Die Kulturpflanze

, Volume 32, Issue 1, pp 35–65 | Cite as

Modellvorstellungen zur Kohlenstoff-Isotopendiskriminierung bei der Photosynthese von C3- und C4-Pflanzen

  • Martin Peisker
Originalarbeiten

Zusammenfassung

In der vorliegenden Arbeit werden Modelle für die Anreicherung des Kohlenstoffatoms12C gegenüber13C bei der Photosynthese von C3- und C4-Pflanzen beschrieben und im Zusammenhang mit Literaturdaten zum Einfluß innerer und äußerer Faktoren auf diesen Vorgang diskutiert. Bei den C3-Pflanzen wird die Isotopendiskriminierung durch das Verhältnis der Geschwindigkeiten des Transports von CO2 in das Blatt und der Carboxylierung des Ribulose-1,5-bisphosphats (RuBP) bestimmt. Ausdruck dieses Zusammenhangs ist eine vonFarquhar (1980) angegebene lineare Beziehung zwischen δ13C-Wert und dem Verhältnis von interzellulärer und äußerer CO2-Konzentration. Die Isotopendiskriminierung bei der C4-Photosynthese ist ebenfalls von den Geschwindigkeiten der Diffusion von CO2 in das Blatt und der primären Carboxylierung von Phosphoenolpyruvat (PEP) abhängig. Wegen der unvermeidlichen Verluste von CO2 aus den Leitbündelscheidenzellen sind aber auch die sekundäre Carboxylierung von RuBP und der CO2-Transport innerhalb des Blattes zu berücksichtigen. Ein auf dieser Grundlage entwickeltes Modell, das im Gegensatz zu einem früher beschriebenen Ansatz auch die CO2-Abhängigkeit der Geschwindigkeitsparameter für die Carboxylierungsschritte einschließlich der Limitierung der PEP-Carboxylierung durch die PEP-Regenerationsrate berücksichtigt, wird ausführlich dargestellt. Nach diesem Modell können die wenigen bisher bekannten Befunde zum Einfluß hoher CO2-Konzentrationen auf die Isotopendiskriminierung von C4-Pflanzen als Hinweis darauf interpretiert werden, daß der Isotopeneffekt der RuBP-Carboxylierung bei den C4-Pflanzen möglicherweise größer ist als bei den C3-Pflanzen.

Liste der Abkürzungen

PEP

Phosphoenolpyruvat

PEPC

Phosphoenolpyruvat-Carboxylase

RuBP

Ribulose-1,5-bisphosphat

RuBPC

Ribulose-1,5-bisphosphat-Carboxylase

RuBPCO

Ribulose-1,5-bisphosphat-Carboxylase/Oxygenase Decarboxylierende Enzyme der drei Typen von C4-Pflanzen:

NADP-Me

NADP-abhängiges Malat-Enzym

NAD-Me

NAD-abhängiges Malat-Enzym

PCK

Phosphoenolpyruvat-Carboxykinase

Models of carbon isotope discrimination during photosynthesis of C3 and C4 plants

Summary

Models of fractionation of carbon atoms12C and13C during photosynthesis of C3 and C4 plants are described and discussed with regard to the influence of internal and external factors on this process. Isotope discrimination in C3 plants is determined by the ratio of the rates of CO2 transfer into the leaf and carboxylation of ribulose-1,5-bisphosphate (RuBP). This relationship can be expressed by a linear equation between δ13C value and the ratio of intercellular and external CO2 concentration as proposed byFarquhar (1980). Carbon isotope discrimination by C4 photosynthesis also depends on the rates of CO2 diffusion into the leaf and of primary carboxylation of phosphoenolpyruvate (PEP). However, because of the unavoidable losses of CO2 from bundle sheath cells the secondary carboxylation of RuBP as well as CO2 transfer processes within the leaf must be taken into account, too. A model based on these features is explained in some detail. In contrast to a model proposed earlier the presented model considers the dependence on CO2 concentration of rate parameters for the carboxylation steps including limitation of PEP carboxylation by PEP regeneration rate. According to the model the spare experimental results on the influence of high concentrations of CO2 on carbon isotope discrimination in C4 plants may be interpreted as an indication that the isotope effect of RuBP carboxylation in C4 plants could be higher than in C3 plants.

