Ilektra Sperdouli

Interaction of proline, sugars, and anthocyanins during photosynthetic acclimation of Arabidopsis thaliana to drought stress

The relationships among photosynthetic acclimation, proline (Pro), soluble sugar (SS), and anthocyanin (An) accumulation in Arabidopsis thaliana leaves to the onset of drought stress (OnDS), mild (MiDS) and moderate drought stress (MoDS), were evaluated. As leaf water content (LWC) decreased, metabolic concentrations (Pro, SS, and An) increased and were negatively and significantly correlated with LWC. Thus, these metabolites may have an important role in the acclimation process to drought stress (DS). No correlations among Pro, SS and An accumulation with the quantum efficiency of PSII photochemistry (Φ(PSII)) and the excitation pressure (1-q(P)) were observed under DS. This implies that, while metabolites increased in a drought-dependent way, PSII activity did not decrease in the same pattern. Our results indicated that, under MoDS, A. thaliana leaves were able to maintain oxidative compounds such as malondialdeyde, an end product of lipid peroxidation, within the range of control leaves, and to cope with oxidative damage, as was evident by the decreased excitation pressure (1-q(P)) and similar (ns difference) Φ(PSII) to that of control leaves. In addition, a statistically significant increased accumulation of Pro, SS and An was recorded only under MoDS compared to controls. The better PSII functioning of MoDS Arabidopsis leaves may reflect the greater capacity of these leaves to undertake key metabolic adjustments, including increased Pro, SS and An accumulation, to maintain a higher antioxidant protection and a better balance between light capture and energy use.

Spatio-temporal heterogeneity in Arabidopsis thaliana leaves under drought stress.

Using chlorophyll (chl) fluorescence imaging, we studied the effect of mild (MiDS), moderate (MoDS) and severe (SDS) drought stress on photosystem II (PSII) photochemistry of 4-week-old Arabidopsis thaliana. Spatio-temporal heterogeneity in all chl fluorescence parameters was maintained throughout water stress. After exposure to drought stress, maximum quantum yield of PSII photochemistry (F(v)/F(m)) and quantum efficiency of PSII photochemistry (Φ(PSΙΙ)) decreased less in the proximal (base) than in the distal (tip) leaf. The chl fluorescence parameter F(v) /F(m) decreased less after MoDS than MiDS. Under MoDS, the antioxidant mechanism of A. thaliana leaves seemed to be sufficient in scavenging reactive oxygen species, as evident by the decreased lipid peroxidation, the more excitation energy dissipated by non-photochemical quenching (NPQ) and decreased excitation pressure (1-q(p)). Arabidopsis leaves appear to function normally under MoDS, but do not seem to have particular metabolic tolerance mechanisms under MiDS and SDS, as revealed by the level of lipid peroxidation and decreased quantum yield for dissipation after down-regulation in PSII (Φ(NPQ)), indicating that energy dissipation by down-regulation did not function and electron transport (ETR) was depressed. The simultaneous increased quantum yield of non-regulated energy dissipation (Φ(NO)) indicated that both the photochemical energy conversion and protective regulatory mechanism were insufficient. The non-uniform photosynthetic pattern under drought stress may reflect different zones of leaf anatomy and mesophyll development. The data demonstrate that the effect of different degrees of drought stress on A. thaliana leaves show spatio-temporal heterogeneity, implying that common single time point or single point leaf analyses are inadequate.

Leaf developmental stage modulates metabolite accumulation and photosynthesis contributing to acclimation of Arabidopsis thaliana to water deficit

We examined whether young and mature leaves of Arabidopsis thaliana in their response to mild water deficit (MiWD) and moderate water deficit (MoWD), behave differentially, and whether photosynthetic acclimation to water deficit correlates with increased proline and sugar accumulation. We observed that with increasing water deficit, leaf relative water content decreased, while proline and sugar accumulation increased in both leaf-developmental stages. Under both MiWD and MoWD, young leaves showed less water loss and accumulated higher level of metabolites compared to mature leaves. This, leaf age-related increase in metabolite accumulation that was significantly higher under MoWD, allowed young leaves to cope with oxidative damage by maintaining their base levels of lipid peroxidation. Thus, acclimation of young leaves to MoWD, involves a better homeostasis of reactive oxygen species (ROS), that was achieved among others by (1) increased sugar accumulation and (2) either increased proline synthesis and/or decreased proline catabolism, that decrease the NADPH/NADP+ ratio, resulting in a higher level of oxidized state of quinone A and thus in a reduced excitation pressure, and by (3) stimulation of the photoprotective mechanism of non-photochemical quenching, that reflects the dissipation of excess excitation energy in the form of harmless heat, thus protecting the plant from the damaging effects of ROS.

