide controls nitrate and ammonium assimilation in Chlamydomonas reinhardtii J Exp Bot 64 3373 3383 23918969
  65. Scott SA Davey MP Dennis JS Horst I Howe CJ Lea-Smith DJ Smith AG 2010 Biodiesel from algae: challenges and prospects Curr Opin Biotechnol 21 277 286 20399634
  66. Seelert H Poetsch A Dencher NA Engel A Stahlberg H Muller DJ 2000 Structural biology. Proton-powered turbine of a plant motor Nature 405 418 419 10839529
  67. Shlomi T Cabili MN Herrgard MJ Palsson BO Ruppin E 2008 Network-based prediction of human tissue-specific metabolism Nat Biotechnol 26 1003 1010 18711341
  68. Small I Peeters N Legeai F Lurin C 2004 Predotar: A tool for rapidly screening proteomes for N-terminal targeting sequences Proteomics 4 1581 1590 15174128
  69. Spalding MH 2008 Microalgal carbon-dioxide-concentrating mechanisms: Chlamydomonas inorganic carbon transporters J Exp Bot 59 1463 1473 17597098
  70. Stock D Gibbons C Arechaga I Leslie AG Walker JE 2000 The rotary mechanism of ATP synthase Curr Opin Struct Biol 10 672 679 11114504
  71. Terashima M Specht M Hippler M 2011 The chloroplast proteome: a survey from the Chlamydomonas reinhardtii perspective with a focus on distinctive features Curr Genet 57 151 168 21533645
  72. Terashima M Specht M Naumann B Hippler M 2010 Characterizing the anaerobic response of Chlamydomonas reinhardtii by quantitative proteomics Mol Cell Proteomics 9 1514 1532 20190198 PMC2938099
  73. Theiss C Bohley P Voigt J 2002 Regulation by polyamines of ornithine decarboxylase activity and cell division in the unicellular green alga Chlamydomonas reinhardtii Plant Physiol 128 1470 1479 11950995 PMC154274
  74. Urzica EI Adler LN Page MD Linster CL Arbing MA Casero D Pellegrini M Merchant SS Clarke SG 2012 Impact of oxidative stress on ascorbate biosynthesis in Chlamydomonas via regulation of the VTC2 gene encoding a GDP-L-galactose phosphorylase J Biol Chem 287 14234 14245 22393048 PMC3340187
  75. Urzica EI Casero D Yamasaki H Hsieh SI Adler LN Karpowicz SJ Blaby-Haas CE Clarke SG Loo JA Pellegrini M Merchant SS 2012 Systems and trans-system level analysis identifies conserved iron deficiency responses in the plant lineage Plant Cell 24 3921 3948 23043051 PMC3517228
  76. Urzica EI Vieler A Hong-Hermesdorf A Page MD Casero D Gallaher SD Kropat J Pellegrini M Benning C Merchant SS 2013 Remodeling of membrane lipids in iron-starved Chlamydomonas J Biol Chem 288 30246 30258 23983122 PMC3798491
  77. van Lis R Atteia A Mendoza-Hernandez G Gonzalez-Halphen D 2003 Identification of novel mitochondrial protein components of Chlamydomonas reinhardtii. A proteomic approach Plant Physiol 132 318 330 12746537 PMC166977
  78. Wijffels RH Barbosa MJ 2010 An outlook on microalgal biofuels Science 329 796 799 20705853
  79. Zur H Ruppin E Shlomi T 2010 iMAT: an integrative metabolic analysis tool Bioinformatics 26 3140 3142 21081510
  80. Zwiener I Frisch B Binder H 2014 Transforming RNA-Seq data to improve the performance of prognostic gene signatures PLoS One 9 e85150 24416353 PMC3885686

Cited by

Sorted by citations

Authors
Imam, S., Schäuble, S., Valenzuela, J., López García de Lomana, A., Carter, W., Price, N. D., & Baliga, N. S.
Affiliations
Institute for Systems Biology 401 Terry Ave N Seattle WA 98109 USA
Venue
The Plant Journal
Publisher
Wiley
Published
2015-11-30
Volume
84
Issue
6
Pages
1239-1256
ISSN
  • 0960-7412
  • 1365-313X
DOI
10.1111/tpj.13059
PMC
PMC4715634
Cited by
92
References
1
Language
en
Licence
onlinelibrary.wiley.com/termsAndConditions#vor (vor)
Funders
National Institutes of Health (2P50 GM076547)

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