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Chloroplast DNA

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1103:. It has been established that some plants have linear cpDNA, such as maize, and that more still contain complex structures that scientists do not yet understand; however, the predominant view today is that most cpDNA is circular. When the original experiments on cpDNA were performed, scientists did notice linear structures; however, they attributed these linear forms to broken circles. If the branched and complex structures seen in cpDNA experiments are real and not artifacts of concatenated circular DNA or broken circles, then a D-loop mechanism of replication is insufficient to explain how those structures would replicate. At the same time, homologous recombination does not explain the multiple A → G gradients seen in plastomes. This shortcoming is one of the biggest for the linear structure theory. 1240: 1611: 1919: 602: 1087:
replication forks were most likely present and the direction that they initially opened (the highest gradient is most likely nearest the start site because it was single stranded for the longest amount of time). This mechanism is still the leading theory today; however, a second theory suggests that most cpDNA is actually linear and replicates through homologous recombination. It further contends that only a minority of the genetic material is kept in circular chromosomes while the rest is in branched, linear, or other complex structures.
1078: 34: 963: 1031: 573:. The existence of chloroplast DNA was identified biochemically in 1959, and confirmed by electron microscopy in 1962. The discoveries that the chloroplast contains ribosomes and performs protein synthesis revealed that the chloroplast is genetically semi-autonomous. The first complete chloroplast genome sequences were published in 1986, 1610: 804:
In most plant species, the chloroplast genome encodes approximately 120 genes. The genes primarily encode core components of the photosynthetic machinery and factors involved in their expression and assembly. Across species of land plants, the set of genes encoded by the chloroplast genome is fairly
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is the insertion, deletion, and substitution of nucleotides in a mRNA transcript prior to translation to protein. The highly oxidative environment inside chloroplasts increases the rate of mutation so post-transcription repairs are needed to conserve functional sequences. The chloroplast editosome
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In cpDNA, there are several A → G deamination gradients. DNA becomes susceptible to deamination events when it is single stranded. When replication forks form, the strand not being copied is single stranded, and thus at risk for A → G deamination. Therefore, gradients in deamination indicate that
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coded by the chloroplast's own genome, which is related to RNA polymerases found in bacteria. Chloroplasts also contain a mysterious second RNA polymerase that is encoded by the plant's nuclear genome. The two RNA polymerases may recognize and bind to different kinds of
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flipped one of its inverted repeats (making them direct repeats). It is possible that the inverted repeats help stabilize the rest of the chloroplast genome, as chloroplast DNAs which have lost some of the inverted repeat segments tend to get rearranged more.
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Ohyama, K.; Fukuzawa, H.; Kohchi, T.; Shirai, H.; Sano, T.; Sano, S.; Umesono, K.; Shiki, Y.; Takeuchi, M.; Chang, Z. & Aota, S. (1986). "Chloroplast gene organization deduced from complete sequence of liverwort Marchantia polymorpha chloroplast DNA".
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Of the approximately three-thousand proteins found in chloroplasts, some 95% of them are encoded by nuclear genes. Many of the chloroplast's protein complexes consist of subunits from both the chloroplast genome and the host's nuclear genome. As a result,
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Over time, base changes in the DNA sequence can arise from deamination mutations. When adenine is deaminated, it becomes hypoxanthine, which can pair with cytosine. During replication, the cytosine will pair with guanine, causing an A → G base
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moves through the circular DNA, it adopts a theta intermediary form, also known as a Cairns replication intermediate, and completes replication with a rolling circle mechanism. Replication starts at specific points of origin. Multiple
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substitutes C -> U and U -> C at very specific locations on the transcript. This can change the codon for an amino acid or restore a non-functional pseudogene by adding an AUG start codon or removing a premature UAA stop codon.
1918: 1892:. They vary in the length of their A-domains, which is completely gone in Toc90. Toc132, Toc120, and Toc90 seem to have specialized functions in importing stuff like nonphotosynthetic preproteins, and can't replace Toc159. 2299:; the mature form probably undergoes processing that trims it down to something that weighs 56 thousand daltons when it gets imported into the chloroplast. Tic56 is largely embedded inside the 1 million dalton complex. 1005:
The editosome recognizes and binds to cis sequence upstream of the editing site. The distance between the binding site and editing site varies by gene and proteins involved in the editosome. Hundreds of different
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than the one that phosphorylates Toc34. Its M-domain forms part of the tunnel that chloroplast preproteins travel through, and seems to provide the force that pushes preproteins through, using the energy from
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Shaw J, Lickey EB, Schilling EE, Small RL (March 2007). "Comparison of whole chloroplast genome sequences to choose noncoding regions for phylogenetic studies in angiosperms: the tortoise and the hare III".
1687:. This loss of GTP makes the Toc34 protein release the chloroplast preprotein, handing it off to the next TOC protein. Toc34 then releases the depleted GDP molecule, probably with the help of an unknown 1867:. This suggests that it might act as a shuttle that finds chloroplast preproteins in the cytosol and carries them back to the TOC complex. There isn't a lot of direct evidence for this behavior though. 1010:
from the nuclear genome are involved in the RNA editing process. These proteins consist of 35-mer repeated amino acids, the sequence of which determines the cis binding site for the edited transcript.
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open up, allowing replication machinery to replicate the DNA. As replication continues, the forks grow and eventually converge. The new cpDNA structures separate, creating daughter cpDNA chromosomes.
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are added to the N-termini of the polypeptides, which are used to help direct the polypeptide to the chloroplast for import (N-terminal transit peptides are also used to direct polypeptides to plant
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to contain as few as four or as many as over 150 genes. While a given pair of inverted repeats are rarely completely identical, they are always very similar to each other, apparently resulting from
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The mechanism for chloroplast DNA (cpDNA) replication has not been conclusively determined, but two main models have been proposed. Scientists have attempted to observe chloroplast replication via
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Sun YJ, Forouhar F, Li Hm HM, Tu SL, Yeh YH, Kao S, Shr HL, Chou CC, Chen C, Hsiao CD (February 2002). "Crystal structure of pea Toc34, a novel GTPase of the chloroplast protein translocon".
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Ohyama, Kanji; Fukuzawa, Hideya; Kohchi, Takayuki; Shirai, Hiromasa; Sano, Tohru; Sano, Satoshi; Umesono, Kazuhiko; Shiki, Yasuhiko; Takeuchi, Masayuki; Chang, Zhen; Aota, Shin-ichi (1986).
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in the chloroplast are now synthesized in the cytoplasm. This means that these proteins must be directed back to the chloroplast, and imported through at least two chloroplast membranes.
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lineages. Even if a chloroplast is eventually lost, the genes it donated to the former host's nucleus persist, providing evidence for the lost chloroplast's existence. For example, while
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Chloroplast DNA has long been thought to have a circular structure, but some evidence suggests that chloroplast DNA more commonly takes a linear shape. Over 95% of the chloroplast DNA in
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Kikuchi S, BĂ©dard J, Hirano M, Hirabayashi Y, Oishi M, Imai M, Takase M, Ide T, Nakai M (February 2013). "Uncovering the protein translocon at the chloroplast inner envelope membrane".
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Basal land plants such as liverworts, mosses and ferns have hundreds of different editing sites while flowering plants typically have between thirty and forty. Parasitic plants such as
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The heat shock protein and the 14-3-3 proteins together form a cytosolic guidance complex that makes it easier for the chloroplast polypeptide to get imported into the chloroplast.
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long each. Inverted repeats in plants tend to be at the upper end of this range, each being 20,000–25,000 base pairs long. The inverted repeat regions usually contain three
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wide at the ends, and shrinks to about 1.4–1.6 nanometers in diameter at its narrowest point—wide enough to allow partially folded chloroplast preproteins to pass through.
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prematurely. This is important because it prevents chloroplast proteins from assuming their active form and carrying out their chloroplast functions in the wrong place—the
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Chloroplast DNA replication via multiple D loop mechanisms. Adapted from Krishnan NM, Rao BJ's paper "A comparative approach to elucidate chloroplast genome replication."
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compared to that of free-living cyanobacteria. Chloroplasts may contain 60–100 genes whereas cyanobacteria often have more than 1500 genes in their genome. The parasitic
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must be coordinated between the chloroplast and the nucleus. The chloroplast is mostly under nuclear control, though chloroplasts can also give out signals regulating
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Shinozaki, K.; Ohme, M.; Tanaka, M.; Wakasugi, T.; Hayashida, N.; Matsubayashi, T.; Zaita, N.; Chunwongse, J.; Obokata, J.; Yamaguchi-Shinozaki, K.; Ohto, C. (1986).
901:, corresponding to about 4,500 protein-coding genes. There have been a few recent transfers of genes from the chloroplast DNA to the nuclear genome in land plants. 561:) is the DNA located in chloroplasts, which are photosynthetic organelles located within the cells of some eukaryotic organisms. Chloroplasts, like other types of 3129:
Shinozaki, K.; Ohme, M.; Tanaka, M.; Wakasugi, T.; Hayashida, N.; Matsubayashi, T.; Zaita, N.; Chunwongse, J.; Obokata, J. & Yamaguchi-Shinozaki, K. (1986).
