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Calvin cycle

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837: 1135:). Each G3P molecule is composed of 3 carbons. For the Calvin cycle to continue, RuBP (ribulose 1,5-bisphosphate) must be regenerated. So, 5 out of 6 carbons from the 2 G3P molecules are used for this purpose. Therefore, there is only 1 net carbon produced to play with for each turn. To create 1 surplus G3P requires 3 carbons, and therefore 3 turns of the Calvin cycle. To make one glucose molecule (which can be created from 2 G3P molecules) would require 6 turns of the Calvin cycle. Surplus G3P can also be used to form other carbohydrates such as starch, sucrose, and cellulose, depending on what the plant needs. 686: 23: 1196:, helps this carbamylation process by removing one proton from the lysine and making the binding of the carbon dioxide molecule possible. Even then the RuBisCo enzyme is not yet functional, as it needs a magnesium ion bound to the lysine to function. This magnesium ion is released from the thylakoid lumen when the inner pH drops due to the active pumping of protons from the electron flow. RuBisCo activase itself is activated by increased concentrations of ATP in the stroma caused by its 1035: 814: 678: 3320: 3344: 312: 174: 409: 846: 3356: 3332: 537:, separating the reactions. They are activated in the light (which is why the name "dark reaction" is misleading), and also by products of the light-dependent reaction. These regulatory functions prevent the Calvin cycle from being respired to carbon dioxide. Energy (in the form of ATP) would be wasted in carrying out these reactions when they have no 1073:. The rate of photorespiration is higher at high temperatures. Photorespiration turns RuBP into 3-PGA and 2-phosphoglycolate, a 2-carbon molecule that can be converted via glycolate and glyoxalate to glycine. Via the glycine cleavage system and tetrahydrofolate, two glycines are converted into serine plus 1167:
The thioredoxin/ferredoxin system activates the enzymes glyceraldehyde-3-P dehydrogenase, glyceraldehyde-3-P phosphatase, fructose-1,6-bisphosphatase, sedoheptulose-1,7-bisphosphatase, and ribulose-5-phosphatase kinase, which are key points of the process. This happens when light is available, as the
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bond found in all these enzymes. This is a dynamic process as the same bond is formed again by other proteins that deactivate the enzymes. The implications of this process are that the enzymes remain mostly activated by day and are deactivated in the dark when there is no more reduced ferredoxin
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The regeneration stage of Calvin's cycle. Substances and their parts are outlined in colors: green - carbon accepting aldoses, pink - ketoses-donors of three-carbon groups, yellow - parts of ketoses remaining after donation of two-carbon keto-groups highlighted in orange. Enzymes are also
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evolved to circumvent photorespiration, but can occur only in certain plants native to very warm or tropical climates—corn, for example. Furthermore, RuBisCOs catalyzing the light-independent reactions of photosynthesis generally exhibit an improved specificity for
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Thus, of six G3P produced, five are used to make three RuBP (5C) molecules (totaling 15 carbons), with only one G3P available for subsequent conversion to hexose. This requires nine ATP molecules and six NADPH molecules per three
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that is captured by enediol in second step produces an unstable six-carbon compound called 2-carboxy 3-keto 1,5-biphosphoribotol (CKABP) (or 3-keto-2-carboxyarabinitol 1,5-bisphosphate) that immediately splits into 2 molecules of
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The immediate products of one turn of the Calvin cycle are 2 glyceraldehyde-3-phosphate (G3P) molecules, 3 ADP, and 2 NADP. (ADP and NADP are not really "products". They are regenerated and later used again in the
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complex of the thylakoid electron chain when electrons are circulating through it. Ferredoxin then binds to and reduces the thioredoxin protein, which activates the cycle enzymes by severing a
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The results suggested that the integrity of thylakoid membranes may be essential for the organisation of sequential enzymes of the Calvin cycle in vivo and facilitate their functioning.
786: 646:, where six-carbon sugars are oxidized in mitochondria. The carbohydrate products of the Calvin cycle are three-carbon sugar phosphate molecules, or "triose phosphates", namely, 518:. In the following biochemical equations, the chemical species (phosphates and carboxylic acids) exist in equilibria among their various ionized states as governed by the 486:
Photosynthesis occurs in two stages in a cell. In the first stage, light-dependent reactions capture the energy of light and use it to make the energy-storage molecule
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Though it is also called the "dark reaction", the Calvin cycle does not actually occur in the dark or during night time. This is because the process requires
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The next stage in the Calvin cycle is to regenerate RuBP. Five G3P molecules produce three RuBP molecules, using up three molecules of ATP. Since each CO
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These reactions do not occur in the dark or at night. There is a light-dependent regulation of the cycle enzymes, as the third step requires NADPH.
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Schulz, L; Guo, Z; Zarzycki, J; Steinchen, W; Schuller, JM; Heimerl, T; Prinz, S; Mueller-Cajar, O; Erb, TJ; Hochberg, GKA (14 October 2022).
73:. The Calvin cycle is present in all photosynthetic eukaryotes and also many photosynthetic bacteria. In plants, these reactions occur in the 1916: 1620:
Konwarh, Rocktotpal; Abda, Ebrahim M.; Haregu, Simatsidk; Singh, Anand Pratap (2022-01-01), Khan, Raju; Murali, S.; Gogoi, Satyabrat (eds.),
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reserves to provide energy for the plant. The Calvin cycle thus happens when light is available independent of the kind of photosynthesis (
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molecules produce six G3P molecules, of which five are used to regenerate RuBP, leaving a net gain of one G3P molecule per three CO
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electron transport chain, as the energy required to reduce the carbon dioxide is provided by NADPH produced during the
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The enzymes in the Calvin cycle are functionally equivalent to most enzymes used in other metabolic pathways such as
1909: 1891:, Fifth Edition by Jeremy M. Berg, John L. Tymoczko and Lubert Stryer. Published by W. H. Freeman and Company (2002). 1402: 1226: 395: 265: 355: 225: 203: 3387: 612:(six-carbon) sugars are not products of the Calvin cycle. Although many texts list a product of photosynthesis as 3360: 290:, also known as C2 cycle, is also coupled to the Calvin cycle, as it results from an alternative reaction of the 1295:"Involvement of thylakoid membranes in supramolecular organisation of Calvin cycle enzymes inAnacystis nidulans" 3382: 3239: 2401: 942: 931: 333: 199: 362: 232: 2682: 2436: 329: 195: 113:
to a sugar. There are three phases to the light-independent reactions, collectively called the Calvin cycle:
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are transferred to one of the G3P, giving another Xu5P. This leaves one G3P as the product of fixation of 3
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Ruuska, Sari A.; Andrews, T. John; Badger, Murray R.; Price, G. Dean; von Caemmerer, Susanne (2000-02-01).
