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Chapter 4 Regulation and Functions of the Chlorophyll Cycle

  • Chapter
The Chloroplast

Part of the book series: Advances in Photosynthesis and Respiration ((AIPH,volume 31))

Summary

The chlorophyll cycle refers to the interconversion of chlorophyll a and chlorophyll b that occurs within the chloroplasts of higher plants. The forward reaction that converts chlorophyll a to b is catalyzed by chlorophyllide a oxygenase. The backward reaction from chlorophyll b to a is catalyzed by a recently identified enzyme, chlorophyll b reductase, and an unidentified enzyme, which is hypothetically named 7-hydroxymethyl chlorophyll a reductase. The chlorophyll cycle plays two important physiological roles: (a) the chlorophyll cycle adjusts the ratio of the peripheral antenna to the core antenna, and (b) it facilitates the degradation of chlorophyll b, light-harvesting complexes, and thylakoid membranes during leaf senescence. In this article, we summarize the research history, the functions, and the regulation of the chlorophyll cycle. Furthermore, we discuss the evolution of the chlorophyll cycle. Recent progresses in evolutionary aspect are emphasized: those include (a) the discovery of the Prochlorococcus gene for chlorophyllide a oxygenase, (b) the presumable gene separation which occurred in the Prasinophyceae, and (c) the duplication and diversification of the genes encoding chlorophyll b reductase. In addition, our current understanding of the regulatory mechanisms of chlorophyll a to b conversion is described and covers topics such as: (a) transcriptional regulation of chlorophyllide a oxygenase in response to light intensities, (b) the mechanism of chlorophyll b-dependent feedback regulation of chlorophyllide a oxygenase which mechanism involves Clp protease, and (c) the close coordination of the chlorophyll cycle activity and the construction of the photosynthetic machinery. Finally, recent progress in the study of chlorophyll b to a conversion is summarized, the major topic of which is the identification of the genes encoding two isoforms of chlorophyll b reductase.

Epilogue

Although the study on the interconversion of Chl a and b has a long history, our understanding on its reaction chemistry, its regulatory mechanisms and its evolution is limited. Identification of HM-Chl a reductase might be the first step toward gaining a better understanding of the all aspects of the Chl cycle. Subsequently, biochemical analyses of the interaction of the Chl cycle enzymes and photosynthetic proteins may reveal how the photosynthetic machinery acclimates to different light conditions. Furthermore, studies of the diversity of the enzymes involved in the Chl cycle may reveal a part of the evolutionary history of photosynthetic organisms.

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Abbreviations

BChl:

bacteriochlorophyll;

CAO:

chlorophyllide a oxygenase;

Chl:

chlorophyll;

CP1:

the core complex of the photosystem I;

GFP:

green fluorescent protein;

HM:

7-hydroxymethyl;

LHC:

light-harvesting complexSAM S-adenosyl-L-methionine;

PS:

photosystem PCB prochlorophytes chlorophyll b binding proteins

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Acknowledgements

We thank Prof. Isamu Inouye and Ms. Tomoko Chikuni for helpful discussion and for sharing unpublished results with us. We are grateful to Ms. Junko Kishimoto for illustrations. We acknowledge the financial support from the Grant-in-Aid for Creative Scientific Research (17GS0314) to AT and the Grant-in-Aid for Scientific Research (19687003) to RT from the Japanese Ministry of Education, Culture, Sports, Science and Technology.

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Authors and Affiliations

  1. Institute of Low Temperature Science, Hokkaido University, N19W8, Kita-ku, Sapporo, 060-0819, Japan

    Ryouichi Tanaka, Hisashi Ito & Ayumi Tanaka

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  1. Ryouichi Tanaka
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  2. Hisashi Ito
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  3. Ayumi Tanaka
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Correspondence to Ayumi Tanaka .

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Editors and Affiliations

  1. Rebeiz Foundation for Basic Research, Champaign, USA

    Constantin A. Rebeiz

  2. Michigan State University, East Lansing, USA

    Christoph Benning

  3. University of Illinois at Urbana-Champaign, Urbana, USA

    Hans J. Bohnert

  4. Central Florida University, Orlando, USA

    Henry Daniell

  5. Arizona State University, Tempe, USA

    J. Kenneth Hoober

  6. University of Karlsruhe, Germany

    Hartmut K. Lichtenthaler

  7. University of Illinois at Urbana-Champaign, Urbana, USA

    Archie R. Portis

  8. Jawaharlal Nehru University, New Delhi, India

    Baishnab C. Tripathy

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Tanaka, R., Ito, H., Tanaka, A. (2010). Chapter 4 Regulation and Functions of the Chlorophyll Cycle. In: Rebeiz, C.A., et al. The Chloroplast. Advances in Photosynthesis and Respiration, vol 31. Springer, Dordrecht. https://doi.org/10.1007/978-90-481-8531-3_4

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