Drought effects on volatile organic compound emissions from Scots pine stems

Tree stems have been identified as sources of volatile organic compounds (VOCs) that play important roles in tree defence and atmospheric chemistry. Yet, we lack understanding on the magnitude and environmental drivers of stem VOC emissions in various forest ecosystems. Due to the increasing importance of extreme drought, we studied drought effects on the VOC emissions from mature Scots pine (Pinus sylvestris L.) stems. We measured monoterpenes, acetone, acetaldehyde and methanol emissions with custom-made stem chambers, online PTR-MS and adsorbent sampling in a drought-prone forest over the hot-dry summer of 2018 and compared the emission rates and dynamics between trees in naturally dry conditions and under long-term irrigation (drought release). The pine stems were significant monoterpene sources. The stem monoterpene emissions potentially originated from resin, based on their similar monoterpene spectra. The emission dynamics of all VOCs followed temperature at a daily scale, but monoterpene and acetaldehyde emission rates decreased nonlinearly with drought over the summer. Despite the dry conditions, large peaks of monoterpene, acetaldehyde and acetone emissions occurred in late summer potentially due to abiotic or biotic stressors. Our results highlight the potential importance of stem emissions in the ecosystem VOC budget, encouraging further studies in diverse environments.     

Effects of drought stress on the evergreen Quercus ilex L., the deciduous Q. robur L. and their hybrid Q. × turneri Willd.

Five-year-old trees of deciduous Quercus robur L., evergreen Q. ilex L., and their semideciduous hybrid, Q. × turneri Willd. (var. pseudoturneri), growing in pots, were subjected to drought stress by withholding water for 18–22 days, until leaf water potentials decreased below ?2 MPa. Gas-exchange rates, oxygen evolution, and modulated chlorophyll (Chl) fluorescence measurements revealed that by strong stomata closure and declining photosynthetic capacity down to approximately 50%, all three taxa responded with strongly reduced photosynthesis rates. In Q. robur, photochemical quenching of the drought-stressed plants was much lower than in nonstressed controls. Dissection of the occurring events in the photosynthetic electron transport chain by fast Chl fluorescence induction analysis with the JIP-test were discussed.     

Increased nitrogen deposition alleviated the adverse effects of drought stress on <Emphasis Type="Italic">Quercus variabilis</Emphasis> and <Emphasis Type="Italic">Quercus mongolica</Emphasis> seedlings

The plasticity response of Quercus variabilis and Quercus mongolica seedlings to combined nitrogen (N) deposition and drought stress was evaluated, and their performance in natural niche overlaps was predicted. Seedlings in a greenhouse were exposed to four N deposition levels (0, 4, 8, and 20 g N m^?2 year^?1) and two water levels (80 and 50 % field-water capacity). Plant traits associated with growth, biomass production, leaf physiology, and morphology were determined. Results showed that drought stress inhibited seedling performance, altered leaf morphology, and decreased fluorescence parameters in both species. By contrast increased N supply had beneficial effects on the nutritional status and activity of the PSII complex. The two species showed similar responses to drought stress. Contrary to the effects in Q. mongolica, N deposition promoted leaf N concentration, PSII activity, leaf chlorophyll contents, and final growth of Q. variabilis under well-watered conditions. Thus, Q. variabilis was more sensitive to N deposition than Q. mongolica. However, excessive N supply (20 g N m^?2 year^?1) did not exert any positive effects on the two species. Among the observed plasticity of the plant traits, plant growth was the most plastic, and leaf morphology was the least plastic. Therefore, drought stress played a primary role at the whole-plant level, but N supply significantly alleviated the adverse effects of drought stress on plant physiology. A critical N deposition load around 20 g N m^?2 year^?1 may exist for oak seedlings, which may more adversely affect Q. variabilis than Q. mongolica.     

Do changes in flood pulse duration disturb soil carbon dioxide emissions in semi-arid floodplains?

