Comparative sensitivity to gamma radiation at the organismal, cell and DNA level in young plants of Norway spruce, Scots pine and Arabidopsis thaliana.

MAIN CONCLUSION: Persistent DNA damage in gamma-exposed Norway spruce, Scots pine and Arabidopsis thaliana, but persistent adverse effects at the organismal and cellular level in the conifers only. Gamma radiation emitted from natural and anthropogenic sources may have strong negative impact on plants, especially at high dose rates. Although previous studies implied different sensitivity among species, information from comparative studies under standardized conditions is scarce. In this study, sensitivity to gamma radiation was compared in young seedlings of the conifers Scots pine and Norway spruce and the herbaceous Arabidopsis thaliana by exposure to 60Co gamma dose rates of 1-540 mGy h-1 for 144 h, as well as 360 h for A. thaliana. Consistent with slightly less prominent shoot apical meristem, in the conifers growth was significantly inhibited with increasing dose rate ≥ 40 mGy h-1. Post-irradiation, the conifers showed dose-rate-dependent inhibition of needle and root development consistent with increasingly disorganized apical meristems with increasing dose rate, visible damage and mortality after exposure to ≥ 40 mGy h-1. Regardless of gamma duration, A. thaliana showed no visible or histological damage or mortality, only delayed lateral root development after ≥ 100 mGy h-1 and slightly, but transiently delayed post-irradiation reproductive development after ≥ 400 mGy h-1. In all species dose-rate-dependent DNA damage occurred following ≥ 1-10 mGy h-1 and was still at a similar level at day 44 post-irradiation. In conclusion, the persistent DNA damage (possible genomic instability) following gamma exposure in all species may suggest that DNA repair is not necessarily mobilized more extensively in A. thaliana than in Norway spruce and Scots pine, and the far higher sensitivity at the organismal and cellular level in the conifers indicates lower tolerance to DNA damage than in A. thaliana.

Transcriptome analysis of shade avoidance and shade tolerance in conifers.

MAIN CONCLUSION: Gymnosperms respond differently to light intensity and R:FR; although some aspects of shade response appear conserved, yet underlying mechanisms seem to be diverse in gymnosperms as compared to angiosperms. Shade avoidance syndrome (SAS) is well-characterized in the shade intolerant model species Arabidopsis thaliana whereas much less is known about shade tolerance response (STR), yet regulation of SAS and STR with reference to conifers remains poorly understood. We conducted a comparative study of two conifer species with contrasting responses to shade, Scots pine (shade-intolerant) and Norway spruce (shade-tolerant), with the aim to understand mechanisms behind SAS and STR in conifers. Pine and spruce seedlings were grown under controlled light and shade conditions, and hypocotyl and seedling elongation following different light treatments were determined in both species as indicators of shade responses. Red to far-red light ratio (R:FR) was shown to trigger the shade response in Norway spruce. In Scots pine, we observed an interaction between R:FR and light intensity. RNA sequencing (RNA-Seq) data revealed that SAS and STR responses included changes in expression of genes involved primarily in hormone signalling and pigment biosynthesis. From the RNA-Seq analysis, we propose that although some aspects of shade response appear to be conserved in angiosperms and gymnosperms, yet the underlying mechanisms may be different in gymnosperms that warrants further research.

Light-induced gradual activation of photosystem II in dark-grown Norway spruce seedlings.

Gymnosperms, unlike angiosperms, are able to synthesize chlorophyll and form photosystems in complete darkness. Photosystem I (PSI) formed under such conditions is fully active, but photosystem II (PSII) is present in its latent form with inactive oxygen evolving complex (OEC). In this work we have studied light-induced gradual changes in PSII function in dark-grown cotyledons of Norway spruce (Picea abies) via the measurement of chlorophyll a fluorescence rise, absorption changes at 830 nm, thermoluminescence glow curves (TL) and protein analysis. The results indicate that in dark-grown cotyledons, alternative reductants were able to act as electron donors to PSII with inactive OEC. Illumination of cotyledons for 5 min led to partial activation of PSII, which was accompanied by detectable oxygen evolution, but still a substantial number of PSII centers remained in the so called PSII-Q(B)-non-reducing form. Interestingly, even 24 h long illumination was not sufficient for the full activation of PSII centers. This was evidenced by a weak attachment of PsbP protein and the absence of PsbQ protein in PSII particles, the absence of PSII supercomplexes, the suboptimal maximum yield of PSII photochemistry, the presence of C band in TL curve and also the presence of up-shifted Q band in TL in DCMU-treated cotyledons. This slow light-induced activation of PSII in dark-grown cotyledons could contribute to the prevention of PSII overexcitation before the light-induced increase in PSI/PSII ratio allows effective operation of linear electron flow.

