Influence of near-ultraviolet light enhancement and photosynthetic photon flux density during photoperiod extension on the morphology and lignin content of black spruce seedlings

When containerized black spruce seedlings (Picea mariana (Mill.) B.S.P.) are grown rapidly in greenhouse culture, they sometimes bend over, grow horizontally and become deformed. This phenomenon has been known to affect between 5% and 10% of a winter greenhouse crop. In this study, near-ultraviolet lamps were used to supplement the artificial light received from high-pressure sodium lamps and the effects on seedling morphology and lignin contents were examined. Neither height to diameter ratios nor lignin concentrations were significantly affected by UV radiation flux density. However, seedling biomass, height, root collar diameter, lignin content, and lignin to cellulose ratios of stems were significantly correlated with total photosynthetic photon flux density (PPFD) received during photoperiod extension. Height to diameter ratios were negatively correlated with PPFD during photoperiod enhancement because of a greater relative increase in diameter growth compared with height growth. Neither UV nor PAR flux density affected the percentage of black spruce seedlings having stem deformations greater than 30 ° from the vertical.

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Résumé Lorsque des semis d'épinette noire (Picea mariana (Mill.) B.S.P.) en conteneur sont forcés d'avoir une croissance rapide en serre, leur tige est parfois courbée ou déformée. Ce phénomène peut affecter entre 5% et 10% d'une culture serricole hivernale. Dans la présente étude, un enrichissement du rayonnement près de l'ultraviolet a été appliqué sur des semis en complément à l'éclairage artificiel procuré par des lampes au sodium à haute pression. La morphologie et le contenu en lignine des semis ont alors été examinés. Le rapport entre la hauteur et le diamètre des semis, et la concentration en lignine n'ont pas été significativement affectés par la densité du flux photonique ultraviolet. Cependant, la biomasse, la hauteur, le diamètre au collet, le contenu en lignine et le ratio lignine: cellulose des tiges des semis ont été significativement corrélés à la densité du flux photonique photosynthétique (PPFD) reçue pendant l'extension de la photopriode. Le rapport entre la hauteur et le diamètre a été négativement corrélé à la PPFD pendant l'augmentation de la photopériode à cause de la plus grande croissance en diamètre des semis comparativement à leur croissance en hauteur. La densité du flux photonique au niveau de l'ultraviolet comme au niveau de la radiation photosynthétiquement active n'a pas affecté le pourcentage de semis d'épinette noire ayant des déformations de leur tige plus grande que 30 ° comparativement à la verticale.

     

The effect of photosynthetic photon flux density on development of frost hardiness in top and roots of Larix leptolepis seedlings

Seedlings of Larix leptolepis were grown in a growth chamber under short day conditions (8 h day) at 4 different photosynthetic photon flux densities (PPFD's). After a period of 8 weeks frost hardiness of the shoot tips and roots, dry matter content, dry weight (dw), content of glucose and starch were determined. The frost hardiness in the shoot tips increased from ‐15°C at a PPFD of 55μmol m^‐2s^‐1 to about ‐35°C at 440 μmol m^‐2s^‐1. No effect of PPFD was found on frost hardiness of the roots. A high PPFD results in a high dry matter content. The effect on dry matter content was most pronounced for the shoot tips and less pronounced for the roots. The total dry weight increased for both root and top with increasing PPFD. The height of the plants increased when the PPFD increased up to 220 μmol m^‐2s^‐1.     

Interacting effects of irradiance and water stress on dry weight and biomass partitioning in Fagus sylvatica seedlings

Abstract

Actual climate models for central Europe predict prolonged summer droughts. Knowledge on how the interaction between light and water availability affects regeneration will hence be of major importance. In an experiment carried out under controlled conditions, newly emerged beech seedlings were grown in pots with sand during 54 days. Three treatments applying three different light levels (2, 9 and 43% relative light intensity) were combined with two soil water treatments (control and drought). At the end of the experiment, seedlings were separated into leaves, stem and root and the seedlings’ dry mass, leaf area and stem length was determined. Low irradiance (2%) had a strong negative effect on dry weights of seedling components, leaf area and specific leaf mass. Drought clearly affected biomass partitioning for seedlings at high irradiance levels (43%). An interaction between irradiance and drought on biomass partitioning in beech seedlings was observed at medium irradiance level (9%). Within a wide range of light levels in the forests, both light and drought may affect biomass partitioning in young seedlings.     