In semi-arid floodplains the average times between floods have been cited to drive metabolic and biogeochemical responses during the subsequent flooding pulse. However, the interaction effects of flood pulse duration and the length of time between floods on the carbon budget are not well understood. Using field experiments, flood pulses—dry cycles were simulated (SF plots—short flood/dry cycles: 15 flood days + 7 dry + 15 flood and LF plots—long flood/dry cycles: 21 flood + 14 dry + 21 flood) in a semi-arid floodplain in Central Spain, in order to study the effects on soil CO₂ emissions. Differences on soil water content among SF, LF and control plots were statistically significant throughout the experiment (p < 0.01). Soil CO₂ emission rates during drying time were significantly related with the duration of previous flooding and inter-flooding intervals (R ² = 0.52–0.64, p = 0.03). During the first stage of desiccation, the high soil water content appears to limit aerobic metabolism. Soil respiration rates similar to those of control plots measurements occurred 1–2 weeks later. Then, soil respiration increased to a maximum rate which was delayed 5–8 weeks, as high soil water content limited microbial activity. While more than 7 days of inundation promoted denitrification, organic nutrients supplied by flood water increased 1% soil respiration during drying. Differences between SF and LF plots in soil CO₂ emissions only appeared after floodplain soil had been subjected to two consecutive flood-dry cycles; 70 days after the second inundation ended, CO₂ fluxes achieved similar values in all treatments. Daily soil CO₂ emission rates during the entire study period (117 days) were comparable, independently of the flood duration and the time between floods (75.76 ± 1.59 and 77.94 ± 0.45 mmol CO₂ m^?2 day^?1, in SF and LF, respectively). Flood disturbance affects site-specific microbial processes, but only during very short time periods. The mechanism by which soil microbial communities cope or adapt to new conditions needs to be reassessed in future research in order to determine the long-term effects of hydrological changes in the soil carbon balance of semi-arid floodplains.     

Arbuscular Mycorrhizal Fungi Alter Fractal Dimension Characteristics of Robinia pseudoacacia L. Seedlings Through Regulating Plant Growth,Leaf Water Status,Photosynthesis, and Nutrient Concentration Under Drought Stress

The influence of arbuscular mycorrhizal fungi (AMF), Funneliformis mosseae and Rhizophagus intraradices, on plant growth, leaf water status, chlorophyll concentration, photosynthesis, nutrient concentration, and fractal dimension (FD) characteristics of black locust (Robinia pseudoacacia L.) seedlings was studied in pot culture under well-watered, moderate drought stress, and severe drought stress treatments. Mycorrhizal seedlings had higher dry biomass, leaf relative water content (RWC), and water use efficiency (WUE) compared with non-mycorrhizal seedlings. Under all treatments, AMF colonization notably enhanced net photosynthetic rate, stomatal conductance, and transpiration rate, but decreased intercellular CO₂ concentration. Leaf chlorophyll a and total chlorophyll concentrations were higher in AM seedlings than those in non-AM seedlings although there was no significant difference between AMF species. AMF colonization improved leaf C, N, and P concentrations, but decreased C:N, C:P, and N:P ratios. Mycorrhizal seedlings had a larger FD value than non-mycorrhizal seedlings. The FD value was positively and significantly correlated to the plant growth parameters, photosynthesis, RWC, WUE, and nutrient concentration but negatively correlated to leaf/stem ratio, C:N and C:P ratios, and intercellular CO₂ concentration. We conclude that AMF lead to an improvement of growth performance of black locust seedlings under all growth conditions, including drought stress via improving leaf water status, chlorophyll concentration, photosynthesis, and nutrient uptake. Moreover, FD technology proved to be a powerful non-destructive method to characterize the effect of AMF on the physiology of host plants during drought stress.     

The cultivation of oak seedlings inoculated with Tuber aestivum Vittad. in the boreal region of Finland

Despite recent findings, truffles are rarely found in Finland. In 2006, we began to explore the cultivation potential of Tuber aestivum/uncinatum in Finland. In 2006–2008, roughly 1,200 Quercus robur seedlings and 200 Q. pubescens seedlings were planted in 20 orchards. We aimed to challenge the Southern European (France) tree provenances of oak seedlings in a boreal climate. Additional winter coverings made up of fabric or plastic and twigs prevented the seedlings’ mortality even when the air temperature was below ?30 °C during the second winter. The results showed that the top soil temperature at 15 cm depth has to be above ?5 °C to guarantee the survival of seedlings. Q. pubescens was more sensitive to low soil temperatures than Q. robur. Morphological and PCR analysis of root samples collected over 2007–2010 confirmed the presence of T. aestivum in all orchards despite unfavorable temperatures during the winter time. The first T. aestivum sporocarps were found under Q. robur in October 2012 in the orchards established in 2006 on old agricultural land, showing truffle cultivation to be successful in the boreal climate.     

