Variations in virulence and hyphal growth of four Raffaelea quercivora isolates within Quercus crispula

Mass mortality of fagaceous trees caused by Japanese oak wilt has occurred widely in Japan. Although virulence of the causal fungus, Raffaelea quercivora, appeared to differ among isolates, its relation to the fungal growth within trees was unknown. To clarify the differences in fungal virulence against susceptible Quercus crispula, we examined fungal growth of four R. quercivora isolates within trees and the resulting virulence. In our study, the isolates were multiple‐inoculated in seedlings and single‐inoculated in twigs of mature trees. In the multiple‐inoculation test, mortality rates were examined by the observation of external symptoms. In the single‐inoculation test, water conductance and hyphal growth within the trees were examined by applying aqueous dyes and fluorescence microscopy, respectively. Mortality rates, the proportion of the cross‐sectional area comprising non‐conductive sapwood and horizontal hyphal growth differed significantly among the isolates. Univariate logistic regression analyses showed that both the proportion of non‐conductive sapwood and hyphal growth were significantly positively related to mortality rates. For three isolates, hyphal growth was significantly positively correlated with the proportion of non‐conductive sapwood. These results suggested that the virulence against Q. crispula varies among R. quercivora isolates and that the extent of fungal colonization of the tree determines fungal virulence.     

Variation in leaf morphology of Quercus crispula and Quercus dentata assemblages among contact zones: a method for detection of probable hybridization

I describe variation in leaf morphology of Quercus crispula and Quercus dentata assemblages within and among their contact zones. Such relationships provide a relatively fast and accurate method for detecting hybridization. Using discriminant function (DF) analysis based on leaf characteristics, I was able to differentiate between these two species in pure zones. In three of five contact zones, the DF scores showed broad, continuous distributions with three or more peaks. The trees showing intermediate leaf morphology are considered to be hybrids. In the other two contact zones, the DF scores showed distinct bimodal distributions, suggesting that Q. crispula and Q. dentata rarely hybridize in these sites. Thus, the method described in this study should be effective for detecting probable hybridization between these two species in contact zones, and the relative frequency of hybridization among contact zones. However, there was likely to be intraspecific variation of Q. crispula between geographic ranges, restricting interpretations to patterns occurring within sites or limited geographic ranges.     

Morphological and photosynthetic responses of Quercus crispula seedlings to high-light conditions

We investigated morphological and photosynthetic responses of current-year seedlings of oak (Quercus crispula Blume) under high-light conditions. Quercus crispula seedlings were grown from seed in a relative photosynthetically active photon flux density (RPPFD) of 100, 10 or 2%. There was no difference in total dry mass between 100 and 10% RPPFD. At the end of the growing season, plants grown in 2% RPPFD had a lower total dry mass than those grown in 100 or 10% RPPFD. Seedlings grown in 100% RPPFD showed morphological acclimation, i.e., high root/shoot ratios and high leaf mass per area. De-epoxidation level in the xanthophyll cycle and activity of an antioxidant enzyme were highest in 100% RPPFD, but total chlorophyll concentration and photosynthetic rate were highest in 10% RPPFD. These results indicate that excess photons were generated in 100% RPPFD, leading to increased capacities for dissipation of received light energy through the xanthophyll cycle and for scavenging of reactive oxygen species through the water-water cycle. Nevertheless, a midday decrease in dark-adapted quantum yield of photosystem II (F(v)/F(m)) indicated that seedlings grown in 100% RPPFD were suffering from photoinhibition. We conclude that Q. crispula current-year seedlings have high morphological acclimation to high light but that photosynthetic efficiency cannot be maintained under high-light conditions even with a photoprotection system.     

Seasonal changes in the temperature response of photosynthesis in canopy leaves of Quercus crispula in a cool-temperate forest

Understanding seasonal changes in photosynthetic characteristics of canopy leaves is indispensable for modeling the carbon balance in forests. We studied seasonal changes in gas exchange characteristics that are related to the temperature dependence of photosynthesis in canopy leaves of Quercus crispula Blume, one of the most abundant species in cool-temperate forests in Japan. Photosynthetic rate and ribulose-1,5-bisphosphate (RuBP) carboxylation capacity (V(cmax)) at 20 degrees C increased from June to August and then decreased in September. The activation energy of V(cmax), a measure of the temperature dependence of V(cmax), was highest in summer, indicating that V(cmax) was most sensitive to leaf temperature at this time. The activation energy of V(cmax) was significantly correlated with growth temperature. Other parameters related to the temperature dependence of photosynthesis, such as intercellular CO(2) partial pressure and temperature dependence of RuBP regeneration capacity, showed no clear seasonal trend. It was suggested that leaf senescence affected the balance between carboxylation and regeneration of RuBP. The model simulation showed that photosynthetic rate and its optimal temperature were highest in summer.     

