The effect of elevated CO2 on photosynthesis is modulated by nitrogen supply and reduced water availability in Picea abies.

It is assumed that the stimulatory effects of elevated CO2 concentration ([CO2]) on photosynthesis and growth may be substantially reduced by co-occurring environmental factors and the length of CO2 treatment. Here, we present the study exploring the interactive effects of three manipulated factors ([CO2], nitrogen supply and water availability) on physiological (gas-exchange and chlorophyll fluorescence), morphological and stoichiometric traits of Norway spruce (Picea abies) saplings after 2 and 3 years of the treatment under natural field conditions. Such multifactorial studies, going beyond two-way interactions, have received only limited attention until now. Our findings imply a significant reduction of [CO2]-enhanced rate of CO2 assimilation under reduced water availability which deepens with the severity of water depletion. Similarly, insufficient nitrogen availability leads to a down-regulation of photosynthesis under elevated [CO2] being particularly associated with reduced carboxylation efficiency of the Rubisco enzyme. Such adjustments in the photosynthesis machinery result in the stimulation of water-use efficiency under elevated [CO2] only when it is combined with a high nitrogen supply and reduced water availability. These findings indicate limited effects of elevated [CO2] on carbon uptake in temperate coniferous forests when combined with naturally low nitrogen availability and intensifying droughts during the summer periods. Such interactions have to be incorporated into the mechanistic models predicting changes in terrestrial carbon sequestration and forest growth in the future.

The Possibility of Consensus Regarding Climate Change Adaptation Policies in Agriculture and Forestry among Stakeholder Groups in the Czech Republic.

As ongoing research efforts contribute to elucidating the consequences of climate change as well as adaptation and mitigation options, aligning the current research knowledge with stakeholder opinions and perceptions remains critical for adopting effective climate change policies. This paper utilizes an interactive survey to (1) address the aforementioned gap in studies involving three groups of stakeholders and opinion makers and (2) perform a comparative primary study of the climate change assumptions, risk perceptions, policy preferences, observations, and knowledge of Czech farmers, governmental policy-makers and researchers. This study shows that the stakeholder groups agree that the climate is clearly changing, attribute this change mostly to man-made causes and expect the negative effects to either prevail or be unevenly geographically distributed. The large majority of all three groups consider unmitigated climate change a major threat even by 2050 and agree that preparing in advance is the best sectoral strategy. Importantly, while investment in adaptation measures is considered the most efficient tool for accelerating the implementation of adaptation measures, the CAP and EU rules (as valid in 2016) are believed to hinder such measures. The results of this study have ramifications for the wider region of Central Europe.

Extreme warm temperatures alter forest phenology and productivity in Europe.

Recent climate warming has shifted the timing of spring and autumn vegetation phenological events in the temperate and boreal forest ecosystems of Europe. In many areas spring phenological events start earlier and autumn events switch between earlier and later onset. Consequently, the length of growing season in mid and high latitudes of European forest is extended. However, the lagged effects (i.e. the impact of a warm spring or autumn on the subsequent phenological events) on vegetation phenology and productivity are less explored. In this study, we have (1) characterised extreme warm spring and extreme warm autumn events in Europe during 2003-2011, and (2) investigated if direct impact on forest phenology and productivity due to a specific warm event translated to a lagged effect in subsequent phenological events. We found that warmer events in spring occurred extensively in high latitude Europe producing a significant earlier onset of greening (OG) in broadleaf deciduous forest (BLDF) and mixed forest (MF). However, this earlier OG did not show any significant lagged effects on autumnal senescence. Needleleaf evergreen forest (NLEF), BLDF and MF showed a significantly delayed end of senescence (EOS) as a result of extreme warm autumn events; and in the following year's spring phenological events, OG started significantly earlier. Extreme warm spring events directly led to significant (p=0.0189) increases in the productivity of BLDF. In order to have a complete understanding of ecosystems response to warm temperature during key phenological events, particularly autumn events, the lagged effect on the next growing season should be considered.

Does long-term cultivation of saplings under elevated CO2 concentration influence their photosynthetic response to temperature?