Seasonal and within-canopy variation in shoot-scale resource-use efficiency trade-offs in a Norway spruce stand.

Previous leaf-scale studies of carbon assimilation describe short-term resource-use efficiency (RUE) trade-offs where high use efficiency of one resource requires low RUE of another. However, varying resource availabilities may cause long-term RUE trade-offs to differ from the short-term patterns. This may have important implications for understanding canopy-scale resource use and allocation. We used continuous gas exchange measurements collected at five levels within a Norway spruce, Picea abies (L.) karst., canopy over 3 years to assess seasonal differences in the interactions between shoot-scale resource availability (light, water and nitrogen), net photosynthesis (An ) and the use efficiencies of light (LUE), water (WUE) and nitrogen (NUE) for carbon assimilation. The continuous data set was used to develop and evaluate multiple regression models for predicting monthly shoot-scale An . These models showed that shoot-scale An was strongly dependent on light availability and was generally well described with simple one- or two-parameter models. WUE peaked in spring, NUE in summer and LUE in autumn. However, the relative importance of LUE for carbon assimilation increased with canopy depth at all times. Our results suggest that accounting for seasonal and within-canopy trade-offs may be important for RUE-based modelling of canopy carbon uptake.

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.

Seedling ontogeny and environmental plasticity in two co-occurring shade-tolerant conifers and implications for environment-population interactions.

PREMISE OF THE STUDY: Seedling success is determined by evolved strategies of intrinsic genetic programming and plasticity that are regulated by extrinsic pathways. We tested the relative importance of these mechanisms in red spruce (Picea rubens Sarg.) and balsam fir (Abies balsamea Lin.), which share understory regeneration niches in northeastern North America. Although its reproductive effort is adequate, spruce has decreased in abundance, in relation to fir, in seedling and sapling populations, even in forests that have a predominance of spruce in the overstory. METHODS: To understand the factors that regulate this phenomenon and their implications for tree populations, we compared intrinsic and plastic regulation of first- and second-year seedlings under steady understory irradiance levels and in response to increases in light environment. KEY RESULTS: Both species exhibited interactions of ontogenetic patterns and plasticity in first- and second-year seedlings. Physiologically, spruce had higher photosynthetic capacity, allocation to photoprotective xanthophylls, and greater plasticity in response to light treatments. Although both species demonstrated an inability to plastically increase photosynthetic capacity in the short term, spruce benefited from greater allocation to foliage under increased irradiance. Fir showed a conservative strategy in root-shoot allocation that may better equip seedlings to withstand drought adaptations and attributes associated with greater shade tolerance. CONCLUSIONS: These attributes likely contribute to the relative success of fir seedlings in the current climate. By contrast, they indicate that spruce would be a superior competitor in cooler, moister climates, which suggests that future forest composition will be largely determined by an interaction of disturbance and moisture regimes.

Growth, allometry and shade tolerance of understory saplings of four subalpine conifers in central Japan.

The conifers Abies veitchii, A. mariesii, Picea jezoensis var. hondoensis, Tsuga diversifolia dominate in subalpine forests in central Japan. We expected that species differences in shade tolerance and in aboveground and belowground architecture are important for their coexistence. We examined net production and carbon allocation of understory saplings. Although the four species allocated similar amounts of biomass to roots at a given trunk height, the root-zone area of T. diversifolia was greater than that of the three other species. T. diversifolia often dominates shallow soil sites, such as ridge and rocky slopes, and, therefore, a wide spread of lateral roots would be an adaptation to such edaphic conditions. Crown width and leaf and branch mass were greatest for T. diversifolia and A. mariesii, followed in order by A. veitchii and P. jezoensis var. hondoensis. Although leaf mass of P. jezoensis var. hondoensis was lowest among the four species, species differences were not found in the net production per sapling because net production per leaf mass was greatest for P. jezoensis var. hondoensis. The leaf lifespan was longer in the order A. mariesii, T. diversifolia, P. jezoensis var. hondoensis and A. veitchii. The minimum rate of net production per leaf mass required to maintain the current sapling leaf mass (MRNP(LM)) was lowest in A. mariesii and T. diversifolia, and increased in the order of A. veitchii and P. jezoensis var. hondoensis. A. mariesii and T. diversifolia may survive in shade conditions by a lower MRNP(LM) than the two other species. Therefore, species differences in aboveground and belowground architecture and MRNPLM reflected their shade tolerance and regeneration strategies, which contribute to their coexistence.

