Abstract
Senescence processes were investigated in attached seedling leaves of barley (Hordeum vulgare L. cv. Steffi) grown under standardized conditions in growth chambers. Even a few
days after reaching full length, total chlorophyll and protein content of first foliage
leaves start to decrease, indicating degradation of photosynthetic units. About five
weeks later, the final stage of leaf senescence is reached, with almost no chlorophyll
and only about 30 % of the protein remaining. In spite of the early decrease in total
protein and chlorophyll content, efficiency of remaining photosystem II units stays
high for about 3 weeks. Then it rapidly decreases, reaching values close to zero two
weeks later. The second leaves show similar changes in photosynthetic parameters as
the primary leaves, but the kinetics are delayed by about 1 week. Immunological analyses
reveal specific changes in the composition of the photosynthetic units during leaf
senescence. The level of photosystem I reaction centre protein sharply decreases at
35 d after sowing. Levels of photosystem II reaction centre protein D1 and of cytochrome
f also decrease at this developmental stage, but not as markedly as the photosytem
I reaction centre. Lower levels of these proteins can still be detected in later stages
of senescence. A preferential loss of photosystem I reaction centre at this developmental
stage could also be shown by spectroscopic measurements of concentrations of reaction
centre pigments P700 (photosystem I) and Qa (photosystem II, existing in a 1 : 1 stochiometry to P680). In contrast to reaction
centre proteins and cytochrome f, inner light-harvesting complex LHCI of photosystem I is more stable, still showing
high levels even in later stages of senescence. Analyses of transcript levels of the
corresponding genes show that changes in protein levels are, in part, due to differential
down regulation of gene expression during senescence. Whereas the proportions of psaA/B and petA transcripts start to decrease in primary leaves about 3 weeks after sowing, proportions
of psbA and Lhcb4 transcripts stay high, even in late stages of senescence.
Key words
Hordeum vulgare
- leaf senescence - photosynthesis-related genes - photosynthetic units
References
- 01
Becker, W., and Apel, K..
(1993);
Differences in gene expression between mature and artificially induced leaf senescence.
Planta.
189
74-79
- 02
Bricker, T. M., and Newman, D. W..
(1980);
Quantitative changes in the chloroplast thylakoid polypeptide complement during senescence.
Z. Pflanzenphys..
98
339-346
- 03
Bruce, B. D., and Malkin, R. D..
(1991);
Biosynthesis of the chloroplast b6 f complex: studies in a photosynthetic mutant of
Lemna.
.
The Plant Cell.
3
203-212
- 04
Chirgwin, J. M.,, Pryzybyla, A. E.,, MacDonald, R. J.,, and Rutter, W. J..
(1979);
Isolation of biological active ribonucleic acid from sources enriched in ribonuclease.
Biochemistry.
18
5294-5299
- 05
Droillard, M. J.,, Bate, N. J.,, Rothstein, S. J.,, and Thompson, J. E..
(1992);
Active translation of the D-1 protein of photosystem II in senescing leaves.
Plant Phys..
99
589-594
- 06
Gan, S., and Amasino, R. M..
(1997);
Making sense of senescence. Molecular genetic regulation and manipulation of leaf
senescence.
Plant Physiol..
113
313-319
- 07 Gepstein, S.. (1988)
Photosynthesis. Senescence and aging in plants. Noodén, L. D. and Leopold, L. C., eds. San Diego;
Academic Press pp. 85-109
- 08 Grabau, L. J.. (1995)
Physiological mechanisms of plant senescence. Handbook of plant and crop physiology. Pessarakli, M., ed. Cleveland; CRC Press pp.
483-496
- 09 Hoyer-Hansen, G.. (1987)
Characterization of monoclonal antibodies. Handbook of immunoblotting of proteins. Bjerum, O. J. and Heegaard, N. H. H., eds.
Cleveland; CRC Press pp. 457-478
- 10
Hoyer-Hansen, G.,, Bassi, R.,, Honberg, L. S.,, and Simpson, D. J..
(1988);
Immunological characterization of chlorophyll a/b-binding proteins of barley thylakoids.
Planta.
173
12-21
- 11
Humbeck, K.,, Melis, A.,, and Krupinska, K..
