Constant ratio of Cc to Ci under various CO2 concentrations and light intensities, and during progressive drought, in seedlings of Japanese white birch.

Constant mesophyll conductance (gm), and two-resistance gm model (involved in resistances of cell wall and chloroplast), where gm reaches maximum under higher CO2 concentrations, cannot describe the phenomenon that gm decreases with increasing intercellular CO2 concentration (Ci) under relatively higher CO2 concentrations. Yin et al. (2020) proposed a gm model, according to which the ratio of chloroplastic CO2 concentration (Cc) to Ci is constant in the two-resistance gm model, which can describe the decreasing gm with increasing Ci. In the present study, we investigated the relationship between Cc and Ci in leaves of Japanese white birch by using simultaneous measurements of gas exchange and chlorophyll fluorescence under various CO2 concentrations, light intensities, and during progressive drought. Across the range of ambient CO2 from 50 to 1000 µmol mol-1, and light intensities of 50 to 2000 µmol m-2 s-1, measured under well irrigation, the ratio of Cc to Ci kept constant. During the progressive drought, overestimated Ci due to stomatal patchiness and/or cuticular transpiration was empirically corrected (threshold: stomatal conductance < 0.08 mol H2O m-2 s-1) from the A/Ci response measured under adequate irrigation. The ratio of Cc to Ci during progressive drought (predawn leaf potential reached ≈ - 2 MPa) also remained constant irrespective of soil drying rate in various pot sizes. The present study suggests the involvement of some physiologically regulative mechanisms to keep Cc:Ci ratio constant, which might act on gm in addition to the physical interaction of diffusive resistances in the cell components.

Molecular cloning and functional analysis of a UV-B photoreceptor gene, BpUVR8 (UV Resistance Locus 8), from birch and its role in ABA response.

As a photoreceptor specifically for UV-B light, UVR8 gene plays an important role in the photomorphogenesis and developmental growth of plants. In this research, we isolated the UVR8 gene from birch, named BpUVR8 (AHY02156). BpUVR8 overexpression rescued the uvr8 mutant phenotype using functional complementation assay of BpUVR8 in Arabidopsis uvr8 mutants, which showed that the function of UVR8 is conserved between Arabidopsis and birch. The expression analysis of BpUVR8 indicated that this gene is expressed in various tissues, but its expression levels in leaves are higher than in other organs. Moreover, abiotic stress factors, such as UV-B, salinity, and abscisic acid (ABA) can induce the expression of BpUVR8 gene. Interestingly, the analysis of promoter activity indicated that BpUVR8 promoter not only has the promoting activity but can also respond to the induction of abiotic stress and ABA signal. So, we analyzed its function in ABA response via transgenic UVR8 overexpression in Arabidopsis. The BpUVR8 enhances the susceptibility to ABA, which indicates that BpUVR8 is regulated by ABA and can inhibit seed germination. The root length of 20-day-old 35S::BpUVR8/WT transgenic plants was 18% reduced as compared to the wild-type under the ABA treatment. The membrane of the BpUVR8-overexpressing in Arabidopsis thaliana was the most damaged after ABA treatment and 35S::BpUVR8/WT transgenic plant was more sensitive to ABA than the wild type. These results showed that BpUVR8 is a positive regulator in the ABA signal transduction pathway. In the presence of low dose of UV-B, the sensitivity of wild-type and 35S::BpUVR8/WT plants to ABA was reduced. Moreover, BpUVR8 regulates the expression of a subset of ABA-responsive genes, both in Arabidopsis and Betula platyphylla, under the ABA treatment. Our data provide evidence that BpUVR8 is a positive regulator in the UV-B-induced photomorphogenesis in plants. Moreover, we propose from this research that BpUVR8 might have an important role in integrating plant growth and ABA signaling pathway.

Linking fine root morphology, hydraulic functioning and shade tolerance of trees.