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that can be used by the organism (and by animals that feed on it). This set of reactions is also called
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Farazdaghi H (2009). "Modeling the Kinetics of Activation and Reaction of Rubisco from Gas Exchange".
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and perform further chemical processes on them. The Calvin cycle uses the chemical energy of ATP and
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Besse, I; Buchanan, B (1997). "Thioredoxin-linked animal and plant processes: the new generation".
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catalyses the phosphorylation of 3-PGA by ATP (which was produced in the light-dependent stage).
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and leads to a non-functional state if left uncarbamylated. A specific activase enzyme, called
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The enzyme RuBisCo has its own, more complex activation process. It requires that a specific
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Campbell, and Reece Biology: 8th Edition, page 198. Benjamin Cummings, December 7, 2007.
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There are two regulation systems at work when the cycle must be turned on or off: the
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molecules. The equation of the overall Calvin cycle is shown diagrammatically below.
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for the plant to use. These substrates are used in a series of reduction-oxidation (
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Bassham JA (2003). "Mapping the carbon reduction cycle: a personal retrospective".
1703: 1687: 1592: 1584: 1520: 1472: 1423: 1351: 1306: 1265: 1257: 1070: 743:(also written as 3-phosphoglyceric acid, PGA, 3PGA, or 3-PGA), a 3-carbon compound. 503: 287: 1501:"Evolution of increased complexity and specificity at the dawn of form I Rubiscos" 941:(one of only three enzymes of the Calvin cycle that are unique to plants) cleaves 3206: 3162: 3157: 3051: 3027: 2861: 2824: 2677: 2667: 2550: 2321: 2286: 1867: 1855: 1427: 1197: 526: 413: 1293:
Sainis, Jayashree Krishna; Dani, Diksha Narhar; Dey, Gautam Kumar (2003-01-01).
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enzyme activation, active in the Calvin cycle, which involves its own activase.
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The overall equation of the Calvin cycle (black circles represent carbon atoms)
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in their vacuoles every night and release it by day to make this process work.
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of 1,3BPGA by NADPH (which is another product of the light-dependent stage).
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molecules (as would be expected from the number of carbon atoms involved).
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The Biochemistry of the Calvin Cycle at Rensselaer Polytechnic Institute
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are the products. (However, note that two 3-PGAs are produced for every
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activation system, which activates some of the cycle enzymes; and the
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generates two more G3P. The ketose S7P has two carbons removed by
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The regeneration stage can be broken down into a series of steps.
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molecule is incorporated into one of two three-carbon molecules (
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Lorimer, G.H.; Andrews, T.J.; Pierce, J.; Schloss, J.V. (1986).
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The reactions of the Calvin cycle are closely coupled to the
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power of NADPH from the light dependent reactions to produce
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Campbell, Neil A.; Brad Williamson; Robin J. Heyden (2006).
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highlighted: aldolases in purple and transketolases in red.
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The sum of reactions in the Calvin cycle is the following:
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Rubisco Activase, from the Plant Physiology Online website
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E4P and a DHAP (formed from one of the G3P from the second
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Calvin cycle step 1 (black circles represent carbon atoms)
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that enters the cycle, so this step utilizes two ATP per
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Thioredoxins, from the Plant Physiology Online website
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Twenty-First Century Books. p. 21. 21: 2423: 2383:Timeline of biology and organic chemistry 1707: 1596: 1269: 490:and the moderate-energy hydrogen carrier 396:Learn how and when to remove this message 266:Learn how and when to remove this message 1555:Biology: Exploring the Diversity of Life 1444: 1033: 972:(R5P), and the two carbons remaining on 860:converts one of the G3P reversibly into 844: 835: 812: 787:glyceraldehyde 3-phosphate dehydrogenase 684: 676: 407: 27:The internal structure of a chloroplast 1924: 1241: 3375: 1791:Portis, Archie; Parry, Martin (2007). 1337:"The path of carbon in photosynthesis" 1335:Bassham J, Benson A, Calvin M (1950). 908:are added to a G3P, giving the ketose 817:Regeneration stage of the Calvin cycle 457:. The cycle was discovered in 1950 by 132:. In the dark, plants instead release 128:, which is short-lived and comes from 2397: 2378:Nobel Prize in Physiology or Medicine 1898: 1168:ferredoxin protein is reduced in the 1014:. This requires the input of one ATP. 164:Coupling to other metabolic pathways 85:. These reactions take the products ( 3331: 1628:, Academic Press, pp. 155–173, 334:adding citations to reliable sources 305: 200:adding citations to reliable sources 167: 69:and hydrogen-carrier compounds into 3355: 1626:Carbon Dots in Agricultural Systems 689:Calvin cycle steps 2 and 3 combined 13: 1842: 471:University of California, Berkeley 154:crassulacean acid metabolism (CAM) 14: 3404: 1873: 864:(DHAP), also a 3-carbon molecule. 754:(glycerate-1,3-bisphosphate) and 442:reactions that take place in the 425:Calvin–Benson–Bassham (CBB) cycle 412:Overview of the Calvin cycle and 3354: 3342: 3330: 3319: 3318: 1097:, this process leads to loss of 999:. Xu5P is converted into RuP by 939:Sedoheptulose-1,7-bisphosphatase 310: 172: 1667: 1647: 1613: 1560: 1547: 1045:also reacts competitively with 896:F6P has two carbons removed by 698:catalyses the carboxylation of 321:needs additional citations for 301: 77:, the fluid-filled region of a 51:photosynthetic carbon reduction 1773:"Photosynthetic Dark Reaction" 1577:Journal of Experimental Botany 1492: 1483: 1411: 1384: 1328: 1286: 1235: 1210: 943:sedoheptulose-1,7-bisphosphate 932:sedoheptulose-1,7-bisphosphate 874:convert a G3P and a DHAP into 1: 2683:Interface and colloid science 2437:Glossary of chemical formulae 1203: 930:fixation) are converted into 1609:– via Oxford Academic. 1457:Phil. Trans. R. Soc. Lond. B 1428:10.1007/978-1-4020-9237-4_12 7: 2960:Bioorganometallic chemistry 2447:List of inorganic compounds 1299:Journal of Plant Physiology 1217:Silverstein, Alvin (2008). 1125: 872:fructose-1,6-bisphosphatase 117:, reduction reactions, and 39:light-independent reactions 10: 3409: 2886:Dynamic covalent chemistry 2857:Enantioselective synthesis 2837:Physical organic chemistry 2790:Organolanthanide chemistry 1142: 1139:Light-dependent regulation 904:(E4P). The two carbons on 862:dihydroxyacetone phosphate 858:Triose phosphate isomerase 795:Glyceraldehyde 3-phosphate 664:glyceraldehyde 3-phosphate 648:glyceraldehyde-3-phosphate 583:glyceraldehyde-3-phosphate 3314: 3217: 2978: 2894: 2815: 2765: 2641: 2584: 2475:Electroanalytical methods 2460: 2432: 2365: 1932: 1812:10.1007/s11120-007-9225-6 1242:Cushman, John C. (2001). 1145:Light-dependent reactions 1133:light-dependent reactions 947:sedoheptulose-7-phosphate 934:(7C) by aldolase enzyme. 881:Then fixation of another 700:ribulose-1,5-bisphosphate 531:pentose phosphate pathway 284:light dependent reactions 130:light-dependent reactions 119:ribulose 1,5-bisphosphate 95:light-dependent reactions 3230:Nobel Prize in Chemistry 3146:Supramolecular chemistry 2785:Organometallic chemistry 1771:Diwan, Joyce J. (2005). 