BACKGROUND AND AIMS: Plants growing under elevated atmospheric CO2 concentrations often have reduced stomatal conductance and subsequently increased leaf temperature. This study therefore tested the hypothesis that under long-term elevated CO2 the temperature optima of photosynthetic processes will shift towards higher temperatures and the thermostability of the photosynthetic apparatus will increase. METHODS: The hypothesis was tested for saplings of broadleaved Fagus sylvatica and coniferous Picea abies exposed for 4-5 years to either ambient (AC; 385 µmol mol(-1)) or elevated (EC; 700 µmol mol(-1)) CO2 concentrations. Temperature response curves of photosynthetic processes were determined by gas-exchange and chlorophyll fluorescence techniques. KEY RESULTS: Initial assumptions of reduced light-saturated stomatal conductance and increased leaf temperatures for EC plants were confirmed. Temperature response curves revealed stimulation of light-saturated rates of CO2 assimilation (Amax) and a decline in photorespiration (RL) as a result of EC within a wide temperature range. However, these effects were negligible or reduced at low and high temperatures. Higher temperature optima (Topt) of Amax, Rubisco carboxylation rates (VCmax) and RL were found for EC saplings compared with AC saplings. However, the shifts in Topt of Amax were instantaneous, and disappeared when measured at identical CO2 concentrations. Higher values of Topt at elevated CO2 were attributed particularly to reduced photorespiration and prevailing limitation of photosynthesis by ribulose-1,5-bisphosphate (RuBP) regeneration. Temperature response curves of fluorescence parameters suggested a negligible effect of EC on enhancement of thermostability of photosystem II photochemistry. CONCLUSIONS: Elevated CO2 instantaneously increases temperature optima of Amax due to reduced photorespiration and limitation of photosynthesis by RuBP regeneration. However, this increase disappears when plants are exposed to identical CO2 concentrations. In addition, increased heat-stress tolerance of primary photochemistry in plants grown at elevated CO2 is unlikely. The hypothesis that long-term cultivation at elevated CO2 leads to acclimation of photosynthesis to higher temperatures is therefore rejected. Nevertheless, incorporating acclimation mechanisms into models simulating carbon flux between the atmosphere and vegetation is necessary.

Morphological, biochemical and physiological traits of upper and lower canopy leaves of European beech tend to converge with increasing altitude.

The present work has explored for the first time acclimation of upper versus lower canopy leaves along an altitudinal gradient. We tested the hypothesis that restrictive climatic conditions associated with high altitudes reduce within-canopy variations of leaf traits. The investigated beech (Fagus sylvatica L.) forest is located on the southern slope of the Hrubý Jeseník Mountains (Czech Republic). All measurements were taken on leaves from upper and lower parts of the canopy of mature trees (>85 years old) growing at low (400 m above sea level, a.s.l.), middle (720 m a.s.l.) and high (1100 m a.s.l.) altitudes. Compared with trees at higher altitudes, those growing at low altitudes had lower stomatal conductance, slightly lower CO(2) assimilation rate (A(max)) and leaf mass per area (LMA), and higher photochemical reflectance index, water-use efficiency and Rubisco content. Given similar stand densities at all altitudes, the different growth conditions result in a more open canopy and higher penetration of light into lower canopy with increasing altitude. Even though strong vertical gradients in light intensity occurred across the canopy at all altitudes, lower canopy leaves at high altitudes tended to acquire the same morphological, biochemical and physiological traits as did upper leaves. While elevation had no significant effect on nitrogen (N) and carbon (C) contents per unit leaf area, LMA, or total content of chlorophylls and epidermal flavonoids in upper leaves, these increased significantly in lower leaves at higher altitudes. The increases in N content of lower leaves were coupled with similar changes in A(max). Moreover, a high N content coincided with high Rubisco concentrations in lower but not in upper canopy leaves. Our results show that the limiting role of light in lower parts of the canopy is reduced at high altitudes. A great capacity of trees to adjust the entire canopy is thus demonstrated.

Impact of elevated CO2 concentration on dynamics of leaf photosynthesis in Fagus sylvatica is modulated by sky conditions.

It has been suggested that atmospheric CO2 concentration and frequency of cloud cover will increase in future. It remains unclear, however, how elevated CO2 influences photosynthesis under complex clear versus cloudy sky conditions. Accordingly, diurnal changes in photosynthetic responses among beech trees grown at ambient (AC) and doubled (EC) CO2 concentrations were studied under contrasting sky conditions. EC stimulated the daily sum of fixed CO2 and light use efficiency under clear sky. Meanwhile, both these parameters were reduced under cloudy sky as compared with AC treatment. Reduction in photosynthesis rate under cloudy sky was particularly associated with EC-stimulated, xanthophyll-dependent thermal dissipation of absorbed light energy. Under clear sky, a pronounced afternoon depression of CO2 assimilation rate was found in sun-adapted leaves under EC compared with AC conditions. This was caused in particular by stomata closure mediated by vapour pressure deficit.