Background and Aims: Understanding root traits and their trade-off with other plant processes is important for understanding plant functioning in natural ecosystems as well as agricultural systems. The aim of the present study was to determine the relationship between root morphology and the hydraulic characteristics of several orders of fine roots (<2 mm) for species differing in shade tolerance (low, moderate and high). Methods: The morphological, anatomical and hydraulic traits across five distal root orders were measured in species with different levels of shade tolerance and life history strategies. The species studied were Acer negundo, Acer rubrum, Acer saccharum, Betula alleghaniensis, Betula lenta, Quercus alba, Quercus rubra, Pinus strobus and Pinus virginiana. Key Results: Compared with shade-tolerant species, shade-intolerant species produced thinner absorptive roots with smaller xylem lumen diameters and underwent secondary development less frequently, suggesting that they had shorter life spans. Shade-tolerant species had greater root specific hydraulic conductance among these roots due to having larger diameter xylems, although these roots had a lower calculated critical tension for conduit collapse. In addition, shade-intolerant species exhibited greater variation in hydraulic conductance across different root growth rings in woody transport roots of the same root order as compared with shade-tolerant species. Conclusions: Plant growth strategies were extended to include root hydraulic properties. It was found that shade intolerance in trees was associated with conservative root hydraulics but greater plasticity in number of xylem conduits and hydraulic conductance. Root traits of shade-intolerant species were consistent with the ability to proliferate roots quickly for rapid water uptake needed to support rapid shoot growth, while minimizing risk in uncertain environments.

Short-term effects of light quality on leaf gas exchange and hydraulic properties of silver birch (Betula pendula).

Leaves have to acclimatize to heterogeneous radiation fields inside forest canopies in order to efficiently exploit diverse light conditions. Short-term effects of light quality on photosynthetic gas exchange, leaf water use and hydraulic traits were studied on Betula pendula Roth shoots cut from upper and lower thirds of the canopy of 39- to 35-year-old trees growing in natural forest stand, and illuminated with white, red or blue light in the laboratory. Photosynthetic machinery of the leaves developed in different spectral conditions acclimated differently with respect to incident light spectrum: the stimulating effect of complete visible spectrum (white light) on net photosynthesis is more pronounced in upper-canopy layers. Upper-canopy leaves exhibit less water saving behaviour, which may be beneficial for the fast-growing pioneer species on a daily basis. Lower-canopy leaves have lower stomatal conductance resulting in more efficient water use. Spectral gradients existing within natural forest stands represent signals for the fine-tuning of stomatal conductance and tree water relations to afford lavish water use in sun foliage and enhance leaf water-use efficiency in shade foliage sustaining greater hydraulic limitations. Higher sensitivity of hydraulic conductance of shade leaves to blue light probably contributes to the efficient use of short duration sunflecks by lower-canopy leaves.

Leaf traits and photosynthetic responses of Betula pendula saplings to a range of ground-level ozone concentrations at a range of nitrogen loads.

Ground-level ozone (O3) concentrations and atmospheric nitrogen (N) deposition rates have increased strongly since the 1950s. Rising ground-level O3 concentrations and atmospheric N deposition both affect plant physiology and growth, however, impacts have often been studied in isolation rather than in combination. In addition, studies are often limited to a control treatment and one or two elevated levels of ozone and/or nitrogen supply. In the current study, three-year old Betula pendula saplings were exposed to seven different O3 profiles (24h mean O3 concentration of 36-68ppb in 2013, with peaks up to an average of 105ppb) in precision-controlled hemispherical glasshouses (solardomes) and four different N loads (10, 30, 50 or 70kgNha-1y-1) in 2012 and 2013. Here we report on the effects of enhanced O3 concentrations and N load on leaf traits and gas exchange in leaves of varying age and developmental stage in 2013. The response of leaf traits to O3 (but not N) vary with leaf developmental stage. For example, elevated O3 did not affect the chlorophyll content of the youngest fully expanded leaf, but it reduced the chlorophyll content and photosynthetic parameters in aging leaves, relatively more so later than earlier in the growing season. Elevated O3 enhanced the N content of senesced leaves prior to leaf fall, potentially affecting subsequent N cycling in the soil. Enhanced N generally stimulated the chlorophyll content and photosynthetic capacity. Whilst elevated O3 reduced the light-saturated rate of photosynthesis (Asat) in aging leaves, it did not affect stomatal conductance (gs). This suggests that photosynthesis and gs are not closely coupled at elevated O3 under-light saturating conditions. We did not observe any interactions between O3 and N regarding photosynthetic parameters (Vc,max, Jmax, Asat), chlorophyll content, gs, N content in senesced leaves and leaf number. Hence, the sensitivity of these leaf traits to O3 in young silver birch trees is neither reduced nor enhanced by N load.

Photosynthetic limitation of several representative subalpine species in the Catalan Pyrenees in summer.