Модели изотопной дискриминации углерода в фотосинтезе С3 - и С4 -растений

Краткое содержание

В предлагаемой работе описываются модели обогащения углеродного атома12С в сравнении13С в фотосинтезе растений типа С3 и С4 и, в связи с литературным данными, обсуждается влияние внутренних и внешних факторов на этот процесс. У С3-растений изотопная дискриминация определяется соотношением скоростей транспорта СО2 к листьям и карбоксилирования рибулозо-1,5-дифосфата (РДФ). Выражением этой связи является, предложенная Фаркуаром (1980), линейная зависимость величины δ13 С от соотношения межклеточной и внешней концентрацнн СО2. Изотопная дискриминация в синтезе типа С4 зависит такзе от скоростей диффузии СО2 к листу и первичного карбоксилирования фосфоэнолпирувата (ФЭП). Благодаря неизбежной потере СО2 из клеток обкладки проводящих пучков, вадо учитыватв вторичное карбоксилирование РДФ и транспорт СО2 внутри листа. Подробно описывается модель, построенная на этой основе, которая, в противоположность ранее предложенной, учитывает также параметры скорости протекания Фаз карбоксилирования в зависимости от концентрации СО2, включая лимитирование карбоксилирования ФЭП его регенеративной скороствю. Применяя описанную модель, можно немногие, до сих пор известные, данные о влиянии высоких концентраций СО2 на изотопную дискриминацию у С4-растений, интерпретировать как указание на то, что изотопный эффект карбоксилирования ФЭП у С4 -растений, возможно, оказывается более сильным, чем у С3-растений.