2012:. Chloroplast polypeptide chains probably often travel through the two complexes at the same time, but the TIC complex can also retrieve preproteins lost in the 3528:"Evolutionary analysis of Arabidopsis, cyanobacterial, and chloroplast genomes reveals plastid phylogeny and thousands of cyanobacterial genes in the nucleus" 1042:
since the 1970s. The results of the microscopy experiments led to the idea that chloroplast DNA replicates using a double displacement loop (D-loop). As the
4620:"A transit peptide-like sorting signal at the C terminus directs the Bienertia sinuspersici preprotein receptor Toc159 to the chloroplast outer membrane" 1512:
Alternatively, if a chloroplast preprotein's transit peptide is not phosphorylated, a chloroplast preprotein can still attach to a heat shock protein or
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among land plants, and accumulate few mutations. Similar inverted repeats exist in the genomes of cyanobacteria and the other two chloroplast lineages (
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might be the exchange factor that carry out the GDP removal. The Toc34 protein can then take up another molecule of GTP and begin the cycle again.
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The first three proteins form a core complex that consists of one Toc159, four to five Toc34s, and four Toc75s that form four holes in a disk 13
861:. Contrarily, there are only a few known instances where genes have been transferred to the chloroplast from various donors, including bacteria. 488: 725:
Each chloroplast contains around 100 copies of its DNA in young leaves, declining to 15–20 copies in older leaves. They are usually packed into
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Palmer JD, Thompson WF (June 1982). "Chloroplast DNA rearrangements are more frequent when a large inverted repeat sequence is lost".
2141: 2056: 1739: 1735: 1346:. They can be from 20 to 150 amino acids long—an unusually long length, suggesting that transit peptides are actually collections of 2303: 2253: 2201: 2182: 2178: 2162: 2158: 2048: 2044: 2028: 1879: 1875: 1871: 1785: 1696: 1574: 1513: 2284: 2230: 2186: 2166: 2137: 2097: 2067: 2052: 2036: 2032: 1960: 1925: 1883: 1821: 1797: 1634: 1586: 1582: 1578: 1570: 1389:
Not all chloroplast proteins include a N-terminal cleavable transit peptide though. Some include the transit sequence within the
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genes in the diatom nucleus provide evidence that the diatom ancestor (probably the ancestor of all chromalveolates too) had a
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and are accessible via the NCBI organelle genome database. The first chloroplast genomes were sequenced in 1986, from tobacco (
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Curiously, around half of the protein products of transferred genes aren't even targeted back to the chloroplast. Many became
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Huang CY, Ayliffe MA, Timmis JN (March 2003). "Direct measurement of the transfer rate of chloroplast DNA into the nucleus".
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Berry, J. O.; Yerramsetty, P.; Zielinski, A. M. & Mure, C. M. (2013). "Photosynthetic gene expression in higher plants".
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Tillich M, Krause K (July 2010). "The ins and outs of editing and splicing of plastid RNAs: lessons from parasitic plants".
740:, a histone-like chloroplast protein (HC) coded by the chloroplast DNA that tightly packs each chloroplast DNA ring into a 2268:, and since the mature protein probably doesn't lose any amino acids when itself imported into the chloroplast (it has no 1506:. At the same time, they have to keep just enough shape so that they can be recognized and imported into the chloroplast. 920: 3526:
Martin W, Rujan T, Richly E, Hansen A, Cornelsen S, Lins T, Leister D, Stoebe B, Hasegawa M, Penny D (September 2002).
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Hedtke B, Börner T, Weihe A (August 1997). "Mitochondrial and chloroplast phage-type RNA polymerases in Arabidopsis".
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Agne B, Andrès C, Montandon C, Christ B, Ertan A, Jung F, Infanger S, Bischof S, Baginsky S, Kessler F (July 2010).
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molecule, inhibiting the protein's activity. This might provide a way to regulate protein import into chloroplasts.
1620:. Toc34 has three almost identical molecules (shown in slightly different shades of green), each of which forms a 2569:"Chloroplast gene organization deduced from complete sequence of liverwort Marchantia polymorpha chloroplast DNA" 2273: 2013: 889: 467: 3943:
Koussevitzky S, Nott A, Mockler TC, Hong F, Sachetto-Martins G, Surpin M, Lim J, Mittler R, Chory J (May 2007).
2272:), it was named Tic100. Tic100 is found at the edges of the 1 million dalton complex on the side that faces the 4513:"Functional characterization of sequence motifs in the transit peptide of Arabidopsis small subunit of rubisco" 4210: 4077:
Takenaka M, Zehrmann A, Verbitskiy D, Härtel B, Brennicke A (2013). "RNA editing in plants and its evolution".
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of Toc34 because it can be turned off by phosphorylation. AtToc34 on the other hand cannot be phosphorylated.
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instead. Most of the polypeptides that lack N-terminal targeting sequences are the ones that are sent to the
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which can contain several identical chloroplast DNA rings. Many nucleoids can be found in each chloroplast.
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In land plants, some 11–14% of the DNA in their nuclei can be traced back to the chloroplast, up to 18% in
3131:"The complete nucleotide sequence of the tobacco chloroplast genome: its gene organization and expression" 3060:"Detection and localization of a chloroplast-encoded HU-like protein that organizes chloroplast nucleoids" 3058:
Kobayashi T, Takahara M, Miyagishima SY, Kuroiwa H, Sasaki N, Ohta N, Matsuzaki M, Kuroiwa T (July 2002).
2512:"The complete nucleotide sequence of the tobacco chloroplast genome: its gene organization and expression" 2348:
Stocking, C. R. & Gifford, E. M. (1959). "Incorporation of thymidine into chloroplasts of Spirogyra".
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Most chloroplasts have their entire chloroplast genome combined into a single large ring, though those of
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long. They can have a contour length of around 30–60 micrometers, and have a mass of about 80–130 million
2323: 843: 768: 586: 1600:. The other two proteins, Toc64 and Toc12, are associated with the core complex but are not part of it. 751:, the chloroplast DNA nucleoids are clustered in the center of a chloroplast, while in green plants and 2242: 2177:
and nearly all chloroplast lineages, suggesting it evolved before the first chloroplast endosymbiosis.
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attaches to Toc34, the protein becomes much more able to bind to many chloroplast preproteins in the
1445:(both very common in chloroplast transit sequences—making up 20–30% of the sequence) are often the 1208: 1096: 933: 798: 519: 3640:"The case of horizontal gene transfer from bacteria to the peculiar dinoflagellate plastid genome" 1188:. In those cases, chloroplast-targeted proteins do initially travel along the secretory pathway). 4997: 3738:
Robison, TA, Grusz AL, Wolf PG, Mower, JP, Fauskee BD, Sosa K, and Schuettpelz E (October 2018).
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Toc75 can also bind to chloroplast preproteins, but is a lot worse at this than Toc34 or Toc159.
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One of the main competing models for cpDNA asserts that most cpDNA is linear and participates in
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among land plants, but they don't resemble any protein whose function is known. Neither has any
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In addition to the early microscopy experiments, this model is also supported by the amounts of
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Toc159 is not always found as part of the TOC complex—it has also been found dissolved in the
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chloroplasts has been observed to be in branched linear form rather than individual circles.
581: 17: 4116:. Special Issue: Biotechnology Annual Review 2010RNA Basics and Biotechnology Applications. 702:), suggesting that they predate the chloroplast, though some chloroplast DNAs like those of 4992: 4982: 4902: 3956: 3899: 3803: 3539: 3479: 3428: 3368: 3188: 2914: 2663: 2580: 2468: 2220:
end. Tic214 is notable for being coded for by chloroplast DNA, more specifically the first
2149: 2132: 2075: 1951: 1888: 1726: 1145: 607: 3014: 1074:, and when the HC base pair is replicated, it becomes a GC (thus, an A → G base change). 664:, which separate a long single copy section (LSC) from a short single copy section (SSC). 8: 1758: 1688: 1684: 1343: 1185: 1039: 1014: 948: 684: 4906: 4400:
Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences
4090: 3960: 3903: 3807: 3543: 3483: 3432: 3372: 3192: 2918: 2667: 2584: 2472: 1937: 4926: 4853: 4828: 4800: 4705: 4644: 4619: 4571:"14-3-3 proteins form a guidance complex with chloroplast precursor proteins in plants" 4537: 4512: 4420: 4395: 4376: 4186: 4159: 3925: 3827: 3764: 3739: 3715: 3688: 3664: 3639: 3615: 3586: 3503: 3452: 3332: 3305: 3240: 3224:"Complete genome structure of the unicellular cyanobacterium Synechocystis sp. PCC6803" 3223: 3204: 3155: 3146: 3130: 2988: 2742: 2604: 2544: 2527: 2511: 2492: 2401: 2376: 2221: 2170: 1829: 1809: 1481: 1458: 1351: 1165: 1141: 814: 691: 495: 474: 460: 439: 418: 285: 271: 229: 138: 131: 108: 4753: 4595: 4570: 4506: 4504: 4502: 4500: 4051: 4026: 3944: 3562: 3527: 3418: 3084: 3059: 2937: 2902: 2873: 2848: 4951: 4918: 4858: 4792: 4757: 4697: 4649: 4600: 4542: 4484: 4450: 4425: 4368: 4191: 4129: 4094: 4056: 4007: 3972: 3917: 3868: 3819: 3788: 3769: 3720: 3669: 3620: 3567: 3495: 3444: 3396: 3391: 3356: 3337: 3286: 3245: 3160: 3089: 3018: 2980: 2976: 2942: 2878: 2814: 2770: 2734: 2691: 2686: 2651: 2596: 2549: 2531: 2484: 2441: 2437: 2406: 2361: 2328: 2307: 2086: 1774: 1731: 1161: 1117:
The movement of so many chloroplast genes to the nucleus means that many chloroplast
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show a loss of RNA editing resulting in a loss of function for photosynthesis genes.
