Cavitation fatigue in conifers: a study on eight European species.

After drought-induced embolism and repair, tree xylem may be weakened against future drought events (cavitation fatigue). As there are few data on cavitation fatigue in conifers available, we quantified vulnerability curves (VCs) after embolism/repair cycles on eight European conifer species. We induced 50% and 100% loss of conductivity (LC) with a cavitron, and analyzed VCs. Embolism repair was obtained by vacuum infiltration. All species demonstrated complete embolism repair and a lack of any cavitation fatigue after 50% LC . After 100% LC, European larch (Larix decidua), stone pine (Pinus cembra), Norway spruce (Picea abies), and silver fir (Abies alba) remained unaffected, while mountain pine (Pinus mugo), yew (Taxus baccata), and common juniper (Juniperus communis) exhibited 0.4-0.9 MPa higher vulnerability to embolism. A small cavitation fatigue observed in Scots pine (Pinus sylvestris) was probably biased by incomplete embolism repair, as indicated by a correlation of vulnerability shifts and conductivity restoration. Our data demonstrate that cavitation fatigue in conifers is species-specific and depends on the intensity of preceding LC. The lack of fatigue effects after moderate LC, and relevant effects in only three species after high LC, indicate that conifers are relatively resistant against cavitation fatigue. This is remarkable considering the complex and delicate conifer pit architecture and may be important considering climate change projections.

An Abies procera-derived tetracyclic triterpene containing a steroid-like nucleus core and a lactone side chain attenuates in vitro survival of both Fasciola hepatica and Schistosoma mansoni.

Two economically and biomedically important platyhelminth species, Fasciola hepatica (liver fluke) and Schistosoma mansoni (blood fluke), are responsible for the neglected tropical diseases (NTDs) fasciolosis and schistosomiasis. Due to the absence of prophylactic vaccines, these NTDs are principally managed by the single class chemotherapies triclabendazole (F. hepatica) and praziquantel (S. mansoni). Unfortunately, liver fluke resistance to triclabendazole has been widely reported and blood fluke insensitivity/resistance to praziquantel has been observed in both laboratory settings as well as in endemic communities. Therefore, the identification of new anthelmintics is necessary for the sustainable control of these NTDs in both animal and human populations. Here, continuing our work with phytochemicals, we isolated ten triterpenoids from the mature bark of Abies species and assessed their anthelmintic activities against F. hepatica and S. mansoni larval and adult lifecycle stages. Full 1H and 13C NMR-mediated structural elucidation of the two most active triterpenoids revealed that a tetracyclic steroid-like nucleus core and a lactone side chain are associated with the observed anthelmintic effects. When compared to representative mammalian cell lines (MDBK and HepG2), the most potent triterpenoid (700015; anthelmintic EC50s range from 0.7 µM-15.6 µM) displayed anthelmintic selectivity (selectivity indices for F. hepatica: 13 for newly excysted juveniles, 46 for immature flukes, 2 for mature flukes; selectivity indices for S. mansoni: 14 for schistosomula, 9 for immature flukes, 4 for adult males and 3 for adult females) and induced severe disruption of surface membranes in both liver and blood flukes. S. mansoni egg production, a process responsible for pathology in schistosomiasis, was also severely inhibited by 700015. Together, our results describe the structural elucidation of a novel broad acting anthelmintic triterpenoid and support further investigations developing this compound into more potent analogues for the control of both fasciolosis and schistosomiasis.

Suppression of growth and death of meristematic tissues in Abies sachalinensis under strong shading: comparisons between the terminal bud, the terminally lateral bud and the stem cambium.

The suppression of apical growth and radial trunk growth in trees under shade is a key factor in the competition mechanism among individuals in natural and artificial forests. However, the timing of apical and radial growth suppression after shading and the physiological processes involved have not been evaluated precisely. Twenty-one Abies sachalinensis seedlings of 5-years-old were shaded artificially under a relative light intensity of 5% for 70 days from August 1, and the histological changes of the terminal bud and terminally lateral bud of terminal leader and the cambial zone of the trunk base were analyzed periodically. In shade-grown trees, cell death of the leaf primordia in a terminal bud of terminal leader was observed in one of the three samples after 56 and 70 days of shading, whereas the leaf primordia in a terminal bud of terminal leader in all open-grown trees survived until the end of the experiment. In addition, the leaf primordia of the terminally lateral buds of terminal leader retained their cell nuclei until the end of the experiment. No histological changes were observed in the cambial cells after shading, but the shade-grown trees had less cambial activity than the open-grown trees through the experiment. Strong shading appeared to inhibit the formation and survival of cells in the terminal bud of terminal leader rather than the terminally lateral buds of terminal leader and the cambium. The suppression of the terminal bud growth and elongation of the surviving lateral buds would result in an umbrella-shaped crown under shade.