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Literatur

  1. Abdulrahman, F. S., andG. J. Williams, 1981: Temperature and salinity regulation of growth and gas exchange ofSalicornia fruticosa (L.) L. — Oecologia48, 346–352.CrossRefGoogle Scholar
  2. Apel, P., undM. Peisker, 1978: Einfluß hoher Sauerstoffkonzentrationen auf den CO2-Kompensationspunkt von C4-Pflanzen. — Kulturpflanze26, 99–103.CrossRefGoogle Scholar
  3. --, und --, 1980: CO2-Versorgung der photosynthetischen Carboxylierungszentren. — In: Biophysik, Biochemie und Physiologie der Photosynthese (Hrsg.P. Hoffmann undB. Hieke), pp. 181–204. Colloquia Pflanzenphysiologie Nr. 3. Humboldt-Universität zu Berlin.Google Scholar
  4. Bauwe, H., 1979: Biochemische und gaswechselphysiologische Untersuchungen zur Charakterisierung von Pflanzenarten mit C3–C4 intermediären Eigenschaften sowie zur Prüfung mathematischer Modellvorstellungen des photosynthetischen CO2-Stoffwechsels. Dissertation A. Martin-Luther-Universität Halle — Wittenberg.Google Scholar
  5. ——, 1984: Photosynthetic enzyme activities and immunofluorescence studies on the localization of ribulose-1,5-bisphosphate carboxylase/oxygenase in leaves of C3, C4, and C3–C4 intermediate species ofFlaveria (Asteraceae) — Biochem. Physiol. Pflanzen179, 253–268.Google Scholar
  6. Berry, J. A., andG. D. Farquhar, 1978: The CO2 concentrating function of C4 photosynthesis. A biochemical model. — In: Photosynthesis 77 (Eds.D. O. Hall, J. Coombs, T. W. Goodwin), pp. 119–131. The Biochemical Society, London.Google Scholar
  7. Boag, T. S., 1982: Characterization of C4 photosynthesis in sodium-deficient plants. — Ph. D. Thesis, Australian National University, Canberra.Google Scholar
  8. Bradford, K. J., T. D. Sharkey, andG. D. Farquhar, 1983: Gas exchange, stomatal behavior, and δ13C values of theflacca tomato mutant in relation to abscisic acid. — Plant Physiol.72, 245–250.PubMedGoogle Scholar
  9. Brown, W. V., 1977: The Kranz syndrome and its subtypes in grass systematics. — Mem. Torrey Bot. Club23 (2), 1–97.Google Scholar
  10. Card, K. A., B. Mahall, andJ. H. Troughton, 1974: Salinity and carbon isotope ratios in C3 and C4 plants. — Carnegie Institution Year Book73, 784–785.Google Scholar
  11. Chartier, P., etO. Bethenod, 1977: La productivité primaire à l'échelle de la feuille. — In: Les processus de la production végétale primaire (Ed.A. Moyse), pp. 77–112. Gauthier-Villars, Paris.Google Scholar
  12. Craig, H., 1957: Isotopic standards for carbon and oxygen and correction factors for mass spectrometric analysis of carbon dioxide. — Geochim. Cosmochim. Acta12, 133–149.CrossRefGoogle Scholar
  13. Edwards, G., andD. Walker, 1983: C3, C4: mechanisms, and cellular and environmental regulation, of photosynthesis. — Blackwell Scientific Publications, Oxford - London - Edinburgh - Boston - Melbourne.Google Scholar
  14. Ehleringer, J., andR. W. Pearcy, 1983: Variation in quantum yield for CO2 uptake among C3 and C4 plants. — Plant Physiol.73, 555–559.PubMedGoogle Scholar
  15. Farquhar, G. D., 1980: Carbon isotope discrimination by plants: Effects of carbon dioxide concentration and temperature via the ratio of intercellular and atmospheric CO2 concentrations. — In: Carbon Dioxide and Climate: Australian Research (Ed.G. I. Pearman), pp. 105–110. Australian Academy of Science, Canberra.Google Scholar
  16. ——, 1983: On the nature of carbon isotope discrimination in C4 species. — Aust. J. Plant Physiol.10, 205–226.Google Scholar
  17. ——, andZ. Roksandic, 1982a: Effect of salinity and humidity on δ13C value of halophytes-evidence for diffusional isotope fractionation determined by the ratio of intercellular/atmospheric partial pressure of CO2 under different environmental conditions. — Oecologia52, 121–124.CrossRefGoogle Scholar
  18. —— andJ. A. Berry, 1982b: On the relationship between carbon isotope discrimination and the intercellular carbon dioxide concentration in leaves. — Aust. J. Plant Physiol.9, 121–137.Google Scholar