Changes in shoot properties in relation to vertical positions within the crown of mature canopy trees of Abies mariesii and Abies veitchii

We investigated current shoot properties in two contrasting vertical positions (leader crown; LC, and lower branch; LB) within the crowns of mature trees of two subalpine conifer species, Abies mariesii and A. veitchii. For both LCs and LBs, shoot length decreased with increasing branching order. However, shoot properties were different between LCs and LBs. Shoots in LCs had more needle biomass per unit of shoot length. Shoots in sunny conditions pack needles closer along the shoot and intercept incoming light more completely. This causes the shoots in the LCs to have more needles. In contrast, less needle packing per unit shoot length in LBs results in the avoidance of mutual shading among needles in order to intercept limited light more effectively. Because branch systems in lower layers tend to be more shaded, the quantity of irradiance received by the shoots in LBs is smaller. Thus, reduced needle amounts on the shoots in LBs reflect the needle arrangement acclimating to the lower light availability. This study suggests the importance of changes in the properties of individual shoot as a component of a branch system and accordingly a whole-crown system in mature canopy trees of A. mariesii and A. veitchii.     

Vegetation and weather explain variation in crown damage within a large mixed-severity wildfire

The 2002 Biscuit Fire burned through more than 200,000 ha of mixed-conifer/evergreen hardwood forests in southwestern Oregon and northwestern California. The size of the fire and the diversity of conditions through which it burned provided an opportunity to analyze relationships between crown damage and vegetation type, recent fire history, geology, topography, and regional weather conditions on the day of burning. We measured pre- and post-fire vegetation cover and crown damage on 761 digital aerial photo-plots (6.25 ha) within the unmanaged portion of the burn and used random forest and regression tree models to relate patterns of damage to a suite of 20 predictor variables. Ninety-eight percent of plots experienced some level of crown damage, but only 10% experienced complete crown damage. The median level of total crown damage was 74%; median damage to conifer crowns was 52%. The most important predictors of total crown damage were the percentage of pre-fire shrub-stratum vegetation cover and average daily temperature. The most important predictors of conifer damage were average daily temperature and “burn period,” an index of fire weather and fire suppression effort. The median level of damage was 32% within large conifer cover and 62% within small conifer cover. Open tree canopies with high levels of shrub-stratum cover were associated with the highest levels of tree crown damage, while closed canopy forests with high levels of large conifer cover were associated with the lowest levels of tree crown damage. Patterns of damage were similar within the area that burned previously in the 1987 Silver Fire and edaphically similar areas without a recent history of fire. Low-productivity sites on ultramafic soils had 92% median crown damage compared to 59% on non-ultramafic sites; the proportion of conifer cover damaged was also higher on ultramafic sites. We conclude that weather and vegetation conditions — not topography — were the primary determinants of Biscuit Fire crown damage.     

Preformation in vegetative buds of pistachio (Pistacia vera): relationship to shoot morphology, crown structure and rootstock vigor

Effects of rootstock, shoot carbohydrate status, crop load and crown position on the number of preformed leaf primordia in the dormant terminal and lateral buds of mature and immature 'Kerman' pistachio (Pistacia vera L.) trees were investigated to determine if rootstock vigor is associated with greater shoot preformation. There was no significant variation in preformation related to the factors studied, suggesting strong genetic control of preformation in 'Kerman' pistachio. The growth differences observed among trees on different rootstocks were associated with greater stimulation of neoformed growth in trees on the more vigorous rootstocks. However, most annual extension growth in mature tree crowns was preformed, contrasting with the relatively high rate of neoformation found in young tree crowns. Large amounts of neoformed growth in young trees may allow the trees to become established quickly and secure resources, whereas predominantly preformed growth in mature trees may allow for continued crown expansion without outgrowing available resources. We hypothesized that the stimulation of neoformed growth by the more vigorous rootstocks is associated with greater resource uptake or transport, or both. Understanding the source of variation in shoot extension growth on different rootstocks has important implications for orchard management practices.     

