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Structural changes in spruce and fir needles.

Needles from spruce and fir trees were analyzed for histological changes induced by long-term exposure in open-top chambers to SO(2) and/or O(3) combined with acid rain. Light and electron microscopical evaluation revealed initial structural changes in the vascular bundle of fir needles, with an increased number of crushed sieve cells in the phloem. In addition the walls of young, adaxial sieve cells lacked the typical thickening usually observed in naturally aged needles. These findings may indicate restricted assimilate translocation. The presence of SO(2) in any treatment led to thylakoidal swellings and membrane reductions in the chloroplasts of mesophyll cells near the vascular bundle. This damage pattern resembled alterations caused by nutrient deficiency rather than by the direct action of gaseous pollutants. In general, fir appears to be more sensitive to environmental stress than spruce; this substantiates the findings of previous studies.

Journal Article↗

Decline of sacred fir (Abies religiosa) in a forest park south of Mexico City.

Decline of sacred fir (Abies religiosa) trees in the high elevation forest park, Desierto de los Leones, located south of Mexico City, is described. Trees located in the windward zone (exposed to air masses from Mexico City) were the most severely affected, especially trees at the distal ends of ravines. Examination of tree growth rings indicated decreases in ring widths for the past 30 years. Polluted air from Mexico City may be an important causal factor in fir decline. Drought, due to excessive removal of soil water, insects, mites and pathogens, and poor forest management are possible contributing and interactive factors in fir decline.

Journal Article↗

Chemical and biological characterization of dissolved organic matter from silver fir and beech forest soils.

Despite a growing attention to the dissolved organic matter (DOM) in terrestrial ecosystems and evidence of the fact that vegetation affects the quality of both undissolved and dissolved organic matter in soil, the role of DOM as a biological indicator is still poorly understood. In this work, the fertility of 59 sites, divided into eight key alliances of the order Fagetalia sylvaticae Pawl., was studied considering chemical and biological parameters such as soil DOM, hormone-like activity, low-molecular-weight (LMW) aliphatic and phenolic acids, and floristic data. Both non-parametric tests and principal component analysis (PCA) revealed differences between silver fir and beech forests and within each type of forest. There were also differences between neutrophilous and acidophilous types. What's more, PCA reveals the dominance of the auxin (IAA)-like activity, and of some phenolic acids in distinguishing the acidophilous beeches (ACI) form the other types, whereas the gibberellin (GA)-like activity is more relevant in neutrophilous conditions such as thermophilous (THE) and mesophilous (MESO) beeches and montane (MO), high montane (HMA), high montane (HMC) silver fir forests. The GA-like activity is also related to the succinic, fumaric, malonic, and l-malic acids in the MO, HMA and HMC silver fir forests. Moreover, the role of LMW aliphatic acids in mobilizing the hormone-like activity, which improves forest growth, is stressed. The growth of seedlings of Picea abies was influenced by the phenolic acid content. At concentrations between 1 and 100 microM, phenylacetic and protocatechuic acids inhibited root growth to the same extent as indoleacetic acid, while p-hydroxybenzoic acid had a stimulating effect comparable to that of gibberellic acid. The aliphatic and phenolic acids appear to be related to plant strategies that influence soil fertility affecting plant growth through rhizodeposition. The role of LMW aliphatic and phenolic acids as molecular markers of ecosystem function is noted.

Abies↗

Artifact reduction for simultaneous EEG/fMRI recording: adaptive FIR reduction of imaging artifacts.

