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Global warming affects the abiotic and biotic growth environment of plants, including the spread of fungal diseases such as Dutch elm disease (DED). Dutch elm disease-resistance of different Ulmus species varies, but how this is reflected in leaf-level physiological pathogen responses has not been investigated. We studied the impacts of mechanical injury alone and mechanical injury plus inoculation with the DED-causing pathogens Ophiostoma novo-ulmi subsp. novo-ulmi and O. novo-ulmi subsp. americana on Ulmus glabra , a more vulnerable species, and U. laevis , a more resistant species. Plant stress responses were evaluated for 12 days after stress application by monitoring leaf net CO2 assimilation rate (A), stomatal conductance (g s), ratio of ambient to intercellular CO2 concentration (C a/ C i) and intrinsic water-use efficiency (A / g s), and by measuring biogenic volatile (VOC) release by plant leaves. In U. glabra and U. laevis , A was not affected by time, stressors or their interaction. Only in U. glabra , g s and C a/ C i decreased in time, yet recovered by the end of the experiment. Although the emission compositions were affected in both species, the stress treatments enhanced VOC emission rates only in U. laevis. In this species, mechanical injury especially when combined with the pathogens increased the emission of lipoxygenase pathway volatiles and dimethylallyl diphosphate and geranyl diphosphate pathway volatiles. In conclusion , the more resistant species U. laevis had a more stable photosynthesis, but stronger pathogen-elicited volatile response, especially after inoculation by O. novo-ulmi subsp. novo-ulmi. Thus, stronger activation of defenses might underlay higher DED-resistance in this species. [ABSTRACT FROM AUTHOR]

Synergistic regulation in leaf architecture and photosynthesis is essential for salt tolerance. However, how plant sex and inorganic nitrogen sources alter salt stress-dependent photosynthesis remains unknown. Leaf anatomical characteristics and photosynthesis of Populus cathayana Rehder females and males were investigated under salt stress conditions combined with nitrate NO3− and ammonium NH4+ supplies to clarify the underlying mechanisms. In salt-stressed females, we observed an increased mesophyll spongy cell density, a reduced chloroplast density, a decreased surface area of chloroplasts adjacent to the intercellular air space (S c/ S) and an increased mesophyll cell area per transverse section width (S / W), consequently causing mesophyll conductance (g m) and photosynthesis inhibition, especially under NH4+ supply. Conversely, males with a greater mesophyll palisade tissue thickness and chloroplast density, but a lower spongy cell density had lower S / W and higher S c/ S , and higher g m and photosynthesis. NH4+-fed females had a lower CO2 conductance through cell wall and stromal conductance perpendicular to the cell wall, but a higher chloroplast conductance from the cell wall (g cyt1) than females supplied with NO3−, whereas males had a higher chloroplast conductance and lower CO2 conductance through cell wall when supplied with NO3− instead of NH4+ under salt stress. These findings indicate sex-specific strategies in coping with salt stress related to leaf anatomy and g m under both types of nitrogen supplies, which may contribute to sex-specific CO2 capture and niche segregation. [ABSTRACT FROM AUTHOR]

Highly host-specific eriophyoid gall- and erineum-forming mites infest a limited range of broadleaf species, with the mites from the genus Eriophyes particularly widespread on Alnus spp. and Tilia spp. Once infected, the infections can be massive, covering a large part of leaf area and spreading through the plant canopy, but the effects of Eriophyes mite gall formation on the performance of host leaves are poorly understood. We studied the influence of three frequent Eriophyes infections, E. inangulis gall-forming mites on Alnus glutinosa , and E. tiliae gall-forming and E. exilis erineum-forming mites on Tilia cordata , on foliage morphology, chemistry, photosynthetic characteristics, and constitutive and induced volatile emissions. For all types of infections, leaf dry mass per unit area, net assimilation rate per area and stomatal conductance strongly decreased with increasing severity of infection. Mite infections resulted in enhancement or elicitation of emissions of fatty acid-derived volatiles, isoprene, benzenoids and carotenoid breakdown products in an infection severity-dependent manner for all different infections. Monoterpene emissions were strongly elicited in T. cordata mite infections, but these emissions were suppressed in E. inangulis -infected A. glutinosa. Although the overall level of mite-induced emissions was surprisingly low, these results highlight the uniqueness of the volatile profiles and offer opportunities for using volatile fingerprints and overall emission rates to diagnose infections by Eriophyes gall- and erineum-forming mites on temperate trees and assess their impact on the physiology of the affected trees. [ABSTRACT FROM AUTHOR]