A better energy allocation of absorbed light in photosystem II and less photooxidative damage contribute to acclimation of Arabidopsis thaliana young leaves to water deficit.

Water deficit stress promotes excitation pressure and photooxidative damage due to an imbalance between light capture and energy use. Young leaves (YL) of Arabidopsis thaliana plants acclimate better to the onset of water deficit (OnsWD) than do mature leaves (ML). To obtain a better understanding of this differential response, we evaluated whether YL and ML of A. thaliana exposed to the OnsWD, mild water deficit (MiWD) and moderate water deficit (MoWD), show differences in their photosynthetic performance, and whether photosynthetic acclimation correlates with leaf developmental stage. Water deficit (WD) resulted in greater photooxidative damage in ML compared to YL, but the latter could not be protected under the OnsWD or MiWD, but only under MoWD. YL of A. thaliana with signs of photosynthetic acclimation under MoWD retained higher maximum quantum yield (Fv/Fm) and decreased reactive oxygen species (ROS) formation. YL under MoWD, show a reduced excitation pressure and a better balance between light capture and photochemical energy use, which contributed to their photoprotection, but only under low light intensity (LL, 130μmolphotonsm(-2)s(-1)) and not under high light (HL, 1200μmolphotonsm(-2)s(-1)). In conclusion, leaf developmental stage was correlated with photo-oxidative damage and a differential allocation of absorbed light energy in photosystem II (PSII) of Arabidopsis leaves under WD.

Differential blockage of photosynthetic electron flow in young and mature leaves of Arabidopsis thaliana by exogenous proline

Responses of the photosynthetic electron transport system of chloroplasts to exogenous proline application were evaluated in young and mature leaves of Arabidopsis thaliana plants under optimal growth conditions. Exogenous proline application (10 mM) during the 4th week of growth increased proline accumulation in young leaves more than in mature leaves, and possibly due to its degradation producing NADPH, decreased significantly the ratio of NADP+/NADPH in both leaf types compared with controls (without proline). However, the ratio of NADP+/NADPH remained significantly higher in the young leaves, suggesting lower proline degradation which resulted in less reduced plastoquinone pool than that in the mature leaves, under both low light [130 μmol(photon) m−2 s−1] and high light [1,200 μmol(photon) m−2 s−1] treatments. The young leaves seemed to adjust nonphotochemical fluorescence quenching in order to maintain a better PSII quantum yield. We concluded that under optimal growth conditions exogenous proline results in overreduction of the plastoquinone pool and blockage of photosynthetic electron flow due to accumulation of NADPH. We suggest that optimum concentrations of proline are required for optimal PSII photochemistry.

Photosystem II Is More Sensitive than Photosystem I to Al3+ Induced Phytotoxicity

Aluminium (Al) the most abundant metal in the earth’s crust is toxic in acid soils (pH < 5.5) mainly in the ionic form of Al3+ species. The ability of crops to overcome Al toxicity varies among crop species and cultivars. Here, we report for a first time the simultaneous responses of photosystem II (PSII) and photosystem I (PSI) to Al3+ phytotoxicity. The responses of PSII and PSI in the durum wheat (Triticum turgidum L. cv. ‘Appulo E’) and the triticale (X Triticosecale Witmark cv. ‘Dada’) were evaluated by chlorophyll fluorescence quenching analysis and reflection spectroscopy respectively, under control (−Al, pH 6.5) and 148 μM Al (+Al, pH 4.5) conditions. During control growth conditions the high activity of PSII in ‘Appulo E’ led to a rather higher electron flow to PSI, which induced a higher PSI excitation pressure in ‘Appulo E’ than in ‘Dada’ that presented a lower PSII activity. However, under 148 μM Al the triticale ‘Dada’ presented a lower PSII and PSI excitation pressure than ‘Appulo E’. In conclusion, both photosystems of ‘Dada’ displayed a superior performance than ‘Appulo E’ under Al exposure, while in both cultivars PSII was more affected than PSI from Al3+ phytotoxicity.