Information on the photosynthetic process and its limitations is essential in order to predict both the capacity of species to adapt to conditions associated with climate change and the likely changes in plant communities. Considering that high-mountain species are especially sensitive, three species representative of subalpine forests of the Central Catalan Pyrenees: mountain pine (Pinus uncinata Mill.), birch (Betula pendula Roth) and rhododendron (Rhododendron ferrugineum L.) were studied under conditions associated with climate change, such as low precipitation, elevated atmospheric [CO2 ] and high solar irradiation incident at Earth's surface, in order to detect any photosynthetic limitations. Short-term high [CO2 ] increased photosynthesis rates (A) and water use efficiency (WUE), especially in birch and mountain pine, whereas stomatal conductance (gs ) was not altered in either species. Birch showed photosynthesis limitation through stomatal closure related to low rainfall, which induced photoinhibition and early foliar senescence. Rhododendron was especially affected by high irradiance, showing early photosynthetic saturation in low light, highest chlorophyll content, lowest gas exchange rates and least photoprotection. Mountain pine had the highest A, photosynthetic capacity (Amax ) and light-saturated rates of net CO2 assimilation (Asat ), which were maintained under reduced precipitation. Furthermore, maximum quantum yield (Fv /Fm ), thermal energy dissipation, PRI and SIPI radiometric index, and ascorbate content indicated improved photoprotection with respect to the other two species. However, maximum velocity of carboxylation of RuBisco (Vcmax ) indicated that N availability would be the main photosynthetic limitation in this species.

Light compensation points in shade-grown seedlings of deciduous broadleaf tree species with different successional traits raised under elevated CO2.

We measured leaf photosynthetic traits in shade-grown seedlings of four tree species native to northern Japan, raised under an elevated CO2 condition, to investigate the effects of elevated CO2 on shade tolerance of deciduous broadleaf tree species with different successional traits. We considered Betula platyphylla var. japonica and Betula maximowicziana as pioneer species, Quercus mongolica var. crispula as a mid-successional species, and Acer mono as a climax species. The plants were grown under shade conditions (10% of full sunlight) in a CO2 -regulated phytotron. Light compensation points (LCPs) decreased in all tree species when grown under elevated CO2 (720 µmol·mol(-1) ), which were accompanied by higher apparent quantum yields but no photosynthetic down-regulation. LCPs in Q. mongolica and A. mono grown under elevated CO2 were lower than those in the two pioneer birch species. The LCP in Q. mongolica seedlings was not different from that of A. mono in each CO2 treatment. However, lower dark respiration rates were observed in A. mono than in Q. mongolica, suggesting higher shade tolerance in A. mono as a climax species in relation to carbon loss at night. Thus, elevated CO2 may have enhanced shade tolerance by lowering LCPs in all species, but the ranking of shade tolerance related to successional traits did not change among species under elevated CO2 , i.e. the highest shade tolerance was observed in the climax species (A. mono), followed by a gap-dependent species (Q. mongolica), while lower shade tolerance was observed in the pioneer species (B. platyphylla and B. maximowicziana).

Climate change alters leaf anatomy, but has no effects on volatile emissions from Arctic plants.

Biogenic volatile organic compound (BVOC) emissions are expected to change substantially because of the rapid advancement of climate change in the Arctic. BVOC emission changes can feed back both positively and negatively on climate warming. We investigated the effects of elevated temperature and shading on BVOC emissions from arctic plant species Empetrum hermaphroditum, Cassiope tetragona, Betula nana and Salix arctica. Measurements were performed in situ in long-term field experiments in subarctic and high Arctic using a dynamic enclosure system and collection of BVOCs into adsorbent cartridges analysed by gas chromatography-mass spectrometry. In order to assess whether the treatments had resulted in anatomical adaptations, we additionally examined leaf anatomy using light microscopy and scanning electron microscopy. Against expectations based on the known temperature and light-dependency of BVOC emissions, the emissions were barely affected by the treatments. In contrast, leaf anatomy of the studied plants was significantly altered in response to the treatments, and these responses appear to differ from species found at lower latitudes. We suggest that leaf anatomical acclimation may partially explain the lacking treatment effects on BVOC emissions at plant shoot-level. However, more studies are needed to unravel why BVOC emission responses in arctic plants differ from temperate species.

