Warming and drought alter C and N concentration,allocation and accumulation in a Mediterranean shrubland

We investigated the effects of warming and drought on C and N concentrations, nitrogen use efficiency (NUE), and C and N accumulation in different ecosystem compartments. We conducted a 6-year (1999–2005) field experiment to simulate the climate conditions projected by IPCC models for the coming decades in a Mediterranean shrubland. We studied the two dominant species, Globularia alypum and Erica multiflora, and an N-fixing species, Dorycnium pentaphyllum, also abundant in this shrubland. Warming (1 °C) decreased N leaf concentrations by 25% and increased N stem concentrations by 40% in G. alypum. Although warming changed the available ammonium in soil in some seasons, it did not increase total soil N contents. Drought (19% average reduction in soil moisture) decreased leaf N concentrations in the two dominant shrub species, E. multiflora and G. alypum by 16% and 19%, respectively, and increased stem N concentrations by 56% and 40%, respectively. Neither warming nor drought changed the leaf N concentrations in the N-fixing species D. pentaphyllum, although warming increased stem N concentration by 9%. In G. alypum, the increase of stem N concentrations contributed to the observed increase of N accumulation in stem biomass in drought treatments with respect to control plots (8 kg N ha⁻¹). Neither warming nor drought changed NUE in the period 1999–2005. Warming increased soil organic C relative to drought. The effects of warming and drought on C and N concentrations, on N accumulation and on leaf/stem N distribution were not the result of dilution or concentration effects produced by changes in biomass accumulation. Other factors such as the changes in soil N availability, photosynthetic capacity, and plant internal C and N remobilization must be involved. These changes which differed depending on the species and the plant tissue show that the climate change projected for the coming decades will have significant effects on the C and N cycle and stoichiometry, with probable implications for ecosystem structure and function, such as changes in plant–herbivore relationships, decomposition rates or community species composition.     

Effects of long-term experimental night-time warming and drought on photosynthesis, <Emphasis Type="Italic">F</Emphasis>v/<Emphasis Type="Italic">F</Emphasis>m and stomatal conductance in the dominant species of a Mediterranean shrubland

We conducted a night-time warming and drought field experiment for 7 years (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.     

Effects of an Experimental Increase of Temperature and Drought on the Photosynthetic Performance of Two Ericaceous Shrub Species Along a North–South European Gradient

Plant ecophysiological changes in response to climatic change may be different in northern and southern European countries because different abiotic factors constrain plant physiological activity. We studied the effects of experimental warming and drought on the photosynthetic performance of two ericaceous shrubs (Erica multiflora and Calluna vulgaris) along a European gradient of temperature and precipitation (UK, Denmark, The Netherlands, and Spain). At each site, a passive warming treatment was applied during the night throughout the whole year, whereas the drought treatment excluded rain events over 6–10 weeks during the growing season. We measured leaf gas exchange, chlorophyll a fluorescence, and leaf carbon isotope ratio (¹³C) during the growing seasons of 1999 and 2000. Leaf net photosynthetic rates clearly followed a gradient from northern to southern countries in agreement with the geographical gradient in water availability. Accordingly, there was a strong correlation between net photosynthetic rates and the accumulated rainfall over the growing season. Droughted plants showed lower leaf gas exchange rates than control plants in the four sites. Interestingly, although leaf photosynthetic rates decreased along the precipitation gradient and in response to drought treatment, droughted plants were able to maintain higher leaf photosynthetic rates than control plants in relation to the accumulated rainfall over the months previous to the measurements. Droughted plants also showed higher values of potential photochemical efficiency (F _v/F _m) in relation to controls, mainly at midday. The warming treatment did not affect significantly any of the studied instantaneous ecophysiological variables..     

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. Published in Russian in Fiziologiya Rastenii, 2009, Vol. 56, No. 1, pp. 35–45. The text was submitted by the authors in English.     

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 (10 kg 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 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 7 years (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.     