1420:Photosynthesis in silico 1001:phosphopentose epimerase 997:phosphopentose isomerase 653: 3388:Carbohydrate metabolism 3168:Combinatorial chemistry 3079:Food physical chemistry 3042:Environmental chemistry 2926:Bioorthogonal chemistry 2852:Retrosynthetic analysis 2673:Chemical thermodynamics 2656:Spectroelectrochemistry 2599:Computational chemistry 1800:Photosynthesis Research 1748:10.1023/A:1024929725022 1525:10.1126/science.abq1416 1394:Biology: Exploring Life 1311:10.1078/0176-1617-00849 893:generates two more G3P. 752:1,3-Bisphosphoglycerate 748:phosphoglycerate kinase 514:of the cycle is called 3240:of element discoveries 3086:Agricultural chemistry 3074:Carbohydrate chemistry 2965:Bioinorganic chemistry 2830:Alkane stereochemistry 2775:Coordination chemistry 2604:Mathematical chemistry 2470:Instrumental chemistry 1992:Biological engineering 1477:10.1098/rstb.1986.0046 1039: 991:R5P is converted into 850: 842: 818: 690: 682: 416: 3383:Biochemical reactions 3235:Timeline of chemistry 3132:Post-mortem chemistry 3117:Clandestine chemistry 3047:Atmospheric chemistry 2970:Biophysical chemistry 2802:Solid-state chemistry 2752:Equilibrium chemistry 2661:Photoelectrochemistry 2077:Developmental biology 2052:Computational biology 2032:Cellular microbiology 1657:Bot. Bull. Acad. Sin. 1553:Russell, Wolfe et al. 1037: 902:erythrose-4-phosphate 848: 839: 816: 688: 680: 670:and two molecules of 411: 121:(RuBP) regeneration. 3225:History of chemistry 3180:Chemical engineering 2955:Bioorganic chemistry 2705:Structural chemistry 2442:List of biomolecules 2297:Reproductive biology 2182:Mathematical biology 2107:Evolutionary biology 2057:Conservation biology 1692:10.1104/pp.122.2.491 993:ribulose-5-phosphate 952:Fixation of a third 910:xylulose-5-phosphate 876:fructose 6-phosphate 330:improve this article 196:improve this section 156:); CAM plants store 3248:The central science 3202:Ceramic engineering 3127:Forensic toxicology 3100:Chemistry education 2998:Radiation chemistry 2980:Interdisciplinarity 2933:Medicinal chemistry 2871:Fullerene chemistry 2747:Microwave chemistry 2616:Molecular mechanics 2611:Molecular modelling 2082:Ecological genetics 1926:Branches of biology 1517:2022Sci...378..155S 1469:1986RSPTB.313..397L 1008:phosphoribulokinase 644:aerobic respiration 83:thylakoid membranes 43:bio synthetic phase 3291:Chemical substance 3153:Chemical synthesis 3122:Forensic chemistry 3003:Actinide chemistry 2945:Clinical chemistry 2626:Molecular geometry 2621:Molecular dynamics 2576:Elemental analysis 2529:Separation process 2373:History of biology 2307:Structural biology 2292:Relational biology 2117:Generative biology 2112:Freshwater biology 1866:2008-06-16 at the 1854:2009-02-18 at the 1589:10.1093/jxb/erz078 1111:C4 carbon fixation 1040: 970:ribose-5-phosphate 851: 843: 819: 741:3-phosphoglycerate 691: 683: 417: 296:glyceraldehyde-3-P 150:C4 carbon fixation 146:C3 carbon fixation 3370: 3369: 3306:Quantum mechanics 3271:Chemical compound 3254:Chemical reaction 3192:Materials science 3110:General chemistry 3105:Amateur chemistry 3033:Photogeochemistry 3018:Stellar chemistry 2988:Nuclear chemistry 2909:Molecular biology 2876:Polymer chemistry 2847:Organic synthesis 2842:Organic reactions 2807:Ceramic chemistry 2797:Cluster chemistry 2727:Chemical kinetics 2715:Molecular physics 2594:Quantum chemistry 2507:Mass spectrometry 2391: 2390: 2312:Synthetic biology 2192:Molecular biology 2047:Cognitive biology 1635:978-0-323-90260-1 1511:(6616): 155–160. 1463:(1162): 397–407. 1437:978-1-4020-9236-7 1262:10.1104/pp.010818 504:organic compounds 406: 405: 398: 380: 286:. The process of 276: 275: 268: 250: 3400: 3358: 3357: 3346: 3334: 3333: 3322: 3321: 3266:Chemical element 2921:Chemical biology 2780:Magnetochemistry 2757:Mechanochemistry 2710:Chemical physics 2651:Electrochemistry 2556:Characterization 2418: 2411: 2404: 2395: 2394: 2037:Chemical biology 1919: 1912: 1905: 1896: 1895: 1838: 1836: 1830:. Archived from 1797: 1787: 1785: 1784: 1767: 1722: 1721: 1711: 1680:Plant Physiology 1671: 1665: 1664: 1651: 1645: 1644: 1643: 1642: 1617: 1611: 1610: 1600: 1571:(1 March 2019). 1567:Schreier, Tina; 1564: 1558: 1551: 1545: 1544: 1496: 1490: 1487: 1481: 1480: 1448: 1442: 1441: 1415: 1409: 1408: 1388: 1382: 1381: 1379: 1378: 1372: 1366:. Archived from 1341: 1332: 1326: 1325: 1290: 1284: 1283: 1273: 1256:(4): 1439–1448. 