Effects of CO2 enrichment at varying photon flux density on the growth of picea abies (L) karst. seedlings

Seedlings of Norway spruce (Picea abies (L.)) were grown at 335 and 1000 μl CO₂ 1^?1 for 118 days in growth rooms at different irradiance levels. Photon flux density ranging from 8.6 to 34.6 mol m^?2 day^?t (PAR) was given either as constant light or as alternating levels in intervals of two or six hours. CO₂ enrichment increased the plant dry weight from 36% to 105% by increasing photon flux density from 8.6 to 25.9 mol m^?2 day^?1. At constant light the dry weight apparently reached its maximum at a photon flux density of 25.9 mol m^?2 day^?t. At the lower radiation levels alternating in CO₂ enriched air gave slightly higher dry weights compared to constant light levels. At the highest radiations the effect on dry weight was the opposite. High CO₂ concentration and 300 μmol m^?2 s^?1 constant light (25.9 mol m^?2 day^?1) gave the best growth and quality of plants. Top, root, stem and foliage weight were proportionally affected. Shoot length was enhanced by CO₂ enrichment. Shoot weight per cm was substantially increased both by CO₂ enrichment and increasing photon flux density.     

Impacts of drought on mineral macro- and microelements in provenances of beech (Fagus sylvatica L.) seedlings

Beech seedlings originating from 11 German provenances with different climatic conditions were grown in pots and cultivated in a greenhouse. The composition of macro- and microelements in roots, axes and leaves was measured after half of the seedlings were subjected to a simulated summer drought. The recently described sensitivity of these provenances to drought was compared with drought-mediated changes in the elemental and ionic composition in organs of the seedlings; in addition, partitioning between roots and shoots was evaluated. A number of element concentrations were decreased in roots due to drought (K 94% of control, Mg 94%, Mn 75% and Zn 85%). However, chloride concentration increased in all organs (115-125%) and was the only element affected in leaves. Some changes in ionome can be related to sensitivity of provenances, but it is difficult to decide whether these changes are a result of, or a reason for, drought tolerance or sensitivity. Observed increases in chloride concentration in all plant parts of drought-treated beech seedlings can be explained by its function in charge balance, in particular since the level of phosphate was reduced. As a result of chloride accumulation, the sum of added charges of anions (and cations) in water extracts of leaf and root material was similar between drought and control plants. Since only the partitioning of Ca and Al (both only in axis) as well as Mn was affected and other elements (together with previously observed effects on C, N, S and P) remained unaffected by drought in all provenances, it can be concluded that direct effects by means of mass flow inhibition in xylem and phloem are unlikely. Secondary effects, for example on the pH of transport sap and the apoplastic space, cannot be excluded from the present study. These effects may affect partitioning between the apoplast and symplast and therefore may be significant for drought sensitivity.     

Growth and survival of Fagus sylvatica seedlings in relation to light intensity and soil water content

Previous investigations have shown different growth and root/shoot ratio increases of beech seedlings (Fagus sylvatica) with increased light intensity. In the present investigation both light intensity and soil water content were regulated on four levels in a factorial experiment. At the highest level of soil water content the seedling growth increased linearly with light intensity while the root/shoot ratio decreased with light intensity. At low levels of soil water content both the growth and root/shoot ratio slightly increased with increased light. The soil water content was also affected by the light intensity indicating that the soil water content may have caused unobserved effects in responding to increased light in previous investigations. It is concluded that both light intensity and soil water content should be quantified or manipulated even if only the effect of one of the two growth factors are investigated.     

The applicability of a color acetate film for estimating photosynthetic photon flux density in a forest understory

The suitability of a color acetate film for estimating photosynthetic photon flux density (PPFD) in a forest understory was examined. The fading ratio of the film (F), the total PPFD (PPFD_total) to which the film was exposed, and the average daily maximum temperature during exposure (T) were obtained from measurements at multiple sampling points throughout an entire year within a natural secondary forest (n = 42). The ranges of the recorded values were as follows: F 35%–99%, PPFD_total 1.4–28.3molm^–2, and T 6°–32°C. PPFD_total was regressed by F and T with a high r ² (=0.94; P < 0.0001): PPFD_total = (100 – F)/(1.085 + 0.051T). The absolute error (|estimated PPFD_total – measured PPFD_total|) averaged 1.3molm^–2 with a maximum of 5.7molm^–2, indicating a good fit. These results indicated broad applicability of the film, both spatially and temporally, for estimating forest understory PPFD.     