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Moustafa A, Beszteri B, Maier UG, Bowler C, Valentin K, Bhattacharya D (June 2009).
3456: 3306:"Chloroplast genomes: diversity, evolution, and applications in genetic engineering" 2992: 2746: 601: 4930: 4910: 4848: 4840: 4829:"The acidic A-domain of Arabidopsis TOC159 occurs as a hyperphosphorylated protein" 4784: 4749: 4687: 4639: 4631: 4590: 4582: 4532: 4524: 4497: 4415: 4407: 4360: 4222: 4181: 4171: 4121: 4086: 4046: 4042: 4038: 3999: 3964: 3907: 3858: 3811: 3759: 3751: 3710: 3700: 3659: 3651: 3610: 3602: 3557: 3547: 3507: 3487: 3436: 3386: 3376: 3327: 3317: 3276: 3235: 3208: 3196: 3150: 3142: 3079: 3071: 2972: 2932: 2922: 2868: 2860: 2726: 2681: 2671: 2608: 2588: 2539: 2523: 2496: 2476: 2433: 2396: 2388: 2357: 1813: 1638: 1383: 1100: 1048: 877: 848: 680: 4345: 3845:
Nowack EC, Vogel H, Groth M, Grossman AR, Melkonian M, Glöckner G (January 2011).
2241:. Tic20 is buried inside the complex while Tic214 is exposed on both sides of the 1843:
Toc159 probably works a lot like Toc34, recognizing proteins in the cytosol using
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Proceedings of the National Academy of Sciences of the United States of America
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Proceedings of the National Academy of Sciences of the United States of America
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Proceedings of the National Academy of Sciences of the United States of America
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long. Each minicircle contains one to three genes, but blank plasmids, with no
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Jarvis P, Soll J (December 2001). "Toc, Tic, and chloroplast protein import".
4125: 3912: 3887: 3838: 3705: 3440: 3322: 2758: 2756: 4971: 4176: 3281: 3264: 2600: 2535: 2459:
Heber, U. (1962). "Protein synthesis in chloroplasts during photosynthesis".
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Lyttleton, J. W. (1962). "Isolation of ribosomes from spinach chloroplasts".
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long. The 871 amino acids collectively weigh slightly less than 100 thousand
2234: 2174: 2093: 2082: 2040: 1929: 1621: 1430: 1379: 1375: 1204: 1181: 1173: 1169: 1157: 1133: 672: 623: 411: 390: 376: 292: 236: 201: 187: 45: 4914: 4692: 4675: 4436: 4018: 3968: 3863: 3846: 3815: 3381: 2676: 2377:"Ultrastructure of DNA-containing areas in the chloroplast of Chlamydomonas" 2204:
is another TIC core complex protein, named because it weighs just under 214
544:, segments with labels on the outside are on the A strand. Notches indicate 4987: 4922: 4862: 4796: 4761: 4676:"Phosphorylation of the transit sequence of chloroplast precursor proteins" 4653: 4635: 4611: 4604: 4546: 4429: 4411: 4372: 4195: 4133: 4098: 3976: 3921: 3872: 3823: 3773: 3724: 3673: 3624: 3571: 3552: 3521: 3519: 3517: 3499: 3448: 3341: 3290: 3164: 3128: 3093: 2946: 2882: 2753: 2738: 2568: 2553: 2488: 2445: 2410: 1462: 1393:
of the protein itself. A few have their transit sequence appended to their
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that often results in base changes. When adenine is deaminated, it becomes
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Lee DW, Lee S, Lee GJ, Lee KH, Kim S, Cheong GW, Hwang I (February 2006).
4060: 4011: 3400: 3249: 3177: 2984: 2695: 2124:), it has been proposed to be part of the TIC import channel. There is no 2078: 797:
origin of the chloroplast. It also demonstrated the significant extent of
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Sato, S.; Nakamura, Y.; Kaneko, T.; Asamizu, E. & Tabata, S. (1999).
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The inverted repeats vary wildly in length, ranging from 4,000 to 25,000
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are a notable exception—their genome is broken up into about forty small
615: 243: 215: 4586: 4226: 3945:"Signals from chloroplasts converge to regulate nuclear gene expression" 3780: 3514: 3491: 2842: 2840: 2838: 2836: 2834: 2832: 2830: 2295:
its gene encodes is 527 amino acids long, weighing close to 62 thousand
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is the most abundant protein on the outer chloroplast envelope. It is a
1907: 1160:. Some transferred chloroplast DNA protein products get directed to the 962: 837:
Over time, many parts of the chloroplast genome were transferred to the
4669: 4667: 4665: 4663: 3591:(Apodanthaceae) Each Retain Just Five or Six Possibly Functional Genes" 3075: 2864: 2292: 2217: 2089: 2004: 1941: 1906:
The general shape of a β-barrel is a hollow cylinder lined by multiple
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M-domain, which anchors the protein to the outer chloroplast membrane.
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to the Toc34 protein, preventing it from being able to receive another
1679:. The chloroplast preprotein's presence causes Toc34 to break GTP into 1645: 1593: 1541: 1394: 1310: 1302: 1269: 1254: 892:
at some point, which was subsequently replaced by the red chloroplast.
853: 668: 642: 638: 619: 38: 3655: 3265:"Complete structure of the chloroplast genome of Arabidopsis thaliana" 1472:
Phosphorylation changes the polypeptide's shape, making it easier for
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Clegg, M. T.; Gaut, B. S.; Learn, G. H. & Morton, B. R. (1994).
1030: 4788: 3057: 2126: 1956: 1418: 1294: 1118: 1071: 1063: 952: 741: 712: 634: 3789:"Genomic footprints of a cryptic plastid endosymbiosis in diatoms" 1812:
and takes up about half the protein length. The A-domain is often
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Chloroplast transit peptides exhibit huge variation in length and
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in the outer chloroplast membrane that's anchored into it by its
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for secretion), the new chloroplast host had to develop a unique
1177: 873: 832: 822: 733: 562: 545: 59: 4396:"The endosymbiotic origin, diversification and fate of plastids" 4076: 3303: 821:
complex that are involved in plastid gene expression. The large
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have since lost the inverted repeats. Others, like the red alga
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surrounded by some loosely associated peripheral proteins like
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synthesize polypeptides from the N-terminus to the C-terminus.
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outside of the cell, because to reach the chloroplast from the
1043: 566: 540:. Segments with labels on the inside reside on the B strand of 4892: 4553: 4387: 3879: 1480:
only bind to chloroplast preproteins. It is also bound by the
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Biochimica et Biophysica Acta (BBA) - Molecular Cell Research
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because they are located at the "front" end of a polypeptide—
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to avoid having chloroplast proteins being sent to the wrong
1168:
are bounded by an outermost membrane derived from the host's
810: 502: 334: 299: 264: 194: 94: 87: 66: 2645: 2643: 2641: 2008:
is another protein complex that imports proteins across the
4208: 3942: 3786: 3053: 3051: 2509: 1762: 1153: 858: 762: 703: 676: 649: 605:
The 154 kb chloroplast DNA map of a model flowering plant (
453: 446: 425: 397: 383: 369: 362: 348: 278: 208: 159: 145: 124: 117: 73: 3680: 3637: 3034: 2958: 2956: 2783: 2566: 2140:
proteins in the outer chloroplast membrane, there are two
1710:
drifting around on the outer chloroplast membrane can use
1437:
to many (but not all) of them in their transit sequences.
4826: 4339: 4337: 4335: 4333: 4331: 4329: 4327: 4325: 4323: 4321: 4319: 4317: 4315: 4313: 4311: 4309: 4307: 4305: 4303: 4301: 4299: 4297: 4295: 4293: 4291: 4289: 4287: 4285: 4283: 4281: 2711: 2709: 2707: 2705: 2638: 1461:
for chloroplast polypeptides, and ignores ones meant for
1317:. For many (but not all) chloroplast proteins encoded by 785:). Comparison of the gene sequences of the cyanobacteria 541: 4279: 4277: 4275: 4273: 4271: 4269: 4267: 4265: 4263: 4261: 3888:"Algal genomics: exploring the imprint of endosymbiosis" 3844: 3587:"The Plastomes of Two Species in the Endoparasite Genus 3048: 2903:"Inverted repeats in chloroplast DNA from higher plants" 2715: 1932:
tube that forms most of the TOC pore itself. Toc75 is a
1648:
tail. Most of the protein, however, including its large
1219:
Cytoplasmic translation and N-terminal transit sequences
801:
from the cyanobacterial ancestor to the nuclear genome.