A mechanistic model for the light response of photosynthetic electron transport rate based on light harvesting properties of photosynthetic pigment molecules.

Models describing the light response of photosynthetic electron transport rate (ETR) are routinely used to determine how light absorption influences energy, reducing power and yields of primary productivity; however, no single model is currently able to provide insight into the fundamental processes that implicitly govern the variability of light absorption. Here we present development and application of a new mechanistic model of ETR for photosystem II based on the light harvesting (absorption and transfer to the core 'reaction centres') characteristics of photosynthetic pigment molecules. Within this model a series of equations are used to describe novel biophysical and biochemical characteristics of photosynthetic pigment molecules and in turn light harvesting; specifically, the eigen-absorption cross-section and the minimum average lifetime of photosynthetic pigment molecules in the excited state, which describe the ability of light absorption of photosynthetic pigment molecules and retention time of excitons in the excited state but are difficult to be measured directly. We applied this model to a series of previously collected fluorescence data and demonstrated that our model described well the light response curves of ETR, regardless of whether dynamic down-regulation of PSII occurs, for a range of photosynthetic organisms (Abies alba, Picea abies, Pinus mugo and Emiliania huxleyi). Inherent estimated parameters (e.g. maximum ETR and the saturation irradiance) by our model are in very close agreement with the measured data. Overall, our mechanistic model potentially provides novel insights into the regulation of ETR by light harvesting properties as well as dynamical down-regulation of PSII.

UV-B exposure of indoor-grown Picea abies seedlings causes an epigenetic effect and selective emission of terpenes.

Terpenoids are involved in various defensive functions in plants, especially conifers. Epigenetic mechanisms, for example DNA methylation, can influence plant defence systems. The purpose of the present study was to investigate the influence of UV-B exposure on the release of terpenoids from spruce seedlings and on needle DNA methylation. Ten-week-old seedlings grown indoors were exposed to UV-B radiation during 4 h, and the volatile compounds emitted from the seedlings were analysed. Analysis of the volatiles 1, 3, and 22 d after this UV-B exposure showed that bornyl acetate, borneol, myrcene, and limonene contents increased during the first 3 days, while at day 22 the level of emission had returned to the control level. UV-B exposure decreased the level of DNA methylation in needles of young seedlings, reflected in methylation changes in CCGG sequences. Exposure of young seedlings to UV-B radiation might be a way to potentiate the general defensive capacity, improving their ability to survive in outdoor conditions. UV-B-induced defence is discussed in the light of epigenetic mechanisms.

Light and temperature sensing and signaling in induction of bud dormancy in woody plants.

In woody species cycling between growth and dormancy must be precisely synchronized with the seasonal climatic variations. Cessation of apical growth, resulting from exposure to short photoperiod (SD) and altered light quality, is gating the chain of events resulting in bud dormancy and cold hardiness. The relative importance of these light parameters, sensed by phytochromes and possibly a blue light receptor, varies with latitude. Early in SD, changes in expression of light signaling components dominate. In Populus active shoot elongation is linked to high expression of FLOWERING LOCUS T (FT) resulting from coincidence of high levels of CONSTANS and light at the end of days longer than a critical one. In Picea, PaFT4 expression increases substantially in response to SD. Thus, in contrast to Populus-FT, PaFT4 appears to function in inhibition of shoot elongation or promotion of growth cessation. Accordingly, different FT-genes appear to have opposite effects in photoperiodic control of shoot elongation. Reduction in gibberellin under SD is involved in control of growth cessation and bud formation, but not further dormancy development. Coinciding with formation of a closed bud, abscisic acid activity increases and cell-proliferation genes are down-regulated. When dormancy is established very few changes in gene expression occur. Thus, maintenance of dormancy is not dependent on comprehensive transcriptional regulation. In some species low temperature induces growth cessation and dormancy, in others temperature affects photoperiod requirement. The temperature under SD affects both the rate of growth cessation, bud formation and depth of dormancy. As yet, information on the molecular basis of these responses to temperature is scarce.