Warming and drought alter soil phosphatase activity and soil P availability in a Mediterranean shrubland

We conducted a field experiment simulating the warming and drought in a Mediterranean shrubland dominated by Erica multiflora and Globularia alypum with the aim to simulate the next future climate conditions predicted by the IPCC and ecophysiological models. As P is frequently a limiting nutrient in Mediterranean ecosystems, we investigated the drought and warming effects on soil phosphatases activities, soil P contents and availability, litter and leaf P concentration, and the capacity of this community to maintain soil P reserves and retain this nutrient in the ecosystem. Warming treatment increased soil and air temperature (an average of 1°C) and drought treatment decreased soil water content in one of the seasons analysed (28% in autum 2004). Warming increased (68%) the activities of soil acid phosphatases in summer and alkaline phosphatase activity (22%) in spring 2004, and increased P concentrations in E. multiflora. Instead, warming decreased P concentrations in litterfall of this same species, E. multiflora, and soil HCO₃-extractable P_i (Olsen-P_i) in some seasons, decreasing total P soil concentration (37%) after 6 years of treatment. The drought treatment did not change soil phosphatase activities, nor available P_i. The effects of climate change on soil P dynamics in Mediterranean areas will thus be strongly dependent on whether the main variable involved in the local change is warming or drought. If warming is the main change without significant changes in water availability, the increases of biological activity can accelerate plant growth, P capture by plants and increase soil-phosphatase activity, altogether decreasing P contents in soil. If drought is the main change, a reduction in P demands by plants is expected, increasing P stocks in soils.     

Precipitation-dependent flowering of Globularia alypum and Erica multiflora in Mediterranean shrubland under experimental drought and warming, and its inter-annual variability

Background and Aims

Relationships 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.

Methods

Beginning 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°C) and reduced soil moisture (–17%) relative to untreated plots.

Key Results

Flowering 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.

Conclusions

Autumn 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.Key words: Globularia alypum, Erica multiflora, autumn flowering, drought, global warming, Mediterranean     

Inter-annual variability of seed rain and seedling establishment of two woody Mediterranean species under field-induced drought and warming

We aimed to assess the impact of warmer and drier climate change conditions on the seed rain and seedling establishment of Globularia alypum L. and Erica multiflora L., two dominant species in Western coastal Mediterranean shrublands. We performed a non-intrusive field experiment in which we increased the night-time temperatures and excluded spring and autumn rainfall. We monitored the seed rain over 5 years and the seedling recruitment over 9 years on these experimental plots. Seed rain of E. multiflora was enhanced by warming treatment in relation to control, and higher annual rainfall, while seed rain of G. alypum was increased by drought treatment in relation to control, dry years and higher minimum annual temperature. Annual rainfall enhanced the seedling emergence of both species, which also positively correlated with annual mean temperatures. Drought treatment significantly decreased seedling emergence for both species, which was higher in open areas than below vegetation cover. The seedling survival of both species diminished at closer distances to competing neighbours, and in G. alypum seedling survival was higher with lower annual mean temperatures and higher annual rainfall, but also in drought treatment, which have experienced vegetation cover decline. The study confirms that the increasing aridity in Mediterranean ecosystems would constrain the early stages of development in typical co-occurring shrubs. However, there are contrasting responses to climatic conditions between species recruitment, which might favour changes in vegetation through modification of species relative abundance.     

Drought advances spring growth phenology of the Mediterranean shrub Erica multiflora

Current climate projections predict drier and warmer conditions in the Mediterranean basin over the next century. While advanced spring growth due to warming has been described in the literature, few data are available on the effects of drought on phenology. Hence, the phenology and growth of two Mediterranean shrubs, Erica multiflora and Globularia alypum, was studied in a rainfall exclusion field experiment to simulate spring drought in a natural shrubland. We estimated the onset of growth in spring by monitoring the appearance of new stems, and the end of growth in summer by following the elongation of stems. Drought treatment caused earlier onset of the spring growing season in E. multiflora, whereas no advance was observed in G. alypum. However, growth cessation was not affected in E. multiflora. Drought reduced the growth of both shrubs, as reflected in less stem elongation. The results show that a drier climate might affect not only growth but also spring phenology of some Mediterranean species. We suggest that a reduction in the cooling effect of transpiration may have analogous effects to warming and might advance the start of growth in E. multiflora, a species whose phenology has been described as warming‐sensitive. The lengthening of the growing season resulting from advanced growth did not imply higher productivity, as growth was restricted by drought.     