Effect of photon flux density on carbon assimilation and chlorophyll a fluorescence of cold-stored white spruce and lodgepole pine seedlings

White spruce (Picea glauca (Moench.) Voss) and lodgepole pine (Pinus contorta Dougl. var. latifolia Engelm.) seedlings previously held in dark, frozen storage (-2 degrees C) for 2.5 or 6 months, and nursery-grown white spruce seedlings lifted in summer were exposed to photon flux densities (PFDs) similar to those that might be encountered at planting. Photosynthetic gas exchange and chlorophyll a (chl a) fluorescence were examined in cold-stored and summer-lifted seedlings before and after a 9 h-exposure to artificial illumination of high PFD (2000 micro mol m(-2) s(-1)) or low PFD (ca. 500 micro mol m(-2) s(-1)), and during exposure to 400 micro mol m(-2) s(-1) for 4-9 days. In the 2.5-month-stored and summer-lifted seedlings, the high-PFD treatment caused a small decrease in carbon fixation and a large decrease in the ratio of variable to maximum fluorescence (F(v)/F(m)) relative to the effect of the low-PFD treatment. In contrast, in the 6-month-stored seedlings the high-PFD treatment caused a significant decrease in rate of light-saturated carbon fixation but little decrease in F(v)/F(m) relative to the effect of the low-PFD treatment, indicating that the mechanisms for maintaining integrity of the photochemical apparatus had changed during the storage interval.     

The effects of a regeneration felling on photosynthetic photon flux density and regeneration growth in a <Emphasis Type="Italic">Nothofagus pumilio</Emphasis> forest

Photosynthetic photon flux density (PPFD) during the growing season and regeneration growth (height and base stem diameter) were investigated in two natural stands in the Patagonian region of Chile, one without silvicultural management and another with a regenerative felling under a shelterwood system. PPFD was measured by means of fifteen sensors (quantum Li-190SA) installed in each stand and distributed within three canopy openness grades. Four regeneration plots (1 m²) were established around each sensor. In each of the plots, the height and base diameter of ten labelled plants within the upper regeneration layer were measured in the growing seasons 2001–2002 and 2002–2003. In the stand with regeneration felling total PPFD in the growing season was 2.5–2.9 times higher than in the stand without intervention. In both stands, total PPFD in the growing season increased by about 420 mmol/m² when the relative canopy, which was in the range between 30 and 70%, was reduced by 10%. An identical behaviour was observed for the height and base stem diameter increases reflecting a clear effect of intervention and canopy coverage on the magnitude of PPFD received by the regeneration and its development.     

Effects of selective thinning on growth and development of beech (Fagus sylvatica L.) forest stands in south-eastern Slovenia

We studied the effects of two types of selective thinning on beech stands formed by a shelterwood cut in 1910 — with lower number of crop trees and higher thinning intensity (T1) and higher number of crop trees with lower thinning intensity (T2). The stands were thinned in 1980, 1991 and 2001. Despite a lower stand density after thinning, the annual basal area increments of thinned stands in both thinning periods (1980–1991 and 1991–2002) were around 20% higher compared to those of the control (unthinned) stands. The mean annual basal area increment of dominant trees was 30–56% larger in the thinned plots compared to the control plots. Of 176 initial crop trees in the T1, 72% were chosen again during the last thinning. In the T2, 258 crop trees were chosen in the first thinning, and only 62% of these trees were chosen again during the last thinning. Only crown suppression and diameter classes of crop trees significantly influenced their basal area increment when diameter classes, crown size, crown suppression, and social status were tested. In the thinned stands, the dominant trees are more uniformly distributed if compared to the dominant trees in the control plots. Finally, the herbaceous cover and the species diversity were higher in the thinned plots.     

Biomass, basic density and biomass expansion factor functions for European beech (Fagus sylvatica L.) in Denmark

The objective of this study was to analyse the within-tree allocation of biomass and to develop biomass functions for above- and below-ground components of European beech in Denmark. Separate functions were developed for stem, branches, below-ground stump and root system, total above-ground biomass and total tree biomass. For each of these components or aggregate components, models were also developed for the average basic density of wood and bark. To enhance the versatility of the models, a function for estimating the biomass expansion factor (BEF) was also developed. The functions were based on 66 trees measured for total biomass. Model performance was evaluated based on 74 trees measured only for above-ground biomass. The trees were sampled in 18 different forest stands covering a wide range of tree sizes and stand treatments. Models were estimated using a linear mixed-effects procedure to account for within-stand correlations. The functions for biomass and BEFs included only diameter at breast height and total tree height for individual trees as predictor variables. Inclusion of additional variables reflecting site quality or stand density did not improve model performance. The functions for basic density included individual tree diameter, tree height and quadratic mean diameter as predictor variables, indicating an effect of stand density on the basic density of wood and bark.     