(1994 a);
Effects of chilling on chloroplast development in barley primary foliage leaves.
J. Plant Physiol..
143
744-749
- 12
Humbeck, K.,, Kloppstech, K.,, and Krupinska, K..
(1994 b);
Expression of early light-inducible proteins in flag leaves of field-grown barley.
Plant Physiol..
105
1217-1222
- 13
Humbeck, K.,, Quast, S.,, and Krupinska, K..
(1996);
Functional and molecular changes in the photosynthetic apparatus during senescence
of flag leaves from field-grown barley plants.
Plant, Cell and Environment.
19
337-344
- 14
Jenkins, G. I.,, Baker, N. R.,, and Woolhouse, H. W..
(1981);
Changes in chlorophyll content and organization during senescence of primary leaves
of Phaseolus vulgaris (L.) in relation to photosynthetic electron transport.
J. Exp. Bot..
32
1009-1020
- 15
Jenkins, G. I., and Woolhouse, H. W..
(1981);
Photosynthetic electron transport during senescence of primary leaves of Phaseolus vulgaris L. II. The activity of photosystems one and two, and a note on the site of reduction
of ferricyanide.
J. Exp. Bot..
32
989-997
- 16
Kleber-Janke, T., and Krupinska, K..
(1997);
Isolation of cDNA clones for genes showing enhanced expression in barley leaves during
dark-induced senescence as well as during senescence under field conditions.
Planta.
203
332-340
- 17
Krause, K.,, Falk, J.,, Humbeck, K.,, and Krupinska, K..
(1998);
Responses of the transcriptional apparatus of barley chloroplasts to a prolonged dark
period and to subsequent reillumination.
Physiol. Plant..
104
143-152
- 18 Krupinska, K., and Humbeck, K.. (2000)
Photosynthesis and chloroplast breakdown. Cell death in plants. Noodén, L. D., ed. San Diego; Academic Press in press
- 19
Mattoo, A. K.,, Marder, J. B.,, and Edelman, M..
(1989);
Dynamics of the photosystem II reaction center.
Cell.
56
241-246
- 20
Matile, P.,, Hörtensteiner, S.,, and Thomas, H..
(1999);
Chlorophyll degradation.
Annu. Rev. Plant Physiol. Plant Mol. Biol..
50
67-95
- 21
Melis, A..
(1991);
Dynamics of photosynthetic membrane composition and function.
Biochim. Biophys. Acta.
1058
87-106
- 22
Noodén, L. D.,, Guiamét, J. J.,, and John, I..
(1997);
Senescence mechanisms.
Physiol. Plant..
101
746-753
- 23 Noodén, L. D., and Leopold, A. C.. (1978)
Phytohormones and the endogenous regulation of senescence and abscission. Phytohormones and related compounds. Letham, D., Goodwin, T., and Higgins, T., eds.
New York; Elsevier pp. 329-369
- 24
Smart, C. M..
(1994);
Gene expression during leaf senescence.
New Phytol..
126
419-448
- 25
Smith, B. M.,, Morrissey, P. J.,, Guenther, J. E.,, Nemson, J. A.,, Harrison,
M. A.,, Allen, J. F.,, and Melis, A..
(1990);
Response of the photosynthetic apparatus in Dunaliella salina (green algae) to irradiance stress.
Plant Phys..
93
1433-1440
- 26
Souza, P. I.,, Egli, D. B.,, and Bruening, W. P..
(1997);
Water stress during seed filling and leaf senescence in soybean.
Agron. J..
89
807-812
- 27
Wittenbach, V. A..
(1977);
Induced senescence of intact wheat seedlings and its reversibility.
Plant Phys..
59
1039-1042
- 28
Yamasaki, T.,, Kudoh, T.,, Kamimura, Y.,, and Katoh, S..
(1996);
A vertical gradient of the chloroplast abundance among leaves of Chenopodium album.
.
Plant Cell Physiol..
37
43-48
K. Humbeck
Institut für Pflanzenphysiologie
Martin-Luther-Universität Halle-Wittenberg
Weinbergweg 10
06099 Halle (Saale)
Germany
Email: humbeck@pflanzenphys.uni-halle.de
Section Editor: U. Lüttge