Seasonal contrasting changes of foliar concentrations of terpenes and other volatile organic compound in four dominant species of a Mediterranean shrubland submitted to a field experimental drought and warming

To test the effect of forecasted drought and warming conditions for the next decades by GCM and ecophysiological models on foliar concentrations of volatile organic compounds (VOCs) and especially of volatile terpenes, we studied four typical Mediterranean woody plants (Pinus halepensis L., Pistacia lentiscus L., Rosmarinus officinalis L. and Globularia alypum L.) under a field experimental drought and warming generated using automatically sliding curtains. Terpenes were detected in the four studied species (R. officinalis L., P halepensis L., Pistacia lentiscus L. and G. alypum L.). In general, maximum concentrations of terpenes were found in the coldest periods and minimum concentrations in the summer. Their concentrations ranged between 0.003 mg g^?1 DM (eugenol) in G. alypum under drought conditions and 37 mg g^?1 DM in R. officinalis under control conditions. Main volatile terpenes found in all studied species except in G. alypum were α‐pinene, camphene, β‐pinene, β‐phellandrene and caryophyllene. In general, VOC leaf concentrations increased when soil moisture increased and decreased when air temperature increased. However, contrasting not consistent responses to the drought and warming treatments were found among species, seasons and years. For example, in P. halepensis, the concentrations decreased in response to drought in winter and instead increased in summer. Contrarily, drought decreased concentrations in summer and increased them in winter in Pistacia lentiscus. In any case, the data on seasonal VOC concentration in Mediterranean woody species provided here will add new knowledge of seasonal variation in essential oil contents of these species. These data might help in the study of flammability of Mediterranean ecosystems and in improving prediction algorithms, inventories and modelling of monoterpene emissions in response to climate change, which mostly do not consider the changes in concentration under drought stress. However, the lack of general and consistent response patterns to increasing drought and warming among species, seasons and years found here makes this task difficult.     

Physiological and antioxidant responses of <Emphasis Type="Italic">Erica multiflora</Emphasis> to drought and warming through different seasons

Climate change projections forecast a warming and an associated change in the distribution and intensity of rainfalls. In the case of the Mediterranean area, this will be reflected in more frequent and severe drought periods in the future. Within a long-term (9 years) manipulation experiment, we aimed to study the effect of the soil drought (15–20% decreased soil moisture) and warming conditions (+1°C warming) projected for the next decades onto photosynthetic rates and water relations, and onto the antioxidant and anti-stress defense capacity of Erica multiflora, a common species of the dry Mediterranean coastal scrublands, in two different seasons, spring and summer. Results indicated that none of the applied treatments was severe enough to induce a pronounced negative response of photosynthesis in this species well adapted to harsh Mediterranean conditions, but also highlighted important seasonal differences. Photosynthesis was limited by photoinhibition in spring and by stomatal closure in summer. Isoprenoid emission and the level of enzymatic and non-enzymatic antioxidants were lower in summer than in spring, whereas pigment and total phenolic contents were generally higher in summer. Volatile isoprenoid emissions were largely inhibited by drought and were not stimulated by warming. Drought and warming increased the oxidation state of ascorbate and reduced total glutathione levels in spring, but not in summer. It is concluded that E. multiflora plants can adapt to prolonged drought and warming, at least at the level simulated by our manipulative experiment, through changes in the seasonal physiological and antioxidant response of leaves.     