Pine (Pinus) species exhibit extensive variation in needle shape and size between juvenile (primary) and adult (secondary) needles (heteroblasty), but few studies have quantified the changes in needle morphological, anatomical and chemical traits upon juvenile-to-adult transition. Mediterranean pines keep juvenile needles longer than most other pines, implying that juvenile needles play a particularly significant role in seedling and sapling establishment in this environment. We studied needle anatomical, morphological and chemical characteristics in juvenile and different-aged adult needles in Mediterranean pines Pinus halepensis Mill., Pinus pinea L. and Pinus nigra J. F. Arnold subsp. salzmannii (Dunal) Franco hypothesizing that needle anatomical modifications upon juvenile-to-adult transition lead to a trade-off between investments in support and photosynthetic tissues and that analogous changes occur with needle aging albeit to a lower degree. Compared with adult needles, juvenile needles of all species were narrower with 1.6- to 2.4-fold lower leaf dry mass per unit area, and had ∼1.4-fold thinner cell walls, but needle nitrogen content per dry mass was similar among plant ages. Juvenile needles also had ∼1.5-fold greater mesophyll volume fraction, ∼3- fold greater chloroplast volume fraction and ∼1.7-fold greater chloroplast exposed to mesophyll exposed surface area ratio, suggesting overall greater photosynthetic activity. Changes in needle traits were similar in aging adult needles, but the magnitude was generally less than the changes upon juvenile to adult transition. In adult needles, the fraction in support tissues scaled positively with known ranking of species tolerance of drought (P. halepensis > P. pinea > P. nigra). Across all species and needle and plant ages, a negative correlation between volume fractions of mesophyll and structural tissues was observed, manifesting a trade-off between biomass investments in different needle functions. These results demonstrate that within the broad trade-off, juvenile and adult needle morphophysiotypes are separated by varying investments in support and photosynthetic functions. We suggest that the ecological advantage of the juvenile morphophysiotype is maximization of carbon gain of establishing saplings, while adult needle physiognomy enhances environmental stress tolerance of established plants. [ABSTRACT FROM AUTHOR]

The sex ratios of co-existing Populus and Salix vary depending on altitude and species: 1:1 equlibrium sex ratios are observed at mid-altitude but skewed ones at high altitudes, where Populus shows male-biased and Salix female-biased sex ratios. However, the underlying ecological mechanisms are poorly known. Reproductive investments of Populus purdomii Rehd. and Salix magnifica Hemsl. were assessed at altitudes of 2000 and 2600 m in the Gongga Mountain by different metrics, including biomass, carbon (C), nitrogen (N) and phosphorus (P) concentrations and construction cost, and by estimating the payback time that combines energy gain and associated costs. Reproductive investment measured as C, N and P concentrations, and construction cost was higher in P. purdomii females at 2600 m. However, in S. magnifica, no difference was observed for biomass, C and N at 2600 m, but the investments for P and construction cost were even greater in males. The payback time showed no significant differences for the sexes at 2000 m, but it was shorter for P. purdomii males and S. magnifica females at 2600 m. We concluded that nutrient- and construction cost-based estimates of reproductive allocation can provide more informative insight into the cost of reproduction than does biomass, and together with the payback time they can supply an explanation for divergent sex ratios in Populus and Salix. Consequently, our results improve our understanding of the causes and consequences of sexual dimorphism in dioecious species. [ABSTRACT FROM AUTHOR]

Leaf dry mass per unit area (LMA) has been suggested to negatively affect the mesophyll conductance to CO2 (gm), the most limiting factor for photosynthesis per unit leaf area (AN) in many evergreens. Several anatomical traits (i.e., greater leaf thickness and thicker cell walls) constraining gm could explain the negative scaling of gm and AN with LMA across species. However, the Mediterranean sclerophyll Quercus ilex L. shows a major within-species variation in functional traits (greater LMA associated with higher nitrogen content and AN) that might contrast the worldwide trends. The objective of this study was to elucidate the existence of variations in other leaf anatomical parameters determining gm and/or biochemical traits improving the capacity of carboxylation (Vc,max) that could modulate the relationship of AN with LMA across this species. The results revealed that gm was the most limiting factor for AN in all the studied Q. ilex provenances from Spain and Italy. The within-species differences in gm can be partly attributed to the variation in several leaf anatomical traits, mainly cell-wall thickness (Tcw), chloroplast thickness (Tchl) and chloroplast exposed surface area facing intercellular air spaces (Sc/S). A positive scaling of gm and AN with Vc,max was also found, associated with an increased nitrogen content per area. A strong correlation of maximum photosynthetic electron transport (Jmax) with AN further indicated a coordination between the carboxylase activity and the electron transport chain. In conclusion, we have confirmed the strong ecotypic variation in the photosynthetic performance of individual provenances of Q. ilex. Thus, the withinspecies increases found in AN for Q. ilex with increasing foliage robustness can be explained by a synergistic effect among anatomical (at the subcellular and cellular level) and biochemical traits, which markedly improved gm and Vc,max. [ABSTRACT FROM AUTHOR]