Characterization of photosystem I in rice (Oryza sativa L.) seedlings upon exposure to random positioning machine

To gain a better understanding of how photosynthesis is adapted under altered gravity forces, photosynthetic apparatus and its functioning were investigated in rice (Oryza sativa L.) seedlings grown in a random positioning machine (RPM). A decrease in fresh weight and dry weight was observed in rice seedlings grown under RPM condition. No significant changes were found in the chloroplast ultrastructure and total chlorophyll content between the RPM and control samples. Analyses of chlorophyll fluorescence and thermoluminescence demonstrate that PSII activity was unchanged under RPM condition. However, PSI activity decreased significantly under RPM condition. 77 K fluorescence emission spectra show a blue-shift and reduction of PSI fluorescence emission peak in the RPM seedlings. In addition, RPM caused a significant decrease in the amplitude of absorbance changes of P700 at 820 nm (A ₈₂₀) induced by saturated far-red light. Moreover, the PSI efficiency (Φ _I) decreased significantly under RPM condition. Immunoblot and blue native gel analyses further illustrate that accumulation of PSI proteins was greatly decreased in the RPM seedlings. Our results suggest that PSI, but not PSII, is down-regulated under RPM condition.     

Drought stress amelioration in wheat through inoculation with Burkholderia phytofirmans strain PsJN

Plant growth promoting endophytic bacteria Burkholderia phytofirmans PsJN was used to investigate the potential to ameliorate the effects of drought stress on growth, physiology and yield of wheat (Triticum aestivum L.) under natural field conditions. Inoculated and uninoculated (control) seeds of wheat cultivar Sahar 2006 was sown in the field. The plants were exposed to drought stress at different stages of growth (tillering stage and flowering stage) by skipping the respective irrigation. The results showed that drought stress adversely affected the physiological, biochemical and growth parameters of wheat seedlings. It decreased the CO₂ assimilation, stomatal conductance, relative water content, transpiration rate and chlorophyll contents in wheat. Inoculation of wheat with PsJN significantly diluted the adverse effects of drought on relative water contents and CO₂ assimilation rate thus improving the photosynthetic rate, water use efficiency and chlorophyll content over the uninoculated control. Grain yield was also decreased when plants were exposed to drought stress at the tillering and flowering stage, but inoculation resulted in better grain yield (up to 21 and 18 % higher, respectively) than the respective uninoculated control. Similarly, inoculation improved the ionic balance, antioxidant levels, and also increased the nitrogen, phosphorus, potassium and protein concentration in the grains of wheat. The results suggested that B. phytofirmans strain PsJN could be effectively used to improve the growth, physiology and quality of wheat under drought conditions.     

Effects of drought on photosynthesis,chlorophyll fluorescence and photoinhibition susceptibility in intact willow leaves

Plants from clonal cuttings of Salix sp. were subjected to a drying cycle of 10 d in a controlled environment. Gas exchange and fluorescence emission were measured on attached leaves. The light-saturated photosynthetic CO₂ uptake became progressively inhibited with decreased leaf water potential both at high, and especially, at low intercellular CO₂ pressure. The maximal quantum yield of CO₂ uptake was more resistant. The inhibition of light-saturated CO₂ uptake at leaf water potentials around-10 bar, measured at a natural ambient CO₂ concentration, was equally attributable to stomatal and non-stomatal factors, but the further inhibition below this water-stress level was caused solely by non-stomatal factors. The kinetics of fluorescence emission was changed at severe water stress; the slow secondary oscillations of the induction curve were attenuated, and this probably indicates perturbations in the carbon reduction cycle. The influence of light level during the drought period was also studied. Provided the leaves were properly light-acclimated, drought at high and low light levels produced essentially the same effects on photosynthesis. However, low-light-acclimated leaves became more susceptible to photoinhibitory treatment under severe water stress, as compared with well-watered conditions.     

