Abstract
Most chloroplast proteins are synthesized as preproteins with N-terminal transit sequences andimported from the cytosol. Protein translocons at the outer (Toc) and inner membranes of the chloroplast(Tic) recognize the presequence and enable the transfer of the polypeptide across the dual membraneenvelope. After the initial characterization of the translocon components, research focused on theimport mechanisms and their regulation. Recent research taking advantage of the Arabidopsissystem has demonstrated the existence of substrate specific import sub-pathways in the general importpathway. New discoveries indicate that glycosylated proteins may take an entirely different pathwayvia the endoplasmic reticulum and the Golgi apparatus. This review will discuss the known importcomponents in the light of the exciting new discoveries.
Access this chapter
Tax calculation will be finalised at checkout
Purchases are for personal use only
Preview
Unable to display preview. Download preview PDF.
References
Aronsson H, Combe J, Jarvis P (2003b) Unusual nucleotide-binding properties of the chloroplast protein import receptor, atToc33. FEBS Lett 544:79–85
Aronsson H, Combe J, Patel R, Jarvis P (2006) In vivo assessment of the significance of phosphorylation of the Arabidopsis chloroplast protein import receptor, atToc33. FEBS Lett 580:649–655
Aronsson H, Sohrt K, Soll J (2000) NADPH:Protochlorophyllide oxidoreductase uses the general import route into chloroplasts. Biol Chem Hoppe Seyler 381:1263–1267
Aronsson H, Sundqvist C, Dahlin C (2003a) POR - import and membrane association of a key element in chloroplast development. Physiol Plant 118:1–9
Asano T, Yoshioka Y, Machida Y (2004) A defect in atToc159 of Arabidopsis thaliana causes severe defects in leaf development. Genes Genet Syst 79:207–212
Baldwin A, Wardle A, Patel R, Dudley P, Park SK, Twell D, Inoue K, Jarvis P (2005) A molecular-genetic study of the Arabidopsis Toc75 gene family. Plant Physiol 138:715–733
Bauer J, Chen K, Hiltbunner A, Wehrli E, Eugster M, Schnell D, Kessler F (2000) The major protein import receptor of plastids is essential for chloroplast biogenesis. Nature 403:203–207
Bauer J, Hiltbrunner A, Weibel P, Vidi PA, Alvarez-Huerta M, Smith MD, Schnell DJ, Kessler F (2002) Essential role of the G-domain in targeting of the protein import receptor atToc159 to the chloroplast outer membrane. J Cell Biol 159:845–854
Becker T (2005) Preprotein recognition and translocation by the Toc complex. Dissertation der Fakultät für Biologie der Ludwig-Maximilians-Universität München
Becker T, Hritz J, Vogel M, Caliebe A, Bukau B, Soll J, Schleiff E (2004b) Toc12, a novel subunit of the intermembrane space preprotein translocon of chloroplasts. Mol Biol Cell 15:5130–5144
Becker T, Jelic M, Vojta A, Radunz A, Soll J, Schleiff E (2004a) Preprotein recognition by the Toc complex. EMBO J 23:520–530
Bedard J, Jarvis P (2005) Recognition and envelope translocation of chloroplast preproteins. J Exp Bot 56:2287–2320
Bhushan S, Kuhn C, Berglund AK, Roth C, Glaser E (2006) The role of the N-terminal domain of chloroplast targeting peptides in organellar protein import and miss-sorting. FEBS Lett 580:3966–3972
Bolter B, May T, Soll J (1998) A protein import receptor in pea chloroplasts, Toc86, is only a proteolytic fragment of a larger polypeptide. FEBS Lett 441:59–62