3525: 2953: 2896: 2894: 2892: 1667:
and hand them off to the rest of the TOC complex. When
3686: 2849:"Circular chloroplast chromosomes: the grand illusion" 2702: 2233:
together probably make up the part of the one million
1350:
with different functions. Transit peptides tend to be
904: 27:
DNA located in cellular organelles called chloroplasts
4258: 2769:. Cambridge: Cambridge university press. p. 62. 2620: 2618: 1025: 864:
Endosymbiotic gene transfer is how we know about the
618:
DNAs are circular, and are typically 120,000–170,000
4241:"Effect of chemical mutagens on nucleotide sequence" 4024: 3304:
Daniell, H.; Lin, C.; Yu, M. & Chang, W (2016).
3221: 2889: 2650:
Clegg MT, Gaut BS, Learn GH, Morton BR (July 1994).
2649: 955:
in chloroplasts are similar to bacterial ribosomes.
829:
proteins are encoded within the chloroplast genome.
4025:Harris EH, Boynton JE, Gillham NW (December 1994). 3740:"Mobile Elements Shape Plastome Evolution in Ferns" 3631: 2350:
Biochemical and Biophysical Research Communications
1090: 942:Protein synthesis within chloroplasts relies on an 732:Though chloroplast DNA is not associated with true 611:: Brassicaceae) showing genes and inverted repeats. 4774: 3737: 3638:Mackiewicz P, BodyĹ‚ A, MoszczyĹ„ski K (July 2013). 3469: 3006: 2810:The Chloroplast: Interactions with the Environment 2615: 2193:chloroplasts, suggesting that they evolved later. 3989: 3357:"Rates and patterns of chloroplast DNA evolution" 3044:. Benjamin Cummings (Pearson). 2009. p. 516. 2999: 2652:"Rates and patterns of chloroplast DNA evolution" 1979:The translocon on the inner chloroplast membrane 1528:The translocon on the outer chloroplast membrane 1144:. A few chloroplast genes found new homes in the 847:. As a result, the chloroplast genome is heavily 4969: 2423: 2374: 1596:across. The whole core complex weighs about 500 1331:). N-terminal transit sequences are also called 1132:, taking on new functions like participating in 4072: 4070: 1301:. The two ends of a polypeptide are called the 1106: 4888: 4886: 4884: 4882: 4880: 4878: 4876: 4874: 4872: 4735: 4733: 4731: 4729: 4727: 4725: 4723: 4721: 4719: 3100: 3013:(3rd ed.). Academic Press. 2005. p.  2962: 2900: 1624:with one of its adjacent molecules. Part of a 833:Chloroplast genome reduction and gene transfer 4943: 4673: 4233: 4111: 4027:"Chloroplast ribosomes and protein synthesis" 3171: 1828:binding domain, which is very similar to the 1628:molecule binding site is highlighted in pink. 773:More than 5000 chloroplast genomes have been 755:, the nucleoids are dispersed throughout the 534:Interactive gene map of chloroplast DNA from 4768: 4510: 4343: 4209:Heinhorst, Gordon C. Cannon, Sabine (1993). 4067: 3687:Leliaert F, Lopez-Bautista JM (March 2015). 3585:Bellot, Sidonie; Renner, Susanne S. (2016). 3584: 3414: 3412: 3410: 2417: 2039:. The core complex weighs about one million 1565:of the TOC complex have been identified—two 1386:, the transit sequence forms a random coil. 4869: 4716: 4157: 4153: 4151: 4149: 4147: 4145: 4143: 3731: 3297: 2341: 4739: 4617: 4476: 3215: 3122: 3029:number of copies of ctDNA per chloroplast. 1917: 1609: 1409:Phosphorylation, chaperones, and transport 1238: 1058:seen in cpDNA. Deamination occurs when an 993: 4852: 4691: 4643: 4594: 4536: 4419: 4202: 4185: 4175: 4050: 3911: 3885: 3862: 3763: 3714: 3704: 3663: 3614: 3561: 3551: 3407: 3390: 3380: 3348: 3331: 3321: 3280: 3239: 3154: 3083: 2936: 2926: 2872: 2806: 2766:An introduction to plant cell development 2685: 2675: 2543: 2458: 2400: 1757:), which are each about 60% identical in 1659:Toc34's job is to catch some chloroplast 1476:to attach to the polypeptide. In plants, 1191:Because the cell acquiring a chloroplast 4449:. Benjamin Cummings. 2008. p. 340. 4447:Biology 8th edition—Campbell & Reece 4140: 3256: 3042:Biology 8th Edition Campbell & Reece 2452: 2136:, it is known that for about every five 1520:on the outer chloroplast membrane using 1457:that carries out the phosphorylation is 1076: 1029: 817:, and 4 subunits of the plastid-encoded 763:Gene content and plastid gene expression 600: 585:(liverwort) by Ozeki et al. Since then, 579:(tobacco) by Sugiura and colleagues and 4568: 4393: 2846: 2762: 1184:, just like if you were headed for the 857:have even lost their plastid genes for 690:The inverted repeat regions are highly 14: 4970: 4353:Nature Reviews. Molecular Cell Biology 2901:Kolodner R, Tewari KK (January 1979). 2368: 2085:TIC complex. Because it is similar to 2023:, the TIC translocon has a large core 1808:end is the A-domain, which is rich in 1769:Toc34). AtToc33 is the most common in 1099:and replication structures similar to 789:to those of the chloroplast genome of 596: 4947:Mitochondrial Function and Biogenesis 2154:) in the inner chloroplast membrane. 4483:. Berlin: Springer. pp. 53–74. 3222:Kaneko, T. & Tabata, S. (1997). 2624: 1910:. Note that the protein depicted is 1832:GTP-binding domain in Toc34. At the 1253:linked together. At the left is the 957: 927: 4680:The Journal of Biological Chemistry 4091:10.1146/annurev-genet-111212-133519 951:within the chloroplast genome. The 905:Proteins encoded by the chloroplast 679:genes, but they can be expanded or 655: 24: 4674:Waegemann K, Soll J (March 1996). 4480:Structure and function of plastids 4346:"Protein import into chloroplasts" 3241:10.1093/oxfordjournals.pcp.a029103 3147:10.1002/j.1460-2075.1986.tb04464.x 2634:. Green DNA: BIOSCIENCE EXPLAINED. 2528:10.1002/j.1460-2075.1986.tb04464.x 1516:. These complexes can bind to the 1026:Leading model of cpDNA replication 1020: 660:Many chloroplast DNAs contain two 587:a great number of chloroplast DNAs 25: 5009: 4618:Lung SC, Chuong SD (April 2012). 4344:Soll J, Schleiff E (March 2004). 4211:"DNA replication in chloroplasts" 3108:"NCBI Organelle Genome Resources" 1940:. The hole it forms is about 2.5 4158:Krishnan NM, Rao BJ (May 2009). 2375:Ris, H. & Plaut, W. (1962). 2212:long and is thought to have six 1702:Toc34 can be turned off through 1671:, an energy molecule similar to 1498:that keeps the polypeptide from 1401:, plus at least one sent to the 1091:Alternative model of replication 961: 32: 4937: 4105: 3851:Molecular Biology and Evolution 3578: 3463: 2274:chloroplast intermembrane space 2081:. It is found in the 1 million 1975:is the most abundant of these. 1603: 890:green algal derived chloroplast 589:from various species have been 569:separate from that in the cell 4944:Curran SP, Koehler CM (2004). 4569:May T, Soll J (January 2000). 4043:10.1128/MMBR.58.4.700-754.1994 3886:Archibald JM (December 2006). 3110:. National Institute of Health 2560: 2503: 1847:. It can be regulated through 1656:projects out into the stroma. 