OBJECTIVE: We present a new method of effectively removing imaging artifacts of electroencephalography (EEG) and extensively conserving the time-frequency features of EEG signals during simultaneous functional magnetic resonance imaging (fMRI) scanning under conventional conditions. METHODS: Under the conventional conditions of a 5000 Hz EEG sampling rate, but in the absence of the MRI slice-timing signals, the imaging artifact during each slice scanning is theoretically inferred to be a linear combination of the average artifact waveform and its derivatives, deduced by band-limited Taylor's expansion. Technically, the imaging artifact reduction algorithm is equivalent to an adaptive finite impulse response (FIR) filter. RESULTS: The capability of this novel method removing the imaging artifacts of EEG recording during fMRI scanning has been demonstrated by a phantom experiment. Moreover, the effectiveness of this method in conserving the time-frequency features of EEG activity has been evaluated by both visually evoked experiments and alpha waves. CONCLUSIONS: The adaptive FIR method is an effective method of removing the imaging artifacts under conventional conditions, and also conserving the time-frequency EEG signals. SIGNIFICANCE: The proposed adaptive FIR method, removing the imaging artifacts, combined with the wavelet-based non-linear noise reduction (WNNR) method [Wan X, Iwata K, Riera J, Ozaki T, Kitamura M, Kawashima R. Artifact reduction for EEG/fMRI recording: Nonlinear reduction of ballistocardiogram artifacts. Clin Neurophysiol 2006;117:668-80], reducing the ballistocardiogram artifacts (BAs), makes it feasible to obtain accurate EEG signals from the simultaneous EEG recordings during fMRI scanning.

Adult↗

Preparation of highly porous carbon from fir wood by KOH etching and CO2 gasification for adsorption of dyes and phenols from water.

Fir wood was first carbonized for 1.5 h at 450 degrees C, then soaked in a KOH solution KOH/char ratio of 1, and last activated for 1 h at 780 degrees C. During the last hour CO2 was poured in for further activation for 0, 15, 30, and 60 min, respectively. Carbonaceous adsorbents with controllable surface area and pore structure were chemically activated from carbonized fir wood (i.e., char) by KOH etching and CO2 gasification. The pore properties, including the BET surface area, pore volume, pore size distribution, and pore diameter, of these activated carbons were first characterized by the t-plot method based on N2 adsorption isotherms. Fir-wood carbon activated with CO2 gasification from 0 to 60 min exhibited a BET surface area ranging from 1371 to 2821 m2 g(-1), with a pore volume significantly increased from 0.81 to 1.73 m2 g(-1). Scanning electron microscopic (SEM) results showed that the surfaces of honeycombed holes in these carbons were significantly different from those of carbons without CO2 gasification. The adsorption of methylene blue, basic brown 1, acid blue 74, p-nitrophenol, p-chlorophenol, p-cresol, and phenol from water on all the carbons studied was examined to check their chemical characteristics. Adsorption kinetics was in agreement with the Elovich equation, and all equilibrium isotherms were in agreement with the Langmuir equation. These results were used to compare the Elovich parameter (1/b) and the adsorption quantity of the unit area (q(mon)/Sp) of activated carbons with different CO2 gasification durations. This work facilitated the preparation of activated carbon by effectively controlling pore structures and the adsorption performance of the activated carbon on adsorbates of different molecular forms.

Journal Article↗

Specific biochemical marker-based techniques for the identification of damage to Douglas-fir seed resulting from feeding by the western conifer seed bug, Leptoglossus occidentalis Heidemann (Hemiptera: Coreidae).

Specific biochemical marker-based techniques were tested for their ability to distinguish between seeds of Douglas-fir, Pseudotsuga menziesii (Mirbel) Franco, that were filled or unfilled (aborted) at maturity and those that were damaged or emptied by the western conifer seed bug, Leptoglossus occidentalis Heidemann. A polyclonal antibody raised against salivary gland extracts from L. occidentalis successfully identified residual salivary proteins on Western blots containing proteins from Douglas-fir seeds that had sustained various degrees of seed bug feeding damage. In a single blind experiment, the polyclonal antibody correctly identified 100% of undamaged control, 97% of unfilled control (aborted), and 98% of seed bug damaged seeds. Polyclonal antibodies raised against insoluble alfalfa crystalloid storage protein (11S globulin) detected the depletion of 11S globulin and the subsequent appearance of its hydrolyzed fragments in the soluble protein fraction of Douglas-fir seeds that were fed-upon by the seed bug. Feeding by L. occidentalis nymphs caused ca. 98% depletion of insoluble protein, but only ca. 53% reduction in the amount of soluble protein in seeds that appeared empty on radiographs. By comparison, unfilled (aborted) seeds contained significantly less insoluble and soluble protein than empty seeds that were fed-upon by L. occidentalis; moreover, no crystalloid (11S globulin) breakdown products were generated. The biochemical markers described in this study are reliable tools that can be used to identify conifer seeds that have sustained light to severe damage from L. occidentalis feeding.