Acclimation effects of heat waves and elevated [CO2] on gas exchange and chlorophyll fluorescence of northern red oak (Quercus rubra L.) seedlings

Heat wave frequency and intensity are predicted to increase. We investigated whether repeated exposure to heat waves would induce acclimation in Quercus rubra seedlings and considered [CO₂] as an interacting factor. We measured gas exchange and chlorophyll fluorescence of seedlings grown in 380 (C _A) or 700 (C _E) μmol CO₂ mol^?1, and three temperature treatments (ambient, ambient +3 °C, and an ambient +12 °C heat wave every fourth week). Measurements were performed during the third and fourth +12 °C heat waves (July and August 2010) at Whitehall Forest, GA, USA. Additionally, previously unexposed seedlings were subjected to the August heat wave to serve as a control to determine acclimation of seedlings which were previously exposed. Seedlings with a history of heat wave exposure showed lower net photosynthesis (A _net) and stomatal conductance (on average ?47 and ?38 %, respectively) than seedlings with no such history, when both were subjected to the same +12 °C heat wave. During both heat waves, A _net significantly declined in the +12 °C treatment compared with the other treatments. Additionally, the A _net decline during the August compared with the July heat wave was stronger in C _E than in C _A, suggesting that elevated [CO₂] might have had a negative effect on acclimation capacity. We conclude that seedlings subjected to consecutive heat waves will moderate stomatal conductance outside the heat wave, to reduce water usage at lower temperatures, increasing survival at the expense of carbon assimilation.     

Using common mycorrhizal networks for controlled inoculation of Quercus spp. with Tuber melanosporum: the nurse plant method

The high cost and restricted availability of black truffle spore inoculum for controlled mycorrhiza formation of host trees produced for truffle orchards worldwide encourage the search for more efficient and sustainable inoculation methods that can be applied globally. In this study, we evaluated the potential of the nurse plant method for the controlled inoculation of Quercus cerris and Quercus robur with Tuber melanosporum by mycorrhizal networks in pot cultures. Pine bark compost, adjusted to pH?7.8 by liming, was used as substrate for all assays. Initially, Q. robur seedlings were inoculated with truffle spores and cultured for 12 months. After this period, the plants presenting 74 % mycorrhizal fine roots were transferred to larger containers. Nurse plants were used for two treatments of two different nursling species: five sterilized acorns or five 45-day-old, axenically grown Q. robur or Q. cerris seedlings, planted in containers around the nurse plant. After 6 months, colonized nursling plant root tips showed that mycorrhiza formation by T. melanosporum was higher than 45 % in the seedlings tested, with the most successful nursling combination being Q. cerris seedlings, reaching 81 % colonization. Bulk identification of T. melanosporum mycorrhizae was based on morphological and anatomical features and confirmed by sequencing of the internal transcribed spacer region of the ribosomal DNA of selected root tips. Our results show that the nurse plant method yields attractive rates of mycorrhiza formation by the Périgord black truffle and suggest that establishing and maintaining common mycorrhizal networks in pot cultures enables sustained use of the initial spore inoculum.     

Effects of light quality on the chloroplastic ultrastructure and photosynthetic characteristics of cucumber seedlings

To understand how light quality influences plant photosynthesis, we investigated chloroplastic ultrastructure, chlorophyll fluorescence and photosynthetic parameters, Rubisco and chlorophyll content and photosynthesis-related genes expression in cucumber seedlings exposed to different light qualities: white, red, blue, yellow and green lights with the same photosynthetic photon flux density of 100 μmol m^?2 s^?1. The results revealed that plant growth, CO₂ assimilation rate and chlorophyll content were significantly reduced in the seedlings grown under red, blue, yellow and green lights as compared with those grown under white light, but each monochromatic light played its special role in regulating plant morphogenesis and photosynthesis. Seedling leaves were thickened and slightly curled; Rubisco biosynthesis, expression of the rca, rbcS and rbcL, the maximal photochemical efficiency of PSII (Fv/Fm) and quantum yield of PSII electron transport (Ф_PSII) were all increased in seedlings grown under blue light as compared with those grown under white light. Furthermore, the photosynthetic rate of seedlings grown under blue light was significantly increased, and leaf number and chlorophyll content of seedlings grown under red light were increased as compared with those exposed to other monochromatic lights. On the contrary, the seedlings grown under yellow and green lights were dwarf with the new leaves etiolated. Moreover, photosynthesis, Rubisco biosynthesis and relative gene expression were greatly decreased in seedlings grown under yellow and green light, but chloroplast structural features were less influenced. Interestingly, the Fv/Fm, Ф_PSII value and chlorophyll content of the seedlings grown under green light were much higher than those grown under yellow light.     