In this study, intra- and interspecific competition were investigated in early successional Salix rehderiana Schneider and laterappearing Populus purdomii Rehder under non-fertilized (control) and nitrogen (N)-fertilized conditions in the Hailuogou glacier retreat area. Our aim was to discover whether N is a key factor in plant-plant competition and whether N drives the primary succession process in a glacier retreat area. We analyzed differences in responses to intra- and interspecific competition and N fertilization between P. purdomii and S. rehderiana, including parameters such as biomass accumulation, nutrient absorption, non-structural carbohydrates, photosynthetic capacity, hydrolysable amino acids and leaf ultrastructure. In the control treatments, S. rehderiana individuals subjected to interspecific competition benefited from the presence of P. purdomii plants, as indicated by higher levels of biomass accumulation, photosynthetic capacity, N absorption, amino acid contents and photosynthetic N-use efficiency. However, in the N-fertilized treatments, P. purdomii individuals exposed to interspecific competition benefited from the presence of S. rehderiana plants, as shown by a higher growth rate, enhanced carbon gain capacity, greater amino acid contents, and elevated water-use efficiency, whereas the growth of S. rehderiana was significantly reduced. Our results demonstrate that N plays a pivotal role in determining the asymmetric competition pattern among Salicaceae species during primary succession. We argue that the interactive effects of plant-plant competition and N availability are key mechanisms that drive primary succession in the Gongga Mountain glacier retreat area. [ABSTRACT FROM AUTHOR]

Sexual competition among dioecious plants affects sex ratios and the spatial distribution of the sexes in different environments. At present, little is known about sexual dimorphisms induced by different competition patterns under salinity stress. We employed Populus deltoides as a model to investigate sex-related growth as well as physiological and biochemical responses to salinity stress under conditions of intrasexual and intersexual competition. Potted seedlings (two seedlings per pot; two females, two males, or one female and one male) were exposed to two salt levels (0 and 50 mM NaCl) and salinity-and competition-driven differences in growth, assimilation rate, water use, contents of leaf pigments and osmotica, hydrogen peroxide (H2O2), and antioxidant enzyme and nitrate reductase activity were examined. In the absence of salinity, no significant differences in competitive ability between males and females subjected to intrasexual competition were observed, although the growth of females was moderately greater under intersexual competition. The salinity treatment significantly increased the sex differences in competitive ability, especially under intersexual competition. Under salinity stress, males showed decreased height, but displayed greater capacity for osmotic adjustment, enhancement of long-term water-use efficiency and increase in antioxidant enzyme activities. The absolute values of these traits were greater in salt-stressed males than in females under intersexual competition. In addition, salt-stressed males accumulated less Cl-and had lower H2O2 contents than females. These data collectively demonstrate that the competitive advantage of females in non-stressed conditions is lost under salinity. Greater salinity resistance of males growing intermixed with females under salt stress can importantly affect the sex ratio of P. deltoides populations. [ABSTRACT FROM AUTHOR]

Although increasing attention has been paid to plant adaptation to soil heavy metal contamination, competition and neighbor effects have been largely overlooked, especially in dioecious plants. In this study, we investigated growth as well as biochemical and ultrastructural responses of Populus cathayana Rehder females and males to cadmium (Cd) stress under different sexual competition patterns. The results showed that competition significantly affects biomass partitioning, photosynthetic capacity, leaf and root ultrastructure, Cd accumulation, the contents of polyphenols, and structural and nonstructural carbohydrates. Compared with single-sex cultivation, plants of opposite sexes exposed to sexual competition accumulated more Cd in tissues and their growth was more strongly inhibited, indicating enhanced Cd toxicity under sexual competition. Under intrasexual competition, females showed greater Cd accumulation, more serious damage at the ultrastructural level and greater reduction in physiological activity than under intersexual competition, while males performed better under intrasexual competition than under intersexual competition. Males improved the female microenvironment by greater Cd uptake and lower resource consumption under intersexual competition. These results demonstrate that the sex of neighbor plants and competition affect sexual differences in growth and in key physiological processes under Cd stress. The asymmetry of sexual competition highlighted here might regulate population structure, and spatial segregation and phytoremediation potential of both sexes in P. cathayana growing in heavy metal-contaminated soils. [ABSTRACT FROM AUTHOR]