Systematic variation in xylem hydraulic capacity within the crown of white ash (Fraxinus americana)

A 7-m tall white ash tree (Fraxinus americana Marsh.) was dissected, and hydraulic parameters of the xylem were determined by inducing a steady-state flow of water through the stem segments and monitoring volume and velocity flow rates. Leaf-specific conductivity (LSC) was highest in the main stem and lowest in some of the leaf-bearing lateral shoots. The LSC was higher in the main stem than in branches and higher in primary than in secondary branch axes. Terminal leaf-bearing shoots were larger and had a significantly greater mean LSC than subjacent lateral shoots. A significant reduction in LSC was associated with the transition between 1- and 2-year-old growth. In branches of the same age, there was a close correspondence among LSC, branch position and branch size. The average LSC of leaf-bearing shoots from south-facing branches was 43% greater than that of shoots from north-facing branches. Within-crown variation in LSC was associated with variation in velocity flow rate (V). By contrast, the ratio of potentially functional xylem area to supported leaf area (A(pf)/A(l)) was relatively stable throughout the crown. Stratification of stems by Strahler order accounted for approximately 70% of the total variation in LSC. These results suggest that (1) there exists a systematic pattern of variation in LSC distribution within the crown of white ash, (2) within-crown variability in LSC is primarily the result of variation in mean vessel diameter, and (3) there is a physiological linkage between LSC and crown morphology that is maintained through a positive feedback mechanism during branch ontogeny.     

Observation of the scale of patchy stomatal behavior in leaves of Quercus crispula using an Imaging-PAM chlorophyll fluorometer

Patchy stomatal closure occurs in plants with heterobaric leaves, in which vertical extensions of bundle sheath cells delimit the mesophyll and restrict the diffusion of CO(2). The scale of patchy stomatal behavior was investigated in this study. The distribution of PSII quantum yield (Φ(II)) obtained from chlorophyll fluorescence images was used to evaluate the scale of stomatal patchiness and its relationship with leaf photosynthesis in the sun leaves of 2-year-old saplings of Quercus crispula Blume. Fluorescent patches were observed only during the day with low stomatal conductance. Comparison of numerical simulation of leaf gas exchange and chlorophyll fluorescence images showed that heterogeneous distribution of electron transport rate through PSII (J) was observed following stomatal closure with a bimodal manner under both natural and saturated photosynthetic photon flux densities. Thus, fluorescence patterns can be interpreted in terms of patchy stomatal closure. The mapping of J from chlorophyll fluorescence images showed that the scale of stomatal patchiness was approximately 2.5-fold larger than that of anatomical patches (lamina areas bounded by bundle sheath extensions within lamina). Our results suggest the spatial scale of stomatal patches in Q. crispula leaves.     

Relative shoot height and irradiance and the shoot and leaf properties of Quercus serrata saplings

Effects of relative shoot height and irradiance on shoot and leaf properties of Quercus serrata Thunb. saplings growing in the understory and in gaps were investigated. Photosynthetic photon flux (PPF) at the location of the shoot relative to that in the open (relative PPF; rPPF) and the height of the shoot base relative to tree height (relative height; rHeight) were measured for all current-year shoots of each sapling. Current-year shoot properties (length, leaf area, number of daughter shoots) and mortality, and leaf properties (mass per area (LMA) and nitrogen content per area (N(area))) were examined in relation to rPPF and rHeight. N(area) was used as a proxy for area-based assimilative capacity. Shoot length, leaf area per shoot and number of daughter shoots increased with increasing rHeight, especially in well-lit conditions. Shoot mortality decreased with increasing rHeight and rPPF. Both LMA and N(area) were positively correlated with rPPF, but not rHeight.     

Steep slopes promote downhill dispersal of Quercus crispula seeds and weaken the fine-scale genetic structure of seedling populations

The seed dispersal patterns and genetic structure of plant populations in mountainous forests may differ from those on flat sites, because some seeds that fall from adults are likely to roll downhill, and thus cause the seed shadows from different mother trees to merge. In the study reported here we used six polymorphic microsatellite markers to track seed dispersal and examine the fine-scale spatial genetic structure of adults and first-year seedlings of Quercus crispula in 2500 m² plots on four slopes. In each of the four plots, leaves of adults, seedlings and endocarps of hypogeal cotyledons attached to the seedlings were genotyped to identify the seedlings’ mother trees. The results showed that steeper slopes result in larger dispersions and smaller genetic structure of seedlings. These findings are a crucial step towards an understanding of the effect of topography on tree regeneration.     