CO<Subscript>2</Subscript> enrichment affects eco-physiological growth of maize and alfalfa under different water stress regimes in the UAE

Water stress has been reported to alter morphology and physiology of plants affecting chlorophyll content, stomatal size and density. In this study, drought stress mitigating effects of CO₂ enrichment was assessed in greenhouse conditions in the hot climate of UAE. Commercially purchased maize (Zea mays L.) and alfalfa (Medicago sativa L.) were seeded in three different custom-built cage structures, inside a greenhouse. One cage was kept at 1000 ppm CO₂, the second at 700 ppm CO₂, and the third at ambient greenhouse CO₂ environment (i.e. 435 ppm). Three water stress treatments HWS (200 ml per week), MWS (400 ml per week), and CWS (600 ml per week) were given to each cage so that five maize pots and five alfalfa pots in each cage received same water stress treatments. In maize, total chlorophyll content was similar or higher in water stress treatments compared to control for all CO₂ concentrations. Stomatal lengths were higher in enriched CO₂ environments under water stress. At 700 ppm CO₂, stomatal widths decreased as water stress increased from MWS to HWS. At both enriched CO₂ environments, stomatal densities decreased compared to ambient CO₂ environment. In alfalfa, there was no significant increase in total chlorophyll content under enriched CO₂ environments, even though a slight increase was noticed.     

Interactive effects of elevated CO2, phosphorus deficiency, and soil drought on nodulation and nitrogenase activity in Alnus hirsuta and Alnus maximowiczii

We examined the interactive effects of elevated CO₂, soil phosphorus (P) availability, and soil drought on nodulation, nitrogenase activity, and biomass allocation in Alnus hirsuta and Alnus maximowiczii. Potted seedlings were grown in either ambient or elevated CO₂ (36 Pa and 72 Pa CO₂), with different levels of P (7.7 and 0.77 mgP pot^?1 week^?1 for high-P and low-P, respectively) and water supply in a natural daylight phytotron. Measurements of nitrogenase activity by an acetylene reduction assay failed to reveal significant effects of the treatments in any species. In high-P, nodule biomass increased under elevated CO₂ and decreased under drought. In low-P, nodule biomass decreased substantially compared to high-P, but the effect of elevated CO₂ on nodule biomass was unclear. Soil drought increased the partitioning of biomass into nodules, especially in A. hirsuta. These results suggest that with high P availability, elevated CO₂ could promote N₂ fixation by increasing nodule biomass even under drought. On the other hand, if soil P is limiting, elevated CO₂ may not enhance N₂ fixation because of the suppression of growth.     

Photosynthesis,stomatal conductance and terpene emission response to water availability in dry and mesic Mediterranean forests

Key message

Warmer summer conditions result in increased terpene emissions except under severe drought, in which case they strongly decrease.

Abstract

Water stress results in a reduction of the metabolism of plants and in a reorganization of their use of resources geared to survival. In the Mediterranean region, periods of drought accompanied by high temperatures and high irradiance occur in summer. Plants have developed various mechanisms to survive in these conditions by resisting, tolerating or preventing stress. We used three typical Mediterranean tree species in Israel, Pinus halepensis L., Quercus calliprinos and Quercus ithaburensis Webb, as models for studying some of these adaptive mechanisms. We measured their photosynthetic rates (A), stomatal conductance (g _s), and terpene emission rates during spring and summer in a geophysical gradient from extremely dry to mesic from Yatir (south, arid) to Birya (north, moist) with intermediate conditions in Solelim. A and g _s of P. halepensis were threefold higher in Birya than in Yatir where they remained very low both seasons. Quercus species presented 2–3-fold higher A and g _s but with much more variability between seasons, especially for Q. ithaburensis with A and g _s that decreased 10–30-fold from spring to summer. Terpene emission rates for pine were not different regionally in spring but they were 5–8-fold higher in Birya than in Yatir in summer (P < 0.05). Higher emissions were also observed in Solelim for the drought resistant Q. ithaburensis (P < 0.001) but not for Q. calliprinos. α-Pinene followed by limonene and 3-carene were the dominant terpenes. Warmer summer conditions result in increased Terpene emission rates except under severe drought, in which case they strongly decrease.