Animals↗

Geographic biotype and host-associated local adaptation in a polyphagous species, Lambdina fiscellaria (Lepidoptera: Geometridae) feeding on balsam fir on Anticosti Island, Canada.

The debate about mechanisms underlying the evolution of host specialization by herbivorous insects remains open. Natural selection may act locally and lead to different patterns of geographic variation in life history traits of polyphagous herbivores. The hypothesis of genetically-based trade-offs in offspring performance on different hosts has been proposed but this has rarely been demonstrated. Under laboratory conditions, the biological performance of two populations of the hemlock looper Lambdina fiscellaria (Guenée), a highly polyphagous lepidopteran, was compared when reared on three different tree host species: balsam fir, eastern hemlock and sugar maple. One population originated from Anticosti Island, Québec, Canada, where the insect has evolved without having access to two of the three tree species tested, the other being from the mainland where all tree species are present. When reared on balsam fir foliage, which was naturally available to each population, larvae from Anticosti Island underwent four instars compared with five for the mainland population, indicating the existence of geographic biotypes in L. fiscellaria. When reared on the foliage of non-naturally available host trees, larvae from Anticosti Island had a higher incidence of supernumerary instars. This is a unique example where local adaptation to environmental conditions of an insect herbivore is expressed through a differential number of larval instars. Moreover, the Anticosti Island population showed a higher growth related index on the host available to both populations indicating that a fitness trade-off was the evolutionary process underlying the local adaptation of this population on balsam fir.

Abies↗

Abietadiene synthase from grand fir (Abies grandis): characterization and mechanism of action of the "pseudomature" recombinant enzyme.

The oleoresin secreted by grand fir (Abies grandis) is composed of resin acids derived largely from the abietane family of diterpene olefins as precursors which undergo subsequent oxidation of the C18-methyl group to a carboxyl function, for example, in the conversion of abieta-7,13-diene to abietic acid. A cDNA encoding abietadiene synthase has been isolated from grand fir and the heterologously expressed bifunctional enzyme shown to catalyze both the protonation-initiated cyclization of geranylgeranyl diphosphate to the intermediate (+)-copalyl diphosphate and the ionization-dependent cyclization of (+)-copalyl diphosphate, via a pimarenyl intermediate, to the olefin end products. Abietadiene synthase is translated as a preprotein bearing an N-terminal plastidial targeting sequence, and this form of the recombinant protein expressed in Escherichia coli proved to be unsuitable for detailed structure-function studies. Since the transit peptide-mature protein cleavage site could not be determined directly, a truncation series was constructed to delete the targeting sequence and prepare a "pseudomature" form of the enzyme that resembled the native abietadiene synthase in kinetic properties. Both the native synthase and the pseudomature synthase having 84 residues deleted from the preprotein converted geranylgeranyl diphosphate and the intermediate (+)-copalyl diphosphate to a nearly equal mixture of abietadiene, levopimaradiene, and neoabietadiene, as well as to three minor products, indicating that this single enzyme accounts for production of all of the resin acid precursors of grand fir. Kinetic evaluation of abietadiene synthase with geranylgeranyl diphosphate and (+)-copalyl diphosphate provided evidence for two functionally distinct active sites, the first for the cyclization of geranylgeranyl diphosphate to (+)-copalyl diphosphate and the second for the cyclization of (+)-copalyl diphosphate to diterpene end products, and demonstrated that the rate-limiting step of the coupled reaction sequence resides in the second cyclization process. The structural implications of these findings are discussed in the context of primary sequence elements considered to be responsible for binding the substrate and intermediate and for initiating the respective cyclization steps.