Where to shoot your pilodyn: within tree variation in basic density in plantation Eucalyptus globulus and E. nitens in Tasmania

Longitudinal patterns of within tree variation for basic density were determined for plantation Eucalyptus globulus (ages 5 and 10 years) and E. nitens (ages 5, 10 and 15 years) growing in three geographic areas in Tasmania. Each tree was sampled by taking discs from a combination of percentage heights (0, 10, 20 ___ 70%) and fixed height samples (0.5 m, 0.7 m ___ 1.5 m). At each of the fixed heights, a single pilodyn reading was taken from each of 4 aspects. Weighted whole tree density was calculated from the percentage height samples and used for correlation and regression analysis with the fixed height samples. Both species showed an initial drop in density between the felling cut (zero height) and 0.5 m, followed by a linear increase in density between 10% and 70% of tree height. Slope of fitted regressions were parallel within species but differed significantly between the species (1.14 kg/m³ and 1.62 kg/m³ per 1% increase in height above 10% for E. globulus and E. nitens, respectively). Density at all fixed heights was highly correlated with whole tree values for E. globulus, but results were variable across sites for E. nitens. Both species were found to contain trees which produced aberrant pilodyn readings. For E. globulus, the optimal sampling height was 1.3 m above ground and the mean pilodyn reading was found to predict whole tree density with an accuracy of ±21 kg/m³. For E. nitens, optimal sampling height was 1.5 m above ground. However, pilodyn readings around the stem were not very repeatable and correlations with whole tree density were lower, resulting in the accuracy of prediction of whole tree density being ±26 kg/m³.     

Measurements of the bioelectrical potential of a Japanese oak (Quercus crispula Blume) sapling: Effect of the radial distribution of inorganic ingredients within a tree stem on the diurnal change in resting potential

This study investigated the relation between the resting potential of a 2-year-old Japanese oak (Quercus crispula Blume) sapling and its physiology, especially the radial transport of water containing inorganic ingredients in the stem using the scanning electron microscope-energy dispersive X-ray microanalyzer (SEM-EDXA) method. The resting potential of a sapling could be monitored continuously with our measuring apparatus. Changes in resting potential were due to the light. The hyperpolarization and depolarization peaks of the resting potential, whose absolute voltage was about 10mV, occurred right after lights-off and lights-on, respectively. The resting potential was found to show periodic responses for each day unit. At night (lights-off), the resting potential tended to depolarize with an increase in tangential strain. On the other hand, during the daytime (lights-on) the resting potential tended to hyperpolarize, depolarize, or show a nearly constant value for the tangential strain. The water containing inorganic ingredients was transported, via transpiration, in both directions between the mature xylem zone and the phloem zone through differentiating xylem cells. This water transport within a tree stem had a significant effect on the diurnal changes in resting potential.Part of this paper was presented at the 47th annual meeting of the Japan Wood Research Society, Kochi, April 3–5, 1997     

The Timing of Canopy and Epicormic shoot growth in Quercus robur L.

The sequence and timing of shoot development from buds on thebranches and trunk of Quercus robur were observed over 2 years.Shoots on crown branches began development before those on existingepicormic branches but both stopped growing at about the sametime. Epicormic buds on the trunk began expanding 3–4weeks after crown buds, and finished growth 3 weeks later. Theseobservations do not support previous claims that new epicormicshoots grow before development of the canopy.     

Response of endophytic Biscogniauxia mediterranea to variation in leaf water potential of Quercus cerris

Endophytic behaviour of Biscogniauxia mediterranea, the causal agent of charcoal disease of oak, was studied over two growing seasons on Quercus cerris in a forest in central Italy. Isolation of the fungus from asymptomatic tissues varied with tissue type, period of sampling and year. Presence of the fungus in asymptomatic plants was significantly higher in fall, compared with that in spring and summer. However, a significant effect of tissue water content on the presence of B. mediterranea was recognized. The results suggest that proliferation of B. mediterranea is favoured during the endophytic phase by a decrease in host water potential.     

Drought susceptibility and recovery of transplanted Quercus rubra seedlings in relation to root system morphology

? Transplant shock, implicated by depressed seedling physiological status associated with moisture stress immediately following planting, limits early plantation establishment. Large root volume (Rv) has potential to alleviate transplant shock because of higher root growth potential and greater access to soil water.

? We investigated impacts of drought and transplant Rv on photosynthetic assimilation (A), transpiration (E), stomatal conductance (g _s ), predawn leaf xylem water potential (Ψ_L), and growth of northern red oak (Quercus rubra L.) seedlings to explain mechanisms associated with susceptibility to transplant shock. One year-old barerooot seedlings were graded into four Rv categories and either well watered or subjected to drought consisting of low, medium, or high moisture stress by discontinuing irrigation at 22-day intervals for 3 months. Thereafter, all treatments were re-watered to examine recovery.

