Summary 1.When a plant species is introduced into a new range, it may differentiate genetically from the original populations in the home range. This genetic differentiation may influence the extent to which the invasion of the new range is successful. We tested this hypothesis by examining Senecio pterophorus , a South African shrub that was introduced into NE Spain about 40 years ago. We predicted that in the introduced range invasive populations would perform better and show greater plasticity than native populations. 2. Individuals of S. pterophorus from four Spanish (invasive) and four South African (native) populations were grown in Catalonia, Spain, in a common garden in which disturbance and water availability were manipulated. Fitness traits and several ecophysiological parameters were measured. 3. The invasive populations of S. pterophorus survived better throughout the summer drought in a disturbed (unvegetated) environment than native South African populations. This success may be attributable to the lower specific leaf area (SLA) and better water content regulation of the invasive populations in this treatment. 4. Invasive populations displayed up to three times higher relative growth rate than native populations under conditions of disturbance and non-limiting water availability. 5. The reproductive performance of the invasive populations was higher in all treatments except under the most stressful conditions (i.e. in non-watered undisturbed plots), where no plant from either population flowered. 6. The results for leaf parameters and chlorophyll fluorescence measurements suggested that the greater fitness of the invasive populations could be attributed to more favourable ecophysiological responses. 7. Synthesis . Spanish invasive populations of S. pterophorus performed better in the presence of high levels of disturbance, and displayed higher plasticity of fitness traits in response to resource availability than native South African populations. Our results suggest that genetic differentiation from source populations associated with founding may play a role in invasion success.

Transglutaminases (TGases, EC 2.3.2.13) are intra- and extra-cellular enzymes that catalyze post-translational modification of proteins by establishing epsilon-(gamma-glutamyl) links and covalent conjugation of polyamines. In chloroplast it is well established that TGases specifically polyaminylate the light-harvesting antenna of Photosystem (PS) II (LHCII, CP29, CP26, CP24) and therefore a role in photosynthesis has been hypothesised (Della Mea et al. [23] and refs therein). However, the role of TGases in chloroplast is not yet fully understood. Here we report the effect of the over-expression of maize (Zea mays) chloroplast TGase in tobacco (Nicotiana tabacum var. Petit Havana) chloroplasts. The transglutaminase activity in over-expressers was increased 4 times in comparison to the wild-type tobacco plants, which in turn increased the thylakoid associated polyamines about 90%. Functional comparison between Wt tobacco and tgz over-expressers is shown in terms of fast fluorescence induction kinetics, non-photochemical quenching of the singlet excited state of chlorophyll a and antenna heterogeneity of PSII. Both in vivo probing and electron microscopy studies verified thylakoid remodeling. PSII antenna heterogeneity in vivo changes in the over-expressers to a great extent, with an increase of the centers located in grana-appressed regions (PSIIalpha) at the expense of centers located mainly in stroma thylakoids (PSIIbeta). A major increase in the granum size (i.e. increase of the number of stacked layers) with a concomitant decrease of stroma thylakoids is reported for the TGase over-expressers.

Holm oak (Quercus ilex L.) is native to hot, dry Mediterranean forests where limited water availability often reduces photosynthesis in many species, and forest fires are frequent. Holm oaks resprout after a disturbance, with improved photosynthetic activity and water relations compared with unburned plants. To better understand the role of water availability in this improvement, watering was withheld from container-grown plants, either intact (controls) or resprouts after excision of the shoot, to gradually obtain a wide range of soil water availabilities. At high water availability, gas exchange rates did not differ between controls and resprouts. At moderate soil dryness, net photosynthesis of control plants decreased as a result of increased stomatal limitation, whereas gas exchange rates of resprouts, which had higher midday and predawn leaf water potentials, were unchanged. Under severe drought, resprouts showed a less marked decline in gas exchange than controls and maintained photosystem II integrity, as indicated by chlorophyll fluorescence measurements. Photosynthesis was down-regulated in both plant types in response to reduced CO2 availability caused by high stomatal limitation. Lower non-stomatal limitations in resprouts than in control plants, as evidenced by higher carboxylation velocity and the capacity for ribulose-1,5-bisphosphate regeneration, conferred greater drought resistance under external constraints similar to summer conditions at midday.

Plants that resprout after fires often have higher rates of photosynthesis than before a fire. To elucidate the mechanism of this response, we studied gas exchange and chlorophyll fluorescence in Quercus ilex L. plants growing on control (unburned) sites and on sites that had been burned the preceding summer. In early July, photosynthetic rates and stomatal conductance were similar in plants on unburned and burned plots, and in young and old foliage within unburned plots. At this time, photochemical efficiency of photosystem II (PSII), nonphotochemical quenching of chlorophyll fluorescence (NPQ), and the de-epoxidation of violaxanthin to zeaxanthin were also similar among leaves of different ages and treatments. In late July, photosynthetic rates and stomatal conductances were much greater in resprouts on the burned areas than in unburned plants. From early to late July, unburned plants showed an increase in NPQ and the de-epoxidation of violaxanthin to zeaxanthin, indicating increased photoprotection as a result of enhanced nonradiative dissipation of excess light energy. Plants on the burned plots did not show these changes. Leaves of all ages and treatments showed no substantial reduction in potential quantum yield of PSII (F(v)/F(m)) at midday or predawn, indicating that there was little or no photoinhibition. Leaf nitrogen and soluble protein contents varied with leaf age during July, but did not vary between treatments. We conclude that the primary effect of burning is an increase in water availability to resprouting plants that eliminates the need for photoprotection, at least in the short term. The decrease in photosynthetic rates of unburned leaves in late July was the result of reduced stomatal conductance. We suggest that lowered stomatal conductance is the primary limiting factor in Q. ilex leaves, governing the regulation of carboxylation activity and energy dissipation processes.