Bourne HR, Sanders DA, McCormick F (1990) The GTPase superfamily: a conserved switch for diverse cell functions. Nature 348:125–132
Bruce BD (2000) Chloroplast transit peptides: structure, function and evolution. Trends Cell Biol 10:440–447
Bruce BD (2001) The paradox of plastid transit peptides: conservation of function despite divergence in primary structure. Biochim Biophys Acta 1541:2–21
Caliebe A, Grimm R, Kaiser G, Lubeck J, Soll J, Heins L (1997) The chloroplastic protein import machinery contains a Rieske-type iron-sulfur cluster and a mononuclear iron-binding protein. EMBO J 16:7342–7350
Chen K, Chen X, Schnell DJ (2000a) Initial binding of preproteins involving the Toc159 receptor can be bypassed during protein import into chloroplasts. Plant Physiol 122:813–822
Chen KY, Li HM (2007) Precursor binding to an 880-kDa Toc complex as an early step during active import of protein into chloroplasts. Plant J 49:149–158
Chen MH, Huang LF, Li HM, Chen YR, Yu SM (2004) Signal peptide-dependent targeting of a rice alpha-amylase and cargo proteins to plastids and extracellular compartments of plant cells. Plant Physiol 135:1367–1377
Chen X, Smith MD, Fitzpatrick L, Schnell DJ (2002) In vivo analysis of the role of atTic20 in protein import into chloroplasts. Plant Cell 14:641–654
Cherfils J, Chardin P (1999) GEFs: structural basis for their activation of small GTP-binding proteins. Trends Biochem Sci 24:306–311
Chew O, Lister R, Qbadou S, Heazlewood JL, Soll J, Schleiff E, Millar AH, Whelan J (2004) A plant outer mitochondrial membrane protein with high amino acid sequence identity to a chloroplast protein import receptor. FEBS Lett 557:109–114
Chigri F, Hormann F, Stamp A, Stammers DK, Bolter B, Soll J, Vothknecht UC (2006) Calcium regulation of chloroplast protein translocation is mediated by calmodulin binding to Tic32. Proc Natl Acad Sci USA 103:16051–16056
Chigri F, Soll J, Vothknecht UC (2005) Calcium regulation of chloroplast protein import. Plant J 42:821–831
Chou ML, Chu CC, Chen LJ, Akita M, Li HM (2006) Stimulation of transit-peptide release and ATP hydrolysis by a cochaperone during protein import into chloroplasts. J Cell Biol 175:893–900
Chou ML, Fitzpatrick LM, Tu SL, Budziszewski G, Potter-Lewis S, Akita M, Levin JZ, Keegstra K, Li HM (2003) Tic40, a membrane-anchored co-chaperone homolog in the chloroplast protein translocon. EMBO J 22:2970–2980
Constan D, Froehlich JE, Rangarajan S, Keegstra K (2004b) A stromal Hsp100 protein is required for normal chloroplast development and function in Arabidopsis. Plant Physiol 136:3605–3615
Constan D, Patel R, Keegstra K, Jarvis P (2004a) An outer envelope membrane component of the plastid protein import apparatus plays an essential role in Arabidopsis. Plant J 38:93–106
Duy D, Wanner G, Meda AR, von Wiren N, Soll J, Philippar K (2007) PIC1, an ancient permease in Arabidopsis chloroplasts, mediates iron transport. Plant Cell 19:986–1006
Friso G, Giacomelli L, Ytterberg AJ, Peltier JB, Rudella A, Sun Q, Wijk KJ (2004) In-depth analysis of the thylakoid membrane proteome of Arabidopsis thaliana chloroplasts: new proteins, new functions, and a plastid proteome database. Plant Cell 16:478–499
Fulgosi H, Soll J (2002) The chloroplast protein import receptors Toc34 and Toc159 are phosphorylated by distinct protein kinases. J Biol Chem 277:8934–8940
Gentle IE, Burri L, Lithgow T (2005) Molecular architecture and function of the Omp85 family of proteins. Mol Microbiol 58:1216–1225