Effects of vegetation canopy and climate on seedling establishment in Mediterranean shrubland

Abstract. Question: Does the influence of plant canopy on seedling establishment interact with climate conditions, and particularly, do intensified drought conditions, enhance a positive effect of the vegetation canopy on seedlings in Mediterranean‐type ecosystems. Location: Mediterranean shrubland near Barcelona, Spain at 210 m a.s.l. Methods: Over the course of four years we recorded seedling emergence and survival in open areas and below vegetation under control, drier and warmer experimental climatic conditions. Results: Seedling emergence is more sensitive to climate conditions than later stages of growth. When considering the whole set of species, the total number of established seedlings at the end of the experiment was lower in the drought and warming stands than in control ones, and vegetation canopy increased the number of these seedlings in the drought stands. Drought reduced seedling emergence but not warming, while the interaction between climate treatments and vegetation canopy was not significant. Seedling survival was lower in the warming treatment than in the control. Under drought conditions, vegetation canopy increased seedling emergence of the dominant Globularia alypum. In control stands, vegetation canopy reduced their survival. Vegetation canopy increased the survival of the dominant Erica multiflora in warming stands, and it reduced the survival of G alypum in drought stands. No significant effects of drought and warming were observed in the seed rain of these two species. Conclusions: The balance of the facilitation‐competition interactions between vegetation canopy and seedling establishment in Mediterranean‐type ecosystems determined by water availability, and drought conditions enhance the positive effect of vegetation canopy. This interaction is species‐specific and shows important between‐year variability.     

Photosynthesis and growth reduction with warming are driven by nonstomatal limitations in a Mediterranean semi‐arid shrub

Whereas warming enhances plant nutrient status and photosynthesis in most terrestrial ecosystems, dryland vegetation is vulnerable to the likely increases in evapotranspiration and reductions in soil moisture caused by elevated temperatures. Any warming‐induced declines in plant primary production and cover in drylands would increase erosion, land degradation, and desertification. We conducted a four‐year manipulative experiment in a semi‐arid Mediterranean ecosystem to evaluate the impacts of a ~2°C warming on the photosynthesis, transpiration, leaf nutrient status, chlorophyll content, isotopic composition, biomass growth, and postsummer survival of the native shrub Helianthemum squamatum. We predicted that warmed plants would show reduced photosynthetic activity and growth, primarily due to the greater stomatal limitation imposed by faster and more severe soil drying under warming. On average, warming reduced net photosynthetic rates by 36% across the study period. Despite this strong response, warming did not affect stomatal conductance and transpiration. The reduction of peak photosynthetic rates with warming was more pronounced in a drought year than in years with near‐average rainfall (75% and 25–40% reductions relative to controls, respectively), with no indications of photosynthetic acclimation to warming through time. Warmed plants had lower leaf N and P contents, δ¹³C, and sparser and smaller leaves than control plants. Warming reduced shoot dry mass production by 31%. However, warmed plants were able to cope with large reductions in net photosynthesis, leaf area, and shoot biomass production without changes in postsummer survival rates. Our findings highlight the key role of nonstomatal factors (biochemical and/or nutritional) in reducing net carbon assimilation rates and growth under warming, which has important implications for projections of plant carbon balance under the warmer and drier climatic scenario predicted for drylands worldwide. Projected climate warming over the coming decades could reduce net primary production by about one‐third in semi‐arid gypsum shrublands dominated by H. squamatum.     

Comparative measurements of chlorophyll a fluorescence, acid accumulation and gas exchange in exposed and shaded plants of Clusia minor L. and Clusia multiflora H. B. K. in the field