Morpho-anatomical and physiological differences between sun and shade leaves in Abies alba Mill. (Pinaceae, Coniferales): a combined approach.

Morphology, anatomy and physiology of sun and shade leaves of Abies alba were investigated and major differences were identified, such as sun leaves being larger, containing a hypodermis and palisade parenchyma as well as possessing more stomata, while shade leaves exhibit a distinct leaf dimorphism. The large size of sun leaves and their arrangement crowded on the upper side of a plagiotropic shoot leads to self-shading which is explainable as protection from high solar radiation and to reduce the transpiration via the lamina. Sun leaves furthermore contain a higher xanthophyll cycle pigment amount and Non-Photochemical Quenching (NPQ) capacity, a lower amount of chlorophyll b and a total lower chlorophyll amount per leaf, as well as an increased electron transport rate and an increased photosynthesis light saturation intensity. However, sun leaves switch on their NPQ capacity at rather low light intensities, as exemplified by several parameters newly measured for conifers. Our holistic approach extends previous findings about sun and shade leaves in conifers and demonstrates that both leaf types of A. alba show structural and physiological remarkable similarities to their respective counterparts in angiosperms, but also possess unique characteristics allowing them to cope efficiently with their environmental constraints.

Why are the seed cones of conifers so diverse at pollination?

Background and Aims: Form and function relationships in plant reproductive structures have long fascinated biologists. Although the intricate associations between specific pollinators and reproductive morphology have been widely explored among animal-pollinated plants, the evolutionary processes underlying the diverse morphologies of wind-pollinated plants remain less well understood. Here we study how this diversity may have arisen by focusing on two conifer species in the pine family that have divergent reproductive cone morphologies at pollination. Methods: Standard histology methods, artificial wind pollination assays and phylogenetic analyses were used in this study. Key Results: A detailed study of cone ontogeny in these species reveals that variation in the rate at which their cone scales mature means that pollination occurs at different stages in their development, and thus in association with different specific morphologies. Pollination experiments nevertheless indicate that both species effectively capture pollen. Conclusions: In wind-pollinated plants, morphological diversity may result from simple variation in development among lineages rather than selective pressures for any major differences in function or performance. This work also illustrates the broader importance of developmental context in understanding plant form and function relationships; because plant reproductive structures perform many different functions over their lifetime, subtle differences in development may dramatically alter the specific morphologies that they use to meet these demands.

Stand Composition, Tree Proximity and Size Have Minimal Effects on Leaf Function of Coexisting Aspen and Subalpine Fir.

Forest structural heterogeneity due to species composition, spatial relationships and tree size are widely studied patterns in forest systems, but their impacts on tree function are not as well documented. The objective of this study was to examine how stand composition, tree proximity relationships and tree size influence the leaf functional traits of aspen, an early successional species, and subalpine fir, a climax species. We measured foliar nutrients, nonstructural carbohydrates (aspen only), defense chemistry and xylem water potential of aspen and subalpine fir trees in three size classes growing in close proximity or independently from other trees under three stand conditions: aspen dominant, aspen-conifer mixed, and conifer dominant stands. Close proximity of subalpine fir to aspen reduced aspen's storage of starch in foliar tissue by 17% suggesting that competition between these species may have small effects on carbon metabolism in aspen leaves. Simple sugar (glucose + sucrose) concentrations in aspen leaves were slightly higher in larger aspen trees than smaller trees. However, no differences were found in stem water potential, foliar concentrations of nitrogen, phosphorus, or secondary defense chemicals of aspen or subalpine fir across the gradients of stand composition, tree proximity or tree size. These results suggest that mechanisms of coexistence allow both aspen and subalpine fir to maintain leaf function across a wide range of stand structural characteristics. For aspen, resource sharing through its clonal root system and high resource storage capacity may partially contribute to its functional stability in mixed aspen-conifer stands.

