Etioplasts and Their Significance in Chloroplast Biogenesis
Summary
Etioplasts are considered as convenient but not completely adequate laboratory models of proplastid-to-chloroplast development. These plastids are formed in light-deprived tissues of angiosperm plants that would become chlorenchyma in the light. Etioplasts have a unique inner membrane consisting of highly regular, paracrystalline prolamellar bodies (PLBs) and of lamellar prothylakoids (PTs). First, we recall different situations where etioplasts or PLBs do appear and play an important role during normal leaf ontogenesis and chloroplast biogenesis under natural light conditions. These structures appear almost exclusively in young tissues with not completely differentiated chloroplasts and photosynthetic apparatus, and under conditions where light is either temporally and/or spatially limited during development. PLBs can be formed in young leaves during the dark phase of the light–dark cycles (LDC); or in young seedlings developing in the soil from seeds; in water plants or inside special structures, where a decreasing light gradient is naturally formed, e.g. buds, enveloping sheaths of outer leaves. Having discussed the relevance of etioplasts in chloroplast biogenesis, we then outline the structure, organization and assembly of etioplast inner membranes in etiolated seedlings. Furthermore, the different factors important for PLB formation, and in parallel, the molecular composition of the PLBs are reviewed in details. A special lipid composition, a high lipid per protein ratio, the presence of oligomers of NADPH:protochlorophyllide (Pchlide) oxidoreductase (LPOR) proteins binding Pchlide, NADPH and carotenoids may all be important for the stabilization and formation of the special cubic membrane of the PLBs. Therefore, the biosynthesis of pigments in etioplasts is also discussed. The last part focuses on the etioplast-to-chloroplast transition during greening of etiolated seedlings, and summarizes the ultrastructural, molecular and physiological changes observed during this process. Finally, the significance of PLBs in plant development and leaf ontogeny is outlined.
Keywords
Carotenoid Biosynthesis Spectral Form Fluorescence Emission Maximum Chloroplast Biogenesis Chloroplast DifferentiationAbbreviations:
- ALA –
5-aminolevulinic acid;
- Chl(s) –
Chlorophyll(s);
- Chlide –
Chlorophyllide;
- Coprogen III –
Coproporphyrinogen III;
- DGDG –
Digalactosyl diacylglycerol;
- DPOR –
Dark-operative NADPH: Pchlide oxidoreductase;
- GG – Geranylgeraniol; GGPP –
Geranylgeraniol diphosphate;
- GSA –
Glutamate-1-semialdehyde;
- IPP –
Isopentenyl diphosphate;
- LDC –
Light–dark cycle;
- LPOR –
Light-dependent NADPH: Pchlide oxidoreductase;
- LPOR-A, -B, -C –
Isoforms of LPOR;
- MGDG –
Monogalactosyl diacylglycerol;
- Pchlide –
Protochlorophyllide;
- PGB –
Porphobilinogen;
- PLB –
Prolamellar body;
- Protogen IX –
Porphyrinogen III;
- PSI –
Photosystem I;
- PSII –
Photosystem II;
- PT –
Prothylakoid;
- Urogen III –
Uroporphyrinogen III
Notes
Acknowledgments
The authors are grateful to Csilla Jónás for skilful technical assistance, Dr Beata Mysliwa-Kurdziel for providing original pictures (Fig. 3.6), Prof. Benoit Schoefs for helpful discussion, and the Swedish Research Council VR for financial support (H.A). The electron microscopic examinations and fluorescence spectroscopy (Fig. 3.4) were done as in Solymosi et al. (2006a). Ultrathin sections were examined using Hitachi 7100 and JEOL JEM 1011 transmission electron microscopes. The project was supported by the European Union and co-financed by the European Social Fund (grant agreement no. TAMOP 4.2.1/B-09/1/KMR-2010-0003) (S.K.).
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