Changes in photosynthesis of alpine plant Saussurea superba during leaf expansion
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Abstract
The native alpine plant Saussurea superba is widely distributed in Qinghai–Tibetan Plateau regions. The leaves of S. superba grow in whorled rosettes, and are horizontally oriented to maximize sunlight exposure. Experiments were conducted in an alpine Kobresia humilis meadow near Haibei Alpine Meadow Ecosystem Research Station (37°29′–37°45′N, 101°12′–101°33′E; alt. 3200 m). Leaf growth, photosynthetic pigments and chlorophyll fluorescence parameters were measured in expanding leaves of S. superba. The results indicate that leaf area increased progressively from inner younger leaves to outside fully expanded ones, and then slightly decreased in nearly senescent leaves, due to early unfavorable environmental conditions, deviating from the ordinary growth pattern. The specific leaf area decreased before leaves were fully expanded, and the leaf thickness was largest in mature leaves. There were no significant changes in the content of chlorophylls (Chl) and carotenoids (Car), but the ratios of Chl a/b and Car/Chl declined after full expansion of the leaves. The variation of Chl a/b coincided well with changes in photochemical quenching (q P) and the fraction of open PSII reaction centers (q L). The maximum quantum efficiency of PSII photochemistry after 5 min dark relaxation (F (v)/F (m)) continuously increased from younger leaves to fully mature leaves, suggesting that mature leaves could recover more quickly from photoinhibition than younger leaves. The light-harvesting capacity was relatively steady during leaf expansion, as indicated by the maximum quantum efficiency of open PSII centers (\(F_{\text{v}}^{{\prime }}\)/\(F_{\text{m}}^{{\prime }}\)). UV-absorbing compounds could effectively screen harmful solar radiation, and are a main protection way on the photosynthetic apparatus. The decline of q P and q L during maturation, together with limitation of quantum efficiency of PSII reaction centers (L (PFD)), shows a decrease of oxidation state of QA in PSII reaction centers under natural sunlight. Furthermore, light-induced (Φ NPQ) and non-light-induced quenching (Φ NO) were consistent with variation of L (PFD). It is concluded that the leaves of S. superba could be classified into four functional groups: young, fully expanded, mature, and senescent. Quick recovery from photoinhibition was correlated with protection by screening pigments, and high level of light energy trapping was correlated with preservation of photosynthetic pigments. Increasing of Φ NPQ and Φ NO during leaves maturation indicates that both thermal dissipation of excessive excitation energy in safety and potential threat to photosynthetic apparatus were strengthened due to the declination of q P and q L, and enhancement of L (PFD).
Keywords
Alpine plant Chlorophyll fluorescence parameters Leaf growth Photosynthetic pigments Qinghai–Tibetan Plateau Saussurea superbaAbbreviations
- SLA
Specific leaf area
- LWR
Ratio of dry to fresh weight of leaves
- Chl
Chlorophylls
- Car
Carotenoids
- Chl a/b
Ratio of chlorophyll a to chlorophyll b
- Car/Chl
Ratio of carotenoids to chlorophylls
- qP
Photochemical quenching coefficient
- NPQ
Non-photochemical quenching coefficient
- qL
Fraction of opened PSII reaction centers
- F(v)/F(m)
Maximum quantum efficiency of PSII photochemistry after 5 min dark relaxation
- \(F_{\text{v}}^{{\prime }}\)/\(F_{\text{m}}^{{\prime }}\)
Maximum quantum efficiency of PSII photochemistry at given light
- L(PFD)
Relative limitation of quantum efficiency of PSII reaction centers
- ΦNPQ
Quantum yield of light-induced PSII regulatory energy dissipation
- ΦNO
Quantum yield of non-light-induced PSII non-regulatory energy dissipation
- ΦPSII
Actual quantum yield of PSII photochemistry
- QA
Primary quinone electron acceptor of PSII
Notes
Acknowledgments
This work was supported by the National Natural Science Foundation of China (30670307), the Key Plan of International Cooperation Research in Sciences and Techniques (2002CB714006), and the Natural Science Foundation of Qinghai Province (2013-Z-915). We thank Professor Lars Olof Björn for critical reading and editing of this manuscript.
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