Morphological defects in native Japanese fir trees around the Fukushima Daiichi Nuclear Power Plant.

After the accident at the Fukushima Daiichi Nuclear Power Plant (F1NPP) in March 2011, much attention has been paid to the biological consequences of the released radionuclides into the surrounding area. We investigated the morphological changes in Japanese fir, a Japanese endemic native conifer, at locations near the F1NPP. Japanese fir populations near the F1NPP showed a significantly increased number of morphological defects, involving deletions of leader shoots of the main axis, compared to a control population far from the F1NPP. The frequency of the defects corresponded to the radioactive contamination levels of the observation sites. A significant increase in deletions of the leader shoots became apparent in those that elongated after the spring of 2012, a year after the accident. These results suggest possibility that the contamination by radionuclides contributed to the morphological defects in Japanese fir trees in the area near the F1NPP.

Effects of tree architecture on pollen dispersal and mating patterns in Abies pinsapo Boiss. (Pinaceae).

Plant architecture is crucial to pollination and mating in wind-pollinated species. We investigated the effect of crown architecture on pollen dispersal, mating system and offspring quality, combining phenotypic and genotypic analyses in a low-density population of the endangered species Abies pinsapo. A total of 598 embryos from three relative crown height levels (bottom, middle and top) in five mother plants were genotyped using eleven nuclear microsatellite markers (nSSRs). Paternity analysis and mating system models were used to infer mating and pollen dispersal parameters. In addition, seeds were weighed (N = 16 110) and germinated (N = 736), and seedling vigour was measured to assess inbreeding depression. Overall, A. pinsapo shows a fat-tailed dispersal kernel, with an average pollen dispersal distance of 113-227 m, an immigration rate of 0.84-26.92%, and a number of effective pollen donors (Nep ) ranging between 3.5 and 11.9. We found an effect of tree height and relative crown height levels on mating parameters. A higher proportion of seeds with embryo (about 50%) and a higher rate of self-fertilization (about 60%) were found at the bottom level in comparison with the top level. Seed weight and seedling vigour are positively related. Nevertheless, no differences were found in seed weight or in seedling-related variables such as weight and length of aerial and subterranean parts among the different relative crown height levels, suggesting that seeds from the more strongly inbred bottom level are not affected by inbreeding depression. Our results point to vertical isotropy for outcross-pollen and they suggest that self-pollen may ensure fertilization when outcross-pollen is not available in low-density population.

Automatic identification and characterization of radial files in light microscopy images of wood.

BACKGROUND AND AIMS: Analysis of anatomical sections of wood provides important information for understanding the secondary growth and development of plants. This study reports on a new method for the automatic detection and characterization of cell files in wood images obtained by light microscopy. To facilitate interpretation of the results, reliability coefficients have been determined, which characterize the files, their cells and their respective measurements. METHODS: Histological sections and blocks of the gymnosperms Pinus canariensis, P. nigra and Abies alba were used, together with histological sections of the angiosperm mahogany (Swietenia spp.). Samples were scanned microscopically and mosaic images were built up. After initial processing to reduce noise and enhance contrast, cells were identified using a 'watershed' algorithm and then cell files were built up by the successive aggregation of cells taken from progressively enlarged neighbouring regions. Cell characteristics such as thickness and size were calculated, and a method was developed to determine the reliability of the measurements relative to manual methods. KEY RESULTS: Image analysis using this method can be performed in less than 20 s, which compares with a time of approx. 40 min to produce the same results manually. The results are accompanied by a reliability indicator that can highlight specific configurations of cells and also potentially erroneous data. CONCLUSIONS: The method provides a fast, economical and reliable tool for the identification of cell files. The reliability indicator characterizing the files permits quick filtering of data for statistical analysis while also highlighting particular biological configurations present in the wood sections.

Combined effect of elevated UVB, elevated temperature and fertilization on growth, needle structure and phytochemistry of young Norway spruce (Picea abies) seedlings.