We employed the warm temperate conifer Cunninghamia lanceolata (Lamb.) Hook. as a model of plantation forest species to investigate ecophysiological responses to root treatments (control (0%), and ~25, 50 or 75% of the initial root mass) under well-watered and water-limited conditions. Our results indicated that total root dry mass accumulation was negatively associated with the severity of root pruning, but there was evidence of multiple compensatory responses. The plants exhibited higher instantaneous and long-term (assessed by carbon isotope composition, δ13C) water-use efficiency in pruning treatments, especially under low water availability. Root pruning also increased the fine root/total root mass ratio, specific root length and fine root vitality in both water availability treatments. As a result of the compensatory responses, under well-watered conditions, height, stem dry mass accumulation, leaf/fine root biomass ratio (L/FR), transpiration rate, photosynthetic capacity and photosynthetic nitrogen-use efficiency (EN) were the highest under 25% pruning. Yet, all these traits except L/FR and foliage nitrogen content were severely reduced under 75% pruning. Drought negatively affected growth and leaf gas exchange rates, and there was a greater negative effect on growth, water potential, gas exchange and EN when >25% of total root biomass was removed. The stem/aboveground mass ratio was the highest under 25% pruning in both watering conditions. These results indicate that the responses to root severance are related to the excision intensity and soil moisture content. A moderate root pruning proved to be an effective means to improve stem dry mass accumulation. [ABSTRACT FROM AUTHOR]

Fungal infections result in decreases in photosynthesis, induction of stress and signaling volatile emissions and reductions in constitutive volatile emissions, but the way different physiological processes scale with the severity of infection is poorly known. We studied the effects of infection by the obligate biotrophic fungal pathogen Melampsora larici-populina Kleb., the causal agent of poplar leaf rust disease, on photosynthetic characteristics, and constitutive isoprene and induced volatile emissions in leaves of Populus balsamifera var. suaveolens (Fisch.) Loudon. exhibiting different degrees of damage. The degree of fungal damage, quantified by the total area of chlorotic and necrotic leaf areas, varied between 0 (noninfected control) and ~60%. The rates of all physiological processes scaled quantitatively with the degree of visual damage, but the scaling with damage severity was weaker for photosynthetic characteristics than for constitutive and induced volatile release. Over the whole range of damage severity, the net assimilation rate per area (AA) decreased 1.5-fold, dry mass per unit area 2.4-fold and constitutive isoprene emissions 5-fold, while stomatal conductance increased 1.9-fold and dark respiration rate 1.6-fold. The emissions of key stress and signaling volatiles (methanol, green leaf volatiles, monoterpenes, sesquiterpenes and methyl salicylate) were in most cases nondetectable in noninfested leaves, and increased strongly with increasing the spread of infection. The moderate reduction in AA resulted from the loss of photosynthetically active biomass, but the reduction in constitutive isoprene emissions and the increase in induced volatile emissions primarily reflected changes in the activities of corresponding biochemical pathways. Although all physiological alterations in fungal-infected leaves occurred in a stress severity-dependent manner, modifications in primary and secondary metabolic pathways scaled differently due to contrasting operational mechanisms. [ABSTRACT FROM AUTHOR]

Leaves growing in the forest understory usually present a decreased mesophyll conductance (gm) and photosynthetic capacity. The role of leaf anatomy in determining the variability in gm among species is known, but there is a lack of information on how the acclimation of gm to shade conditions is driven by changes in leaf anatomy. Within this context, we demonstrated that Abies pinsapo Boiss. experienced profound modifications in needle anatomy to drastic changes in light availability that ultimately led to differential photosynthetic performance between trees grown in the open field and in the forest understory. In contrast to A. pinsapo, its congeneric Abies alba Mill. did not show differences either in needle anatomy or in photosynthetic parameters between trees grown in the open field and in the forest understory. The increased gm values found in trees of A. pinsapo grown in the open field can be explained by occurrence of stomata at both needle sides (amphistomatous needles), increased chloroplast surface area exposed to intercellular airspace, decreased cell wall thickness and, especially, decreased chloroplast thickness. To the best of our knowledge, the role of such drastic changes in ultrastructural needle anatomy in explaining the response of gm to the light environment has not been demonstrated in field conditions. [ABSTRACT FROM AUTHOR]