Isomerases↗

(Z,Z)-4,7-tridecadien-(S)-2-yl acetate: sex pheromone of Douglas-fir cone gall midge, Contarinia oregonensis.

Our objectives were to identify and field test the sex pheromone of female Douglas-fir cone gall midge, Contarinia oregonensis (Diptera: Ce-cidomyiidae). Coupled gas chromatographic-electroantennographic detection (GC-EAD) analyses of pheromone extract revealed a single compound (A) that elicited responses from male antennae. Hydrogenation of pheromone extract, followed by renewed GC-EAD analysis, revealed a new EAD-active compound with chromatographic characteristics identical to those of tridecan-2-yl acetate on five fused silica columns (DB-5, DB-210, DB-23, SP-1000, and Cyclodex-B). Syntheses, chromatography, and retention index calculations of all possible tridecen-2-yl acetates suggested that the candidate pheromone A was a tridecadien-2-yl acetate with nonconjugated double bonds. Synthetic candidate pheromone component (Z,Z)-4,7-tridecadien-2-yl acetate (Z4Z7) cochromatographed with A on all analytical columns and elicited comparable antennal activity. In GC-EAD analyses that separated the enantiomers (Z,Z)-4,7-tridecadien-(S)-2-yl acetate (2S-Z4Z7) and (Z,Z)-4,7-tridecadien-(R)-2-yl acetate (2R-Z4Z7) with baseline resolution, only 2S-Z4Z7 as a component in a racemic standard or in pheromone extract elicited antennal responses. In Douglas-fir seed orchards, sticky traps baited with 2S-Z4Z7 captured male C. oregonensis, whereas 2R-Z4Z7 was behaviorally benign. Comparable catches of males in traps baited with racemic Z4Z7 (50 microg) or virgin female C. oregonensis suggested that synthetic pheromone baits could be developed for monitoring C. oregonensis populations in commercial Douglas-fir seed orchards.

Acetates↗

Terpenoid-based defenses in conifers: cDNA cloning, characterization, and functional expression of wound-inducible (E)-alpha-bisabolene synthase from grand fir (Abies grandis).

(E)-alpha-Bisabolene synthase is one of two wound-inducible sesquiterpene synthases of grand fir (Abies grandis), and the olefin product of this cyclization reaction is considered to be the precursor in Abies species of todomatuic acid, juvabione, and related insect juvenile hormone mimics. A cDNA encoding (E)-alpha-bisabolene synthase was isolated from a wound-induced grand fir stem library by a PCR-based strategy and was functionally expressed in Escherichia coli and shown to produce (E)-alpha-bisabolene as the sole product from farnesyl diphosphate. The expressed synthase has a deduced size of 93.8 kDa and a pI of 5. 03, exhibits other properties typical of sesquiterpene synthases, and resembles in sequence other terpenoid synthases with the exception of a large amino-terminal insertion corresponding to Pro81-Val296. Biosynthetically prepared (E)-alpha-[3H]bisabolene was converted to todomatuic acid in induced grand fir cells, and the time course of appearance of bisabolene synthase mRNA was shown by Northern hybridization to lag behind that of mRNAs responsible for production of induced oleoresin monoterpenes. These results suggest that induced (E)-alpha-bisabolene biosynthesis constitutes part of a defense response targeted to insect herbivores, and possibly fungal pathogens, that is distinct from induced oleoresin monoterpene production.

Alkyl and Aryl Transferases↗

Monoterpene synthases from grand fir (Abies grandis). cDNA isolation, characterization, and functional expression of myrcene synthase, (-)-(4S)-limonene synthase, and (-)-(1S,5S)-pinene synthase.