Simultaneously with warming climate, other climatic and environmental factors are also changing. Here, we investigated for the first time the effects of elevated temperature, increased ultraviolet-B (UVB) radiation, fertilization and all combinations of these on the growth, secondary chemistry and needle structure of 1-year-old Norway spruce (Picea abies (L.) Karst.) seedlings in an outdoor experiment. After one growing season, elevated temperature increased root : shoot ratio and concentrations of needle piperidine alkaloids, while concentrations of needle catechins and acetophenones and bark flavonoids decreased compared with ambient temperature seedlings. UVB-radiation increased concentrations of bark condensed tannins, while fertilization increased total biomass and concentrations of needle catechins. In addition to the main effects, concentrations of some individual phenolic compounds showed UV × temperature or UV × temperature × fertilization interactions, and fertilization modified temperature response on root : shoot ratio. All the treatments described here affected the defence chemistry profiles of the seedlings, which may imply some changes in plant-herbivore interactions in connection with changing climate. The interactions between treatments indicate a need for further experiments involving several simultaneously affecting environmental changes.

Tree branching: Leonardo da Vinci's rule versus biomechanical models.

This study examined Leonardo da Vinci's rule (i.e., the sum of the cross-sectional area of all tree branches above a branching point at any height is equal to the cross-sectional area of the trunk or the branch immediately below the branching point) using simulations based on two biomechanical models: the uniform stress and elastic similarity models. Model calculations of the daughter/mother ratio (i.e., the ratio of the total cross-sectional area of the daughter branches to the cross-sectional area of the mother branch at the branching point) showed that both biomechanical models agreed with da Vinci's rule when the branching angles of daughter branches and the weights of lateral daughter branches were small; however, the models deviated from da Vinci's rule as the weights and/or the branching angles of lateral daughter branches increased. The calculated values of the two models were largely similar but differed in some ways. Field measurements of Fagus crenata and Abies homolepis also fit this trend, wherein models deviated from da Vinci's rule with increasing relative weights of lateral daughter branches. However, this deviation was small for a branching pattern in nature, where empirical measurements were taken under realistic measurement conditions; thus, da Vinci's rule did not critically contradict the biomechanical models in the case of real branching patterns, though the model calculations described the contradiction between da Vinci's rule and the biomechanical models. The field data for Fagus crenata fit the uniform stress model best, indicating that stress uniformity is the key constraint of branch morphology in Fagus crenata rather than elastic similarity or da Vinci's rule. On the other hand, mechanical constraints are not necessarily significant in the morphology of Abies homolepis branches, depending on the number of daughter branches. Rather, these branches were often in agreement with da Vinci's rule.

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

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

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

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

Hydraulic traits are associated with the distribution range of two closely related Mediterranean firs, Abies alba Mill. and Abies pinsapo Boiss.

Abies alba and Abies pinsapo are two closely related fir species that occur in the Iberian Peninsula under very different environmental conditions. Abies alba proliferates in the humid European mountains, including the Spanish Pyrenees. In contrast, A. pinsapo is a relict species that occurs in some restricted areas of the Mediterranean mountain ranges in Spain and Morocco, which experience intense summer drought periods. To cope with the high atmospheric evaporative demand during summer, A. pinsapo may either have a high resistance to xylem cavitation or develop a very efficient conducting system to reduce the soil-to-leaf water potential gradient. To investigate such hypotheses, we measured (i) the xylem vulnerability to cavitation for different populations, and (ii) several anatomical and hydraulic parameters indicating xylem sufficiency for -supplying water to the shoot in two contrasting populations of both species. Our results show that the resistance to cavitation was not different between species or populations. However, hydraulic conductivity (K(h)), specific hydraulic conductivity (K(s)), leaf-specific conductivity (LSC) and whole-shoot hydraulic conductance (K(shoot)) were higher in A. pinsapo, indicating a higher efficiency of water transport, which should contribute to maintaining its xylem tension below the threshold for rapidly increasing cavitation. The higher K(s) in A. pinsapo was largely a result of its wider tracheids, suggesting that this species may be much more vulnerable to freeze-thaw-induced cavitation than A. alba. This is consistent with the absence of A. pinsapo in northern mountain ranges with cooler winters. These physiological differences could partly explain the niche segregation and the geographical separation of these two firs.