Effect of season, needle age and elevated CO2 concentration on photosynthesis and Rubisco acclimation in Picea abies.

While downward photosynthetic acclimation in response to elevated CO(2) (EC) is frequently accompanied by reduction in Rubisco (ribulose-1,5-bisphosphate carboxylase/oxygenase), the exact mechanism behind this decrease and its dynamics are not well understood. We comprehensively studied Rubisco adjustment to EC in coniferous Picea abies using an electrophoretic (protein content), spectrophotometric (initial (RA(initial)) and total (RA(total)) in vitro Rubisco activities), and gas-exchange (maximum carboxylation activity in vivo (V(Cmax))) techniques. With respect to differing carbon sink strength and nitrogen remobilization, we hypothesized greater acclimation of photosynthesis in one-year-old as compared to current-year needles and at the end than at the beginning of the vegetation season. EC treatment led to a decrease in V(Cmax) values in current-year needles, but the ribulose-1,5-bisphosphate (RuBP)-limited rate of photosynthesis (J(max)) remained unaffected. Indeed, both V(Cmax) and J(max) were reduced by the EC treatment in one-year-old needles. The extent of photosynthetic acclimation in EC plants did not increase, however, during the vegetation season. EC decreased the activation state of Rubisco (RA(initial)/RA(total)) by 16% and 5% in current-year and one-year-old needles, respectively (averaged over the growing season). While during spring (short-term effect) EC treatment did not influence the Rubisco content per unit leaf area and decreased its specific activity (activity per unit Rubisco mass) in both needle age classes studied, exposure to EC during the entire vegetation season tended to reduce the Rubisco content while increasing its specific activity. Irrespective of CO(2) treatment and needle age, a hyperbolic-decay relationship was observed between Rubisco-specific activity and its content.

The impact of long-term CO2 enrichment on sun and shade needles of Norway spruce (Picea abies): photosynthetic performance, needle anatomy and phenolics accumulation.

Norway spruce (Picea abies L. Karst) grown under ambient (365-377 µmol(CO(2)) mol(-1); AC) and elevated (700 µmol(CO(2)) mol(-1); EC) CO(2) concentrations within glass domes with automatically adjustable windows and on an open-air control site were studied after 8 years of treatment. The effect of EC on photosynthesis, mesophyll structure and phenolics accumulation in sun and shade needles was examined. Photosynthetic assimilation and dark respiration rates were measured gasometrically; the structural parameters of mesophyll were determined using confocal microscopy and stereological methods. The contents of total soluble phenolics and lignin were assessed spectrophotometrically, and localizations of different phenolic groups were detected histochemically on needle cross-sections. EC enhanced the light-saturated CO(2) assimilation rate and reduced dark respiration in the current-year needles. No effects of CO(2) enrichment on mesophyll structural parameters were observed. Similarly, the accumulation and localization of phenolics and lignin remained unaffected by EC treatment. Needles differentiated into sun and shade ecotypes in the same manner and to the same extent irrespective of CO(2) treatment. Based on these results, it is apparent that the EC-induced enhancement of photosynthesis is not related to changes in the examined structural parameters of mesophyll and accumulation of phenolic compounds.

Carbon exchange between ecosystems and atmosphere in the Czech Republic is affected by climate factors.

By comparing five ecosystem types in the Czech Republic over several years, we recorded the highest carbon sequestration potential in an evergreen Norway spruce forest (100%) and an agroecosystem (65%), followed by European beech forest (25%) and a wetland ecosystem (20%). Because of a massive ecosystem respiration, the final carbon gain of the grassland was negative. Climate was shown to be an important factor of carbon uptake by ecosystems: by varying the growing season length (a 22-d longer season in 2005 than in 2007 increased carbon sink by 13%) or by the effect of short- term synoptic situations (e.g. summer hot and dry days reduced net carbon storage by 58% relative to hot and wet days). Carbon uptake is strongly affected by the ontogeny and a production strategy which is demonstrated by the comparison of seasonal course of carbon uptake between coniferous (Norway spruce) and deciduous (European beech) stands.

Near-distance imaging spectroscopy investigating chlorophyll fluorescence and photosynthetic activity of grassland in the daily course.

