How do Mediterranean shrub species cope with shade? Ecophysiological response to different light intensities.

Under natural conditions, light exposure for Mediterranean shrubs can be highly variable, especially during cloudy days or under a canopy, and can interfere with other environmental factors such as temperature and water availability. With the aim of decoupling the effect of radiation and temperature from water availability, we conducted an experiment where two perennial and three summer semi-deciduous shrub species were subjected to different levels of irradiation. In order to follow plant responses to light exposure, we measured gas exchange, photosystem II photochemical efficiency, photosynthetic pigments and leaf mass area in spring and summer. Results showed that all study species presented a plastic response to different light conditions, and that light-related traits varied in a coordinated manner. Summer semi-deciduous species exhibited a more opportunistic response, with higher photosynthesis rates in full sun, but under shade conditions, the two strategies presented similar assimilation rates. Stomatal conductance did not show such a drastic response as photosynthetsis, being related to changes in WUE. Daily cycles of Fv /Fm revealed a slight photoinhibitory response during summer, mainly in perennial species. In all cases photosynthetic pigments adjusted to the radiation level; leaves had lower chlorophyll content, higher pool of xanthophylls and higher proportion of the de-epoxydaded state of xanthophylls under sun conditions. Lutein content increased in relation to the xanthophyll pool under shade conditions. Our results evidenced that radiation is an important driving factor controlling morphological and physiological status of Mediterranean shrub species, independently of water availability. Summer semi-deciduous species exhibit a set of traits with higher response variability, maximising their photosynthetic assimilation under different sun conditions.

Changes in growth, physiological parameters and the hormonal status of Myrtus communis L. plants irrigated with water with different chemical compositions.

Myrtus communis, an important Mediterranean ornamental shrub, was used to study the effect of irrigation water with different chemical compositions in the plant response. A treatment with NaCl was used to establish the plant resistance to high salinity at long term. Plants were subjected to four irrigation treatments with drainage for three months: Control (0.8 dS m(-1)); two treatments using reclaimed water (RWs): RW1 (2.0 dS m(-1)) and RW2 (5.0 dS m(-1)); and NaCl (10.0 dS m(-1)). High levels of electric conductivity of RWs not affected plant growth, while NaCl decreased leaf dry weight. Coinciding with the accumulation of Na(+) and Cl(-) in the roots, soil water potential decreased, which hinders the mobilization of water to the leaves, decreasing leaf water potential. The osmotic adjustment in the NaCl treatment was due to Na(+) and Cl(-) ions, although the proline could contribute as an Osmo compatible solute, increasing the turgor plants. Also changes in cell walls rigidity minimize the negative effects on the water balance; however, a higher lipid peroxidation was observed in these plants. Stomatal closure was associated with a decrease in K(+) and an increase in abscisic acid. NaCl produced an increase in salicylic acid and did not affect jasmonic acid contents at the end of the experiment. Similar behavior in soil and leaf water potentials, although less pronounced than in NaCl, was shown in RW2 plants. The abscisic acid increased in the RW2 with respect to the control and a decrease in stomatal conductance was observed at the end of the experiment. Plants irrigated with RW1 behaved similarly to the control.

NaCl-induced physiological and biochemical adaptative mechanisms in the ornamental Myrtus communis L. plants.

Physiological and biochemical changes in Myrtus communis L. plants after being subjected to different solutions of NaCl (44, and 88 mM) for up to 30 days (Phase I) and after recovery from the salinity period (Phase II) were studied. Myrtle plants showed salinity tolerance by displaying a series of adaptative mechanisms to cope with salt-stress, including controlled ion homeostasis, the increase in root/shoot ratio, the reduction of water potentials and stomatal conductance to limit water loss. In addition, they displayed different strategies to protect the photosynthetic machinery, including limiting toxic ion accumulation in leaves, increase in chlorophyll content, and changes in chlorophyll fluorescence parameters, leaf anatomy and increases in catalase activity. Anatomical modifications in leaves, including a decrease in spongy parenchyma and increased intercellular spaces, allow CO2 diffusion in a situation of reduced stomatal aperture. In spite of all these changes, salinity produced oxidative stress in myrtle plants as monitored by increases in oxidative stress parameter values. The post-recovery period is perceived as a new stress situation, as observed through effects on plant growth and alterations in non-photochemical quenching parameters and lipid peroxidation values.