Insusceptibility of oxygen-evolving complex to high light in Betula platyphylla.

High mountain plants growing at high altitude have to regularly cope with high light and high UV radiation that can lead to photodamage of oxygen-evolving complex (OEC). However, the underlying mechanism of photoprotection for OEC in high mountain plants is unclear. Sun leaves of Betula platyphylla were used to examine whether cyclic electron flow (CEF) around photosystem I (PSI) plays an important role in photoprotection for OEC. Our results indicated that the value of ETRI/ETRII ratio significantly increased under high light. With increasing light intensity, non-photochemical quenching (NPQ) gradually increased, and the fraction of P700 that is oxidized in a given state gradually increased. These results indicated that CEF was significantly activated under high light. After treatment with a high light of 1600 µmol photons m(-2) s(-1) for 8 h, the OEC activity did not decline, but the maximum quantum yield of PSII (F v /F m ) ratio significantly decreased. These results suggested that CEF-dependent generation of proton gradient across thylakoid membrane protected OEC activity against high light. Furthermore, the stability of PSI activity during exposure to high light suggested that the high CEF activity in B. platyphylla played an important role in photoprotection for PSI activity.

How does solar ultraviolet-B radiation improve drought tolerance of silver birch (Betula pendula†Roth.) seedlings?

We hypothesized that solar ultraviolet (UV) radiation would protect silver birch seedlings from the detrimental effects of water stress through a coordinated suite of trait responses, including morphological acclimation, improved control of water loss through gas exchange and hydraulic sufficiency. To better understand how this synergetic interaction works, plants were grown in an experiment under nine treatment combinations attenuating ultraviolet-A and ultraviolet-B (UVB) from solar radiation together with differential watering to create water-deficit conditions. In seedlings under water deficit, UV attenuation reduced height growth, leaf production and leaf length compared with seedlings receiving the full spectrum of solar radiation, whereas the growth and morphology of well-watered seedlings was largely unaffected by UV attenuation. There was an interactive effect of the treatment combination on water relations, which was more apparent as a change in the water potential at which leaves wilted or plants died than through differences in gas exchange. This suggests that changes occur in the cell wall elastic modulus or accumulation of osmolites in cells under UVB. Overall, the strong negative effects of water deficit are partially ameliorated by solar UV radiation, whereas well-watered silver birch seedlings are slightly disadvantaged by the solar UV radiation they receive.

Can spatial data substitute temporal data in phenological modelling? A survey using birch flowering.

In addition to the evaluation of long-term series, the analysis of spatial gradients, such as urbanization gradients, may be helpful in assessing phenological responses to global warming. But are phenological responses of birch (Betula pendula Roth) assessed by temperature variations comparable over time and space and can spatially calibrated models predict long-term phenological data adequately? We calibrated and tested linear regression models and the process-based DORMPHOT model on phenological and temperature data sampled along an urbanization gradient in 2010 and 2011 in the German cities Munich and Ingolstadt (spatial data). Additionally, we analysed data from the German Meteorological Service for the period 1991-2010 (long-term data). The model comparison showed that the DORMPHOT model performed better than the linear model. Therefore, the importance of forcing and chilling sums as well as photoperiod, factors which were all considered in the DORMPHOT model, was evident. Models calibrated on spatial data produced good predictions of spatial data, but they were less adequate for predicting long-term data. Therefore, a time-for-space substitution might not always be appropriate. This finding was also confirmed by a comparison of temperature response rates. The rate of change in the spatial data (-4.4 days °C(-1)) did not match the changes observed in the long-term data (-1.9 days °C(-1)). Consequently, it is important not to generalize results derived from one specific study method, but their inherent methodological, spatial and temporal peculiarities have to be considered.

Combination treatment of elevated UVB radiation, CO2 and temperature has little effect on silver birch (Betula pendula) growth and phytochemistry.