Gutensohn M, Fan E, Frielingsdorf S, Hanner P, Hou B, Hust B, Klosgen RB (2006) Toc, Tic, Tat et al.: structure and function of protein transport machineries in chloroplasts. J Plant Physiol 163:333–347
Gutensohn M, Pahnke S, Kolukisaoglu U, Schulz B, Schierhorn A, Voigt A, Hust B, Rollwitz I, Stockel J, Geimer S, Albrecht V, Flugge UI, Klosgen RB (2004) Characterization of a T-DNA insertion mutant for the protein import receptor atToc33 from chloroplasts. Mol Genet Genomics 272:379–396
Gutensohn M, Schulz B, Nicolay P, Flugge UI (2000) Functional analysis of the two Arabidopsis homologues of Toc34, a component of the chloroplast protein import apparatus. Plant J 23:771–783
Heins L, Mehrle A, Hemmler R, Wagner R, Kuchler M, Hormann F, Sveshnikov D, Soll J (2002) The preprotein conducting channel at the inner envelope membrane of plastids. EMBO J 21:2616–2625
Hiltbrunner A, Bauer J, Alvarez-Huerta M, Kessler F (2001a) Protein translocon at the Arabidopsis outer chloroplast membrane. Biochem Cell Biol 79:629–635
Hiltbrunner A, Bauer J, Vidi PA, Infanger S, Weibel P, Hohwy M, Kessler F (2001b) Targeting of an abundant cytosolic form of the protein import receptor at Toc159 to the outer chloroplast membrane. J Cell Biol 154:309–316
Hiltbrunner A, Grunig K, Alvarez-Huerta M, Infanger S, Bauer J, Kessler F (2004) AtToc90, a new GTP-binding component of the Arabidopsis chloroplast protein import machinery. Plant Mol Biol 54:427–440
Hinnah SC, Hill K, Wagner R, Schlicher T, Soll J (1997) Reconstitution of a chloroplast protein import channel. EMBO J 16:7351–7360
Hinnah SC, Wagner R, Sveshnikova N, Harrer R, Soll J (2002) The chloroplast protein import channel Toc75: pore properties and interaction with transit peptides. Biophys J 83:899–911
Hirohashi T, Hase T, Nakai M (2001) Maize non-photosynthetic ferredoxin precursor is mis-sorted to the intermembrane space of chloroplasts in the presence of light. Plant Physiol 125:2154–2163
Hirsch S, Muckel E, Heemeyer F, von Heijne G, Soll J (1994) A receptor component of the chloroplast protein translocation machinery. Science 266:1989–1992
Hofmann NR, Theg SM (2005a) Chloroplast outer membrane protein targeting and insertion. Trends Plant Sci 10:450–457
Hofmann NR, Theg SM (2005b) Protein- and energy-mediated targeting of chloroplast outer envelope membrane proteins. Plant J 44:917–927
Hormann F, Kuchler M, Sveshnikov D, Oppermann U, Li Y, Soll J (2004) Tic32, an essential component in chloroplast biogenesis. J Biol Chem 279:34756–34762
Horniak L, Pilon M, van 't Hof R, de Kruijff B (1993) The secondary structure of the ferredoxin transit sequence is modulated by its interaction with negatively charged lipids. FEBS Lett 334:241–246
Inaba T, Alvarez-Huerta M, Li M, Bauer J, Ewers C, Kessler F, Schnell DJ (2005) Arabidopsis tic110 is essential for the assembly and function of the protein import machinery of plastids. Plant Cell 17:1482–1496
Inaba T, Li M, Alvarez-Huerta M, Kessler F, Schnell DJ (2003) atTic110 functions as a scaffold for coordinating the stromal events of protein import into chloroplasts. J Biol Chem 278:38617–38627
Ivanova Y, Smith MD, Chen K, Schnell DJ (2004) Members of the Toc159 import receptor family represent distinct pathways for protein targeting to plastids. Mol Biol Cell 15:3379–3392