Characterization of the response of in vitro cultured Myrtus communis L. plants to high concentrations of NaCl.

Effect of salt stress was examined in in vitro shoot cultures of Myrtus communis L. a species of the Mediterranean maquis. To determine the effects of high salt concentrations on myrtle plantlets and contribute toward understanding the mechanisms adopted from this species to counteract soil salinity, in vitro rooted shoots were transferred to a liquid culture medium containing 0, 125 or 250 mM NaCl for 30 days. After 15 and 30 days of in vitro culture, shoot and root growth, chlorosis and necrosis extension, chlorophylls, carotenoids, proline, arginine, cysteine and total sugars content, as well as guaiacol peroxidase (G-POD, EC 1.11.1.7) and ascorbate peroxidase (APX, EC 1.11.1.11) activities were determined. In treated plants shoot and root growth, as well as chlorophyll content, significantly decreased, while carotenoids content was not affected by the NaCl treatment. Among osmolytes, proline did not significantly increase, arginine and cysteine decreased, while total sugars were found to be higher in the treated plants than in the control. Enhancement of G-POD and APX activities was positively related to increasing salt concentrations in the culture media, regardless of the exposure time. Salt-treated plants did not show significant changes in lipid peroxidation or DNA fragmentation after 30 days salt treatment, regardless of the NaCl concentrations applied. The results represent a contribution towards understanding the mechanisms adopted by this species to high salinity.

Scaling Erica arborea transpiration from trees up to the stand using auxiliary micrometeorological information in a wax myrtle-tree heath cloud forest (La Gomera, Canary Islands).

We investigate evapotranspiration, sap flow and top soil water content variations in a wax myrtle-tree heath ('fayal-brezal' in Spanish) cloud forest in the Garajonay National Park (La Gomera, Canary Islands) over a 1-year period. We provide transpiration estimates for one of the representative species, the shrubby needle-like Erica arborea L., present in this relict subtropical forest. An ad hoc tree up to the stand scaling method that combines the sap flow and auxiliary reference evapotranspiration data is illustrated, showing to be useful when sap flow in a limited number of trees has been monitored. Individual daily-based scaling curves of the Gompertz type were necessary to explain the observed sap flow variability in E. arborea during the 1-year period investigated (r(2) ≥ 0.953 with mode of r(2) = 0.9999). The mean daily sap flow of an E. arborea individual amounted to 8.37 ± 5.65 kg day(-1) tree(-1), with a maximum of 20.48 kg day(-1) tree(-1), yielding an annual total of 3052.89 kg tree(-1). A comparison of the computed daily transpiration with the continuous micrometeorological time series monitored in the studied plot suggested that solar radiation was the main driving force of transpiration in E. arborea (cross correlation index = 0.94). Fog may also affect tree transpiration via its reduction of radiation and temperature, such that during foggy periods the mean daily water loss estimate of E. arborea was 5.35 ± 4.30 kg day(-1) tree(-1), which sharply contrasted with the 2.4-fold average transpiration values obtained for fog-free days, i.e., 12.81 ± 4.33 kg day(-1) tree(-1). The annual water balance rendered a 288 mm year(-1) water input to the forest and evidenced the need for accurately quantifying the contribution of fog water dripping from the canopy.

Among-individual variation in pollen limitation and inbreeding depression in a mixed-mating shrub.

