Patricia Prieto

Drought and warming induced changes in P and K concentration and accumulation in plant biomass and soil in a Mediterranean shrubland

A field experiment involving drought and warming manipulation was conducted over a 6-year period in a Mediterranean shrubland to simulate the climate conditions projected by IPCC models for the coming decades (20% decreased soil moisture and 1°C warming). We investigated P and K concentration and accumulation in the leaves and stems of the dominant species, and in soil. Drought decreased P concentration in Globularia alypum leaves (21%) and in Erica multiflora stems (30%) and decreased K concentration in the leaves of both species (20% and 29%, respectively). The general decrease of P and K concentration in drought plots was due to the reduction of soil water content, soil and root phosphatase activity and photosynthetic capacity that decreased plant uptake capacity. Warming increased P concentration in Erica multiflora leaves (42%), but decreased it in the stems and leaf litter of Erica multiflora and the leaf litter (33%) of Globularia alypum, thereby demonstrating that warming improved the P retranslocation and allocation from stem to leaves. These results correlate with the increase in photosynthetic capacity and growth of these two dominant shrub species in warming plots. Drought and warming had no significant effects on biomass P accumulation in the period 1999–2005, but drought increased K accumulation in aboveground biomass (10xa0kg ha−1) in Globularia alypum due to the increase in K concentration in stems. The stoichiometric changes produced by the different responses of the nutrients led to changes in the P/K concentration ratio in Erica multiflora leaves, stems and litter, and in Globularia alypum stems and litter. This may have implications for the nutritional value of these plant species and plant–herbivore relationships. The effects of climate change on P and K concentrations and contents in Mediterranean ecosystems will differ depending on whether the main component of change is drought or warming.

Effects of long-term experimental night-time warming and drought on photosynthesis, Fv/Fm and stomatal conductance in the dominant species of a Mediterranean shrubland.

We conducted a night-time warming and drought field experiment for 7xa0years (1999–2005) in a Mediterranean shrubland. We focused on the two dominant shrub species, Erica multiflora L. and Globularia alypum L. and the tree Pinus halepensis L. and the final years to study the effects of the experimental night-time warming and drought on Fv/Fm, photosynthesis, and stomatal conductance. Warming treatment increased mean air temperature and mean soil temperature through the years by an average of 0.7 and 0.9°C respectively, and drought treatment reduced soil moisture through the years by an average of 19%. Warming tended to increase photosynthetic rates in E. multiflora, G. alypum and P. halepensis mostly in the cold seasons, when plants were more limited by temperature, as shown by the lowest values of Fv/Fm being detected in winter in the three studied species. A negative effect of warming was only detected for E. multiflora in summer 2003. Drought treatment generated different responses of net photosynthetic rates depending on the species, season and year. Stomatal conductance showed the same pattern as photosynthesis for the three studied species, displaying seasonal and inter-annual variability, although with an overall negative effect of drought for P. halepensis. Photosynthetic rates decreased significantly in the dry winter 2005 and spring 2005 in comparison to the same seasons of 2003 and 2004. There were positive correlations between the photosynthetic rates in different seasons for E. multiflora, G. alypum and P. halepensis and the soil moisture of the week prior to measurements. The great variation in the photosynthetic rates was thus explained in a significant part by soil moisture levels. The lowest Fv/Fm values usually corresponded with lowest stomatal conductances suggesting that drought stress could be associated to stress by low temperatures in winter.

Precipitation-dependent Flowering of Globularia alypum and Erica multiflora in Mediterranean Shrubland Under Experimental Drought and Warming, and its Inter-annual Variability

BACKGROUND AND AIMSnRelationships between autumn flowering, precipitation and temperature of plant species of Mediterranean coastal shrublands have been described, but not analysed experimentally. These relationships were analysed for two species of co-occurring, dominant, autumn-flowering shrubs, Globularia alypum and Erica multiflora, over 4 years and in experimentally generated drought and warming conditions. The aim was to improve predictions about the responses and adaptations of flowering of Mediterranean vegetation to climate change.nnnMETHODSnBeginning of anthesis and date of maximum flowering intensity (peak date) were monitored over 4 years (2001-2004) on a garrigue land type in the noth-east of the Iberian Peninsula. Two experimental treatments were applied, increased temperature (+0.73 degrees C) and reduced soil moisture (-17%) relative to untreated plots.nnnKEY RESULTSnFlowering of Globularia alypum and Erica multiflora differed greatly between years depending on the precipitation of the previous months and the date of the last substantial rainfall (>10 mm). Globularia alypum flowered once or twice (unimodal or bimodal) as the result of differences in the distribution and magnitude of precipitation in late-spring and summer (when floral buds develop). The drought treatment delayed and decreased flowering of Globularia alypum in 2001 and delayed flowering in 2002. Warming extended the period between the beginning of flowering and the end of the second peak for autumn flowering in 2001 and also increased peak intensity in 2002. Flowering of Erica multiflora was unaffected by either treatment.nnnCONCLUSIONSnAutumn flowering of Globularia alypum and Erica multiflora is more dependent on water availability than on temperature. Considerable inter-annual plasticity in the beginning of anthesis and peak date and on unimodal or bimodal flowering constitutes a safe strategy for both species in relation to varying precipitation and temperature. However, severe changes in precipitation in spring and summer may severely affect flowering of Globularia alypum but not Erica multiflora, thus affecting development/structure of the ecosystem if such conditions persist.