1249:Plant Physiology 1239: 1233: 1232: 1214: 1194:RuBisCo activase 1108: 1107: 1106: 1096: 1095: 1094: 1084: 1083: 1082: 1071:photorespiration 1068: 1067: 1066: 1056: 1055: 1054: 1030: 1029: 1028: 987: 986: 985: 963: 962: 961: 929: 928: 927: 892: 891: 890: 808: 807: 806: 781: 780: 779: 769: 768: 767: 737: 736: 735: 725: 724: 723: 713: 712: 711: 641: 640: 639: 631: 630: 622: 621: 580: 578: 577: 559: 558: 557: 539:net productivity 401: 394: 390: 387: 381: 379: 338: 314: 306: 288:photorespiration 271: 264: 260: 257: 251: 249: 208: 176: 168: 25: 3408: 3407: 3403: 3402: 3401: 3399: 3398: 3397: 3373: 3372: 3371: 3366: 3310: 3213: 3207:Polymer science 3163:Click chemistry 3158:Green chemistry 3052:Ocean chemistry 3028:Biogeochemistry 2974: 2890: 2862:Total synthesis 2825:Stereochemistry 2811: 2761: 2678:Surface science 2668:Thermochemistry 2637: 2580: 2551:Crystallography 2456: 2428: 2422: 2392: 2387: 2361: 2322:Systems biology 2287:Quantum biology 1928: 1923: 1876: 1868:Wayback Machine 1856:Wayback Machine 1845: 1843:Further reading 1834: 1795: 1782: 1780: 1736:Photosynth. Res 1726: 1725: 1672: 1668: 1652: 1648: 1640: 1638: 1636: 1618: 1614: 1569:Hibberd, Julian 1565: 1561: 1552: 1548: 1497: 1493: 1488: 1484: 1449: 1445: 1438: 1416: 1412: 1405: 1389: 1385: 1376: 1374: 1370: 1339: 1333: 1329: 1291: 1287: 1240: 1236: 1229: 1215: 1211: 1206: 1198:phosphorylation 1147: 1141: 1128: 1121: 1117: 1105: 1102: 1101: 1100: 1098: 1093: 1090: 1089: 1088: 1086: 1081: 1078: 1077: 1076: 1074: 1065: 1062: 1061: 1060: 1058: 1053: 1050: 1049: 1048: 1046: 1027: 1024: 1023: 1022: 1020: 995:(Ru5P, RuP) by 984: 981: 980: 979: 977: 960: 957: 956: 955: 953: 935: 926: 923: 922: 921: 919: 889: 886: 885: 884: 882: 832: 828: 824: 805: 802: 801: 800: 798: 778: 775: 774: 773: 771: 766: 763: 762: 761: 759: 734: 731: 730: 729: 727: 722: 719: 718: 717: 715: 710: 707: 706: 705: 703: 661: 656: 638: 635: 634: 633: 629: 626: 625: 624: 620: 617: 616: 615: 613: 600: 596: 576: 573: 572: 571: 569: 556: 553: 552: 551: 549: 527:gluconeogenesis 508:carbon fixation 435:is a series of 414:carbon fixation 402: 391: 385: 382: 339: 337: 327: 315: 304: 272: 261: 255: 252: 209: 207: 193: 177: 166: 112: 31: 30: 29: 28: 17: 12: 11: 5: 3406: 3396: 3395: 3393:Photosynthesis 3390: 3385: 3368: 3367: 3365: 3364: 3352: 3340: 3328: 3315: 3312: 3311: 3309: 3308: 3303: 3298: 3293: 3288: 3283: 3278: 3273: 3268: 3263: 3262: 3261: 3251: 3244: 3243: 3242: 3232: 3227: 3221: 3219: 3215: 3214: 3212: 3211: 3210: 3209: 3204: 3199: 3189: 3188: 3187: 3177: 3176: 3175: 3170: 3165: 3160: 3150: 3149: 3148: 3137: 3136: 3135: 3134: 3129: 3119: 3114: 3113: 3112: 3107: 3096: 3095: 3094: 3093: 3091:Soil chemistry 3083: 3082: 3081: 3076: 3069:Food chemistry 3066: 3064:Carbochemistry 3061: 3059:Clay chemistry 3056: 3055: 3054: 3049: 3038: 3037: 3036: 3035: 3030: 3020: 3014:Astrochemistry 3010:Cosmochemistry 3007: 3006: 3005: 3000: 2995: 2993:Radiochemistry 2984: 2982: 2976: 2975: 2973: 2972: 2967: 2962: 2957: 2952: 2950:Neurochemistry 2947: 2942: 2941: 2940: 2930: 2929: 2928: 2918: 2917: 2916: 2911: 2900: 2898: 2892: 2891: 2889: 2888: 2883: 2881:Petrochemistry 2878: 2873: 2868: 2859: 2854: 2849: 2844: 2839: 2834: 2833: 2832: 2821: 2819: 2813: 2812: 2810: 2809: 2804: 2799: 2794: 2793: 2792: 2782: 2777: 2771: 2769: 2763: 2762: 2760: 2759: 2754: 2749: 2744: 2742:Spin chemistry 2739: 2737:Photochemistry 2734: 2729: 2724: 2722:Femtochemistry 2719: 2718: 2717: 2707: 2702: 2697: 2692: 2691: 2690: 2680: 2675: 2670: 2665: 2664: 2663: 2658: 2647: 2645: 2639: 2638: 2636: 2635: 2634: 2633: 2623: 2618: 2613: 2608: 2607: 2606: 2596: 2590: 2588: 2582: 2581: 2579: 2578: 2573: 2568: 2563: 2558: 2553: 2548: 2547: 2546: 2541: 2534:Chromatography 2531: 2526: 2525: 2524: 2519: 2514: 2504: 2503: 2502: 2497: 2492: 2487: 2477: 2472: 2466: 2464: 2458: 2457: 2455: 2454: 2452:Periodic table 2449: 2444: 2439: 2433: 2430: 2429: 2421: 2420: 2413: 2406: 2398: 2389: 2388: 2386: 2385: 2380: 2375: 2369: 2367: 2363: 2362: 2360: 2359: 2354: 2349: 2344: 2339: 2334: 2329: 2324: 2319: 2314: 2309: 2304: 2299: 2294: 2289: 2284: 2279: 2274: 2269: 2264: 2259: 2254: 2249: 2244: 2239: 2234: 2229: 2224: 2219: 2214: 2209: 2204: 2199: 2194: 2189: 2184: 2179: 2177:Marine biology 2174: 2169: 2164: 2159: 2154: 2149: 2144: 2139: 2134: 2129: 2124: 2119: 2114: 2109: 2104: 2099: 2094: 2089: 2084: 2079: 2074: 2069: 2064: 2059: 2054: 2049: 2044: 2039: 2034: 2029: 2024: 2019: 2014: 2009: 2004: 1999: 1994: 1989: 1987:Bioinformatics 1984: 1979: 1974: 1969: 1964: 1959: 1954: 1949: 1944: 1939: 1933: 1930: 1929: 1922: 1921: 1914: 1907: 1899: 1893: 1892: 1882: 1875: 1874:External links 1872: 1871: 1870: 1858: 1844: 1841: 1840: 1839: 1837:on 2012-03-12. 