Ivey RA 3rd, Bruce BD (2000) In vivo and in vitro interaction of DnaK and a chloroplast transit peptide. Cell Stress Chaperones 5:62–71
Jackson-Constan D, Keegstra K (2001a) Arabidopsis genes encoding components of the chloroplastic protein import apparatus. Plant Physiol 125:1567–1576
Jackson DT, Froehlich JE, Keegstra K (1998) The hydrophilic domain of Tic110, an inner envelope membrane component of the chloroplastic protein translocation apparatus, faces the stromal compartment. J Biol Chem 273:16583–16588
Jarvis P, Chen LJ, Li H, Peto CA, Fankhauser C, Chory J (1998) An Arabidopsis mutant defective in the plastid general protein import apparatus. Science 282:100–103
Jarvis P, Robinson C (2004) Mechanisms of protein import and routing in chloroplasts. Curr Biol 14:R1064–1077
Jelic M, Soll J, Schleiff E (2003) Two Toc34 homologues with different properties. Biochemistry 42:5906–5916
Jelic M, Sveshnikova N, Motzkus M, Horth P, Soll J, Schleiff E (2002) The chloroplast import receptor Toc34 functions as preprotein-regulated GTPase. Biol Chem Hoppe Seyler 383:1875–1883
Kessler F, Blobel G (1996) Interaction of the protein import and folding machineries of the chloroplast. Proc Natl Acad Sci USA 93:7684–7689
Kessler F, Blobel G, Patel HA, Schnell DJ (1994) Identification of two GTP-binding proteins in the chloroplast protein import machinery. Science 266:1035–1039
Kessler F, Schnell DJ (2002) A GTPase gate for protein import into chloroplasts. Nat Struct Biol 9:81–83
Kessler F, Schnell DJ (2004) Chloroplast protein import: solve the GTPase riddle for entry. Trends Cell Biol 14:334–338
Kikuchi S, Hirohashi T, Nakai M (2006) Characterization of the preprotein translocon at the outer envelope membrane of chloroplasts by blue native PAGE. Plant Cell Physiol 47:363–371
Kilian O, Kroth PG (2005) Identification and characterization of a new conserved motif within the presequence of proteins targeted into complex diatom plastids. Plant J 41:175–183
Kim C, Apel K (2004) Substrate-dependent and organ-specific chloroplast protein import in planta. Plant Cell 16:88–98
Kleffmann T, Russenberger D, von Zychlinski A, Christopher W, Sjolander K, Gruissem W, Baginsky S (2004) The Arabidopsis thaliana chloroplast proteome reveals pathway abundance and novel protein functions. Curr Biol 14:354–362
Kouranov A, Chen X, Fuks B, Schnell DJ (1998) Tic20 and Tic22 are new components of the protein import apparatus at the chloroplast inner envelope membrane. J Cell Biol 143:991–1002
Kouranov A, Schnell DJ (1997) Analysis of the interactions of preproteins with the import machinery over the course of protein import into chloroplasts. J Cell Biol 139:1677–1685
Kovacheva S, Bedard J, Patel R, Dudley P, Twell D, Rios G, Koncz C, Jarvis P (2005) In vivo studies on the roles of Tic110, Tic40 and Hsp93 during chloroplast protein import. Plant J 41:412–428
Kovacheva S, Bedard J, Wardle A, Patel R, Jarvis P (2007) Further in vivo studies on the role of the molecular chaperone, Hsp93, in plastid protein import. Plant J 50:364–379
Kubis S, Baldwin A, Patel R, Razzaq A, Dupree P, Lilley K, Kurth J, Leister D, Jarvis P (2003) The Arabidopsis ppi1 mutant is specifically defective in the expression, chloroplast import, and accumulation of photosynthetic proteins. Plant Cell 15:1859–1871
Kubis S, Patel R, Combe J, Bedard J, Kovacheva S, Lilley K, Biehl A, Leister D, Rios G, Koncz C, Jarvis P (2004) Functional specialization amongst the Arabidopsis Toc159 family of chloroplast protein import receptors. Plant Cell 16:2059–2077