Detection of grassland canopy chlorophyll fluorescence (Chl-F) conducted with an imaging spectroradiometer provided evidence of potential remote sensing estimation of steady-state Chl-F (Chl-Fs). Daily near-nadir views of extremely high spatial resolution hyperspectral images were acquired from a distance of 4 m for temperate montane grassland in the Czech Republic. Simultaneously, measurements of Chl-F and total chlorophyll content (Chla + b) were made on a single leaf at ground level were collected. A specifically designed 'shade removal' experiment revealed the influence of dynamic physiological plant processes on hyperspectral reflectance of three wavelengths: 532, 686 and 740 nm. Based on this information, the vegetation indexes R686/R630, R740/R800 and PRI calculated as (R532-R570)/(R532+R570) were tested for statistical significance with directly measured Chl-F parameters (maximum fluorescence yield, Fv/Fm; steady-state chlorophyll fluorescence, Chl-Fs and actual quantum yield, ФII). The grassland species under investigation were: Festuca rubra agg. (L.), Hieracium sp., Plantago sp., Nardus stricta (L.) and Jacea pseudophrygia (C.A. Meyer). The coefficients of determination (R2) for best-fit relationships between PRI-ФII and PRI-Chl-Fs, measured in the daily course, show a high variability of 0.23-0.78 and 0.20-0.65, respectively. Similarly, R2 for the R686/R630-ФII and R686/R630-Chl-Fs relationships varied between 0.20-0.73 and 0.41-0.70, respectively. The highest average R2 values were found between PRI and Chla + b (0.63) and R686/R630 and Chla + b (0.72). The ratio R740/R800 did not yield a statistically significant relation with Chl-F parameters.

Comparison of photosynthetic induction and transient limitations during the induction phase in young and mature leaves from three poplar clones.

We tested the hypothesis that leaf age affects photosynthetic induction, because conductance to CO2 diffusion usually decreases with increasing leaf age. Photosynthetic inductions, primarily determined by the light modulation of Rubisco activity and stomatal opening, were investigated in both young and mature leaves, as defined by leaf plastochron index (LPI), from three poplar clones: Populus alba L., P. nigra L. and P. x euramericana (Dode) Guinier. In all clones, maximum assimilation rates (A max), maximum stomatal conductance (G Smax) and dark respiration rates (RD) were higher in young leaves (LPI = 3-5) than in mature leaves (LPI = 10-14), and A max decreased from P. alba via P. x euramericana to P. nigra. The clones with high photosynthetic capacity had low induction states 60 s after leaf illumination (IS60; indicating a slow initial induction phase), and required less time to reach 90% photosynthetic induction (T90). In contrast, the clone with the lowest photosynthetic capacity (P. nigra) exhibited high IS60 (high initial induction state) but a long induction time (high T90). A comparison of mature leaves with young leaves revealed significantly (P < 0.01) lower IS60 values in mature leaves of P. nigra only, and significantly higher T90 values in mature leaves of P. alba only. In all clones, young leaves exhibited a lower percentage of maximum transient stomatal limitation during photosynthetic induction (4-9%) compared with mature leaves (16-30%). Transient biochemical limitation, assessed on the basis of the time constants of Rubisco activation (tau), was significantly higher in mature leaves than in young leaves of P. alba; whereas there were no significant differences in tau between young and mature leaves of the other poplar clones. Thus, our hypothesis that leaf age affects photosynthetic induction was confirmed at the level of transient stomatal limitation, which was significantly higher in mature leaves than in young leaves in all clones. For the induction parameters IS60, T90 and tau, photosynthetic induction was more clone-specific and was dependent on leaf age only in some cases, an observation that may apply to other tree species.

Seasonal variation in CO2 efflux of stems and branches of Norway spruce trees.

BACKGROUND AND AIMS: Stem and branch respiration, important components of total forest ecosystem respiration, were measured on Norway spruce (Picea abies) trees from May to October in four consecutive years in order (1) to evaluate the influence of temperature on woody tissue CO2 efflux with special focus on variation in Q10 (change in respiration rate resulting from a 10 degrees C increase in temperature) within and between seasons, and (2) to quantify the contribution of above-ground woody tissue (stem and branch) respiration to the carbon balance of the forest ecosystem. METHODS: Stem and branch CO2 efflux were measured, using an IRGA and a closed gas exchange system, 3-4 times per month on 22-year-old trees under natural conditions. Measurements of ecosystem CO2 fluxes were also determined during the whole experiment by using the eddy covariance system. Stem and branch temperatures were monitored at 10-min intervals during the whole experiment. KEY RESULTS: The temperature of the woody tissue of stems and branches explained up to 68% of their CO2 efflux. The mean annual Q10 values ranged from 2.20 to 2.32 for stems and from 2.03 to 2.25 for branches. The mean annual normalized respiration rate, R10, for stems and branches ranged from 1.71 to 2.12 micromol CO2 m(-2)s (-1) and from 0.24 to 0.31 micromol CO2 m(-2) s(-1), respectively. The annual contribution of stem and branch CO2 efflux to total ecosystem respiration were, respectively, 8.9 and 8.1% in 1999, 9.2 and 9.2% in 2000, 7.6 and 8.6% in 2001, and 8.6 and 7.9% in 2002. Standard deviation for both components ranged from 3 to 8% of the mean. CONCLUSIONS: Stem and branch CO2 efflux varied diurnally and seasonally, and were related to the temperature of the woody tissue and to growth. The proportion of CO2 efflux from stems and branches is a significant component of the total forest ecosystem respiration, approx. 8% over the 4 years, and predictive models must take their contribution into account.