1806:(1): 121–143. 1788: 1768: 1742:(1–3): 35–52. 1730: 1729: 1724: 1723: 1686:(2): 491–504. 1666: 1646: 1634: 1612: 1559: 1546: 1491: 1482: 1443: 1436: 1410: 1403: 1383: 1356:10.2172/910351 1327: 1285: 1234: 1227: 1219:Photosynthesis 1208: 1207: 1205: 1202: 1184:amino acid be 1143:Main article: 1140: 1137: 1127: 1124: 1119: 1115: 1103: 1091: 1079: 1063: 1051: 1025: 1016: 1015: 1004: 989: 982: 958: 950: 924: 918: 917: 916: 913: 894: 887: 879: 865: 830: 826: 822: 811: 810: 803: 789:catalyses the 783: 776: 764: 744: 732: 720: 708: 659: 655: 652: 636: 627: 618: 607: 606: 598: 597:  (P 594: 574: 554: 496:carbon dioxide 452:photosynthetic 404: 403: 345:"Calvin cycle" 318: 316: 309: 303: 300: 274: 273: 215:"Calvin cycle" 180: 178: 171: 165: 162: 110: 67:carbon dioxide 63:photosynthesis 47:dark reactions 26: 20: 19: 18: 15: 9: 6: 4: 3: 2: 3405: 3394: 3391: 3389: 3386: 3384: 3381: 3380: 3378: 3363: 3362: 3353: 3351: 3350: 3345: 3341: 3339: 3338: 3329: 3327: 3326: 3317: 3316: 3313: 3307: 3304: 3302: 3299: 3297: 3296:Chemical bond 3294: 3292: 3289: 3287: 3284: 3282: 3279: 3277: 3274: 3272: 3269: 3267: 3264: 3260: 3257: 3256: 3255: 3252: 3249: 3245: 3241: 3238: 3237: 3236: 3233: 3231: 3228: 3226: 3223: 3222: 3220: 3216: 3208: 3205: 3203: 3200: 3198: 3195: 3194: 3193: 3190: 3186: 3185:Stoichiometry 3183: 3182: 3181: 3178: 3174: 3171: 3169: 3166: 3164: 3161: 3159: 3156: 3155: 3154: 3151: 3147: 3144: 3143: 3142: 3141:Nanochemistry 3139: 3138: 3133: 3130: 3128: 3125: 3124: 3123: 3120: 3118: 3115: 3111: 3108: 3106: 3103: 3102: 3101: 3098: 3097: 3092: 3089: 3088: 3087: 3084: 3080: 3077: 3075: 3072: 3071: 3070: 3067: 3065: 3062: 3060: 3057: 3053: 3050: 3048: 3045: 3044: 3043: 3040: 3039: 3034: 3031: 3029: 3026: 3025: 3024: 3021: 3019: 3015: 3011: 3008: 3004: 3001: 2999: 2996: 2994: 2991: 2990: 2989: 2986: 2985: 2983: 2981: 2977: 2971: 2968: 2966: 2963: 2961: 2958: 2956: 2953: 2951: 2948: 2946: 2943: 2939: 2936: 2935: 2934: 2931: 2927: 2924: 2923: 2922: 2919: 2915: 2912: 2910: 2907: 2906: 2905: 2902: 2901: 2899: 2897: 2893: 2887: 2884: 2882: 2879: 2877: 2874: 2872: 2869: 2867: 2866:Semisynthesis 2863: 2860: 2858: 2855: 2853: 2850: 2848: 2845: 2843: 2840: 2838: 2835: 2831: 2828: 2827: 2826: 2823: 2822: 2820: 2818: 2814: 2808: 2805: 2803: 2800: 2798: 2795: 2791: 2788: 2787: 2786: 2783: 2781: 2778: 2776: 2773: 2772: 2770: 2768: 2764: 2758: 2755: 2753: 2750: 2748: 2745: 2743: 2740: 2738: 2735: 2733: 2730: 2728: 2725: 2723: 2720: 2716: 2713: 2712: 2711: 2708: 2706: 2703: 2701: 2700:Sonochemistry 2698: 2696: 2695:Cryochemistry 2693: 2689: 2688:Micromeritics 2686: 2685: 2684: 2681: 2679: 2676: 2674: 2671: 2669: 2666: 2662: 2659: 2657: 2654: 2653: 2652: 2649: 2648: 2646: 2644: 2640: 2632: 2629: 2628: 2627: 2624: 2622: 2619: 2617: 2614: 2612: 2609: 2605: 2602: 2601: 2600: 2597: 2595: 2592: 2591: 2589: 2587: 2583: 2577: 2574: 2572: 2569: 2567: 2566:Wet chemistry 2564: 2562: 2559: 2557: 2554: 2552: 2549: 2545: 2542: 2540: 2537: 2536: 2535: 2532: 2530: 2527: 2523: 2520: 2518: 2515: 2513: 2510: 2509: 2508: 2505: 2501: 2498: 2496: 2493: 2491: 2488: 2486: 2483: 2482: 2481: 2478: 2476: 2473: 2471: 2468: 2467: 2465: 2463: 2459: 2453: 2450: 2448: 2445: 2443: 2440: 2438: 2435: 2434: 2431: 2427: 2419: 2414: 2412: 2407: 2405: 2400: 2399: 2396: 2384: 2381: 2379: 2376: 2374: 2371: 2370: 2368: 2364: 2358: 2355: 2353: 2350: 2348: 2345: 2343: 2340: 2338: 2335: 2333: 2330: 2328: 2325: 2323: 2320: 2318: 2315: 2313: 2310: 2308: 2305: 2303: 2300: 2298: 2295: 2293: 2290: 2288: 2285: 2283: 2280: 2278: 2275: 2273: 2270: 2268: 2265: 2263: 2260: 2258: 2257:Phylogenetics 2255: 2253: 2250: 2248: 2245: 2243: 2240: 2238: 2235: 2233: 2230: 2228: 2225: 2223: 2220: 2218: 2215: 2213: 2210: 2208: 2205: 2203: 2200: 2198: 2195: 2193: 2190: 2188: 2185: 2183: 2180: 2178: 2175: 2173: 2170: 2168: 2165: 2163: 2160: 2158: 2155: 2153: 2152:Human biology 2150: 2148: 2145: 2143: 2140: 2138: 2135: 2133: 2130: 2128: 2125: 2123: 2120: 2118: 2115: 2113: 2110: 2108: 2105: 2103: 2100: 2098: 2095: 2093: 2090: 2088: 2085: 2083: 2080: 2078: 2075: 2073: 2070: 2068: 2065: 2063: 2060: 2058: 2055: 2053: 2050: 2048: 2045: 2043: 