Kuchler M, Decker S, Hormann F, Soll J, Heins L (2002) Protein import into chloroplasts involves redox-regulated proteins. EMBO J 21:6136–6145
Kutschera U, Niklas KJ (2005) Endosymbiosis, cell evolution, and speciation. Theory Biosci 124:1–24
Lee KH, Kim SJ, Lee YJ, Jin JB, Hwang I (2003) The M domain of atToc159 plays an essential role in the import of proteins into chloroplasts and chloroplast biogenesis. J Biol Chem 278:36794–36805
Leister D, Schneider A (2003) From genes to photosynthesis in Arabidopsis thaliana. Int Rev Cytol 228:31–83
Lopez-Juez E, Pyke KA (2005) Plastids unleashed: their development and their integration in plant development. Int J Dev Biol 49:557–577
Lubeck J, Soll J, Akita M, Nielsen E, Keegstra K (1996) Topology of IEP110, a component of the chloroplastic protein import machinery present in the inner envelope membrane. EMBO J 15:4230–4238
Ma Y, Kouranov A, LaSala SE, Schnell DJ (1996) Two components of the chloroplast protein import apparatus, IAP86 and IAP75, interact with the transit sequence during the recognition and translocation of precursor proteins at the outer envelope. J Cell Biol 134:315–327
Martin T, Sharma R, Sippel C, Waegemann K, Soll J, Vothknecht UC (2006) A protein kinase family in Arabidopsis phosphorylates chloroplast precursor proteins. J Biol Chem 281:40216–40223
Martin W, Rujan T, Richly E, Hansen A, Cornelsen S, Lins T, Leister D, Stoebe B, Hasegawa M, Penny D (2002) Evolutionary analysis of Arabidopsis, cyanobacterial, and chloroplast genomes reveals plastid phylogeny and thousands of cyanobacterial genes in the nucleus. Proc Natl Acad Sci USA 99:12246–12251
May T, Soll J (2000) 14–3-3 proteins form a guidance complex with chloroplast precursor proteins in plants. Plant Cell 12:53–64
McFadden GI (2001) Chloroplast origin and integration. Plant Physiol 125:50–53
Miras S, Salvi D, Ferro M, Grunwald D, Garin J, Joyard J, Rolland N (2002) Non-canonical transit peptide for import into the chloroplast. J Biol Chem 277:47770–47778
Nada A, Soll J (2004) Inner envelope protein 32 is imported into chloroplasts by a novel pathway. J Cell Sci 117:3975–3982
Nakrieko KA, Mould RM, Smith AG (2004) Fidelity of targeting to chloroplasts is not affected by removal of the phosphorylation site from the transit peptide. Eur J Biochem 271:509–516
Nanjo Y, Oka H, Ikarashi N, Kaneko K, Kitajima A, Mitsui T, Munoz FJ, Rodriguez-Lopez M, Baroja-Fernandez E, Pozueta-Romero J (2006) Rice plastidial N-glycosylated nucleotide pyrophosphatase/phosphodiesterase is transported from the ER-golgi to the chloroplast through the secretory pathway. Plant Cell 18:2582–2592
Nassoury N, Morse D (2005) Protein targeting to the chloroplasts of photosynthetic eukaryotes: getting there is half the fun. Biochim Biophys Acta 1743:5–19
Olsen LJ, Keegstra K (1992) The binding of precursor proteins to chloroplasts requires nucleoside triphosphates in the intermembrane space. J Biol Chem 267:433–439
Pain D, Blobel G (1987) Protein import into chloroplasts requires a chloroplast ATPase. Proc Natl Acad Sci USA 84:3288–3292
Paschen SA, Neupert W, Rapaport D (2005) Biogenesis of beta-barrel membrane proteins of mitochondria. Trends Biochem Sci 30:575–582
Perry SE, Keegstra K (1994) Envelope membrane proteins that interact with chloroplastic precursor proteins. Plant Cell 6:93–105