Elevations of carbon dioxide, temperature and ultraviolet-B (UBV) radiation in the growth environment may have a high impact on the accumulation of carbon in plants, and the different factors may work in opposite directions or induce additive effects. To detect the changes in the growth and phytochemistry of silver birch (Betula pendula) seedlings, six genotypes were exposed to combinations of ambient or elevated levels of CO2 , temperature and UVB radiation in top-closed chambers for 7 weeks. The genotypes were relatively similar in their responses, and no significant interactive effects of three-level climate factors on the measured parameters were observed. Elevated UVB had no effect on growth, nor did it alter plant responses to CO2 and/or temperature in combined treatments. Growth in all plant parts increased under elevated CO2 , and height and stem biomass increased under elevated temperature. Increased carbon distribution to biomass did not reduce its allocation to phytochemicals: condensed tannins, most flavonols and phenolic acids accumulated under elevated CO2 and elevated UVB, but this effect disappeared under elevated temperature. Leaf nitrogen content decreased under elevated CO2 . We conclude that, as a result of high genetic variability in phytochemicals, B. pendula seedlings have potential to adapt to the tested environmental changes. The induction in protective flavonoids under UVB radiation together with the positive impact of elevated CO2 and temperature mitigates possible UVB stress effects, and thus atmospheric CO2 concentration and temperature are the climate change factors that will dictate the establishment and success of birch at higher altitudes in the future.

Oak loss increases foliar nitrogen, δ(15)N and growth rates of Betula lenta in a northern temperate deciduous forest.

Oak forests dominate much of the eastern USA, but their future is uncertain due to a number of threats and widespread failure of oak regeneration. A sudden loss of oaks (Quercus spp.) could be accompanied by major changes in forest nitrogen (N) cycles with important implications for plant nutrient uptake and tree species composition. In this study, we measured the changes in N use and growth rates of black birch trees (Betula lenta L.) following oak girdling at the Black Rock Forest in southeastern New York, USA. Data were collected from nine experimental plots composed of three treatments: 100% oaks girdled (OG), 50% oaks girdled (O50) and control (C). Foliar N concentration and foliar (15)N abundance increased significantly in the oak-girdled plots relative to the control, indicating that the loss of oaks significantly altered N cycling dynamics. As mineralization and nitrification rates increase following oak loss, black birch trees increase N absorption as indicated by higher foliar N content and increased growth rates. Foliar N concentration increased by 15.5% in the O50 and 30.6% in the OG plots relative to the control, while O50 and OG plots were enriched in (15)N by 1.08‰ and 3.33‰, respectively (P < 0.0001). A 641% increase in black birch growth rates in OG plots suggests that this species is able to respond to additional N availability and/or increased light availability. The loss of oaks and subsequent increase in black birch productivity may have a lasting impact on ecosystem form and function.

Twenty-two years of warming, fertilisation and shading of subarctic heath shrubs promote secondary growth and plasticity but not primary growth.

Most manipulation experiments simulating global change in tundra were short-term or did not measure plant growth directly. Here, we assessed the growth of three shrubs (Cassiope tetragona, Empetrum hermaphroditum and Betula nana) at a subarctic heath in Abisko (Northern Sweden) after 22 years of warming (passive greenhouses), fertilisation (nutrients addition) and shading (hessian fabric), and compare this to observations from the first decade of treatment. We assessed the growth rate of current-year leaves and apical stem (primary growth) and cambial growth (secondary growth), and integrated growth rates with morphological measurements and species coverage. Primary- and total growth of Cassiope and Empetrum were unaffected by manipulations, whereas growth was substantially reduced under fertilisation and shading (but not warming) for Betula. Overall, shrub height and length tended to increase under fertilisation and warming, whereas branching increased mostly in shaded Cassiope. Morphological changes were coupled to increased secondary growth under fertilisation. The species coverage showed a remarkable increase in graminoids in fertilised plots. Shrub response to fertilisation was positive in the short-term but changed over time, likely because of an increased competition with graminoids. More erected postures and large, canopies (requiring enhanced secondary growth for stem reinforcement) likely compensated for the increased light competition in Empetrum and Cassiope but did not avoid growth reduction in the shade intolerant Betula. The impact of warming and shading on shrub growth was more conservative. The lack of growth enhancement under warming suggests the absence of long-term acclimation for processes limiting biomass production. The lack of negative effects of shading on Cassiope was linked to morphological changes increasing the photosynthetic surface. Overall, tundra shrubs showed developmental plasticity over the longer term. However, such plasticity was associated clearly with growth rate trends only in fertilised plots.

The minimum temperature for budburst in Betula depends on the state of dormancy.