2042:Chronobiology 2040: 2038: 2035: 2033: 2030: 2028: 2025: 2023: 2020: 2018: 2017:Biotechnology 2015: 2013: 2012:Biostatistics 2010: 2008: 2005: 2003: 2000: 1998: 1995: 1993: 1990: 1988: 1985: 1983: 1980: 1978: 1975: 1973: 1970: 1968: 1965: 1963: 1960: 1958: 1955: 1953: 1950: 1948: 1945: 1943: 1940: 1938: 1935: 1934: 1931: 1927: 1920: 1915: 1913: 1908: 1906: 1901: 1900: 1897: 1890: 1886: 1883: 1881: 1878: 1877: 1869: 1865: 1862: 1859: 1857: 1853: 1850: 1847: 1846: 1833: 1829: 1825: 1821: 1817: 1813: 1809: 1805: 1801: 1794: 1789: 1779:on 2005-03-16 1778: 1774: 1769: 1765: 1761: 1757: 1753: 1749: 1745: 1741: 1737: 1732: 1731: 1728: 1727: 1719: 1715: 1710: 1705: 1701: 1697: 1693: 1689: 1685: 1681: 1677: 1670: 1662: 1659: 1658: 1650: 1637: 1631: 1627: 1623: 1616: 1608: 1604: 1599: 1594: 1590: 1586: 1582: 1578: 1574: 1570: 1563: 1556: 1550: 1542: 1538: 1534: 1530: 1526: 1522: 1518: 1514: 1510: 1506: 1502: 1495: 1486: 1478: 1474: 1470: 1466: 1462: 1458: 1454: 1447: 1439: 1433: 1429: 1425: 1421: 1414: 1406: 1404:0-13-250882-6 1400: 1396: 1395: 1387: 1373:on 2009-02-19 1369: 1365: 1361: 1357: 1353: 1349: 1345: 1338: 1331: 1324: 1320: 1316: 1312: 1308: 1304: 1300: 1296: 1289: 1281: 1277: 1272: 1267: 1263: 1259: 1255: 1251: 1250: 1245: 1238: 1230: 1228:9780822567981 1224: 1220: 1213: 1209: 1201: 1199: 1195: 1191: 1187: 1183: 1178: 1175: 1171: 1170:photosystem I 1165: 1163: 1159: 1155: 1150: 1146: 1136: 1134: 1123: 1118:relative to O 1112: 1072: 1044: 1036: 1032: 1013: 1009: 1005: 1002: 998: 994: 990: 975: 974:transketolase 971: 967: 966:transketolase 951: 948: 944: 940: 937: 936: 933: 914: 911: 907: 906:transketolase 903: 899: 898:transketolase 895: 880: 877: 873: 869: 866: 863: 859: 856: 855: 854: 847: 838: 834: 815: 796: 792: 788: 784: 757: 753: 749: 745: 742: 701: 697: 693: 692: 687: 679: 675: 673: 669: 665: 651: 649: 645: 611: 604: 592: 588: 584: 567: 563: 547: 546: 545: 542: 540: 536: 532: 528: 523: 521: 517: 513: 509: 505: 501: 497: 493: 489: 484: 482: 479: 476: 473:by using the 472: 468: 467:Andrew Benson 464: 463:James Bassham 460: 459:Melvin Calvin 456: 453: 449: 445: 441: 438: 434: 430: 426: 422: 415: 410: 400: 397: 389: 386:December 2023 378: 375: 371: 368: 364: 361: 357: 354: 350: 347: â€“  346: 342: 341:Find sources: 335: 331: 325: 324: 319:This section 317: 313: 308: 307: 299: 297: 293: 289: 285: 281: 270: 267: 259: 256:December 2023 248: 245: 241: 238: 234: 231: 227: 224: 220: 217: â€“  216: 212: 211:Find sources: 205: 201: 197: 191: 190: 186: 181:This section 179: 175: 170: 169: 161: 159: 155: 151: 147: 143: 139: 135: 131: 127: 122: 120: 116: 115:carboxylation 108: 104: 100: 96: 92: 88: 84: 80: 76: 72: 68: 64: 60: 56: 52: 48: 44: 40: 36: 24: 3359: 3347: 3335: 3323: 3173:Biosynthesis 3023:Geochemistry 2938:Pharmacology 2914:Cell biology 2904:Biochemistry 2732:Spectroscopy 2631:VSEPR theory 2480:Spectroscopy 2424:Branches of 2302:Sociobiology 2282:Protistology 2247:Photobiology 2242:Pharmacology 2232:Parasitology 2227:Paleontology 2207:Neuroscience 2187:Microbiology 2097:Epidemiology 2067:Cytogenetics 2027:Cell biology 2007:Biosemiotics 1997:Biomechanics 1977:Biogeography 1972:Biochemistry 1967:Bacteriology 1962:Astrobiology 1889:Biochemistry 1888: 1832:the original 1803: 1799: 1781:. Retrieved 1777:the original 1739: 1735: 1683: 1679: 1669: 1660: 1655: 1649: 1639:, retrieved 1625: 1615: 1580: 1576: 1562: 1554: 1549: 1508: 1504: 1494: 1485: 1460: 1456: 1446: 1419: 1413: 1393: 1386: 1375:. Retrieved 1368:the original 1350:(2): 781–7. 1347: 1343: 1330: 1322: 1305:(1): 23–32. 1302: 1298: 1288: 1253: 1247: 1237: 1218: 1212: 1186:carbamylated 1179: 1166: 1151: 1148: 1129: 1041: 1017: 1011: 852: 820: 657: 608: 601:= inorganic 543: 524: 507: 485: 432: 428: 424: 421:Calvin cycle 420: 418: 392: 383: 373: 366: 359: 352: 340: 328:Please help 323:verification 320: 302:Calvin cycle 277: 262: 253: 243: 236: 229: 222: 210: 194:Please help 182: 123: 81:outside the 58: 54: 50: 46: 42: 38: 35:Calvin cycle 34: 32: 3361:WikiProject 2586:Theoretical 2571:Calorimetry 2352:Xenobiology 2347:Virophysics 2317:Systematics 2272:Primatology 2217:Ornithology 2157:Ichthyology 2142:Herpetology 2137:Gerontology 2102:Epigenetics 2062:Cryobiology 1952:Agrostology 1942:Aerobiology 1937:Abiogenesis 1344:J Biol Chem 1177:available. 