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Cuticular wax layer is the first barrier against the outside environment and the first defense encountered by herbivores and pathogens. The effects of environmental factors on cuticular chemistry, and on the formation of glandular trichomes that account for the storage and secretion of lipophilic compounds to the leaf surface are poorly understood. Low vapor pressure deficit (VPD) has shown to reduce the nitrogen (N) status of plants. Thus, we studied the effects of elevated air humidity, indicated as VPD, and the effect of N fertilization on cuticular waxes and glandular trichome density in silver birch (Betula pendula Roth). Experiments were carried out in growth chambers with juvenile plants and in a long-term field experiment with older trees. Low VPD reduced the glandular trichome density in both experiments, in chamber and in field. The contents of the major triterpenoid and flavonoid aglycones correlated positively with glandular trichome density, which supports the role of trichomes in the exudation of secondary compounds to the leaf surface. A closer examination of the cuticular wax chemistry in the chamber experiment revealed that low VPD and N supply affected the composition of cuticular waxes, but not the total wax content. The deposition of different wax compounds followed a co-ordinated pattern in birch leaves, but different compound groups varied in their responses to N fertilization and low VPD. Low VPD reduced the hydrophobicity of cuticular waxes, as demonstrated by lower alkane content and less hydrophobic flavonoid profile in low VPD than in high VPD. Reduced hydrophobicity of the wax layer is presumed to increase leaf wettability. Together with reduced trichome density in low VPD it may enhance the susceptibility of trees to fungal pathogens and herbivores. High N supply under low VPD reduced the effect of low VPD on the cuticular wax composition. Total fatty acid content and the expression of β-amyrin synthase were lower under high N supply than under moderate N supply irrespective of VPD treatment. Nitrogen availability and decreasing VPD will modify leaf surface properties in silver birch and thereby affect tree defence against abiotic and biotic stress factors that emerge under climate change.

Correlations between leaf abscisic acid concentration ([ABA]), stomatal conductance (g s ), rate of stomatal opening in response to an increase in leaf water potential (s i ), shoot hydraulic conductance (L) and photosynthetic characteristics were examined in saplings of six temperate deciduous tree species: Acer platanoides L., Padus avium Mill., Populus tremula L., Quercus robur L., Salix caprea L. and Tilia cordata Mill. Species-specific values of foliar [ABA] were negatively related to the mean values of g s , s i , L and light- and CO 2 - saturated net photosynthesis (P max ), thus providing strong correlative evidence of a scaling of foliar gas exchange and hydraulic characteristics with leaf endogenous [ABA]. In addition, we suggest that mean g s , s i , L and P max for mature leaves may partly be determined by the species-specific [ABA] during leaf growth. The most drought-intolerant species had the lowest [ABA] and the highest g s , suggesting that interspecific differences in [ABA] may be linked to differences in species-specific water-use efficiency. Application of high concentrations of exogenous ABA led to large decreases in g s , s i and P max , further underscoring the direct role of ABA in regulating stomatal opening and photosynthetic rate. Exogenous ABA also decreased L, but the decreases were considerably smaller than the decreases in g s , s i and P max . Thus, exogenous ABA predominantly affected the stomata directly, but modification of L by ABA may also be an important mechanism of ABA action. We conclude that interspecific variability in endogenous [ABA] during foliage growth and in mature leaves provides an important factor explaining observed differences in L, g s , s i and P max among temperate deciduous tree species.