Grand fir (Abies grandis) has been developed as a model system for studying defensive oleoresin formation in conifers in response to insect attack or other injury. The turpentine fraction of the oleoresin is a complex mixture of monoterpene (C10) olefins in which (-)-limonene and (-)-alpha- and (-)-beta-pinene are prominent components; (-)-limonene and (-)-pinene synthase activities are also induced upon stem wounding. A similarity based cloning strategy yielded three new cDNA species from a wounded stem cDNA library that appeared to encode three distinct monoterpene synthases. After expression in Escherichia coli and enzyme assay with geranyl diphosphate as substrate, subsequent analysis of the terpene products by chiral phase gas chromatography and mass spectrometry showed that these sequences encoded a (-)-limonene synthase, a myrcene synthase, and a (-)-pinene synthase that produces both alpha-pinene and beta-pinene. In properties and reaction stereochemistry, the recombinant enzymes resemble the corresponding native monoterpene synthases of wound-induced grand fir stem. The deduced amino acid sequences indicated the limonene synthase to be 637 residues in length (73.5 kDa), the myrcene synthase to be 627 residues in length (72.5 kDa), and the pinene synthase to be 628 residues in length (71.5 kDa); all of these monoterpene synthases appear to be translated as preproteins bearing an amino-terminal plastid targeting sequence. Sequence comparison revealed that these monoterpene synthases from grand fir resemble sesquiterpene (C15) synthases and diterpene (C20) synthases from conifers more closely than other monoterpene synthases from angiosperm species. This similarity between extant monoterpene, sesquiterpene, and diterpene synthases of gymnosperms is surprising since functional diversification of this enzyme class is assumed to have occurred over 300 million years ago. Wound-induced accumulation of transcripts for monoterpene synthases was demonstrated by RNA blot hybridization using probes derived from the three monoterpene synthase cDNAs. The availability of cDNA species encoding these monoterpene synthases will allow an understanding of the regulation of oleoresin formation in conifers and will ultimately permit the transgenic manipulation of this defensive secretion to enhance resistance to insects. These cDNAs also furnish tools for defining structure-function relationships in this group of catalysts that generate acyclic, monocyclic, and bicyclic olefin products.

Acyclic Monoterpenes↗

Growth response of bigcone Douglas fir (Pseudotsuga macrocarpa) to long-term ozone exposure in southern California.

Long-term radial growth of bigcone Douglas fir (Pseudotsuga macrocarpa) was studied throughout its range in the San Bernardino Mountains of southern California, where ambient ozone has been high for approximately the past 40 years. A gradient of both ozone concentration and precipitation exists from west (high) to east (low). Growth rates of bigcone Douglas fir are considerably lower since 1950 throughout the San Bernardino Mountains, with the largest growth reductions in the western part of the range where ozone exposure is highest. Needle retention is also somewhat lower at high ozone sites. Lower annual precipitation since 1950 may have some impact on long-term growth reductions, and short-term growth reductions induced by drought are an important component of long-term growth reductions at sites with high ozone exposure. An ozone-climate stress complex may be responsible for recent reductions in the growth of bigcone Douglas fir.

Air Pollutants↗

Genecology of Douglas fir in western Oregon and Washington.

BACKGROUND AND AIMS: Genecological knowledge is important for understanding evolutionary processes and for managing genetic resources. Previous studies of coastal Douglas fir (Pseudotsuga menziesii var. menziesii) have been inconclusive with respect to geographical patterns of variation, due in part to limited sample intensity and geographical and climatic representation. This study describes and maps patterns of genetic variation in adaptive traits in coastal Douglas fir in western Oregon and Washington, USA. METHODS: Traits of growth, phenology and partitioning were measured in seedlings of 1338 parents from 1048 locations grown in common gardens. Relations between traits and environments of seed sources were explored using regressions and canonical correlation analysis. Maps of genetic variation as related to the environment were developed using a geographical information system (GIS). KEY RESULTS: Populations differed considerably for adaptive traits, in particular for bud phenology and emergence. Variation in bud-set, emergence and growth was strongly related to elevation and cool-season temperatures. Variation in bud-burst and partitioning to stem diameter versus height was related to latitude and summer drought. Seedlings from the east side of the Washington Cascades were considerably smaller, set bud later and burst bud earlier than populations from the west side. CONCLUSIONS: Winter temperatures and frost dates are of overriding importance to the adaptation of Douglas fir to Pacific Northwest environments. Summer drought is of less importance. Maps generated using canonical correlation analysis and GIS allow easy visualization of a complex array of traits as related to a complex array of environments. The composite traits derived from canonical correlation analysis show two different patterns of variation associated with different gradients of cool-season temperatures and summer drought. The difference in growth and phenology between the westside and eastside Washington Cascades is hypothesized to be a consequence of the presence of interior variety (P. menziessii var. glauca) on the eastside.