     

An analysis of the chlorophyll-fluorescence transients from pea leaves generated by changes in atmospheric concentrations of CO2 and O2

Removal of CO₂ from pea leaves, which were either grown in a glasshouse at 15°C or in a controlled environment at 23°C, produced an initial increase in chlorophyll fluorescence emission followed by a slow decrease to steady state. From estimations of the redox state of Q, using a nondestructive in vivo technique, the contributions of photochemical and nonphotochemical quenching processes to these fluorescence transients were determined. The fluorescence changes observed on removal of CO₂ from the two types of pea leaves were mainly attributable to changes in nonphotochemical quenching although markedly different changes in photochemical quenching were also observed. When leaves grown at 23°C were depleted of CO₂, Q immediately became more reduced, whereas in leaves grown at 15°C Q, unexpectedly, became more oxidised. On return of CO₂ to the leaves these phenomena were reversed, i.e., in leaves grown at 23°C Q became more oxidised and in the 15°C grown leaves Q became more reduced. Increased electron transport to O₂ may account for the oxidation of Q on depletion of CO₂ from 15°C grown leaves. The generation of fluorescence transients on removal and return of CO₂ to the leaf required the presence of oxygen. The fast fluorescence kinetics observed on exposure of the leaf at steady state to a second saturating irradiation suggest that O₂ may accept electrons directly from Photosystem II at a site between Q and B.     

5-Aminolevulinic acid enhances photosynthetic gas exchange, chlorophyll fluorescence and antioxidant system in oilseed rape under drought stress

This study evaluates the role of exogenous foliar application of 5-aminolevulinic acid (ALA) on water relations, gas exchange, chlorophyll fluorescence, and the activities and gene expression patterns of antioxidant enzymes in leaves of oilseed rape under drought stress and recovery conditions. Seedlings at four-leaf stage were imposed to well-watered condition (80 % of water-holding capacity) or drought stress (40 % of water-holding capacity) and subsequently foliar sprayed with water or ALA (30 mg l^?1). Drought suppressed the accumulation of plant biomass and decreased chlorophyll content and leaf water status (relative water content and water potential). The actual quantum yield of photosystem II and electron transport rates were hampered in parallel to net photosynthetic rate. However, drought stress induced the accumulation of malondialdehyde (MDA) and hydrogen peroxide, enhanced the activities of catalase (CAT), peroxidase (POD), ascorbate peroxidase (APX), glutathione reductase (GR) and superoxide dismutase and up-regulated the expression of APX and GR. After rehydration for 4 days, the growth of drought-treated seedlings was restored to normal level for most of the physiological parameters. Foliar application of ALA maintained relatively higher leaf water status and enhanced chlorophyll content, net photosynthetic rate, actual quantum yield of photosystem II, photochemical quenching, non-photochemical quenching and electron transport rates in stressed leaves. Exogenous ALA also alleviated the accumulation of MDA and hydrogen peroxide, increased the activities of antioxidant enzymes and enhanced the expression of CAT and POD in drought-treated plants. These results indicate that ALA may effectively protect rapeseed seedlings from damage induced by drought stress.     

Osmotic stress affects physiological responses and growth characteristics of three pistachio cultivars