  19. ——, andS. von Caemmerer, 1982: Modelling of photosynthetic response to environmental conditions. — In: Physiological plant ecology II (Eds.O. L. Lange, P. S. Nobel, C. B. Osmond, andH. Ziegler), pp. 549–587. Encyclopedia of Plant Physiology. New Series, Vol. 12B, Springer-Verlag, Berlin - Heidelberg - New York.Google Scholar
  20. Francey, R. J., andG. D. Farquhar, 1982: An explanation of13C/12C variations in tree rings. — Nature197, 28–31.CrossRefGoogle Scholar
  21. Guy, R. D., D. M. Reid andH. R. Krouse, 1980: Shifts in carbon isotope ratios of two C3 halophytes under natural and artificial conditions. — Oecologia44, 241–247.CrossRefGoogle Scholar
  22. Hatch, M. D., andC. B. Osmond, 1976: Compartmentation and transport in C4 photosynthesis. — In: Transport in plants III (Eds.C. R. Stocking andU. Heber), pp. 143–184. Encyclopedia of Plant Physiology. New Series, Vol. 3. Springer-Verlag, Berlin - Heidelberg - New York.Google Scholar
  23. Hattersley, P. W., 1982: δ13C values of C4 types in grasses. — Austr. J. Plant Physiol.9, 139–154.Google Scholar
  24. ——, andA. J. Browning, 1981: Occurence of the suberized lamella in leaves of grasses of different photosynthetic types. I. In parenchymatous bundle sheaths and PCR („Kranz“) sheaths. — Protoplasma109, 371–401.CrossRefGoogle Scholar
  25. Holtum, J. A. M., M. H. O'Leary, andC. B. Osmond, 1983: Effect of varying CO2 partial pressure on photosynthesis and on carbon isotope composition of carbon-4 of malate from the crassulacean acid metabolism plantKalanchoë daigremontiana Hamet et Perr. — Plant Physiol.71, 602–609.PubMedGoogle Scholar
  26. Keeling, C. D., W. G. Mook, andP. P. Tans, 1979: Recent trends in the13C/12C ratio of atmospheric carbon dioxide. — Nature277, 121–123.CrossRefGoogle Scholar
  27. Lerman, J. C., 1975: How to interpret variations in the carbon isotope ratio of plants: biologic and environmental effects. — In: Environmental and biological control of photosynthesis (Ed.R. Marcelle), pp. 323–335. Dr. W. Junk b. v., Publishers, The Hague.Google Scholar
  28. O'Leary, M. H., 1981: Carbon isotope fractionation in plants. — Phytochem.20, 553–567.CrossRefGoogle Scholar
  29. ——, andC. B. Osmond, 1980: Diffusional contribution to carbon isotope fractionation during dark CO2 fixation in CAM plants. — Plant Physiol.66, 931–934.PubMedGoogle Scholar
  30. Osmond, C. B., 1971: Metabolite transport in C4 photosynthesis. — Aust. J. Biol. Sci.24, 159–163.PubMedGoogle Scholar
  31. ——, 1978: Crassulacean acid metabolism: a curiosity in context. — Ann. Rev. Plant Physiol.29, 379–414.CrossRefGoogle Scholar
  32. Park, R., andS. Epstein, 1960: Carbon isotope fractionation during photosynthesis. — Geochim. Cosmochim. Acta21, 110–126.CrossRefGoogle Scholar
  33. Peisker, M., 1978: Der Einfluß von Sauerstoff auf die CO2-Kompensationskonzentration von C3- und C4-Pflanzen und von Intermediärformen. — Kulturpflanze26, 81–98.CrossRefGoogle Scholar
  34. ——, 1979: Conditions for low, and oxygen-independent, CO2 compensation concentrations in C4 plants as derived from a simple model. — Photosynthetica13, 198–207.Google Scholar
  35. ——, 1982a: The effect of CO2 leakage from bundle sheath cells on carbon isotope discrimination in C4 plants. — Photosynthetica16, 533–541.Google Scholar
  36. --, 1982b: Theoretische Analyse der Änderungen des δ13C-Wertes bei der Evolution des C4-pathway der Photosynthese. — In: Photosynthese: Regulation und Evolution (Hrsg.P. Hoffmann undB. Hieke), pp. 163–167. Colloquia Pflanzenphysiologie Nr. 5. Humboldt-Universität zu Berlin.Google Scholar
  37. ——, andH. Bauwe, 1984: Modelling carbon metabolism in C3–C4 intermediate species. 1. CO2 compensation concentration and its O2 dependence. — Photosynthetica18, 9–19.Google Scholar