Little change in the fir tree-line position on the southeastern Tibetan Plateau after 200 years of warming.

• As one of the world's highest natural tree lines, the Smith fir (Abies georgei var. smithii) tree line on the southeastern Tibetan Plateau is expected to vary as a function of climate warming. However, the spatial patterns and dynamics of the Smith fir tree line are not yet well understood. • Three rectangular plots (30 m × 150 m) were established in the natural alpine tree-line ecotone on two north-facing (Plot N1, 4390 m asl; Plot N2, 4380 m asl) and one east-facing (Plot E1, 4360 m asl) slope. Dendroecological methods were used to monitor the tree-line patterns and dynamics over a 50-yr interval. • The three study plots showed a similar pattern of regeneration dynamics, characterized by increased recruitment after the 1950s and an abrupt increase in the 1970s. Smith fir recruitment was significantly positively correlated with both summer and winter temperatures. However, Smith fir tree lines do not show a significant upward movement, despite warming on the Tibetan Plateau. • The warming in the past 200 yr is already having a significant impact on the population density of the trees, but not on the position of the Smith fir tree line.

Stand-structural effects on Heterobasidion abietinum-related mortality following drought events in Abies pinsapo.

Climate change may affect tree-pathogen interactions. This possibility has important implications for drought-prone forests, where stand dynamics and disease pathogenicity are especially sensitive to climatic stress. In addition, stand structural attributes including density-dependent tree-to-tree competition may modulate the stands' resistance to drought events and pathogen outbreaks. To assess the effects of stand structure on root-rot-related mortality after severe droughts, we focused on Heterobasidion abietinum mortality in relict Spanish stands of Abies pinsapo, a drought-sensitive fir. We compared stand attributes and tree spatial patterns in three plots with H. abietinum root-rot disease and three plots without root-rot. Point-pattern analyses were used to investigate the scale and extent of mortality patterns and to test hypotheses related to the spread of the disease. Dendrochronology was used to date the year of death and to assess the association between droughts and growth decline. We applied a structural equation modelling approach to test if tree mortality occurs more rapidly than predicted by a simple distance model when trees are subjected to high tree-to-tree competition and following drought events. Contrary to expectations of drought mortality, the effect of precipitation on the year of death was strong and negative, indicating that a period of high precipitation induced an earlier tree death. Competition intensity, related to the size and density of neighbour trees, also induced an earlier tree death. The effect of distance to the disease focus was negligible except in combination with intensive competition. Our results indicate that infected trees have decreased ability to withstand drought stress, and demonstrate that tree-to-tree competition and fungal infection act as predisposing factors of forest decline and mortality.

Defining how aging Pseudotsuga and Abies compensate for multiple stresses through multi-criteria assessment of a functional-structural model.

Many hypotheses have been advanced about factors that control tree longevity. We use a simulation model with multi-criteria optimization and Pareto optimality to determine branch morphologies in the Pinaceae that minimize the effect of growth limitations due to water stress while simultaneously maximizing carbohydrate gain. Two distinct branch morphologies in the Pareto optimal space resemble Pseudotsuga menziesii (Mirb.) Franco and Abies grandis (Dougl. ex D. Don) Lindl., respectively. These morphologies are distinguished by their performance with respect to two pathways of compensation for hydraulic limitation: minimizing the mean path length to terminal foliage (Pseudotsuga) and minimizing the mean number of junction constrictions to terminal foliage (Abies). Within these two groups, we find trade-offs between the criteria for foliage display and the criteria for hydraulic functioning, which shows that an appropriate framework for considering tree longevity is how trees compensate, simultaneously, for multiple stresses. The diverse morphologies that are found in a typical old-growth conifer forest may achieve compensation in different ways. The method of Pareto optimization that we employ preserves all solutions that are successful in achieving different combinations of criteria. The model for branch development that we use simulates the process of delayed adaptive reiteration (DAR), whereby new foliage grows from suppressed buds within the established branch structure. We propose a theoretical synthesis for the role of morphology in the persistence of old Pseudotsuga based on the characteristics of branch morphogenesis found in branches simulated from the optimal set. (i) The primary constraint on branch growth for Pseudotsuga is the mean path length; (ii) as has been previously noted, DAR is an opportunistic architecture; and (iii) DAR is limited by the number of successive reiterations that can form. We show that Pseudotsuga morphology is not the only solution to old-growth constraints, and we suggest how the model results should be used to guide future empirical investigation based on the two contrasting morphologies and how the morphological contrast may relate to physiological processes. Our results show that multi-criteria optimization with Pareto optimality has promise to advance the use of models in theory development and in exploration of functional-structural trade-offs, particularly in complex biological systems with multiple limiting factors.