Pistachio (Pistacia vera L.) has a high tolerance to drought and soil salinity. Although adult pistachio trees are well known to be drought tolerant, the studies on physiological adaptation of pistachio cultivars to drought are limited. Therefore, three pistachio cultivars, i.e., Akbari, Kaleghochi, and Ohadi were subjected to three osmotic drought stress treatments: control (?0.1 MPa), moderate (?0.75 MPa) and severe drought (?1.5 MPa) stress using PEG 6000 for a 14-day period. All drought stress treatments decreased net photosynthesis (P _n), stomatal conductance (g _s), intercellular CO₂ concentration (C _i), and transpiration rate (E), but Ohadi maintained better its photosynthetic capacity compared to Akbari and Kaleghochi. Maximum quantum yield of PSII photochemistry (F _v /F _m), effective PSII quantum yield (ΦPSII) and photochemical quenching (qP) were also reduced. The chlorophyll fluorescence parameters indicated that Akbari was more susceptible to the applied drought stress. Drought stress levels decreased chlorophyll pigments, fresh weight, stem elongation, leaf nitrogen content (N), leaf water potential and increased water use efficiency (WUE). Proline increased strongly under drought stress for Akbari. After 2 weeks of stress a recovery of 2 weeks was applied. This period was insufficient to fully restore the negative effects of the applied stress on the studied cultivars. Based on the reduction of photosynthesis and the increase of the proline content Akbari seems more sensitive to the applied drought stress.     

Chlorophyll a Fluorescence and Photosynthetic and Growth Responses of Pinus radiata to Phosphorus Deficiency, Drought Stress, and High CO(2)

Needles from phosphorus deficient seedlings of Pinus radiata D. Don grown for 8 weeks at either 330 or 660 microliters CO₂ per liter displayed chlorophyll a fluorescence induction kinetics characteristic of structural changes within the thylakoid chloroplast membrane, i.e. constant yield fluorescence (F_O) was increased and induced fluorescence ([F_P-F_I]/F_O) was reduced. The effect was greatest in the undroughted plants grown at 660 μl CO₂ L⁻¹. By week 22 at 330 μl CO₂ L⁻¹ acclimation to P deficiency had occurred as shown by the similarity in the fluorescence characteristics and maximum rates of photosynthesis of the needles from the two P treatments. However, acclimation did not occur in the plants grown at 660 μl CO₂ L⁻¹. The light saturated rate of photosynthesis of needles with adequate P was higher at 660 μl CO₂ L⁻¹ than at 330 μl CO₂ L⁻¹, whereas photosynthesis of P deficient plants showed no increase when grown at the higher CO₂ concentration. The average growth increase due to CO₂ enrichment was 14% in P deficient plants and 32% when P was adequate. In drought stressed plants grown at 330 μl CO₂ L⁻¹, there was a reduction in the maximal rate of quenching of fluorescence (R_Q) after the major peak. Constant yield fluorescence was unaffected but induced fluorescence was lower. These results indicate that electron flow subsequent to photosystem II was affected by drought stress. At 660 μl CO₂ L⁻¹ this response was eliminated showing that CO₂ enrichment improved the ability of the seedlings to acclimate to drought stress. The average growth increase with CO₂ enrichment was 37% in drought stressed plants and 19% in unstressed plants.     

Growth and carbon isotopes of Mediterranean trees reveal contrasting responses to increased carbon dioxide and drought

Forest dynamics will depend upon the physiological performance of individual tree species under more stressful conditions caused by climate change. In order to compare the idiosyncratic responses of Mediterranean tree species (Quercus faginea, Pinus nigra, Juniperus thurifera) coexisting in forests of central Spain, we evaluated the temporal changes in secondary growth (basal area increment; BAI) and intrinsic water-use efficiency (iWUE) during the last four decades, determined how coexisting species are responding to increases in atmospheric CO₂ concentrations (C _a) and drought stress, and assessed the relationship among iWUE and growth during climatically contrasting years. All species increased their iWUE (ca. +15 to +21 %) between the 1970s and the 2000s. This increase was positively related to C _a for J. thurifera and to higher C _a and drought for Q. faginea and P. nigra. During climatically favourable years the study species either increased or maintained their growth at rising iWUE, suggesting a higher CO₂ uptake. However, during unfavourable climatic years Q. faginea and especially P. nigra showed sharp declines in growth at enhanced iWUE, likely caused by a reduced stomatal conductance to save water under stressful dry conditions. In contrast, J. thurifera showed enhanced growth also during unfavourable years at increased iWUE, denoting a beneficial effect of C _a even under climatically harsh conditions. Our results reveal significant inter-specific differences in growth driven by alternative physiological responses to increasing drought stress. Thus, forest composition in the Mediterranean region might be altered due to contrasting capacities of coexisting tree species to withstand increasingly stressful conditions.