Vegis has put forward the theory that the range of growth-promoting temperatures changes during the induction and the release of dormancy. We have tested the response of buds of Betula pubescens Ehrh. and B. pendula Roth. on temperature during the induction and release of dormancy. Betula seedlings were exposed to dormancy-inducing high-temperature and short-day conditions and subsequently to dormancy-releasing chilling conditions in darkness. To monitor the dormancy status of the seedlings, subsets of them were transferred to five forcing temperatures and their budburst was observed. The results show that the expression of dormancy was temperature dependent, so that the minimum temperature for 100% budburst rose during the induction and dropped during the release of dormancy. These responses may explain previous contradictions between experimental and modelling studies, but that needs to be verified with more extensive experiments, some of which are identified in this study. The results provide further evidence for the concept of gradual change in bud dormancy. They also suggest that global change studies modelling budburst phenology should address the changing expression of bud dormancy.

Solar UV-B effects on PSII performance in Betula nana are influenced by PAR level and reduced by EDU: results of a 3-year experiment in the High Arctic.

The long-term and diurnal responses of photosystem II (PSII) performance to near-ambient UV-B radiation were investigated in High Arctic Betula nana. We conducted an UV exclusion experiment with five replicated blocks consisting of open control (no filter), photosynthetic active radiation and UV-B transparent filter control (Teflon), UV-B-absorbing filter (Mylar) and UV-AB-absorbing filter (Lexan). Ethylenediurea (EDU), a chemical normally used to protect plants against ozone injury, was sprayed on the leaves both in the field and in an additional laboratory study to investigate if EDU mitigated the effects of UV-B. Chlorophyll-a fluorescence induction curves were used for analysis of OJIP test parameters. Near-ambient UV-B radiation reduced across season maximum quantum yield (TR(o) /ABS = F(v) /F(m)), approximated number of active PSII reaction center (RC/ABS) and the performance index (PI(ABS)), despite improved leaf screening against UV-B with higher content of UV-B-absorbing compounds and a lower specific leaf area. EDU application counteracted the negative impact of UV-B on TR(o) /ABS, RC/ABS and PI(ABS) . This indicates that the mechanisms behind UV-B and ozone damage share some common features. The midday depression was present in all treatments, but TR(o) /ABS and PI(ABS) were persistently lower in near-ambient UV-B compared to UV-B reduction. The recovery phase was particularly impaired in near-ambient UV-B and interactive effects between treatment × hour raised TR(o) /ABS, RC/ABS and PI(ABS) higher in reduced UV-B compared to near-ambient UV-B. This demonstrates current solar UV-B to reduce the PSII performance both on a daily as well as a seasonal basis in this High Arctic species.

Species-specific effect of UV-B radiation on the temporal pattern of leaf growth.

Recent molecular and physiological studies have demonstrated that ultraviolet-B radiation (UV-B) can affect some of the processes involved in leaf growth, but the phases of leaf growth affected have not been clearly delimited. We used functional growth analysis to assess the effects of UV-B radiation on the time course of leaf growth in seedlings of two birch species (Betula pendula and Betula pubescens). Our aim was to identify the phase(s) of leaf development affected by UV-B radiation. In a greenhouse study, 1-year-old birch seedlings were subjected to three daily doses of supplemental UV-B radiation treatments (UV-B⁺) and no UV-B radiation controls (UV-B⁻). Leaf growth measurements every 2 days were complemented by assessment of other functional traits over a 4-week period at the start of the growing season. Using fitted curves, we were able to determine that the rate of leaf expansion was slowed by the UV-B⁺ treatment in leaves of B. pendula because of a slower maximum leaf growth rate compared with plants under the UV-B⁻ controls, but that compensation toward the end of the period of expansion negated this difference when leaves reached their final size. UV-B⁺ had little effect on the rate of B. pubescens leaf growth despite a larger reduction in leaf final size due to UV-B⁺ than occurred in B. pendula leaves. In conclusion, effective regulation ameliorated the effects of UV-B radiation on leaf and seedling growth in B. pendula, whereas in B. pubescens, reductions in leaf final size under UV-B⁺ were consistent with a slightly reduced rate of height growth.

Temporal variation in epidermal flavonoids due to altered solar UV radiation is moderated by the leaf position in Betula pendula.