Green spruce aphid infestations cause larger growth reductions to Sitka spruce under shade.

Light availability and infestation by the green spruce aphid (Elatobium abietinum) are key factors affecting the growth of Sitka spruce (Picea sitchensis) seedlings under a mature tree canopy, but their combined effect on seedling growth has not previously been quantified. A controlled outdoor experiment in which light levels (high light (HL): 100%, intermediate light (IL): 24%) and aphid infestation (absence/presence) were manipulated was conducted over 2 years to look at the effects on seedling growth and biomass distribution patterns. Aphid population assessments showed a significantly increased population density under IL, with three to four times higher cumulative aphid densities than that under HL. Defoliation rates of infested seedlings were directly related to aphid density. Total seedling biomass was strongly reduced in IL, and aphid infestation caused additional reductions in the biomass of particular components of the seedlings. Dry weight (DW) of older (≥1-year-old) needles in infested trees was significantly decreased in both years. Total root DW at the end of the second year was significantly affected by aphid infestation, and the reduction (14-18%) was similar in IL and HL treatments despite large differences in aphid density. Biomass distribution patterns in infested trees were similar to that of uninfested trees within each light treatment, indicating that the relative decreases in root biomass were accompanied by similar reductions in distribution to the above-ground parts of the seedlings. Leader extension growth of infested seedlings was reduced by 15-17% compared with uninfested seedlings under IL, whereas only a 2-3% reduction in leader extension of infested seedlings under HL was observed. The results showed that the response of seedlings to E. abietinum were primarily dependent on the light environment. The significant reduction caused by aphids on the total DW of older needles and roots, and on leader extension growth, does suggest the potential for effects to accumulate over time.

A relationship between galactic cosmic radiation and tree rings.

Here, we investigated the interannual variation in the growth rings formed by Sitka spruce (Picea sitchensis) trees in northern Britain (55 degrees N, 3 degrees W) over the period 1961-2005 in an attempt to disentangle the influence of atmospheric variables acting at different times of year. Annual growth rings, measured along the north radius of freshly cut (frozen) tree discs and climatological data recorded at an adjacent site were used in the study. Correlations were based on Pearson product-moment correlation coefficients between the annual growth anomaly and these climatic and atmospheric factors. Rather weak correlations between these variables and growth were found. However, there was a consistent and statistically significant relationship between growth of the trees and the flux density of galactic cosmic radiation. Moreover, there was an underlying periodicity in growth, with four minima since 1961, resembling the period cycle of galactic cosmic radiation. * We discuss the hypotheses that might explain this correlation: the tendency of galactic cosmic radiation to produce cloud condensation nuclei, which in turn increases the diffuse component of solar radiation, and thus increases the photosynthesis of the forest canopy.

Microwave-assisted rapid plant sample preparation for transmission electron microscopy.

The preparation of plant leaf material for transmission electron microscopical investigations can be a very time- and labour-consuming task as the reagents infiltrate the samples quite slowly and as usually most steps have to be performed manually. Fixation, buffer washes, dehydration, resin infiltration and polymerization of the resin-infiltrated leaf samples can take several days before the specimen can be cut ultrathin and used for ultrastructural investigations. In this study, we present a microwave-assisted automated sample preparation procedure that reduces preparation time from at least 3 days to about 5 h - with only a few steps that have to be performed manually - until the plant sample can be ultrathin sectioned and observed with the transmission electron microscope. For studying the efficiency of this method we have compared the ultrastructure of different leaf material (Arabidopsis thaliana, Nicotiana tabacum and Picea abies) which was prepared with a conventional, well-established chemical fixation and embedding protocol and a commercially available automated microwave tissue processor. Despite the massive reduction in sample preparation time no negative effects on cutting properties of the blocks, stability of the sections in the electron beam, contrast and ultrastructure of the cells were observed under the transmission electron microscope when samples were prepared with the microwave-assisted protocol. Additionally, no negative effects were detected on the dimensions of fine structures of grana stacks (including membranes, inter- and intrathylakoidal spaces), the nuclear envelope and the plasma membrane as the diameter of these structural components did not differ between leaf samples (of the same species) that were processed with the automated microwave tissue processor or by conventional fixation and embedding at room temperature.

Water deficit affects mesophyll limitation of leaves more strongly in sun than in shade in two contrasting Picea asperata populations.