Impact of late frost events on radial growth of common beech (Fagus sylvatica L.) in Southern Germany

Phenological, temperature, and tree-ring data were used in order to identify and quantify the impact of late frosts on common beech (Fagus sylvatica L.) at different altitudes in Southern Germany during the last century. For this intention, dendroecological investigations were made upon trees at the Meteorological Observatory Hohenpeißenberg as well as from seven stands in the Bavarian Forest and 17 stands at the northern fringe of the Alps. From these locations, a considerable number of severe growth minima in the tree-ring series could be related to late frost in the days of or immediately after leaf unfolding. The frequency of frost-related growth minima increases with altitude. In individual years, radial growth can be reduced by more than 90% (stand mean) in relation to the average growth of the ten previous years. Hence, late frosts are considered as important ecological events that strongly affect beech vitality and competitiveness especially at high altitudes. Evidence of significant impacts on radial growth by late frosts distinct before leaf unfolding or with temperatures above ?3°C was not found. Also, increasing frequency and intensity of late frosts during recent decades were not ascertained. Hence, the recently observed decreased vitality of common beech accompanied by growth depressions especially at high altitude sites in Central Europe cannot be explained as a consequence of late frost damage.     

Effects of elevated CO(2) concentration on leaf characteristics and photosynthetic capacity of beech (Fagus sylvatica) during the growing season

Two-year-old beech (Fagus sylvatica L.) saplings were planted directly in the ground at high density (100 per m(2)), in an experimental design that realistically mimicked field conditions, and grown for two years in air containing CO(2) at either ambient or an elevated (ambient + 350 ppm) concentration. Plant dry mass and leaf area were increased by a two-year exposure to elevated CO(2). The saplings produced physiologically distinct types of sun leaves associated with the first and second growth flushes. Leaves of the second flush had a higher leaf mass per unit area and less chlorophyll per unit area, per unit dry mass and per unit nitrogen than leaves of the first flush. Chlorophyll content expressed per unit nitrogen decreased over time in plants grown in elevated CO(2), which suggests that, in elevated CO(2), less nitrogen was invested in machinery of the photosynthetic light reactions. In early summer, the photosynthetic capacity measured at saturating irradiance and CO(2) was slightly but not significantly higher in saplings grown in elevated CO(2) than in saplings grown in ambient CO(2). However, a decrease in photosynthetic capacity was observed after July in leaves of saplings grown in CO(2)-enriched air. The results demonstrate that photosynthetic acclimation to elevated CO(2) can occur in field-grown saplings in late summer, at the time of growth cessation.     

Effects of drought and rewatering on growth and transpiration in European beech seedlings late in the growing season

Drought stress is one of the most important environmental factors affecting plant growth and survival. To date, most studies aim at understanding of post-stress physiological and anatomical adaptation to drought stress; however only few studies focus on plant recovery. In the present study, transpiration, shoot water potential, and anatomical and morphological measurements were performed on 4-year-old European beech seedlings with fully developed leaves. The seedlings were exposed to three levels of soil water potential (well-watered, moderate drought stress and severe drought stress) and followed by rewatering under greenhouse conditions. Reduced transpiration rates were observed in the stressed seedlings as a response to drought stress, whereas anatomical and morphological variables remained unchanged. Three days after rewatering, transpiration rates in both moderately and severely stressed seedlings recovered to the levels of those of well-watered seedlings. Drought stress promoted leaf budding, resulting in higher shoot dry mass of stressed seedlings. Our findings indicate that anatomical and morphological adaptations of European beech seedlings to drought stress are visibly limited during late-season growth stages. These results will help us to further understand factors involved in drought adaptation potential of European beech seedlings faced with expected climate-related environmental changes. To complete our findings, further experiments on plant recovery from drought stress should be focused on different periods of growing season.     

Diurnal and seasonal variability in radial distribution of sap flux density: Implications for estimating stand transpiration

Daily and seasonal patterns in radial distribution of sap flux density were monitored in six trees differing in social position in a mixed coniferous stand dominated by silver fir (Abies alba Miller) and Norway spruce (Picea abies (L.) Karst) in the Alps of northeastern Italy. Radial distribution of sap flux was measured with arrays of 1-cm-long Granier probes. The radial profiles were either Gaussian or decreased monotonically toward the tree center, and seemed to be related to social position and crown distribution of the trees. The ratio between sap flux estimated with the most external sensor and the mean flux, weighted with the corresponding annulus areas, was used as a correction factor (CF) to express diurnal and seasonal radial variation in sap flow. During sunny days, the diurnal radial profile of sap flux changed with time and accumulated photosynthetic active radiation (PAR), with an increasing contribution of sap flux in the inner sapwood during the day. Seasonally, the contribution of sap flux in the inner xylem increased with daily cumulative PAR and the variation of CF was proportional to the tree diameter, ranging from 29% for suppressed trees up to 300% for dominant trees. Two models were developed, relating CF with PAR and tree diameter at breast height (DBH), to correct daily and seasonal estimates of whole-tree and stand sap flow obtained by assuming uniform sap flux density over the sapwood. If the variability in the radial profile of sap flux density was not accounted for, total stand transpiration would be overestimated by 32% during sunny days and 40% for the entire season.     