? Transplant shock was signified by reduced A, E, g _s, and Ψ_L, which generally increased with increasing moisture stress and Rv. Physiological status improved during recovery, though stress was still evident in seedlings exposed to medium or high moisture stress and in larger Rv seedlings. Growth declined with increasing moisture stress but was generally similar among Rv treatments, likely reflecting greater A at the whole plant level and/or reliance upon stored reserves in large Rv seedlings.

? The most effective drought avoidance mechanisms were root growth, stomatal regulation, reduced leaf area, and higher growth allocation to roots relative to shoots. Our results suggest that large initial Rv does not enhance drought avoidance during the first season after transplant in northern red oak seedlings.

     

Factors affecting post-fire crown regeneration in cork oak (Quercus suber L.) trees

Cork oak (Quercus suber) forests are acknowledged for their biodiversity and economic (mainly cork production) values. Wildfires are one of the main threats contributing to cork oak decline in the Mediterranean Basin, and one major question that managers face after fire in cork oak stands is whether the burned trees should be coppiced or not. This decision can be based on the degree of expected crown regeneration assessed immediately after fire. In this study we carried out a post-fire assessment of the degree of crown recovery in 858 trees being exploited for cork production in southern Portugal, 1.5 years after a wildfire. Using logistic regression, we modelled good or poor crown recovery probability as a function of tree and stand variables. The main variables influencing the likelihood of good or poor crown regeneration were bark thickness, charring height, aspect and tree diameter. We also developed management models, including simpler but easier to measure variables, which had a lower predictive power but can be used to help managers to identify, immediately after fire, trees that will likely show good crown regeneration, and trees that will likely die or show poor regeneration (and thus, potential candidates for trunk coppicing).     

Extractives of Quercus crispula sapwood infected by the pathogenic fungus Raffaelea quercivora II: isolation and identification of phenolic compounds from infected sapwood

There has been a mass mortality of oak trees in the area along the coast of the Japan Sea. This phenomenon is caused by the ambrosia beetle Platypus quercivorus, which carries the ambrosia fungus Raffaelea quercivora. Extractives of a necrotic brownish coloration formed in the infected sapwood of Quercus crispula were investigated. The methanol extract of the damaged sapwood of Q. crispula was concentrated in vacuo and centrifuged to yield precipitates and the supernatant. The precipitates were subjected to Sephadex LH-20 column chromatography and preparative HPLC to describe a novel ellagic acid derivative (1). The 10 % methanol water-soluble part of the supernatant was subjected to medium-pressure ODS column chromatography and preparative HPLC, respectively, to analyze a known lignan (2). Sulfuric acid hydrolysis of (1) yielded an ellagic acid and a gallic acid. NMR and LC-TOF/MS indicated that an ellagic acid and a gallic acid bonded to a xylose with glycosidic and ester bonds, respectively. Compound (1) was identified as 4,5-dihydroxy-6-(3,7,8–trihydroxy-5,10-dihydro–chromeno[5,4,3-cde]chromen-2-yloxy)-tetrahydro-pyran-3-yl ester, and compound (2) was identified as (–)-lyoniresinol. The presence of (–)-lyoniresinol from damaged sapwood indicated that infection of R. quercivora may cause the formation of a pseudo-heartwood in the sapwood of Q. crispula.     

Reduced photosynthesis in old oak (Quercus robur): the impact of crown and hydraulic architecture

We tested the hypothesis that changes in crown architecture of old pedunculate oak trees (Quercus robur L. ssp. robur Kl. et Kr. et Rol.) reduce leaf specific hydraulic conductance of shoots, thereby limiting stomatal conductance and assimilation of affected shoots. At the end of summer 1999, hydraulic conductance and leaf specific hydraulic conductance, measured with a high-pressure flow meter in 0.5- to 1.5-m long shoots, were 27 and 39% lower, respectively, in shoots of low vigor compared with vigorously growing shoots in a 165-year- old stand in southeastern Germany. Two types of bottlenecks to water transport can be identified in shoots of old oak trees, namely nodes and abscission zones. The reduction in hydraulic conductance was especially severe in shoots with diameters of less than 2 mm. Maximum stomatal conductance and maximum net assimilation rate increased significantly with hydraulic conductance and leaf specific hydraulic conductance. Our data support the hypothesis that changes in shoot and consequently crown architecture observed in aging trees can limit photosynthesis by reducing shoot hydraulic conductance. Thus, in addition to increasing pathway length and lower conductivity of xylem in old trees, structural changes in shoot and crown architecture need to be considered when analyzing water relations and photosynthesis in mature and declining trees.