  38. —— andJ. Čatsky, 1981: Ontogenetic changes in the internal limitations to bean leaf photosynthesis. 7. Interpretation of the linear correlation between CO2 compensation concentration and CO2 evolution in darkness. — Photosynthetica15, 161–168.Google Scholar
  39. Rathnam, C. K. M., 1978: Metabolic regulation of carbon flux during C4 photosynthesis. I. Evidence for parallel CO2 fixation by mesophyll and bundle sheath cells in situ. — Z. Pflanzenphysiol.87, 65–84.Google Scholar
  40. Raven, J. A., 1977: Ribulose bisphosphate carboxylase activity in terrestrial plants: significance of O2 and CO2 diffusion. — Curr. Adv. Plant Sci.9, 579–590.Google Scholar
  41. ——, andS. M. Glidewell, 1981: Processes limiting photosynthetic conductance. — In: Physiological processes limiting plant productivity (Ed.C. B. Johnson), pp. 109–136. Butterworths, London - Boston - Sydney - Wellington- Durban - Toronto.Google Scholar
  42. Schmidt, H.-L., andF. J. Winkler, 1979: Einige Ursachen der Variationsbreite von δ13C-Werten bei C3- und C4-Pflanzen. — Ber. Deutsch. Bot. Ges.92, 185–191.Google Scholar
  43. Shomer-Ilan, A., S. Beer, andY. Waisel, 1975:Suaeda monoica, a C4 plant without typical bundle sheaths. — Plant Physiol.56, 676–679.PubMedGoogle Scholar
  44. Smith, B. N., andT. W. Boutton, 1911: Environmental influences on13C/12C ratios and C4 photosynthesis. — In: Photosynthesis VI. Photosynthesis and productivity, photosynthesis and environment (Ed.G. Akoyunoglou), pp. 255–262. Balaban International Science Services, Philadelphia, Pa.Google Scholar
  45. —— andC. McMillan, 1976: Influence of carbon source, oxygen concentration, light intensity, and temperature on13C/12C ratios in plant tissues. — Bot. Gaz.137, 99–104.CrossRefGoogle Scholar
  46. Troughton, J. H., 1979: δ13C as an indicator of carboxylation reactions. — In: Photosynthesis II (Eds.M. Gibbs andE. Latzko), pp. 140–149. Encyclopedia of Plant Physiology. New Series, Vol. 6. Springer-Verlag, Berlin- Heidelberg - New York.Google Scholar
  47. Vogel, J. C., 1980: Fractionation of the carbon isotopes during photosynthesis. — Sitzungsberichte der Heidelberger Akad. d. Wiss., Math.-naturwiss. Klasse, Jahrgang 1980, 3. Abhdlg., 111–135.Google Scholar
  48. —— andR. P. Ellis, 1978: The geographical distribution of Kranz grasses in South Africa. — South Afr. J. Sci.74, 209–215.Google Scholar
  49. Winter, K., 1981: CO2 and water vapour exchange, malate content and δ13C value inCicer arietinum grown under two water regimes. — Z. Pflanzenphysiol.101, 421–430.Google Scholar
  50. ——, andM. H. O'Leary, 1982: Effect of low relative humidity on δ13C value in two C3 grasses and inPanicum milioides, a C3–C4 intermediate species. — J. Exp. Bot.33, 88–91.CrossRefGoogle Scholar
  51. —— andJ. S. Pate, 1981: Coping with salinity. — In: Biology of Australian native plants (Eds.J. S. Pate andA. J. McComb), pp. 88–113. University of Western Australia Press, Perth.Google Scholar
  52. ——, andJ. H. Troughton, 1978: Photosynthetic pathways in plants of coastal and inland habitats of Israel and the Sinai. — Flora167, 1–34.Google Scholar
  53. Wong, S. C., I. R. Cowan, andG. D. Farquhar, 1979: Stomatal conductance correlates with photosynthetic capacity. — Nature282, 424–426.CrossRefGoogle Scholar
  54. Yeoh, H.-H., M. R. Badger, andL. Watson, 1980: Variations in Km (CO2) of ribulose-1,5-bisphosphate carboxylase among grasses. — Plant Physiol.66, 1110–1112.PubMedCrossRefGoogle Scholar
  55. Ziegler, H., K. H. Batanouny, N. Sankhla, O. P. Vyas, andW. Stichler, 1981: The photosynthetic pathway types of some desert plants from India, Saudi Arabia, Egypt, and IRAQ. — Oecologia48, 93–99.CrossRefGoogle Scholar

Copyright information

© Akademie-Verlag 1983

Authors and Affiliations

  • Martin Peisker
    • 1
  1. 1.Zentralinstitut für Genetik und Kulturpflanzenforschung der Akademie der Wissenschaften der DDRGaterslebenDDR

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