Functional and morphological (structural) characteristics of Quercus ilex L. leaves under drought stress were studied in the forest and in a nursery. We compared undisturbed individuals (controls) with resprouts emerging after clear-cut or excision. When soil water availability was high, gas-exchange was similar in resprouts and controls, despite higher midday leaf water potential, midday leaf hydration and relative water content (RWC). In moderate drought, stomatal closure was found to limit photosynthesis in controls, and in severe drought non-stomatal limitations of photosynthesis were also greater than in resprouts. Leaf structure and chemical composition changed under drought stress. Leaves tended to be smaller in controls with increasing drought, and resprouts had larger leaves and lower leaf mass area (LMA). The relationship between nitrogen (N) content and LMA implied lower N investment in photosynthetic components in controls, which could be responsible for their increased non-stomatal limitation of photosynthesis. Changes were more apparent in leaf density (D) and thickness (T), components of LMA. Decreases in D were related to reductions in cell wall components: hemicellulose, cellulose and lignin. In resprouts, reduced D and leaf T accounted for the higher mesophyll conductance (gmes) to CO2 measured.

Here, we provide the first report on flavonoid content in holm oak (Quercus ilex L.) leaves, analyzed by HPLC-MS/MS. Flavanols and flavonols were the predominant groups, although proanthocyanidins and many soluble tannins had a relevant presence in all leaf samples. Seasonal variation of flavonoids was determined in extracts from Q. ilex leaves during resprouting after a forest fire in two Mediterranean forests. Similar seasonal trends were observed over 2 years during the two main stress seasons (winter and summer). The most abundant flavonoid was the flavanol epicatechin, which showed similar values during the two seasons. Hexosides of the flavonols, quercetin, kaempferol and rhamnetin showed considerably higher content in winter, especially at the lowest temperatures. These variations in both forests are discussed on the basis of the chlorophyll fluorescence results obtained. Anthocyanins were found practically absent in mature leaves. Nutrient or water availability differences between sites or seasons were not related to changes in leaf flavonolhexoside content.

We examined chloroplast pigment variation in holm oak (Quercus ilex L.) leaves for two periods under two climatic conditions, at midday during summer. We compared variation between control (unburned) plants and plants burned the preceding summer, since post-fire resprouts show higher photosynthetic rates and lower thermal energy dissipation. Principal component (PC) analysis was performed on nine pigment-content variables for the two periods separately. Two PC factors (PC1 and PC2) explained 83 and 84% of the variance of the data for each period. In both periods, PC1 was marked by positive loading of pigments associated with light absorption or structural function namely neoxanthin, lutein, β-carotene, chlorophyll a, and chlorophyll b. These pigments were only affected by leaf age. In contrast, PC2 was marked by high loadings of xanthophyll-cycle pigments (associated with photoprotection), and lutein-5,6-epoxide. Leaf content of these pigments was affected by climatic conditions. In the situations considered in PC analysis (leaf types, periods), the lutein-5,6-epoxide content presented a variation pattern similar to that of violaxanthin, and was significantly correlated with thermal dissipation of excess energy (represented by non-photochemical quenching or NPQ). These results suggest a relationship of lutein and lutein-5,6-epoxide with photoprotection.

Red (retro)-carotenoids accumulate in chloroplasts of Buxus sempervirens leaves during the process of winter leaf acclimation. As a result of their irregular presence, different leaf colour phenotypes can be found simultaneously in the same location. Five different colour phenotypes (green, brown, red, orange, and yellow), with a distinct pattern of pigment distribution and concentration, have been characterized. Leaf reddening due to the presence of anthocyanins or carotenoids, is a process frequently observed in plant species under photoinhibitory situations. Two main hypotheses have been proposed to explain the function of such colour change: antioxidative protection exerted by red-coloured molecules, and green light filtering. The potential photoprotective role of red (retro-) carotenoids as light filters was tested in Buxus sempervirens leaves. In shade leaves of this species the upper (adaxial) mesophyll of the lamina was replaced by the equivalent upper part of a different colour phenotype. These hybrid leaves were exposed to a photoinhibitory treatment in order to compare the photoprotective effect exerted by adaxial parts of phenotypes with a different proportion of red (retro)-carotenoids in the lower mesophyll of a shade leaf. The results indicated that the presence of red (retro)-carotenoids in the upper mesophyll did not increase photoprotection of the lower mesophyll when compared with chlorophyll, and the best protection was achieved by an upper green layer. This was due to the fact that the extent of photoinhibition was proportional to the amount of red light transmitted by the upper mesophyll and/or to the chlorophyll pool located above. These results do not exclude a protective function of carotenoids in the upper leaf layer, but imply that, at least under the conditions of this experiment, the accumulation of red pigments in the outer leaf layers does not increase photoprotection in the lower mesophyll.