1070:(H). Hypoxanthine can bind to 841:of the host, a process called 825:subunit and 28 photosynthetic 805:conserved. This includes four 13: 1: 4754:10.1016/S0167-4889(01)00147-1 2334: 2130:evidence for this though. In 2074:protein thought to have four 1870:A family of Toc159 proteins, 882:red algal derived chloroplast 793:provided confirmation of the 4004:10.1126/science.277.5327.809 3744:Genome Biology and Evolution 3595:Genome Biology and Evolution 2977:10.1016/0092-8674(82)90170-2 2438:10.1016/0014-4827(62)90183-0 2362:10.1016/0006-291X(59)90010-5 1577:, the protein import tunnel 1107:Protein targeting and import 720: 7: 4477:Wise RR, Hoober JK (2007). 2324:List of sequenced plastomes 2317: 2237:TIC complex that spans the 1773:, and it is the functional 1268:) group in green. The blue 1148:—most became nonfunctional 844:endosymbiotic gene transfer 769:List of sequenced plastomes 10: 5014: 2719:American Journal of Botany 2243:inner chloroplast membrane 2059:, possibly three of each. 2010:inner chloroplast envelope 1559:outer chloroplast envelope 1403:inner chloroplast membrane 1399:outer chloroplast membrane 1324:cleavable transit peptides 1222: 1110: 931: 766: 182:replication origin regions 154:replication origin regions 4777:Nature Structural Biology 4394:Keeling PJ (March 2010). 4126:10.1016/j.nbt.2010.02.020 4079:Annual Review of Genetics 3913:10.1016/j.cub.2006.11.008 3706:10.1186/s12864-015-1418-3 3441:10.1007/s11120-013-9880-8 3323:10.1186/s13059-016-1004-2 2270:cleavable transit peptide 2248: 2196: 1780: 1121:that were supposed to be 4950:. Springer. p. 59. 4177:10.1186/1471-2164-10-237 2847:Bendich AJ (July 2004). 2813:. Springer. p. 18. 2279: 2062: 1895: 1209:protein targeting system 1180:, you have to cross the 1156:genes still work in the 1097:homologous recombination 645:, have also been found. 4915:10.1126/science.1229262 4693:10.1074/jbc.271.11.6545 4215:Journal of Cell Science 4031:Microbiological Reviews 3969:10.1126/science.1140516 3816:10.1126/science.1172983 3644:Mobile Genetic Elements 3382:10.1073/pnas.91.15.6795 2677:10.1073/pnas.91.15.6795 1824:. In the middle is its 994:RNA editing in plastids 918:in the nucleus, called 884:, the presence of many 4636:10.1105/tpc.112.096248 4412:10.1098/rstb.2009.0103 3553:10.1073/pnas.182432999 3282:10.1093/dnares/6.5.283 1963:that are named by the 1650:guanosine triphosphate 1429:energy can be used to 1083: 1035: 612: 490:atp-dependent protease 54:acetyl-CoA carboxylase 4845:10.1104/pp.110.158048 4529:10.1104/pp.105.074575 3864:10.1093/molbev/msq209 2807:Sandelius AS (2009). 2312:transmembrane domains 2214:transmembrane domains 2092:transporters and the 1851:, but by a different 1681:guanosine diphosphate 1080: 1033: 783:Marchantia polymorpha 604: 582:Marchantia polymorpha 2928:10.1073/pnas.76.1.41 2731:10.3732/ajb.94.3.275 2628:Bioscience—Explained 2393:10.1083/jcb.13.3.383 2133:Arabidopsis thaliana 2003:hloroplast membrane 1971:that code for them. 1952:Arabidopsis thaliana 1936:channel lined by 16 1889:Arabidopsis thaliana 1727:Arabidopsis thaliana 1581:, plus the proteins 1417:is synthesized on a 1413:After a chloroplast 1358:amino acids such as 1293:, the precursors of 1146:mitochondrial genome 921:retrograde signaling 637:, each 2,000–10,000 608:Arabidopsis thaliana 4907:2013Sci...339..571K 4587:10.1105/tpc.12.1.53 4227:10.1242/jcs.104.1.1 3961:2007Sci...316..715K 3904:2006CBio...16R1033A 3808:2009Sci...324.1724M 3544:2002PNAS...9912246M 3492:10.1038/nature01435 3484:2003Natur.422...72H 3433:2013PhoRe.117...91B 3373:1994PNAS...91.6795C 3193:1986Natur.322..572O 2919:1979PNAS...76...41K 2668:1994PNAS...91.6795C 2585:1986Natur.322..572O 2473:1962Natur.195...91H 2260:protein that's 871 2144:proteins (the main 2014:intermembrane space 1914:Toc75 specifically. 1886:have been found in 1816:off, leaving an 86 1800:. Toc159 has three 1759:amino acid sequence 1689:GDP exchange factor 1685:inorganic phosphate 1555:hloroplast membrane 1344:amino acid sequence 1284:H) is at the right. 1186:extracellular space 1040:electron microscopy 1015:Epifagus virginiana 809:, approximately 30 685:concerted evolution 597:Molecular structure 469:initiation factor 1 3756:10.1093/gbe/evy189 3607:10.1093/gbe/evv251 3228:Plant Cell Physiol 3076:10.1105/tpc.002717 3009:Plant Biochemistry 2865:10.1105/tpc.160771 2763:Burgess J (1989). 2222:open reading frame 1810:acidic amino acids 1569:-binding proteins 1352:positively charged 1166:secondary plastids 1142:disease resistance 1084: 1036: 815:ribosomal proteins 613: 497:ribosomal proteins 476:ribosomal proteins 462:ribosomal proteins 441:nadh dehydrogenase 420:nadh dehydrogenase 287:ribosomal proteins 273:nadh dehydrogenase 231:ribosomal proteins 140:ribosomal proteins 133:nadh dehydrogenase 4490:978-1-4020-6570-5 4456:978-0-321-54325-7 4114:New Biotechnology 3750:(10): 2669–2571. 3656:10.4161/mge.25845 3367:(15): 6795–6801. 3234:(11): 1171–1176. 3187:(6079): 572–574. 2820:978-3-540-68696-5 2776:978-0-521-31611-8 2579:(6079): 572–574. 2329:Mitochondrial DNA 1995:ranslocon on the 1967:positions of the 1374:amino acids like 1162:secretory pathway 1113:Protein targeting 1062:is lost and is a 1049:replication forks 991: 990: 928:Protein synthesis 912:protein synthesis 866:lost chloroplasts 781:) and liverwort ( 779:Nicotiana tabacum 576:Nicotiana tabacum 537:Nicotiana tabacum 512:nicotiana tabacum 434:ribosomal protein 406:ribosomal protein 343:ribosomal protein 322:ribosomal protein 175:ribosomal protein 16:(Redirected from 5005: 4962: 4961: 4941: 4935: 4934: 4890: 4867: 4866: 4856: 4833:Plant Physiology 4824: 4809: 4808: 4772: 4766: 4765: 4737: 4714: 4713: 4695: 4671: 4658: 4657: 4647: 4615: 4609: 4608: 4598: 4566: 4551: 4550: 4540: 4517:Plant Physiology 4508: 4495: 4494: 4474: 4461: 4460: 4443: 4434: 4433: 4423: 4406:(1541): 729–48. 4391: 4385: 4384: 4350: 4341: 4256: 4255: 4253: 4251: 4237: 4231: 4230: 4206: 4200: 4199: 4189: 4179: 4155: 4138: 4137: 4109: 4103: 4102: 4074: 4065: 4064: 4054: 4022: 4016: 4015: 3998:(5327): 809–11. 3987: 3981: 3980: 3940: 3934: 3933: 3915: 3883: 3877: 3876: 3866: 3842: 3836: 3835: 3802:(5935): 1724–6. 3793: 3784: 3778: 3777: 3767: 3735: 3729: 3728: 3718: 3708: 3684: 3678: 3677: 3667: 3635: 3629: 3628: 3618: 3582: 3576: 3575: 3565: 3555: 3538:(19): 12246–51. 3523: 3512: 3511: 3467: 3461: 3460: 3416: 3405: 3404: 3394: 3384: 3352: 3346: 3345: 3335: 3325: 3301: 3295: 3294: 3284: 3260: 3254: 3253: 3243: 3219: 3213: 3212: 3201:10.1038/322572a0 3175: 3169: 3168: 3158: 3141:(9): 2043–2049. 3126: 3120: 3119: 3117: 3115: 3104: 3098: 3097: 3087: 3055: 3046: 3045: 3038: 3032: 3031: 3012: 3003: 2997: 2996: 2960: 2951: 2950: 2940: 2930: 2898: 2887: 2886: 2876: 2844: 2825: 2824: 2804: 2781: 2780: 2760: 2751: 2750: 2713: 2700: 2699: 2689: 2679: 2662:(15): 6795–801. 