The degree to which branches are autonomous in their acclimation responses to alteration in light environment is still poorly understood. We investigated the effects of shading of the sapling crown of Cunninghamia lanceolata (Lamb.) Hook on the wholetree and mid-crown branch growth and current-year foliage structure and physiology. Four treatments providing 0, 50, 75 and 90% shading compared with full daylight (denoted as Treatment0, Treatment50%, Treatment75% and Treatment90%, and Shaded0, Shaded50%, Shaded75% and Shaded90% for the shaded branches and Sunlit0, Sunlit50%, Sunlit75% and Sunlit90% for the opposite sunlit branches under natural light conditions, respectively), were applied over two consecutive growing seasons. Shading treatments decreased the growth of basal stem diameter, leaf dry mass per unit leaf area, stomatal conductance, transpiration rate, the ratio of water-soluble to structural leaf nitrogen content, photosynthetic nitrogen-use efficiency and instantaneous and longterm (estimated from carbon isotope composition) water-use efficiency in shaded branches. Differences between shaded and sunlit branches increased with increasing severity and duration of shading. A non-autonomous, partly compensatory behavior of non-shaded branches was observed for most traits, thus reflecting the dependence between the traits of sunlit branches and the severity of shading of the opposite crown half. The results collectively indicated that tree growth and branch and leaf acclimation responses of C. lanceolata are not only affected by the local light environment, but also by relative within-crown light conditions. We argue that such a non-autonomous branch response to changes in light conditions can improve whole-tree resource optimization. These results contribute to better understanding of tree growth and utilization of water and nitrogen under heterogeneous light conditions within tree canopies. [ABSTRACT FROM AUTHOR]

Acclimation potential of needle photosynthetic capacity varies greatly among pine species, but the underlying chemical, anatomical and morphological controls are not entirely understood. We investigated the light-dependent variation in needle characteristics in individuals of Pinus patula Schlect. & Cham., which has 19–31-cm long pendulous needles, and individuals of P. radiata D. Don., which has shorter (8–17-cm-long) stiffer needles. Needle nitrogen and carbon contents, mesophyll and structural tissue volume fractions, needle dry mass per unit total area (MA) and its components, volume to total area ratio (V/AT) and needle density (D = MA/(V/AT)), and maximum carboxylase activity of Rubisco (Vcmax) and capacity of photosynthetic electron transport (Jmax) were investigated in relation to seasonal mean integrated irradiance (Qint). Increases in Qint from canopy bottom to top resulted in proportional increases in both needle thickness and width such that needle total to projected surface area ratio, characterizing the efficiency of light interception, was independent of Qint. Increased light availability also led to larger MA and nitrogen content per unit area (NA). Light-dependent modifications in MA resulted from increases in bothV/AT and D, whereas NA changed because of increases in both MA and mass-based nitrogen content (NM) (NA = NMMA). Overall, the volume fraction of mesophyll cells increased with increasing irradiance and V/AT as the fraction of hypodermis and epidermis decreased with increasing needle thickness. Increases in MA and NA resulted in enhancedJmax and Vcmax per unit area in both species, but mass-based photosynthetic capacity increased only in P. patula. In addition,Jmax and Vcmax showed greater plasticity in response to light in P. patula. Species differences in mesophyll volume fraction explained most of the variation in mass-based needle photosynthetic capacity between species, demonstrating that differences in plastic adjustments in mass-based photosynthetic activities among these representative individuals were mainly associated with contrasting investments in mesophyll cells. Greater area-based photosynthetic plasticity in P. patula relative to P. radiata was associated with larger increases in MA and mesophyll volume fraction with increasing irradiance. These data collectively demonstrate that light-dependent increases in mass-based nitrogen contents and photosynthetic activities were associated with an increased mesophyll volume fraction in needles at higher irradiances. They also emphasize the importance of light-dependent anatomical modifications in determining needle photosynthetic capacity. [ABSTRACT FROM PUBLISHER]

Leaf hydraulic conductance (Kleaf) 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 Kleaf 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, Kleaf 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 Kleaf 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 Kleaf 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 Kleaf among leaves growing under different environmental conditions. [ABSTRACT FROM AUTHOR]