Philippar K, Geis T, Ilkavets I, Oster U, Schwenkert S, Meurer J, Soll J (2007) Chloroplast biogenesis: The use of mutants to study the etioplast-chloroplast transition. Proc Natl Acad Sci USA 104:678–683
Pollmann S, Neu D, Lehmann T, Berkowitz O, Schafer T, Weiler EW (2006) Subcellular localization and tissue specific expression of amidase 1 from Arabidopsis thaliana. Planta 224:1241–1253
Pollmann S, Neu D, Weiler EW (2003) Molecular cloning and characterization of an amidase from Arabidopsis thaliana capable of converting indole-3-acetamide into the plant growth hormone, indole-3-acetic acid. Phytochemistry 62:293–300
Qbadou S, Becker T, Mirus O, Tews I, Soll J, Schleiff E (2006) The molecular chaperone Hsp90 delivers precursor proteins to the chloroplast import receptor Toc64. EMBO J 25:1836–1847
Radhamony RN, Theg SM (2006) Evidence for an ER to Golgi to chloroplast protein transport pathway. Trends Cell Biol 16:385–387
Reddick LE, Vaughn MD, Wright SJ, Campbell IM, Bruce BD (2007) In vitro comparative kinetic analysis of the chloroplast toc GTPases. J Biol Chem 282:11410–11426
Reinbothe C, Lebedev N, Apel K, Reinbothe S (1997) Regulation of chloroplast protein import through a protochlorophyllide-responsive transit peptide. Proc Natl Acad Sci USA 94:8890–8894
Reinbothe S, Pollmann S, Springer A, James RJ, Tichtinsky G, Reinbothe C (2005) A role of Toc33 in the protochlorophyllide-dependent plastid import pathway of NADPH:protochlorophyllide oxidoreductase (POR) A. Plant J 42:1–12
Reinbothe S, Quigley F, Gray J, Schemenewitz A, Reinbothe C (2004) Identification of plastid envelope proteins required for import of protochlorophyllide oxidoreductase A into the chloroplast of barley. Proc Natl Acad Sci USA 101:2197–2202
Reuber TL, Ausubel FM (1996) Isolation of Arabidopsis genes that differentiate between resistance responses mediated by the RPS2 and RPM1 disease resistance genes. Plant Cell 8:241–249
Reumann S, Inoue K, Keegstra K (2005) Evolution of the general protein import pathway of plastids (review). Mol Membr Biol 22:73–86
Reumann S, Keegstra K (1999) The endosymbiotic origin of the protein import machinery of chloroplastic envelope membranes. Trends Plant Sci 4:302–307
Rial DV, Arakaki AK, Ceccarelli EA (2000) Interaction of the targeting sequence of chloroplast precursors with Hsp70 molecular chaperones. Eur J Biochem 267:6239–6248
Richter S, Lamppa GK (2003) Structural properties of the chloroplast stromal processing peptidase required for its function in transit peptide removal. J Biol Chem 278:39497–39502
Scheffzek K, Ahmadian MR (2005) GTPase activating proteins: structural and functional insights 18 years after discovery. Cell Mol Life Sci 62:3014–3038
Schleiff E, Jelic M, Soll J (2003b) A GTP-driven motor moves proteins across the outer envelope of chloroplasts. Proc Natl Acad Sci USA 100:4604–4609
Schleiff E, Klosgen RB (2001) Without a little help from 'my' friends: direct insertion of proteins into chloroplast membranes? Biochim Biophys Acta 1541:22–33
Schleiff E, Soll J (2005) Membrane protein insertion: mixing eukaryotic and prokaryotic concepts. EMBO Rep 6:1023–1027
Schleiff E, Soll J, Kuchler M, Kuhlbrandt W, Harrer R (2003a) Characterization of the translocon of the outer envelope of chloroplasts. J Cell Biol 160:541–551
Schnell DJ, Blobel G (1993) Identification of intermediates in the pathway of protein import into chloroplasts and their localization to envelope contact sites. J Cell Biol 120:103–115