The aim of this study was to examine the response of internal conductance to CO(2) (g(i)) to soil water deficit and contrasting light conditions, and their consequences on photosynthetic physiology in two Picea asperata Mast. populations originating from wet and dry climate regions of China. Four-year-old trees were subjected to two light treatments (30% and 100% of full sunlight) and two watering regimes (well watered, drought) for 2 years. In both tested populations, drought significantly decreased g(i) and the net photosynthesis rate (A) and increased carbon isotope composition (delta(13)C) values in both light treatments, in particular in the sun. Moreover, drought resulted in a significantly higher relative limitation due to stomatal conductance (L(s)) in both light treatments and higher relative limitation due to internal conductance (L(i)) and abscisic acid (ABA) in the sun plants. The results also showed that L(i) (0.26-0.47) was always greater than L(s) (0.12-0.28). On the other hand, drought significantly decreased the ratio of chloroplastic to internal CO(2) concentration (C(c)/C(i)), photosynthetic nitrogen utilization efficiency (PNUE) and total biomass in the sun plants of the wet climate population, whereas there were no significant changes in these parameters in the dry climate population. Our results also showed that the dry climate population possessed higher delta(13)C values with higher ratio of internal conductance to stomatal conductance (g(i)/g(s)), suggesting that increasing the g(i)/g(s) ratio enhances water-use efficiency (WUE) in plants evolved in arid environments. Thus, we propose that the use of the g(i)/g(s) parameter to screen P. asperata plants with higher water deficit tolerance is certainly worthy of consideration. Furthermore, g(i) is an important variable, which reflects the population differences in PNUE, and it should thus be included in plant physiological investigations related to leaf economics.

The effect of ultraviolet-C and precursor feeding on stilbene biosynthesis in spruce Picea jezoensis.

This study examines the effects of p-coumaric (CoA) and caffeic (CaA) acids on stilbene biosynthesis in one-year-old needles of Picea jezoensis (Lindl. et Gord.) Fisch ex Carr. under control conditions and after ultraviolet (UV-C) irradiation. HPLC analysis revealed that while UV-C irradiation slightly affected the total stilbene content, CoA and CaA feeding increased the total content of stilbenes by 1.2-1.3-fold. UV-C treatment combined with CoA-feeding of the P. jezoensis cuttings exerted the most pronounced stimulatory effect on stilbene accumulation leading to the total stilbene content of 9.18 mg/g of dry weight or DW. This increase correlated with the elevated transcription of the stilbene synthase PjSTS1a and PjSTS1b genes. UV-C treatment in combination with CaA feeding of P. jezoensis did not considerably influence stilbene content. These results revealed a positive effect of UV-C radiation and phenolic precursors on the content of stilbenes in spruce needles.

Climatic control of bud burst in young seedlings of nine provenances of Norway spruce.

Detailed knowledge of temperature effects on the timing of dormancy development and bud burst will help evaluate the impacts of climate change on forest trees. We tested the effects of temperature applied during short-day treatment, duration of short-day treatment, duration of chilling and light regime applied during forcing on the timing of bud burst in 1- and 2-year-old seedlings of nine provenances of Norway spruce (Picea abies (L.) Karst.). High temperature during dormancy induction, little or no chilling and low temperature during forcing all delayed dormancy release but did not prevent bud burst or growth onset provided the seedlings were forced under long-day conditions. Without chilling, bud burst occurred in about 20% of seedlings kept in short days at 12 degrees C, indicating that young Norway spruce seedlings do not exhibit true bud dormancy. Chilling hastened bud burst and removed the long photoperiod requirement, but the effect of high temperature applied during dormancy induction was observed even after prolonged chilling. Extension of the short-day treatment from 4 to 8 or 12 weeks hastened bud burst. The effect of treatments applied during dormancy development was larger than that of provenance; in some cases no provenance effect was detected, but in 1-year-old seedlings, time to bud burst decreased linearly with increasing latitude of origin. Differences among provenances were complicated by different responses of some origins to light conditions under long-day forcing. In conclusion, timing of bud burst in Norway spruce seedlings is significantly affected by temperature during bud set, and these effects are modified by chilling and environmental conditions during forcing.

Efficiency of the photosynthetic apparatus in developing needles of Norway spruce (Picea abies L. Karst.).