To clarify interactions between stomatal responses to two simultaneous environmental changes, the rates of change in stomatal conductance were measured after simultaneously changing two environmental factors from the set of air humidity, leaf water potential (hydraulic environmental factors), air CO(2) concentration and light intensity (photosynthetic environmental factors). The stomatal responses to changes in leaf water potential were not significantly modified by any other simultaneous environmental change. A decrease in air humidity was followed by a decrease in stomatal conductance, and an increase in air humidity was followed by an increase in the conductance, irrespective of the character of the simultaneous change in the photosynthetic environmental factor. If the simultaneous change had an opposite effect on stomatal conductance, the rate of change in stomatal conductance was higher than the theoretical summed rate-the sum of the rate following one environmental change and the rate following another environmental change, measured separately. That is, the stomatal response to air humidity dominated over the responses to photosynthetic environmental factors. Yet, if the simultaneous change in photosynthetic factors had a codirectional effect on stomatal conductance, the rate of stomatal conductance change was lower than the theoretical summed rate. After a simultaneous change of two photosynthetic environmental factors, the rate of stomatal conductance change was very similar to the theoretical rate, if both the environmental changes had a codirectional effect on stomatal conductance. If the changes in the photosynthetic factors had opposite effects on stomatal conductance, the conductance increased, irrespective of the character of the increasing environmental factor. In drought-stressed trees, the rates of change in stomatal conductance tended to differ from the theoretical summed rates more than in well-watered trees. Stomatal closure following an increase in CO(2) concentration was the stomatal response that was most strongly suppressed by the response to another simultaneous environmental change. Six species of temperate deciduous trees were shown to be similar in their relations between the stomatal responses to two simultaneous environmental changes. The mechanism and ecological significance of the interactions between the two signal response pathways of stomata are discussed.

Paper birch (Betula papyrifera Marsh.) and three trembling aspen clones (Populus tremuloides Michx.) were studied to determine if alterations in carbon gain in response to an elevated concentration of CO 2 ([CO 2 ]) or O 3 ([O 3 ]) or a combination of both affected bud size and carbohydrate composition in autumn, and early leaf development in the following spring. The trees were measured for gas exchange, leaf size, date of leaf abscission, size and biochemical characteristics of the overwintering buds and early leaf development during the 8th-9th year of free-air CO 2 and O 3 exposure at the Aspen FACE site located near Rhinelander, WI. Net photosynthesis was enhanced 49-73% by elevated [CO 2 ], and decreased 13-30% by elevated [O 3 ]. Elevated [CO 2 ] delayed, and elevated [O 3 ] tended to accelerate, leaf abscission in autumn. Elevated [CO 2 ] increased the ratio of monosaccharides to di- and oligosaccharides in aspen buds, which may indicate a lag in cold acclimation. The total carbon concentration in overwintering buds was unaffected by the treatments, although elevated [O 3 ] decreased the amount of starch by 16% in birch buds, and reduced the size of aspen buds, which may be related to the delayed leaf development in aspen during the spring. Elevated [CO 2 ] generally ameliorated the effects of elevated [O 3 ]. Our results show that both elevated [CO 2 ] and elevated [O 3 ] have the potential to alter carbon metabolism of overwintering buds. These changes may cause carry-over effects during the next growing season.

Leaf hydraulic conductance (K(leaf)) and several characteristics of hydraulic architecture and physiology were measured during the first 10 weeks of leaf ontogeny in Populus tremula L. saplings growing under control, mild water deficit or elevated temperature conditions. During the initial 3 weeks of leaf ontogeny, most measured characteristics rapidly increased. Thereafter, a gradual decrease in K(leaf) was correlated with a decrease in leaf osmotic potential under all conditions, and with increases in leaf dry mass per area and bulk modulus of elasticity under mild water deficit and control conditions. From about Week 3 onward, K(leaf) was 33% lower in trees subjected to mild water deficit and 33% higher in trees held at an elevated temperature relative to control trees. Mild water deficit and elevated temperature treatment had significant and opposite effects on most of the other characteristics measured. The ontogenetic maximum in K(leaf) was correlated positively with the width of xylem conduits in the midrib, but negatively with the overall width of the midrib xylem, number of lateral ribs, leaf dry mass per area and bulk modulus of elasticity. The ontogenetic maximum in K(leaf) was also correlated positively with the proportion of intercellular spaces and leaf osmotic potential, but negatively with leaf thickness, volume of mesophyll cells and epidermis and number of cells per total mesophyll cell volume, the closest relationships being between leaf osmotic potential and number of cells per total mesophyll cell volume. It was concluded that differences in protoplast traits are more important than differences in xylem or parenchymal cell wall traits in determining the variability in K(leaf) among leaves growing under different environmental conditions.