Convergence of leaf display and photosynthetic characteristics of understory Abies amabilis and Tsuga heterophylla in an old-growth forest in southwestern Washington State, USA.

We compared the morphological and physiological characteristics of understory trees of Abies amabilis (Dougl. ex Loud.) Dougl. ex J. Forbes and Tsuga heterophylla (Raf.) Sarg. growing adjacent to each other in an old-growth forest in southwestern Washington State, USA. We hypothesized that, despite contrasting branching patterns and crown architectures, the two species should exhibit convergence in leaf display and photosynthetic gain per light intercepting area, because these are important properties determining their survival in the light-limited understory. The branching pattern of A. amabilis was regular (normal shoot-length distribution, less variable branching angle and bifurcation ratio), whereas that of T. heterophylla was more plastic (positively skewed shoot-length distribution, more variable branching angle and bifurcation ratio). The two species had similar shoot morphologies: number of leaves per unit shoot length and leaf to axis dry mass ratio. Leaf morphology, in contrast, was significantly different. Leaves of A. amabilis were larger and heavier than those of T. heterophylla, which resulted in lower mass-based photosynthetic rate for A. amabilis. Despite these differences, the two species had similar levels of leaf overlap and area-based photosynthetic characteristics. Needle longevity of A. amabilis was nearly twice that of T. heterophylla. The leaf N contents of current and 1-year-old leaves were lower for A. amabilis than for T. heterophylla. However, the leaf N content of A. amabilis did not change from current leaves to 6-year-old leaves, whereas that of T. heterophylla decreased with increasing leaf age. Abies amabilis had deeper crowns than T. heterophylla and retained branches with low relative growth rates. Longer branch retention may compensate for the lower branch-level assimilation rate of A. amabilis. We inferred that the convergence of leaf display and photosynthetic characteristics between A. amabilis and T. heterophylla may contribute to the persistence of both species in the understory of this forest.

Wood formation in Abies balsamea seedlings subjected to artificial defoliation.

We determined the cambial sensitivity and quantified the anatomical differences in xylem of Abies balsamea (L.) Mill. seedlings subjected to artificial defoliation to simulate spruce budworm feeding. Defoliation was performed by removing two-thirds of needles of all current-year shoots for up to four consecutive growth cycles to account for inter- and intra-annual xylem formation. In Experiment 1, xylem development was studied from May to October 2005 in seedlings defoliated at the end of June. In Experiment 2, anatomical features of the xylem were measured along the tree rings formed in 2005 and 2006 during the four cycles of growth and defoliation. Control and defoliated seedlings showed similar patterns of cambial activity and timing of xylem differentiation, although fewer enlarging cells were observed in August to September in defoliated seedlings. Tree-ring widths were similar in control and defoliated seedlings, with thinner rings produced in the greenhouse in winter. No effect of defoliation on cell lumen area was observed, and effects on radial cell diameter and wall thickness were found only occasionally. The results indicate that the A. balsamea seedlings produced all the resources required to maintain stem growth during the four cycles of defoliation.

Widespread increase of tree mortality rates in the western United States.

Persistent changes in tree mortality rates can alter forest structure, composition, and ecosystem services such as carbon sequestration. Our analyses of longitudinal data from unmanaged old forests in the western United States showed that background (noncatastrophic) mortality rates have increased rapidly in recent decades, with doubling periods ranging from 17 to 29 years among regions. Increases were also pervasive across elevations, tree sizes, dominant genera, and past fire histories. Forest density and basal area declined slightly, which suggests that increasing mortality was not caused by endogenous increases in competition. Because mortality increased in small trees, the overall increase in mortality rates cannot be attributed solely to aging of large trees. Regional warming and consequent increases in water deficits are likely contributors to the increases in tree mortality rates.