Analysis of Variance↗

Phenology and density of balsam twig aphid, Mindarus abietinus Koch (Homoptera: Aphididae) in relation to bud break, shoot damage, and value of fir Christmas trees.

The balsam twig aphid, Mindarus abietinus Koch (Homoptera: Aphididae), is a major insect pest of balsam and Fraser fir grown for Christmas trees. Our objectives in this study were to 1) monitor the phenology of A. abietinus in fir plantations; 2) assess relationships among M. abietinus density, tree phenology, and damage to tree foliage; and 3) develop an esthetic injury level for M. abietinus on Christmas trees. We monitored phenology of M. abietinus and fir trees on three commercial Christmas tree plantations in central and northern Lower Michigan for 3 yr (1999-2001). Phenology of M. abietinus fundatrices and sexuparae was strongly correlated with accumulated degree-days (DD) base 10 degrees C. Fundatrices matured by approximately 83 DD(10 degrees C) and sexuparae were first observed at approximately 83-111 DD(10 degrees C). Trees that broke bud approximately 1 wk later than other trees in the same field escaped M. abietinus damage and shoot expansion rate in spring was generally positively correlated with M. abietinus damage. Retail customers surveyed at a choose-and-cut Christmas plantation in 2 yr did not consistently differentiate between similarly sized trees with no, light, and moderate M. abietinus damage, but heavy damage (>50% damaged shoots) did affect customer perception. Similarly, when wholesale grades were assigned, the high quality Grade 1 trees had up to 40% shoot damage, whereas Grade 2 trees had 32-62% shoot damage. Two trees ranked as unsaleable had sparse canopies and distorted needles on 42% to almost 100% of the shoots.

Abies↗

Absolute and relative growth of Douglas-fir seedlings of different sizes.

Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) seedlings of a single seed source were grown in a bare-root nursery for two years to produce five different stock types by varying spacing and transplanting treatments. They were then planted in the forest together with one-year-old container seedlings of the same seed source, which constituted a sixth treatment. Stem volume mean relative growth rate (R(v)) was low immediately after planting for all stock types except the container seedlings, and increased over the following 7.6 years. An 8-week greenhouse test of the six stock types showed that dry weight mean relative growth rate (R(w)) generally decreased with seedling dry weight, but this effect was less obvious after planting, because only the three smaller stock types showed a decrease in R(w) with size. In another experiment, bare-root Douglas-fir seedlings were grown at five different spacings in a nursery for two years, and seedlings from each spacing treatment were sorted into large or small by height. The resulting 10 treatments were planted in the forest and stem volumes determined over 6.7 years. The linear effect of nursery spacing on stem volume was no longer evident 3.6 years after planting, but large seedlings had greater stem volume than the small seedlings throughout the 6.7 years of the study. There was no indication that R(v) declined with time, but small seedlings had higher R(v) than large seedlings. Relative growth rates of two-year-old Douglas-fir were depressed for a year or two after planting, but then remained relatively constant, or increased during the ensuing 5 years. Relative growth rates of smaller seedlings were greater than those of larger seedlings so that relative biomass differences decreased with time, and the time advantage (the time necessary for small seedlings to reach the present biomass of large stock) of larger stock decreased. Absolute size differences increased with time.

Journal Article↗

Nutrition and bud removal affect biomass and nutrient allocation in Douglas-fir and western red cedar.