2647: 2636: 2635: 2633: 2622: 2613: 2612: 2593:10.1038/322572a0 2564: 2558: 2557: 2547: 2522:(9): 2043–2049. 2516:The EMBO Journal 2507: 2501: 2500: 2481:10.1038/195091a0 2456: 2450: 2449: 2421: 2415: 2414: 2404: 2372: 2366: 2365: 2345: 1982: 1938:β-pleated sheets 1921: 1820:fragment called 1639:integral protein 1613: 1531: 1449:that accept the 1384:aqueous solution 1297:, are chains of 1242: 1172:, and therefore 1101:bacteriophage T4 986: 983: 965: 958: 878:heterokontophyte 744:has been found. 662:inverted repeats 656:Inverted repeats 514: 505: 498: 491: 484: 477: 470: 463: 456: 449: 442: 435: 428: 421: 414: 407: 400: 393: 386: 379: 372: 365: 358: 351: 344: 337: 330: 323: 316: 309: 302: 295: 288: 281: 274: 267: 260: 253: 246: 239: 232: 225: 218: 211: 204: 197: 190: 183: 176: 169: 162: 155: 148: 141: 134: 127: 120: 113: 104: 97: 90: 83: 76: 69: 62: 55: 48: 41: 36: 35: 21: 5013: 5012: 5008: 5007: 5006: 5004: 5003: 5002: 4968: 4967: 4966: 4965: 4958: 4942: 4938: 4901:(6119): 571–4. 4891: 4870: 4825: 4812: 4773: 4769: 4738: 4717: 4686:(11): 6545–54. 4672: 4661: 4616: 4612: 4567: 4554: 4509: 4498: 4491: 4475: 4464: 4457: 4445: 4444: 4437: 4392: 4388: 4365:10.1038/nrm1333 4348: 4342: 4259: 4249: 4247: 4239: 4238: 4234: 4207: 4203: 4156: 4141: 4110: 4106: 4075: 4068: 4023: 4019: 3988: 3984: 3955:(5825): 715–9. 3941: 3937: 3898:(24): R1033-5. 3892:Current Biology 3884: 3880: 3843: 3839: 3791: 3785: 3781: 3736: 3732: 3685: 3681: 3636: 3632: 3583: 3579: 3524: 3515: 3468: 3464: 3421:Photosynth. Res 3417: 3408: 3353: 3349: 3302: 3298: 3261: 3257: 3220: 3216: 3176: 3172: 3127: 3123: 3113: 3111: 3106: 3105: 3101: 3056: 3049: 3040: 3039: 3035: 3025: 3005: 3004: 3000: 2961: 2954: 2899: 2890: 2845: 2828: 2821: 2805: 2784: 2777: 2761: 2754: 2714: 2703: 2648: 2639: 2631: 2625:Dann L (2002). 2623: 2616: 2565: 2561: 2508: 2504: 2457: 2453: 2422: 2418: 2373: 2369: 2346: 2342: 2337: 2320: 2289:nuclear encoded 2282: 2258:nuclear encoded 2251: 2239:entire membrane 2199: 2191:chloroplastidan 2096:import protein 2065: 1984: 1980: 1923: 1922: 1916: 1915: 1898: 1849:phosphorylation 1783: 1716:phosphate group 1704:phosphorylation 1632: 1631: 1630: 1629: 1606: 1533: 1529: 1478:14-3-3 proteins 1474:14-3-3 proteins 1451:phosphate group 1435:phosphate group 1411: 1391:functional part 1288: 1287: 1286: 1285: 1283: 1264: 1235: 1221: 1152:, though a few 1138:protein routing 1115: 1109: 1093: 1028: 1023: 1021:DNA replication 996: 987: 981: 978: 971:needs expansion 940: 930: 916:gene expression 907: 835: 771: 765: 723: 658: 631:dinophyte algae 599: 555:Chloroplast DNA 552: 551: 550: 549: 532:Chloroplast DNA 529: 516: 510: 507: 503: 500: 496: 493: 489: 486: 482: 479: 475: 472: 468: 465: 461: 458: 454: 451: 447: 444: 440: 437: 433: 430: 426: 423: 419: 416: 412: 409: 405: 402: 398: 395: 391: 388: 384: 381: 377: 374: 370: 367: 363: 360: 356: 353: 349: 346: 342: 339: 335: 332: 328: 325: 321: 318: 314: 311: 307: 304: 300: 297: 293: 290: 286: 283: 279: 276: 272: 269: 265: 262: 258: 255: 251: 248: 244: 241: 237: 234: 230: 227: 223: 220: 216: 213: 209: 206: 202: 199: 195: 192: 188: 185: 181: 178: 174: 171: 167: 164: 160: 157: 153: 150: 146: 143: 139: 136: 132: 129: 125: 122: 118: 115: 111: 109: 106: 102: 99: 95: 92: 88: 85: 81: 78: 74: 71: 67: 64: 60: 57: 53: 50: 46: 43: 39: 33: 28: 23: 22: 15: 12: 11: 5: 5011: 5001: 5000: 4998:Photosynthesis 4995: 4990: 4985: 4980: 4964: 4963: 4956: 4936: 4868: 4839:(3): 1016–30. 4810: 4789:10.1038/nsb744 4767: 4748:(1–2): 64–79. 4715: 4659: 4630:(4): 1560–78. 4624:The Plant Cell 4610: 4575:The Plant Cell 4552: 4496: 4489: 4462: 4455: 4435: 4386: 4359:(3): 198–208. 4257: 4232: 4201: 4139: 4104: 4066: 4017: 3982: 3935: 3878: 3837: 3779: 3730: 3679: 3630: 3601:(1): 189–201. 3577: 3513: 3478:(6927): 72–6. 3462: 3406: 3347: 3296: 3275:(5): 283–290. 3255: 3214: 3170: 3121: 3099: 3070:(7): 1579–89. 3064:The Plant Cell 3047: 3033: 3023: 2998: 2952: 2888: 2853:The Plant Cell 2826: 2819: 2782: 2775: 2752: 2701: 2637: 2614: 2559: 2502: 2451: 2432:(1): 312–317. 2416: 2367: 2356:(3): 159–164. 2339: 2338: 2336: 2333: 2332: 2331: 2326: 2319: 2316: 2281: 2278: 2250: 2247: 2198: 2195: 2189:are unique to 2064: 2061: 2021:TOC translocon 1988:TIC translocon 1983: 1977: 1902: 1901: 1900: 1899: 1897: 1894: 1853:protein kinase 1782: 1779: 1708:protein kinase 1652:(GTP)-binding 1615: 1614: 1608: 1607: 1605: 1602: 1532: 1526: 1410: 1407: 1370:, and poor in 1281: 1274:carboxyl group 1262: 1244: 1243: 1237: 1236: 1220: 1217: 1108: 1105: 1092: 1089: 1027: 1024: 1022: 1019: 995: 992: 989: 988: 968: 966: 944:RNA polymerase 929: 926: 906: 903: 870:chromalveolate 839:nuclear genome 834: 831: 819:RNA polymerase 807:ribosomal RNAs 764: 761: 722: 719: 657: 654: 598: 595: 530: 517: 508: 501: 494: 487: 483:RNA polymerase 480: 473: 466: 459: 452: 445: 438: 431: 424: 417: 410: 403: 396: 389: 382: 375: 368: 361: 357:photosystem II 354: 347: 340: 333: 326: 319: 315:RNA polymerase 312: 308:photosystem II 305: 298: 291: 284: 277: 270: 263: 256: 252:photosystem II 249: 242: 235: 228: 224:photosystem II 221: 214: 207: 200: 193: 186: 179: 172: 165: 158: 151: 144: 137: 130: 123: 116: 107: 103:photosystem II 100: 93: 86: 82:photosystem II 79: 72: 65: 58: 51: 44: 37: 31: 30: 29: 26: 9: 6: 4: 3: 2: 5010: 4999: 4996: 4994: 4991: 4989: 4986: 4984: 4981: 4979: 4976: 4975: 4973: 4959: 4957:9783540214892 4953: 4949: 4948: 4940: 4932: 4928: 4924: 4920: 4916: 4912: 4908: 4904: 4900: 4896: 4889: 4887: 4885: 4883: 4881: 4879: 4877: 4875: 4873: 4864: 4860: 4855: 4850: 4846: 4842: 4838: 4834: 4830: 4823: 4821: 4819: 4817: 4815: 4806: 4802: 4798: 4794: 4790: 4786: 4783:(2): 95–100. 4782: 4778: 4771: 4763: 4759: 4755: 4751: 4747: 4743: 4736: 4734: 4732: 4730: 4728: 4726: 4724: 4722: 4720: 4711: 4707: 4703: 4699: 4694: 4689: 4685: 4681: 4677: 4670: 4668: 4666: 4664: 4655: 4651: 4646: 4641: 4637: 4633: 4629: 4625: 4621: 4614: 4606: 4602: 4597: 4592: 4588: 4584: 4580: 4576: 4572: 4565: 4563: 4561: 4559: 4557: 4548: 4544: 4539: 4534: 4530: 4526: 4523:(2): 466–83. 4522: 4518: 4514: 4507: 4505: 4503: 4501: 4492: 4486: 4482: 4481: 4473: 4471: 4469: 4467: 4458: 4452: 4448: 4442: 4440: 4431: 4427: 4422: 4417: 4413: 4409: 4405: 4401: 4397: 4390: 4382: 4378: 4374: 4370: 4366: 4362: 4358: 4354: 4347: 4340: 4338: 4336: 4334: 4332: 4330: 4328: 4326: 4324: 4322: 4320: 4318: 4316: 4314: 4312: 4310: 4308: 4306: 4304: 4302: 4300: 4298: 4296: 4294: 4292: 4290: 4288: 4286: 4284: 4282: 4280: 4278: 4276: 4274: 4272: 4270: 4268: 4266: 4264: 4262: 4246: 4245:Biocyclopedia 4242: 4236: 4228: 4224: 4220: 4216: 4212: 4205: 4197: 4193: 4188: 4183: 4178: 4173: 4169: 4165: 4161: 4154: 4152: 4150: 4148: 4146: 4144: 4135: 4131: 4127: 4123: 4120:(3): 256–66. 4119: 4115: 4108: 4100: 4096: 4092: 4088: 4085:(1): 335–52. 