Schnell DJ, Kessler F, Blobel G (1994) Isolation of components of the chloroplast protein import machinery. Science 266:1007–1012
Seedorf M, Waegemann K, Soll J (1995) A constituent of the chloroplast import complex represents a new type of GTP-binding protein. Plant J 7:401–411
Siddique MA, Grossmann J, Gruissem W, Baginsky S (2006) Proteome analysis of bell pepper (Capsicum annuum L.) Chromoplasts. Plant Cell Physiol 47:1663–1673
Sjogren LL, MacDonald TM, Sutinen S, Clarke AK (2004) Inactivation of the clpC1 gene encoding a chloroplast Hsp100 molecular chaperone causes growth retardation, leaf chlorosis, lower photosynthetic activity, and a specific reduction in photosystem content. Plant Physiol 136:4114–4126
Smith MD, Hiltbrunner A, Kessler F, Schnell DJ (2002) The targeting of the atToc159 preprotein receptor to the chloroplast outer membrane is mediated by its GTPase domain and is regulated by GTP. J Cell Biol 159:833–843
Smith MD, Rounds CM, Wang F, Chen K, Afitlhile M, Schnell DJ (2004) atToc159 is a selective transit peptide receptor for the import of nucleus-encoded chloroplast proteins. J Cell Biol 165:323–334
Sohrt K, Soll J (2000) Toc64, a new component of the protein translocon of chloroplasts. J Cell Biol 148:1213–1221
Soll J (2002) Protein import into chloroplasts. Curr Opin Plant Biol 5:529–535
Stahl T, Glockmann C, Soll J, Heins L (1999) Tic40, a new "old" subunit of the chloroplast protein import translocon. J Biol Chem 274:37467–37472
Sulli C, Fang Z, Muchhal U, Schwartzbach SD (1999) Topology of Euglena chloroplast protein precursors within endoplasmic reticulum to Golgi to chloroplast transport vesicles. J Biol Chem 274:457–463
Sun CW, Chen LJ, Lin LC, Li HM (2001) Leaf-specific upregulation of chloroplast translocon genes by a CCT motif-containing protein, CIA 2. Plant Cell 13:2053–2061
Sun YJ, Forouhar F, Li Hm HM, Tu SL, Yeh YH, Kao S, Shr HL, Chou CC, Chen C, Hsiao CD (2002) Crystal structure of pea Toc34, a novel GTPase of the chloroplast protein translocon. Nat Struct Biol 9:95–100
Sveshnikova N, Soll J, Schleiff E (2000) Toc34 is a preprotein receptor regulated by GTP and phosphorylation. Proc Natl Acad Sci USA 97:4973–4978
Teng YS, Su YS, Chen LJ, Lee YJ, Hwang I, Li HM (2006) Tic21 is an essential translocon component for protein translocation across the chloroplast inner envelope membrane. Plant Cell 18:2247–2257
Theg SM, Bauerle C, Olsen LJ, Selman BR, Keegstra K (1989) Internal ATP is the only energy requirement for the translocation of precursor proteins across chloroplastic membranes. J Biol Chem 264:6730–6736
Timmis JN, Ayliffe MA, Huang CY, Martin W (2004) Endosymbiotic gene transfer: organelle genomes forge eukaryotic chromosomes. Nat Rev Genet 5:123–135
Tu SL, Chen LJ, Smith MD, Su YS, Schnell DJ, Li HM (2004) Import pathways of chloroplast interior proteins and the outer-membrane protein OEP14 converge at Toc75. Plant Cell 16:2078–2088
Tu SL, Li HM (2000) Insertion of OEP14 into the outer envelope membrane is mediated by proteinaceous components of chloroplasts. Plant Cell 12:1951–1960
van den Wijngaard PW, Vredenberg WJ (1999) The envelope anion channel involved in chloroplast protein import is associated with Tic110. J Biol Chem 274:25201–25204
van Wijk KJ (2004) Plastid proteomics. Plant Physiol Biochem 42:963–977