BACKGROUND AND AIMS: Variation in inbreeding depression (δ) among individual plants is considered to play a central role in mating system evolution and population genetics. Moreover, such variation could be linked to individual susceptibility to pollen limitation (PL) because those individuals strongly affected by δ for seed production will require more outcross pollen for setting a given number of fruits or seeds. However, no study has tested explicitly for associations between PL and δ at the individual plant level. This study assesses the extent of among-individual variation in PL and δ, the consistency of δ across life stages, and the relationships between individual PL and δ in the mixed-mating shrub Myrtus communis. METHODS: Controlled hand-pollinations were performed in a natural M. communis population. Marked flowers were monitored until fruit production and a greenhouse experiment was conducted with the seeds produced. KEY RESULTS: Compared with selfing, outcross-pollination enhanced seed number per fruit, germination rate and seedling growth, but did not enhance fruit-set. Only seed number per fruit was pollen limited and, thus, cumulative pollen limitation depended more on pollen quality (outcross pollen) than on quantity. The effects of δ varied considerably across life stages and individual plants. Cumulative δ was high across individuals (mean δ = 0·65), although there were no positive correlations between δ values at different life stages. Interestingly, maternal plants showing stronger δ for seed production were more pollen limited, but they were also less affected by δ for seedling growth because of a seed size/number trade-off. CONCLUSIONS: Results show a general inconsistency in δ across life stages and individuals, suggesting that different deleterious loci are acting at different stages. The association between δ and PL at the individual level corroborates the idea that pollen limitation may be 'genotype-dependent' regardless of other factors.

Seed trait changes in dispersers' guts and consequences for germination and seedling growth.

The effectiveness of a frugivore as a disperser of a plant is greatly determined by how fruits and seeds are handled in its mouth and its digestive tract. Although a number of studies have investigated the effect of avian ingestion on germination, we still know very little about the modifications to seeds during ingestion and the specific consequences on plant fitness. Here we investigate for the first time the different mechanisms by which germination patterns of seeds are modified following ingestion by frugivores. Specifically, we examine changes in seed mass, water content, permeability, seed coat thickness, texture, and resistance in two common Mediterranean fleshy-fruited plants, Phillyrea angustifolia and Myrtus communis, after ingestion by Eurasian Blackbirds, Turdus merula. We found a number of differences between the plant species: Phillyrea seeds lost mass, mainly due to water loss, and had thinner coats after gut passage, but Myrtus seeds did not. Seeds of both species showed increased permeability, while Myrtus seeds in particular became less resistant to breakage. No quantifiable changes in seed coat texture were detected in either species, although this trait was partly associated with differences in germination rate in Phillyrea. High intraspecific plant variation was found for most seed traits measured. Seed passage through birds' guts sped up germination in both species, especially in Myrtus. Increased permeability in seeds of both species following ingestion resulted in a higher germination rate. Moreover, seeds with thick coats (and in the case of Phillyrea, harder coats) germinated at a slower rate and produced seedlings that also grew more slowly, indicating a cost of coat thickness and/or hardness for seedling emergence. Results obtained here contribute to explaining the great heterogeneity in germination responses among and within plant species and the large variety of factors, both intrinsic and extrinsic to the plants, that influence such responses.

Morpho-anatomical, physiological and biochemical adjustments in response to root zone salinity stress and high solar radiation in two Mediterranean evergreen shrubs, Myrtus communis and Pistacia lentiscus.

Salt- and light-induced changes in morpho-anatomical, physiological and biochemical traits were analysed in Myrtus communis and Pistacia lentiscus with a view to explaining their ecological distribution in the Mediterranean basin. In plants exposed to 20 or 100% solar radiation and supplied with 0 or 200 mm NaCl, measurements were conducted for ionic and water relations and photosynthetic performance, leaf morpho-anatomical and optical properties and tissue-specific accumulation of tannins and flavonoids. Net carbon gain and photosystem II (PSII) efficiency decreased less in P. lentiscus than in M. communis when exposed to salinity stress, the former having a superior ability to use Na(+) and Cl(-) for osmotic adjustment. Morpho-anatomical traits also allowed P. lentiscus to protect sensitive targets in the leaf from the combined action of salinity stress and high solar radiation to a greater degree than M. communis. Salt and light-induced increases in carbon allocated to polyphenols, particularly to flavonoids, were greater in M. communis than in P. lentiscus, and appeared to be related to leaf oxidative damage. Our data may conclusively explain the negligible distribution of M. communis in open Mediterranean areas suffering from salinity stress, and suggest a key antioxidant function of flavonoids in response to different stressful conditions.