Azimuthal variations of sap flux density within Japanese cypress xylem trunks and their effects on tree transpiration estimates

Sap flow techniques are practical tools for estimating tree transpiration. Though many previous studies using sap flow techniques did not consider azimuthal variations of sap flux density (F _d) on xylem trunk to estimate tree transpiration, a few studies reported that ignoring the azimuthal variations in F _d could cause large errors in tree transpiration estimates for some tree species. Therefore, examining azimuthal variations in F _d for major plantation tree species is critical for estimating tree transpiration. Using the thermal dissipation method, we examined azimuthal variations in F _d in six trees of Japanese cypress Chamaecyparis obtusa (Sieb. et Zucc.) Endl., which is one of the most common plantation tree species in Japan. We recorded considerable variations among F _d at four different azimuthal directions. The F _d value for one aspect was more than 100% larger than those for the other aspects. We calculated differences between tree transpiration estimates based on F _d for one to three azimuthal directions and those based on F _d for four aspects. The differences relative to tree transpiration estimates based on F _d for four aspects were typically 30, 20, and 10% in accordance with the F _d for one, two, and three measurement aspects, respectively. This finding indicates that ignoring azimuthal variations could cause large errors in tree transpiration estimates for Japanese cypress.     

Influence of nitrogen fertilization on minerals, carbohydrates, amino acids and phenolic compounds in beech (Fagus sylvatica L.) leaves

Beginning in 1985, plots in a 120-year-old beech (Fagus sylvatica L.) stand in the province of Scania, southern Sweden were fertilized annually with ammonium nitrate. In August 1989, leaves from both fertilized and unfertilized trees were analyzed for mineral nutrients, soluble carbohydrates, amino acids and phenolic compounds. Leaf concentrations of total nitrogen were increased by N fertilization. Ammonium was not detected in the leaves, but NO(3) (-) was increased by up to 65% as a result of N fertilization. Nitrogen fertilization decreased foliar concentrations of P and Cu, leading to a nutrient imbalance relative to N. Of the free amino acids, glutamic acid and aspartic acid predominated and, together with glutamine and asparagine, were significantly increased by N fertilization. The concentrations of amides were 2-4 times higher in fertilized trees than in unfertilized trees. Soluble carbohydrates did not change in response to N fertilization, but total phenolic compounds decreased markedly. The reduced concentration of phenolics may explain the greater susceptibility to parasite attacks of trees exposed to N deposition and excess N.     

Different reactions of central and marginal provenances of Fagus sylvatica to experimental drought

Climate extremes are expected to increase in frequency and magnitude as a consequence of global warming, threatening the functioning, services and goods of forest ecosystems. Across Europe, the ecologically and economically important tree species Fagus sylvatica is expected to suffer particularly under such conditions. The regional introduction of provenances from drier and warmer climates is one option to adapt beech forest ecosystems to these adverse effects of climate change. Marginal populations from the drought-prone southern and north-eastern edges of the species’ distribution come into focus in search of suitable candidates for Central European deciduous forests. Here, we test three marginal provenances (Spain, Bulgaria and Poland) and three provenances from the centre of the distribution range (Germany) for their response to drought in two different soil types (sand, loam) in a full factorial common garden experiment in Landau, Germany. Drought impacted all growth parameters negatively (leaf damage +22 % (percentage points), height ?40 % and diameter increment ?41 %), and the sandy substrate exacerbated this effect. However, provenances differed in their response to drought and soil type. Evidence for a local adaptation to summer drought was detected, especially in terms of mortality rates. The Bulgarian and Spanish provenance showed a stable performance under drought conditions (BG ?27 % in diameter increment; ES ?32 %), compared to the Polish (?48 %) or the most sensitive German provenances (?57 %), yet for Bulgaria on a low level of total increment. This may indicate a trade-off between drought tolerance and growth. Therefore, a sole focus on drought-resistant marginal provenances seems to not be conducive, as they might be less adapted to other climatic factors, e.g. frost, as well. However, intermixed with local Central European provenances, these may act as functional insurance in future drought-prone forest stands.