 Changes in chlorophyll a fluorescence during the day and diurnal-changes of net CO₂-exchange and organic acid contents were determined in two species of the genus Clusia during the dry season in Venezuela. The investigations included plants of the C₃/CAM intermediate species Clusia minor and the C₃ species C. multiflora growing at exposed and shaded sites. Both species showed a C₃ pattern of net CO₂-exchange at the exposed site. In the shade under extreme drought stress C. minor showed a weak expression of CAM without CO₂-uptake during the afternoon (phase IV of CAM). C. multiflora growing in the shade exhibited a C₃-pattern of net CO₂-exchange and a small but significant nocturnal accumulation of citrate. Shaded plants of C. minor were able to double their light utilisation for electron transport and to reduce non-photochemical quenching during phase III compared to phase II of CAM. Furthermore, increase of electron transport rate through photosystem II in phase III of CAM is correlated to decarboxylation of malate. At the exposed site C. multiflora was less negatively affected by high PPFD than C. minor. This was shown by a lower reduction of potential electron quantum yield (F_v/F_m) and higher light utilisation of electron transport of C. multiflora compared to C. minor. At the exposed site C. minor did not make use of the CAM option to increase light utilisation of electron transport and to reduce non-photochemical quenching as did the plants growing in the shade. Received: 20 March 1996 / Accepted: 24 June 1996     

Shift in community structure in an early‐successional Mediterranean shrubland driven by long‐term experimental warming and drought and natural extreme droughts

Global warming and recurring drought are expected to accelerate water limitation for plant communities in semiarid Mediterranean ecosystems and produce directional shifts in structure and composition that are not easily detected, and supporting evidence is scarce. We conducted a long‐term (17 years) nocturnal‐warming (+0.6°C) and drought (?40% rainfall) experiments in an early‐successional Mediterranean shrubland to study the changes in community structure and composition, contrasting functional groups and dominant species, and the superimposed effects of natural extreme drought. Species richness decreased in both the warming and drought treatments. Responses to the moderate warming were associated with decreases in herb abundance, and responses to the drought were associated with decreases in both herb and shrub abundances. The drought also significantly decreased community diversity and evenness. Changes in abundance differed between herbs (decreases) and shrubs (increases or no changes). Both warming and drought, especially drought, increased the relative species richness and abundance of shrubs, favoring the establishment of shrubs. Both warming and drought produced significant shifts in plant community composition. Experimental warming shifted the community composition from Erica multiflora toward Rosmarinus officinalis, and drought consistently shifted the composition toward Globularia alypum. The responses in biodiversity (e.g., community biodiversity, changes of functional groups and compositional shifts) were also strongly correlated with atmospheric drought (SPEI) in winter–spring and/or summer, indicating sensitivity to water limitation in this early‐successional Mediterranean ecosystem, especially to natural extreme droughts. Our results suggest that the shifts in species assembles and community diversity and composition are accelerated by the long‐term nocturnal‐warming and drought, combined with natural severe droughts, and that the magnitude of the impacts of climate change is also correlated with the successional status of ecosystem. The results thus highlight the necessity for assessing the impacts on ecosystemic functioning and services and developing effective measures for conserving biodiversity.     

Effects of water stress and differential hardening treatments on photosynthetic characteristics of a xeromorphic shrub,Eucalyptus socialis,F. Muell.

Summary The influence of water stress on photosynthesis of drought hardened, and non-hardened,Eucalyptus socialis plants was examined. Particular attention was given to the effects of low leaf water potential on stomatal and intracellular resistance to CO₂ transport and on the CO₂ compensation point. Though the hardening treatment had a pronounced influence on leaf morphology, there was no apparent difference in the photosynthetic response to drought stress between hardened and non-hardened treatments, or with repeated drought cycles. These results suggest a high degree of genetic preconditioning to drought in this species.     