Effects of experimental warming and drought on biomass accumulation in a Mediterranean shrubland

We studied the effects of experimental warming and drought on the plant biomass of a Mediterranean shrubland. We monitored growth at plant level and biomass accumulation at stand level. The experimentation period stretched over 7xa0years (1999–2005) and we focused on the two dominant shrub species, Erica multiflora L. and Globularia alypum L. and the tree species Pinus halepensis L. The warming treatment increased shoot elongation in E. multiflora, and the drought treatment reduced shoot elongation in G. alypum. The elongation of P. halepensis remained unaffected under both treatments. The balance between the patterns observed in biomass accumulation for the three studied species in the drought plots (reduction in E. multiflora and P. halepensis and increase in G. alypum) resulted in a trend to reduce 33% the biomass of the drought treatment plots with respect to the untreated plots, which almost doubled their biomass from 1998 to 2005. The results also suggest that under drier conditions larger accumulation of dead biomass may occur at stand level, which combined with higher temperatures, may thus increase fire risk in the Mediterranean area.

Changes in Ca, Fe, Mg, Mo, Na, and S content in a Mediterranean shrubland under warming and drought

[1] In an evergreen Mediterranean shrubland we conducted a 6-year field experiment simulating the warming and drought projected by general circulation models and ecophysiological models for the next decades: 20% reduction of soil moisture and 1 Co f temperature increase. We tested whether warming and drought have effects on Ca, Fe, Mg, Mo, Na and S availability, concentrations and accumulation patterns in the three dominant plant species and in soil. Warming increased concentrations of Ca and Mo in leaves in Erica multiflora (42% and 65%, respectively) and in Dorycnium pentaphyllum (38% and 60%, respectively). Warming increased Mo accumulation in leaves and aboveground biomass in Globularia alypum (0.07 and 0.40 g ha � 1 ) and in E. multiflora (0.12 and 0.4 g ha � 1 ), and increased Fe accumulation in stem biomass of G. alypum (600 g ha � 1 ), increasing the capacity to retain these nutrients in the ecosystem. The increase of Fe and Mo capture capacity under warming conditions was greater in G. alypum than in E. multiflora coinciding with its greater increases in photosynthetic capacity. Warming decreased soil total-Fe concentration by 24% and increased Mg accumulation in soil exchange complex by 19%. Drought increased Na leaf and stem concentrations (93% and 50%, respectively) and accumulation in leaf and aboveground biomass (780 and 800 g ha � 1 , respectively) in G. alypum, allowing an increase of osmotic pressure which helps to prevent water losses and is related to its capacity to resist drought. Drought reduced S leaf and Mg leaf-litter concentrations of G. alypum and increased them in leaves of E. multiflora and also increased Mo and Na concentrations in leaves of D. pentaphyllum. Drought increased Fe soil solubility by 65%. The results indicate different effects of climate change on nutrient status in the ecosystem depending on whether the main change is warming or drought. The changes in concentration and biomass accumulation were different depending on the nutrient and the species, changing the stoichiometry among these nutrients and modifying the nutritional quality of plant tissues.

Net ecosystem exchange and whole plant isoprenoid emissions by a mediterranean shrubland exposed to experimental climate change

We tested the effect of forecasted soil drought and warming climate conditions for the next decades on emission rates of isoprenoids by mediterranean shrublands. We measured isoprenoid emissions by whole dominant mediterranean woody plants (Erica multiflora L. and Globularia alypum L.) inhabiting the studied shrublands. Monoterpene emissions were detected in both species, but isoprene was emitted only by E. multiflora. Maximum emission rates were found during the hottest periods (except for G. alypum, in which they occurred in autumn), and minimum emission rates in winter in E. multiflora. Terpene emission rates ranged from 0.08 μg/(g dry wt h) in winter in E. multiflora to 8.8 μg/(g dry wt h) in G. alypum in autumn. In E. multiflora, the terpene emission rates decreased in response to soil drought only in summer, but increased in response to warming in spring and autumn. Isoprene emissions ranged from 0.1 μg/(g dry wt h) in spring to 4.4 μg/(g dry wt h) in summer. The effect of the treatments was only detected in autumn when soil drought and warming had a negative effect on isoprene emission rates. These data might improve our knowledge of isoprenoid emissions at the canopy level and in response to climate change, soil drought, or warming.