Differences in pigment composition, photosynthetic rates and chlorophyll fluorescence images of sun and shade leaves of four tree species.

The differential pigment composition and photosynthetic activity of sun and shade leaves of deciduous (Acer pseudoplatanus, Fagus sylvatica, Tilia cordata) and coniferous (Abies alba) trees was comparatively determined by studying the photosynthetic rates via CO(2) measurements and also by imaging the Chl fluorescence decrease ratio (R(Fd)), which is an in vivo indicator of the net CO(2) assimilation rates. The thicker sun leaves and needles in all tree species were characterized by a lower specific leaf area, lower water content, higher total chlorophyll (Chl) a+b and total carotenoid (Cars) content per leaf area unit, as well as higher values for the ratio Chl a/b compared to the much thinner shade leaves and needles that possess a higher Chl a+b and Cars content on a dry matter basis and higher values for the weight ratio Chls/Cars. Sun leaves and needles exhibited higher rates of maximum net photosynthetic CO(2) assimilation (P(Nmax)) measured at saturating irradiance associated with higher maximum stomatal conductance for water vapor efflux. The differences in photosynthetic activity between sun and shade leaves and needles could also be sensed via imaging the Chl fluorescence decrease ratio R(Fd), since it linearly correlated to the P(Nmax) rates at saturating irradiance. Chl fluorescence imaging not only provided the possibility to screen the differences in P(N) rates between sun and shade leaves, but in addition permitted detection and quantification of the large gradients in photosynthetic rates across the leaf area existing in sun and shade leaves.

Induction of photosynthesis and importance of limitations during the induction phase in sun and shade leaves of five ecologically contrasting tree species from the temperate zone.

We examined the principal differences in photosynthetic characteristics between sun and shade foliage and determined the relative importance of biochemical and stomatal limitations during photosynthetic induction. Temperate-zone broadleaf and conifer tree species, ranging widely in shade tolerance, were investigated from one locality in the Czech Republic. The study species included strongly shade-tolerant Abies alba Mill. and Tilia cordata Mill., less shade-tolerant Fagus sylvatica L. and Acer pseudoplatanus L. and sun-demanding Picea abies (L.) Karst. In the fully activated photosynthetic state, sun foliage of all species had significantly higher maximum CO(2) assimilation rates, maximum stomatal conductance and maximum rates of carboxylation than shade foliage. Compared with shade leaves, sun leaves had significantly higher nocturnal stomatal conductances. In all species, shade foliage tended to have higher induction states 60 s after leaf illumination than sun foliage. Sun and shade foliage did not differ in the rate of disappearance of the transient biochemical limitation during the induction phase. Longer time periods were required to reach 90% photosynthetic induction and 90% stomatal induction in sun foliage than in shade foliage of the less shade-tolerant F. sylvatica and A. pseudoplatanus and in sun-demanding P. abies; however, in sun foliage of the strongly shade-tolerant species T. cordata and A. alba, the time needed for photosynthetic induction was similar to, or less than, that for shade foliage. Shade but not sun needles of P. abies and A. alba had significantly slower induction kinetics than the broadleaf tree species. Among species, the sun-demanding P. abies exhibited the shortest stomatal induction times in both sun and shade leaves. Independently of shade tolerance ranking, the transient stomatal and total limitations that characterize photosynthetic induction were relieved significantly earlier in shade foliage than in sun foliage. Sun foliage generally exhibited a hyperbolic photosynthetic induction response, whereas a sigmoidal induction response was more frequent in shade foliage. The different relative proportions of transient biochemical and stomatal limitations during photosynthetic induction in sun and shade foliage indicate an essential role of stomata in photosynthetic limitation during induction, mainly in shade foliage, with a consequent influence on the shape of the photosynthetic induction curve.