4084: 4080: 4073: 4071: 4062: 4058: 4053: 4048: 4044: 4040: 4037:(4): 700–54. 4036: 4032: 4028: 4021: 4013: 4009: 4005: 4001: 3997: 3993: 3986: 3978: 3974: 3970: 3966: 3962: 3958: 3954: 3950: 3946: 3939: 3931: 3927: 3923: 3919: 3914: 3909: 3905: 3901: 3897: 3893: 3889: 3882: 3874: 3870: 3865: 3860: 3857:(1): 407–22. 3856: 3852: 3848: 3841: 3833: 3829: 3825: 3821: 3817: 3813: 3809: 3805: 3801: 3797: 3790: 3783: 3775: 3771: 3766: 3761: 3757: 3753: 3749: 3745: 3741: 3734: 3726: 3722: 3717: 3712: 3707: 3702: 3698: 3694: 3690: 3683: 3675: 3671: 3666: 3661: 3657: 3653: 3650:(4): e25845. 3649: 3645: 3641: 3634: 3626: 3622: 3617: 3612: 3608: 3604: 3600: 3596: 3592: 3590: 3581: 3573: 3569: 3564: 3559: 3554: 3549: 3545: 3541: 3537: 3533: 3529: 3522: 3520: 3518: 3509: 3505: 3501: 3497: 3493: 3489: 3485: 3481: 3477: 3473: 3466: 3458: 3454: 3450: 3446: 3442: 3438: 3434: 3430: 3427:(1): 91–120. 3426: 3422: 3415: 3413: 3411: 3402: 3398: 3393: 3388: 3383: 3378: 3374: 3370: 3366: 3362: 3358: 3351: 3343: 3339: 3334: 3329: 3324: 3319: 3315: 3311: 3307: 3300: 3292: 3288: 3283: 3278: 3274: 3270: 3266: 3259: 3251: 3247: 3242: 3237: 3233: 3229: 3225: 3218: 3210: 3206: 3202: 3198: 3194: 3190: 3186: 3182: 3174: 3166: 3162: 3157: 3152: 3148: 3144: 3140: 3136: 3132: 3125: 3109: 3103: 3095: 3091: 3086: 3081: 3077: 3073: 3069: 3065: 3061: 3054: 3052: 3043: 3037: 3030: 3026: 3024:9780120883912 3020: 3016: 3011: 3010: 3002: 2994: 2990: 2986: 2982: 2978: 2974: 2971:(2): 537–50. 2970: 2966: 2959: 2957: 2948: 2944: 2939: 2934: 2929: 2924: 2920: 2916: 2912: 2908: 2904: 2897: 2895: 2893: 2884: 2880: 2875: 2870: 2866: 2862: 2859:(7): 1661–6. 2858: 2854: 2850: 2843: 2841: 2839: 2837: 2835: 2833: 2831: 2822: 2816: 2812: 2811: 2803: 2801: 2799: 2797: 2795: 2793: 2791: 2789: 2787: 2778: 2772: 2768: 2767: 2759: 2757: 2748: 2744: 2740: 2736: 2732: 2728: 2725:(3): 275–88. 2724: 2720: 2712: 2710: 2708: 2706: 2697: 2693: 2688: 2683: 2678: 2673: 2669: 2665: 2661: 2657: 2653: 2646: 2644: 2642: 2630: 2629: 2621: 2619: 2610: 2606: 2602: 2598: 2594: 2590: 2586: 2582: 2578: 2574: 2570: 2563: 2555: 2551: 2546: 2541: 2537: 2533: 2529: 2525: 2521: 2517: 2513: 2506: 2498: 2494: 2490: 2486: 2482: 2478: 2474: 2470: 2466: 2462: 2455: 2447: 2443: 2439: 2435: 2431: 2427: 2426:Exp. Cell Res 2420: 2412: 2408: 2403: 2398: 2394: 2390: 2387:(3): 383–91. 2386: 2382: 2378: 2371: 2363: 2359: 2355: 2351: 2344: 2340: 2330: 2327: 2325: 2322: 2321: 2315: 2313: 2309: 2305: 2300: 2298: 2294: 2291:protein. The 2290: 2286: 2277: 2275: 2271: 2267: 2263: 2259: 2255: 2246: 2244: 2240: 2236: 2232: 2229:. Tic214 and 2228: 2227: 2223: 2219: 2215: 2211: 2208:. It is 1786 2207: 2203: 2194: 2192: 2188: 2184: 2180: 2176: 2175:cyanobacteria 2173:relatives in 2172: 2168: 2164: 2160: 2155: 2153: 2152: 2147: 2143: 2139: 2135: 2134: 2129: 2128: 2123: 2122: 2120: 2117:itochondrial 2116: 2112: 2107: 2105: 2099: 2095: 2094:mitochondrial 2091: 2088: 2084: 2080: 2077: 2076:transmembrane 2073: 2069: 2060: 2058: 2054: 2050: 2046: 2043:and contains 2042: 2038: 2034: 2030: 2026: 2022: 2017: 2015: 2011: 2007: 2006: 2002: 1998: 1994: 1989: 1976: 1974: 1970: 1966: 1962: 1958: 1955:has multiple 1954: 1953: 1948: 1945: 1943: 1939: 1935: 1931: 1930:transmembrane 1927: 1920: 1913: 1909: 1905: 1893: 1891: 1890: 1885: 1881: 1877: 1873: 1868: 1866: 1861: 1859: 1854: 1850: 1846: 1841: 1839: 1835: 1831: 1827: 1823: 1819: 1815: 1811: 1807: 1803: 1799: 1795: 1791: 1787: 1778: 1776: 1772: 1768: 1764: 1760: 1756: 1754: 1750: 1745: 1741: 1737: 1733: 1729: 1728: 1723: 1721: 1717: 1713: 1709: 1705: 1700: 1698: 1694: 1690: 1686: 1682: 1678: 1674: 1670: 1666: 1662: 1657: 1655: 1651: 1647: 1644: 1640: 1636: 1627: 1623: 1619: 1616:Toc34 from a 1612: 1601: 1599: 1595: 1590: 1588: 1584: 1580: 1576: 1572: 1568: 1564: 1560: 1556: 1554: 1550: 1546: 1544: 1538: 1525: 1523: 1519: 1515: 1510: 1507: 1505: 1501: 1497: 1494: 1492: 1488: 1484: 1479: 1475: 1470: 1468: 1464: 1460: 1456: 1452: 1448: 1444: 1440: 1436: 1432: 1431:phosphorylate 1428: 1424: 1420: 1416: 1406: 1404: 1400: 1396: 1392: 1387: 1385: 1381: 1380:glutamic acid 1377: 1376:aspartic acid 1373: 1369: 1365: 1361: 1357: 1353: 1349: 1345: 1340: 1338: 1334: 1330: 1326: 1325: 1320: 1316: 1312: 1308: 1304: 1300: 1296: 1292: 1279: 1275: 1271: 1267: 1260: 1256: 1252: 1248: 1241: 1234: 1230: 1226: 1216: 1214: 1210: 1206: 1205:cell membrane 1202: 1198: 1194: 1189: 1187: 1183: 1182:cell membrane 1179: 1175: 1174:topologically 1171: 1170:cell membrane 1167: 1164:(though many 1163: 1159: 1158:mitochondrion 1155: 1151: 1147: 1143: 1139: 1135: 1134:cell division 1131: 1126: 1124: 1120: 1114: 1104: 1102: 1098: 1088: 1079: 1075: 1073: 1069: 1065: 1061: 1057: 1052: 1050: 1045: 1041: 1032: 1018: 1016: 1011: 1009: 1003: 1000: 985: 976: 972: 969:This section 967: 964: 960: 959: 956: 954: 950: 945: 939: 935: 934:Transcription 925: 923: 922: 917: 913: 902: 900: 899: 893: 891: 887: 883: 880:) now have a 879: 875: 871: 867: 862: 860: 856: 855: 850: 846: 845: 840: 830: 828: 824: 820: 816: 812: 808: 802: 800: 799:gene transfer 796: 795:endosymbiotic 792: 788: 787:Synechocystis 784: 780: 776: 770: 760: 758: 754: 750: 747:In primitive 745: 743: 739: 735: 730: 728: 718: 715: 714: 709: 705: 701: 697: 693: 688: 686: 682: 678: 674: 673:ribosomal RNA 670: 665: 663: 653: 651: 646: 644: 640: 636: 632: 627: 625: 621: 617: 610: 609: 603: 594: 592: 588: 584: 583: 578: 577: 572: 568: 564: 560: 556: 547: 543: 539: 538: 533: 528: 527: 522: 521: 515: 513: 506: 499: 492: 485: 478: 471: 464: 457: 450: 443: 436: 429: 422: 415: 413:photosystem I 408: 401: 394: 392:ribosomal RNA 387: 380: 378:ribosomal RNA 373: 366: 359: 352: 345: 338: 331: 324: 317: 310: 303: 296: 294:photosystem I 289: 282: 275: 268: 261: 254: 247: 240: 238:photosystem I 233: 226: 219: 212: 205: 203:ribosomal RNA 198: 191: 189:ribosomal RNA 184: 177: 170: 163: 156: 149: 142: 135: 128: 121: 114: 105: 98: 91: 84: 77: 70: 63: 56: 49: 47:photosystem I 42: 19: 4978:Cell anatomy 4946: 4939: 4898: 4894: 4836: 4832: 4780: 4776: 4770: 4745: 4741: 4683: 4679: 4627: 4623: 4613: 4581:(1): 53–64. 4578: 4574: 4520: 4516: 4479: 4446: 4403: 4399: 4389: 4356: 4352: 4248:. Retrieved 4244: 4235: 4218: 4214: 4204: 4170:(237): 237. 4167: 4164:BMC Genomics 4163: 4117: 4113: 4107: 4082: 4078: 4034: 4030: 4020: 3995: 3991: 3985: 3952: 3948: 3938: 3895: 3891: 3881: 3854: 3850: 3840: 3799: 3795: 3782: 3747: 3743: 3733: 3696: 3693:BMC Genomics 3692: 3682: 3647: 3643: 3633: 3598: 3594: 3588: 3580: 3535: 3531: 3475: 3471: 3465: 3424: 3420: 3364: 3360: 3350: 3313: 3309: 3299: 3272: 3268: 3258: 3231: 3227: 3217: 3184: 3180: 3173: 3138: 3134: 3124: 3112:. Retrieved 3102: 3067: 3063: 3041: 3036: 3028: 3008: 3001: 2968: 2964: 2910: 2906: 2856: 2852: 2809: 2765: 2722: 2718: 2659: 2655: 2627: 2576: 2572: 2562: 2519: 2515: 2505: 2467:(1): 91–92. 