Villarejo A, Buren S, Larsson S, Dejardin A, Monne M, Rudhe C, Karlsson J, Jansson S, Lerouge P, Rolland N, von Heijne G, Grebe M, Bako L, Samuelsson G (2005) Evidence for a protein transported through the secretory pathway en route to the higher plant chloroplast. Nat Cell Biol 7:1224–1231
Vojta A, Alavi M, Becker T, Hormann F, Kuchler M, Soll J, Thomson R, Schleiff E (2004) The protein translocon of the plastid envelopes. J Biol Chem 279:21401–21405
Waegemann K, Soll J (1996) Phosphorylation of the transit sequence of chloroplast precursor proteins. J Biol Chem 271:6545–6554
Wallas TR, Smith MD, Sanchez-Nieto S, Schnell DJ (2003) The roles of toc34 and toc75 in targeting the toc159 preprotein receptor to chloroplasts. J Biol Chem 278:44289–44297
Waller RF, Reed MB, Cowman AF, McFadden GI (2000) Protein trafficking to the plastid of Plasmodium falciparum is via the secretory pathway. EMBO J 19:1794–1802
Wan J, Blakeley SD, Dennis DT, Ko K (1996) Transit peptides play a major role in the preferential import of proteins into leucoplasts and chloroplasts. J Biol Chem 271:31227–31233
Weibel P, Hiltbrunner A, Brand L, Kessler F (2003) Dimerization of Toc-GTPases at the chloroplast protein import machinery. J Biol Chem 278:37321–37329
Wu C, Seibert FS, Ko K (1994) Identification of chloroplast envelope proteins in close physical proximity to a partially translocated chimeric precursor protein. J Biol Chem 269:32264–32271
Yan X, Khan S, Hase T, Emes MJ, Bowsher CG (2006) Differential uptake of photosynthetic and non-photosynthetic proteins by pea root plastids. FEBS Lett 580:6509–6512
Yeh YH, Kesavulu MM, Li HM, Wu SZ, Sun YJ, Konozy EH, Hsiao CD (2007) Dimerization is important for the GTPase activity of chloroplast translocon components atToc33 and psToc159. J Biol Chem 282:13845–13853
Young JC, Hoogenraad NJ, Hartl FU (2003) Molecular chaperones Hsp90 and Hsp70 deliver preproteins to the mitochondrial import receptor Tom70. Cell 112:41–50
Young ME, Keegstra K, Froehlich JE (1999) GTP promotes the formation of early-import intermediates but is not required during the translocation step of protein import into chloroplasts. Plant Physiol 121:237–244
Yu TS, Li H (2001) Chloroplast protein translocon components atToc159 and atToc33 are not essential for chloroplast biogenesis in guard cells and root cells. Plant Physiol 127:90–96
Zhang XP, Glaser E (2002) Interaction of plant mitochondrial and chloroplast signal peptides with the Hsp70 molecular chaperone. Trends Plant Sci 7:14–21
Zhong R, Wan J, Jin R, Lamppa G (2003) A pea antisense gene for the chloroplast stromal processing peptidase yields seedling lethals in Arabidopsis: survivors show defective GFP import in vivo. Plant J 34:802–812
Author information
Authors and Affiliations
Corresponding author
Editor information
Rights and permissions
Copyright information
© 2007 Springer-Verlag Berlin Heidelberg
About this chapter
Cite this chapter
Agne, B., Kessler, F. (2007). Protein import into plastids. In: Bock, R. (eds) Cell and Molecular Biology of Plastids. Topics in Current Genetics, vol 19. Springer, Berlin, Heidelberg. https://doi.org/10.1007/4735_2007_0237
Download citation
DOI: https://doi.org/10.1007/4735_2007_0237
Published:
Publisher Name: Springer, Berlin, Heidelberg
Print ISBN: 978-3-540-75375-9
Online ISBN: 978-3-540-75376-6
eBook Packages: Biomedical and Life SciencesBiomedical and Life Sciences (R0)