The photosynthetic performance of developing spruce (Picea abies L. Karst.) needles was investigated. As revealed by previous reports, the biosynthesis of chlorophylls and carotenoids was not following the characteristic chloroplast ultrastructure building up during needle elongation process. The aim of our study was to investigate photosynthetic capability (evaluated by oxygen evolution and chlorophyll a fluorescence kinetics measurements), the dynamics of chloroplast pigments biosynthesis and the expression of major photosynthetic proteins as well as to find out possible correlation between components of issue. Low amounts of chlorophylls and carotenoids, LHC II and Rubisco LSU were detected in the embryonic shoot of vegetative buds. Although PS II was functional, oxygen production was not sufficient to compensate for respiration in the same developmental stage. The light compensation point of respiration was successively lowered during the needle elongation. Nevertheless the significant increase in photosynthetic pigments as well as the high level of expression of LHC II and Rubisco LSU proteins was observed in the later stages of needle development. Our results suggest that, besides light, some other environmental factors could be critical for producing fully functional chloroplasts in rapidly growing young needles.

Significance of ozone exposure for inter-annual differences in primary metabolites of old-growth beech (Fagus sylvatica L.) and Norway spruce (Picea abies L.) trees in a mixed forest stand.

The influence of long-term free-air ozone fumigation and canopy position on leaf contents of total glutathione, its redox state, non-structural proteins (NSP), soluble amino compounds, and total soluble sugars in old-growth beech (FAGUS SYLVATICA) and spruce (PICEA ABIES) trees were determined over a period of five years. Ozone fumigation had weak effects on the analysed metabolites of both tree species and significant changes in the contents of total glutathione, NSP, and soluble sugars were observed only selectively. Beech leaves were affected by crown position to a higher extent than spruce needles and exhibited lower contents of total glutathione and NSP and total soluble sugars, but enhanced contents of oxidised glutathione and amino compounds in the shade compared to the sun crown. Contents of total soluble sugars generally were decreased in shade compared to sun needles of spruce trees. Interspecific differences between beech and spruce were more distinct in the sun compared to the shade crown. Contents of total glutathione were increased whilst contents of amino compounds and total soluble sugars were lower in spruce needles compared to beech leaves. The metabolites determined showed individual patterns in the course of the five measurement years. Contents of total glutathione and its redox state correlated with air temperature and global radiation, indicating an important role for the antioxidant at low temperatures. Correlations of glutathione with instantaneous ozone concentrations seem to be a secondary effect. Differences in proteins and/or amino compounds in the inter-annual course are assumed to be a consequence of alterations in specific N uptake rates.

[Characteristics of morphogenesis and growth processes of conifers in the Chernobyl nuclear accident zone].

In the article we present data on the study of morphogenesis and of growth processes of Scotch pine (Pinus sylvestris L.) and of Norwey spruce (Picea abies (L.) Karst.) vegetative organs after radiation exposure in the Chernobyl nuclear accident zone. The anomalies in morphogenesis processes at different absorbed doses are described in detail. It is established that the death of pine forest began under absorbed dose 80-100 Gy and more, mass yellowing of needles at 50-60 Gy, and maximal morphosis at 8-12 Gy. Inhibition phenomenon of growth processes under acute irradiation and giantism under durable chronic irradiation were also put under investigation. Features of radiation exposure on pine and fir growth processes at different ontogeny phases were characterized. High radio-sensitivity of Norwey spruce is established.

Long-term exposure to enhanced UV-B radiation has no significant effects on growth or secondary compounds of outdoor-grown Scots pine and Norway spruce seedlings.

The effects of long-term enhanced UV-B radiation on growth and secondary compounds of two conifer species were studied in an outdoor experiment. Scots pine (Pinus sylvestris) seedlings were exposed for two growing seasons and Norway spruce (Picea abies) seedlings for three growing seasons to supplemental UV-B radiation, corresponding to a 30% increase in ambient UV-B radiation. The experiment also included appropriate controls for ambient and increased UV-A radiation. Enhanced UV-B did not affect the growth of the conifer seedlings. In addition, neither the concentrations of terpenes and phenolics in the needles nor the concentrations of terpenes in the wood were affected. However, in the UV-A control treatment the concentrations of diterpenes in the wood of Scots pine decreased significantly compared to the ambient control. Apparently, a small increase in UV-B radiation has no significant effects on the secondary compounds and growth of Scots pine and Norway spruce seedlings.