大气增温对滇西北高原典型湿地湖滨带优势植物的光和CO2利用能力的影响

大气增温对湿地植物光合作用的影响及其作用机制是近年来生态学界关注的热点。采用开顶式生长室(Open-top chambers,OTCs)模拟大气增温((2.0±0.5)℃,(3.5±0.5)℃),研究增温对滇西北高原典型湿地纳帕海湖滨带2种优势植物(茭草Zizania caduciflora,黑三棱Sparganium stoloniferum)的光和CO_2利用以及光合碳同化速率的影响。结果表明,(1)增温对不同植物的光和CO_2利用能力以及碳同化速率的影响存在种间差异。增温显著降低了茭草的光饱和点(LSP)、光补偿点(LCP)、光能利用幅(LSP-LCP)、CO_2饱和点(CSP)、CO_2利用幅(CSP-CCP)以及最大净光合速率(P_(max)),却显著增加了其CO_2补偿点(CCP);相反,增温显著提高了黑三棱的LSP、(LSP-LCP)以及最大净光合速率(P_(nmax)),而显著降低了其LCP,但对其CO_2利用参数无显著影响。(2)2种植物的光和CO_2利用能力对增温的响应存在季节性差异。不同温度处理下,茭草在6、8月的LSP和(LSP-LCP)均显著高于10月的对应值,其P_(nmax)也随生长期的增加而降低;黑三棱的光响应参数在不同生长季间无显著差异,其P_(nmax)在8月最高,而在6月和10月相对较低。(3)温度因子与茭草的光能利用参数均呈负相关,而与黑三棱的光能利用参数均呈正相关。本研究的温度因子中,日间积温对P_(nmax)和(LSP-LCP),年均温和最低温对(CSP-CCP)值的影响最为显著。本研究进一步表明,气候变暖对滇西北高原湿地湖滨带优势植物光合作用存在影响,但不同物种间存在差异响应,这种差异响应可能导致湿地生态系统植被结构的改变,其有关生态过程有待于进一步研究。     

Development of Water Stress under Increased Atmospheric CO2 Concentration

The increase in water use efficiency (the ratio of photosynthetic to transpiration rates) is likely to be the commonest positive effect of long-term elevation in CO₂ concentration (CE). This may not necessarily lead to decrease in long-term water use owing to increased leaf area. However, some plant species seem to cope better with drought stress under CE, because increased production of photosynthates might enhance osmotic adjustment and decreased stomatal conductance and transpiration rate under CE enable plants to maintain a higher leaf water potential during drought. In addition, at the same stomatal conductance, internal CO₂ concentration might be higher under CE which results in higher photosynthetic rate. Therefore plants under CE of the future atmosphere will probably survive eventual higher drought stress and some species may even be able to extend their biotope into less favourable sites. This revised version was published online in July 2006 with corrections to the Cover Date.     

Seasonal changes in net photosynthesis rates and photosynthetic capacity in leaves of Cistus salvifolius,a European mediterranean semi-deciduous shrub

Summary During five different periods between Nov. 1982 and Aug. 1983, the diurnal patterns exhibited in photosynthetic CO₂ uptake and stomatal conductance were observed under natural conditions on twigs of Cistus salvifolius, a Mediterranean semi-deciduous shrub which retains a significant proportion of its leaves through the summer drought. During the same periods, net photosynthesis at saturating CO₂ partial pressure was measured on the same twigs as a function of irradiance at different temperatures. From these data, photosynthetic capacity, defined here as the CO₂- and light-saturated net photosynthesis rate, was obtained as a function of leaf temperature. C. salvifolius is a winter growing species, shoot growth being initiated in Nov. and continuing through May. Photosynthetic capacity was quite high in Nov., March and June, exceeding 40 mol m^-2 s^-1 at optimum temperature. In Dec., photosynthetic capacity was somewhat reduced, perhaps due to low night-time temperatures (<5°C) during the measurement period. In Aug., capacity in oversummering shoots at optimum temperature fell to less than 8 mol m^-2 s^-1, due to water trees and perhaps leaf aging. Seasonal changes in maximal photosynthetic rates under ambient conditions were similar, and like those found in co-occurring evergreen sclerophylls. Like the evergreens, Cistus demonstrated considerable stomatal control of transpirational water loss, particularly in oversummering leaves. During each measurement period except Aug. when capacity was quite low, the maximum rates of net photosynthesis measured under ambient conditions were less than half the measured photosynthetic capacities at comparable temperatures, suggesting an apparent excess nitrogen investment in the photosynthetic apparatus.