1154:thioredoxin 1057:instead of 785:The enzyme 746:The enzyme 694:The enzyme 585:(G3P) + 6 475:radioactive 448:chloroplast 437:biochemical 140:from their 79:chloroplast 3377:Categories 3197:Metallurgy 2896:Biological 2462:Analytical 2337:Toxicology 2332:Teratology 2277:Proteomics 2262:Physiology 2202:Neontology 2167:Lipidology 2162:Immunology 2132:Geobiology 2092:Embryology 2072:Dendrology 2002:Biophysics 1982:Biogeology 1783:2012-10-24 1641:2023-04-22 1377:2013-07-03 1204:References 1158:ferredoxin 510:. The key 356:newspapers 298:molecule. 226:newspapers 158:malic acid 3259:Catalysis 2767:Inorganic 2561:Titration 2426:chemistry 2252:Phycology 2237:Pathology 2222:Osteology 2212:Nutrition 2172:Mammalogy 2147:Histology 1700:1532-2548 1541:252897276 1319:0176-1617 1006:Finally, 968:, giving 900:, giving 791:reduction 603:phosphate 481:carbon-14 455:organisms 280:thylakoid 183:does not 136:into the 3325:Category 3281:Molecule 3218:See also 2643:Physical 2366:See also 2342:Virology 2327:Taxonomy 2267:Pomology 2197:Mycology 2127:Genomics 2122:Genetics 1947:Agronomy 1864:Archived 1852:Archived 1828:39767233 1820:17665149 1764:52854452 1756:16228564 1718:10677442 1607:30916343 1533:36227987 1364:14774424 1280:11743087 1126:Products 868:Aldolase 529:and the 433:C3 cycle 99:reducing 3337:Commons 3301:Alchemy 2817:Organic 2357:Zoology 2087:Ecology 1957:Anatomy 1663:: 1–11. 1598:6436154 1513:Bibcode 1505:Science 1465:Bibcode 1271:1540176 1174:cystine 1162:RuBisCo 1043:RuBisCO 912:(Xu5P). 782:fixed.) 696:RuBisCO 650:(G3P). 535:cytosol 516:RuBisCO 478:isotope 469:at the 370:scholar 292:RuBisCO 240:scholar 204:removed 189:sources 134:sucrose 71:glucose 3349:Portal 2495:UV-Vis 2022:Botany 1826:  1818:  1762:  1754:  1716:  1706:  1698:  1632:  1605:  1595:  1539:  1531:  1434:  1401:  1362:  1317:  1278:  1268:  1225:  1182:lysine 809:fixed. 610:Hexose 512:enzyme 465:, and 444:stroma 372:  365:  358:  351:  343:  242:  235:  228:  221:  213:  152:, and 142:starch 138:phloem 103:sugars 75:stroma 2522:MALDI 2490:Raman 1887:from 1835:(PDF) 1824:S2CID 1796:(PDF) 1760:S2CID 1709:58886 1537:S2CID 1371:(PDF) 1340:(PDF) 1012:cycle 945:into 672:NADPH 654:Steps 593:+ 8 P 562:NADPH 502:into 500:water 492:NADPH 440:redox 377:JSTOR 363:books 247:JSTOR 233:books 126:NADPH 107:redox 93:) of 91:NADPH 59:cycle 49:, or 3276:Atom 2544:HPLC 1816:PMID 1752:PMID 1714:PMID 1696:ISSN 1630:ISBN 1603:PMID 1529:PMID 1432:ISBN 1399:ISBN 1360:PMID 1315:ISSN 1276:PMID 1223:ISBN 1190:RuBP 870:and 589:+ 9 587:NADP 568:+ 5 564:+ 9 560:+ 6 498:and 419:The 349:news 219:news 187:any 185:cite 89:and 33:The 3286:Ion 2517:ICP 2500:NMR 1808:doi 1744:doi 1704:PMC 1688:doi 1684:122 1593:PMC 1585:doi 1521:doi 1509:378 1473:doi 1461:313 1424:doi 1352:doi 1348:185 1307:doi 1303:160 1266:PMC 1258:doi 1254:127 1069:in 756:ADP 714:or 668:ATP 591:ADP 566:ATP 488:ATP 450:in 446:of 431:or 332:by 198:by 87:ATP 61:of 55:PCR 3379:: 3016:/ 3012:/ 2864:/ 2539:GC 2512:EI 2485:IR 1822:. 1814:. 1804:94 1802:. 1798:. 1758:. 1750:. 1740:76 1738:. 1712:. 1702:. 1694:. 1682:. 1678:. 1661:38 1624:, 1601:. 1591:. 1581:70 1579:. 1575:. 1535:. 1527:. 1519:. 1507:. 1503:. 1471:. 1459:. 1455:. 1430:. 1358:. 1346:. 1342:. 1321:. 1313:. 1301:. 1297:. 1274:. 1264:. 1252:. 1246:. 1200:. 1114:CO 1109:. 1099:CO 1087:CO 1075:CO 1059:CO 1021:CO 978:CO 954:CO 920:CO 883:CO 799:CO 772:CO 760:CO 728:CO 704:CO 628:12 581:→ 550:CO 548:3 541:. 522:. 520:pH 483:. 461:, 427:, 423:, 148:, 57:) 45:, 41:, 37:, 3250:" 3246:" 2417:e 2410:t 2403:v 1918:e 1911:t 1904:v 1810:: 1786:. 1766:. 1746:: 1720:. 1690:: 1587:: 1543:. 1523:: 1515:: 1479:. 1475:: 1467:: 1440:. 1426:: 1407:. 1380:. 1354:: 1309:: 1282:. 1260:: 1231:. 1156:/ 1120:2 1116:2 1104:2 1092:2 1080:2 1064:2 1052:2 1047:O 1026:2 1003:. 983:2 959:2 925:2 888:2 831:2 827:2 823:2 804:2 777:2 765:2 733:2 721:2 716:O 709:2 660:2 637:6 632:O 623:H 619:6 614:C 605:) 599:i 595:i 579:O 575:2 570:H 555:2 399:) 393:( 388:) 384:( 374:· 367:· 360:· 353:· 326:. 269:) 263:( 258:) 254:( 244:· 237:· 230:· 223:· 206:. 192:. 111:2 53:(

Index

The internal structure of a chloroplast
photosynthesis
carbon dioxide
glucose
stroma
chloroplast
thylakoid membranes
ATP
NADPH
light-dependent reactions
reducing
sugars
redox
carboxylation
ribulose 1,5-bisphosphate
NADPH
light-dependent reactions
sucrose
phloem
starch
C3 carbon fixation
C4 carbon fixation
crassulacean acid metabolism (CAM)
malic acid

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