2464: 2460: 2454: 2429: 2425: 2419: 2384: 2381:J. Cell Biol 2380: 2370: 2353: 2349: 2343: 2301: 2283: 2252: 2224: 2200: 2169:, Tic20 has 2156: 2150: 2148:of Tic20 in 2131: 2125: 2118: 2114: 2110: 2103: 2101: 2066: 2018: 2000: 1996: 1992: 1991: 1985: 1950: 1949: 1946: 1924: 1911: 1903: 1887: 1869: 1862: 1842: 1792:binding TOC 1784: 1770: 1766: 1761:to Toc34 in 1752: 1748: 1747: 1743: 1725: 1724: 1701: 1658: 1633: 1604:Toc34 and 33 1591: 1552: 1548: 1542: 1540: 1534: 1511: 1508: 1490: 1486: 1482: 1471: 1463:mitochondria 1412: 1388: 1356:hydroxylated 1341: 1333:presequences 1332: 1329:mitochondria 1322: 1315:carboxyl end 1314: 1306: 1291:Polypeptides 1289: 1277: 1265: 1197:mitochondria 1190: 1127: 1116: 1094: 1085: 1068:hypoxanthine 1053: 1037: 1012: 1008:PPR proteins 1004: 997: 982:January 2013 979: 975:adding to it 970: 941: 919: 908: 896: 894: 863: 852: 842: 836: 803: 790: 786: 782: 778: 772: 746: 731: 724: 711: 689: 666: 661: 659: 647: 628: 614: 606: 580: 574: 565:, contain a 558: 554: 553: 535: 531: 524: 518: 511: 329:atp synthase 259:atp synthase 4993:Plant genes 4983:Chromosomes 3310:Genome Biol 2913:(1): 41–5. 2306:are highly 2262:amino acids 2210:amino acids 2206:kilodaltons 2151:Arabidopsis 1973:AtToc75 III 1965:chromosomal 1838:hydrophilic 1836:end is the 1788:is another 1771:Arabidopsis 1751:rabidopsis 1746:stands for 1661:preproteins 1643:hydrophobic 1598:kilodaltons 1537:TOC complex 1518:TOC complex 1467:peroxisomes 1447:amino acids 1433:, or add a 1415:polypeptide 1299:amino acids 1272:, with its 1257:, with its 1251:amino acids 1247:polypeptide 1233:translation 1225:Polypeptide 1201:peroxisomes 1150:pseudogenes 1140:, and even 1130:exaptations 1060:amino group 1056:deamination 999:RNA editing 938:translation 898:Arabidopsis 886:green algal 791:Arabidopsis 753:green algae 700:rhodophyceæ 696:glaucophyta 616:Chloroplast 245:cytochromes 217:cytochromes 4972:Categories 4250:24 October 3699:(1): 204. 3589:Pilostyles 3316:(1): 134. 3114:August 18, 2335:References 2302:Tic56 and 2293:preprotein 2287:is also a 2218:N-terminal 2171:homologous 2106:ranslocase 2090:amino acid 2005:translocon 1942:nanometers 1834:C-terminal 1830:homologous 1818:kilodalton 1806:N-terminal 1734:proteins, 1732:homologous 1683:(GDP) and 1646:C-terminal 1594:nanometers 1395:C-terminus 1354:, rich in 1311:C-terminus 1309:, and the 1303:N-terminus 1270:C-terminus 1255:N-terminus 1249:with four 1223:See also: 1123:translated 1111:See also: 932:See also: 854:Pilostyles 767:See also: 706:and a few 669:base pairs 643:coding DNA 639:base pairs 620:base pairs 40:cytochrome 2601:1476-4687 2536:0261-4189 2308:conserved 2087:bacterial 2079:α-helices 2019:Like the 1804:. At the 1714:to add a 1618:pea plant 1545:ranslocon 1443:threonine 1364:threonine 1337:ribosomes 1307:amino end 1213:organelle 953:ribosomes 949:promoters 827:thylakoid 775:sequenced 749:red algae 738:red algae 727:nucleoids 721:Nucleoids 708:red algae 692:conserved 591:sequenced 168:small RNA 110:ribosomal 4923:23372012 4863:20457805 4805:21855733 4797:11753431 4762:11750663 4710:26014578 4654:22517318 4605:10634907 4547:16384899 4430:20124341 4381:32453554 4373:14991000 4196:19457260 4134:20206308 4099:24274753 3977:17395793 3930:17830745 3922:17174910 3873:20702568 3832:11408339 3824:19556510 3774:30165616 3725:25879186 3674:24195014 3625:26660355 3572:12218172 3500:12594458 3457:16536768 3449:23839301 3342:27339192 3291:10574454 3165:16453699 3094:12119376 2993:11571695 2947:16592612 2883:15235123 2747:30501148 2739:21636401 2554:16453699 2489:13905812 2446:14467684 2411:14492436 2318:See also 2127:in vitro 2072:integral 1957:isoforms 1934:β-barrel 1908:β-sheets 1904:β-barrel 1775:analogue 1765:(called 1730:has two 1563:subunits 1524:energy. 1459:specific 1419:ribosome 1382:. In an 1295:proteins 1203:, and a 1119:proteins 1072:cytosine 1064:mutation 868:in many 742:nucleoid 734:histones 713:Porphyra 675:and two 635:plasmids 112:proteins 4931:5062593 4903:Bibcode 4895:Science 4854:2899928 4702:8626459 4645:3398564 4538:1361317 4421:2817223 4221:: 1–9. 4187:2695485 4061:7854253 4012:9242608 3992:Science 3957:Bibcode 3949:Science 3900:Bibcode 3804:Bibcode 3796:Science 3765:6166771 3716:4487195 3665:3812789 3616:4758247 3540:Bibcode 3508:4319507 3480:Bibcode 3429:Bibcode 3401:8041699 3369:Bibcode 3333:4918201 3269:DNA Res 3250:9435137 3209:4311952 3189:Bibcode 3156:1167080 2985:6288261 2915:Bibcode 2696:8041699 2664:Bibcode 2609:4311952 2581:Bibcode 2545:1167080 2497:4265095 2469:Bibcode 2402:2106071 2297:daltons 2266:daltons 2216:on its 2157:Unlike 2142:Tic20 I 2121:embrane 2108:on the 2057:Tic20 I 2041:daltons 2025:complex 1865:cytosol 1814:cleaved 1802:domains 1796:, like 1794:subunit 1755:haliana 1740:AtToc34 1736:AtToc33 1677:cytosol 1665:cytosol 1663:in the 1561:. Five 1547:on the 1504:cytosol 1500:folding 1423:cytosol 1421:in the 1368:proline 1348:domains 1321:genes, 1319:nuclear 1229:protein 1193:already 1178:cytosol 1082:change. 874:diatoms 849:reduced 823:Rubisco 681:reduced 624:daltons 571:nucleus 563:plastid 546:introns 61:rubisco 4954:  4929:  4921:  4861:  4851:  4803:  4795:  4760:  4708:  4700:  4652:  4642:  4603:  4596:140214 4593:  4545:  4535:  4487:  4453:  4428:  4418:  4379:  4371:  4194:  4184:  4132:  4097:  4059:  4052:372988 4049:  4010:  3975:  3928:  3920:  3871:  3830:  3822:  3772:  3762:  3723:  3713:  3672:  3662:  3623:  3613:  3570:  3563:129430 3560:  3506:  3498:  3472:Nature 3455:  3447:  3399:  3389:  3340:  3330:  3289:  3248:  3207:  3181:Nature 3163:  3153:  3135:EMBO J 3092:  3085:150708 3082:  3021:  2991:  2983:  2945:  2938:382872 2935:  2881:  2874:514151 2871:  2817:  2773:  2745:  2737:  2694:  2684:  2607:  2599:  2573:Nature 2552:  2542:  2534:  2495:  2487:  2461:Nature 2444:  2409:  2399:  2304:Tic100 2254:Tic100 2249:Tic100 2235:dalton 2202:Tic214 2197:Tic214 2185:, and 2183:Tic100 2179:Tic214 2163:Tic100 2159:Tic214 2083:dalton 2070:is an 2055:, and 2049:Tic100 2045:Tic214 2035:, and 2029:Tic110 1882:, and 1880:Toc120 1876:Toc132 1872:Toc159 1786:Toc159 1781:Toc159 1697:Toc159 1693:domain 1654:domain 1637:is an 1575:Toc159 1514:Toc159 1493:rotein 1455:enzyme 1453:. 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Index

CpDNA
cytochrome
photosystem I
acetyl-CoA carboxylase
rubisco
tRNAs
tRNA
photosystem II
tRNAs
tRNAs
photosystem II
ribosomal
proteins

tRNA
tRNA
nadh dehydrogenase
ribosomal proteins
tRNA
replication origin regions
tRNA
small RNA
ribosomal protein
replication origin regions
ribosomal RNA
tRNAs
ribosomal RNA
tRNA
cytochromes
photosystem II
ribosomal proteins
photosystem I

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