The physiological mechanisms controlling plant responses to dynamic changes in ambient solar ultraviolet (UV) radiation are not fully understood: this information is important to further comprehend plant adaptation to their natural habitats. We used the fluorimeter Dualex to estimate in vivo the epidermal flavonoid contents by measuring epidermal UV absorbance (A(375) ) in Betula pendula Roth (silver birch) leaves of different ages under altered UV. Seedlings were grown in a greenhouse for 15 days without UV and transferred outdoors under three UV treatments (UV-0, UV-A and UV-A+B) created by three types of plastic film. After 7 and 13 days, Dualex measurements were taken at adaxial and abaxial epidermis of the first three leaves (L1, L2 and L3) of the seedlings. After 14 days, some of the seedlings were reciprocally swapped amongst the treatments to study the accumulation of epidermal flavonoids in the youngest unfolded leaves (L3) during leaf expansion under changing solar UV environments. A(375) of the leaves responded differently to the UV treatment depending on their position. UV-B increased the A(375) in the leaves independently of leaf position. L3 quickly adjusted A(375) in their epidermis according to the UV they received and these adjustments were affected by previous UV exposure. The initial absence of UV-A+B or UV-A, followed by exposure to UV-A+B, particularly enhanced leaf A(375) . Silver birch leaves modulate their protective pigments in response to changes in the UV environment during their expansion, and their previous UV exposure history affects the epidermal-absorbance achieved during later UV exposure.

Ultraviolet-B-induced flavonoid accumulation in Betula pendula leaves is dependent upon nitrate reductase-mediated nitric oxide signaling.

Nitric oxide (NO) is an important signaling molecule involved in many physiological processes in plants. Nitric oxide generation and flavonoid accumulation are two early reactions of plants to ultraviolet-B (UV-B) irradiation. However, the source of UV-B-triggered NO generation and the role of NO in UV-B-induced flavonoid accumulation are not fully understood. In order to evaluate the origin of UV-B-triggered NO generation, we examined the responses of nitrate reductase (NR) activity and the expression levels of NIA1 and NIA2 genes in leaves of Betula pendula Roth (silver birch) seedlings to UV-B irradiation. The data show that UV-B irradiation stimulates NR activity and induces up-regulation of NIA1 but does not affect NIA2 expression during UV-B-triggered NO generation. Pretreatment of the leaves with NR inhibitors tungstate (TUN) and glutamine (Gln) abolishes not only UV-B-triggered NR activities but also UV-B-induced NO generation. Furthermore, application of TUN and Gln suppresses UV-B-induced flavonoid production in the leaves and the suppression of NR inhibitors on UV-B-induced flavonoid production can be reversed by NO via its donor sodium nitroprusside. Together, the data indicate that NIA1 in the leaves of silver birch seedlings is sensitive to UV-B and the UV-B-induced up-regulation of NIA1 may lead to enhancement of NR activity. Furthermore, our results demonstrate that NR is involved in UV-B-triggered NO generation and NR-mediated NO generation is essential for UV-B-induced flavonoid accumulation in silver birch leaves.

Impact of light quality on leaf and shoot hydraulic properties: a case study in silver birch (Betula pendula).

Responses of leaf and shoot hydraulic conductance to light quality were examined on shoots of silver birch (Betula pendula), cut from lower ('shade position') and upper thirds of the crowns ('sun position') of trees growing in a natural temperate forest stand. Hydraulic conductances of leaf blades (K(lb) ), petioles (K(P) ) and branches (i.e. leafless stem; K(B) ) were determined using a high pressure flow meter in steady state mode. The shoots were exposed to photosynthetic photon flux density of 200-250 µmol m⁻² s⁻¹ using white, blue or red light. K(lb) depended significantly on both light quality and canopy position (P<0.001), K(B) on canopy position (P<0.001) and exposure time (P=0.014), and none of the three factors had effect on K(P) . The highest values of K(lb) were recorded under the blue light (3.63 and 3.13×10⁻4 kg m⁻² MPa⁻¹ s⁻¹ for the sun and shade leaves, respectively), intermediate values under white light (3.37 and 2.46×10⁻4 kg m⁻² MPa⁻¹ s⁻¹ , respectively) and lowest values under red light (2.83 and 2.02×10⁻4 kg m⁻² MPa⁻¹ s⁻¹, respectively). Light quality has an important impact on leaf hydraulic properties, independently of light intensity or of total light energy, and the specific light receptors involved in this response require identification. Given that natural canopy shade depletes blue and red light, K(lb) may be decreased both by reduced fluence and shifts in light spectra, indicating the need for studies of the natural heterogeneity of K(lb) within and under canopies, and its impacts on gas exchange.