Seedlings of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) and western red cedar (Thuja plicata J. Donn ex D. Don) were grown at high (250 mg l(-1)) and low (20 mg l(-1)) nitrogen (N) supply for a year. Before the second growing season, half of the seedlings in each nutrient treatment were allocated to the other treatment. Half of the seedlings in each nutrient treatment then had all growing points removed. Biomass and N, phosphorus (P) and potassium (K) concentrations of old and new shoots and roots were measured three times in the second year to test the interaction of current-year and previous-year nutrient supply on biomass and nutrient allocation in these two species with different growth habits. Pruned seedlings served as controls. Unpruned seedlings of both species increased in height throughout the second growing season, except for Douglas-fir in the N250 --> N20 treatment. Repeated pruning did not prevent new shoot growth, but resulted in a 12 to 52% reduction in biomass of new shoots and new and old roots. Seedlings receiving a low N supply in the first growing season were more severely affected by pruning than seedings receiving a high N supply. Growth was reduced more by pruning in western red cedar than in Douglas-fir. Concentrations of N, P and K were higher in pruned seedlings than in unpruned seedlings. Although dry weights of all plant parts in all treatments increased throughout the second growing season, some retranslocation of N, P and K was observed from old shoots of both species in the N250 --> N20 and N20 --> N20 treatments after August. Quantities of N, P and K retranslocated were greatest in seedlings grown the previous year in the high-N treatment.

Journal Article↗

Needle anatomy changes with increasing tree age in Douglas-fir.

Morphological differences between old-growth trees and saplings of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) may extend to differences in needle anatomy. We used microscopy with image analysis to compare and quantify anatomical parameters in cross sections of previous-year needles of old-growth Douglas-fir trees and saplings at the Wind River Canopy Crane site in Washington and at three sites in the Cascade Mountains of Oregon. We also compared needle anatomy across a chronosequence of 10-, 20-, 40- and 450-year-old Douglas-fir trees from the Wind River site. Anatomy differed significantly between needles of old-growth trees and saplings at all sites, suggesting a developmental change in needle anatomy with increasing tree age. Compared with needles of old-growth trees, needles of saplings were longer and had proportionately smaller vascular cylinders, larger resin canals and few hypodermal cells. Astrosclereids, which sequester lignin in their secondary cell walls and occupy space otherwise filled by photosynthetic cells, were scarce in needles of saplings but abundant in needles of old-growth trees. Needles of old-growth trees had an average of 11% less photosynthetic mesophyll area than needles of saplings. The percentage of non-photosynthetic area in needles increased significantly with increasing tree age from the chronosequence of 10-, 20-, 40- and 450-year-old trees at the Wind River site. This reduction in photosynthetic area may contribute to decreased growth rates in old trees.

Microscopy, Electron, Scanning↗

Changes in net ecosystem productivity with forest age following clearcutting of a coastal Douglas-fir forest: testing a mathematical model with eddy covariance measurements along a forest chronosequence.

We hypothesized that changes in net ecosystem productivity (NEP) during aging of coastal Douglas-fir (Pseudotsuga menziesii Mirb. Franco) stands could be explained by (1) changing nutrient uptake caused by different time scales for decomposition of fine, non-woody and coarse woody litter left after harvesting, (2) declines in canopy water status with lengthening of the water uptake pathway during bole and branch growth, and (3) increases in the ratio of autotrophic respiration (R (a)) to gross primary productivity (GPP) with phytomass accumulation. These hypotheses were implemented and tested in the mathematical model ecosys against eddy covariance (EC) measurements of forest CO(2) and energy exchange in a post-clearcut Douglas-fir chronosequence. Hypothesis 1 explained how (a) an initial rise in GPP observed during the first 3 years after clearcutting could be caused by nutrient mineralization from rapid decomposition of fine, non-woody litter with lower C:N ratios (assart effect), (b) a slower rise in GPP during the next 20 years could be caused by immobilization during later decomposition of coarse woody litter, and (c) a rapid rise in GPP between 20 and 40 years after clearcutting could be caused by nutrient mineralization with further decomposition of coarse woody litter and of its decomposition products. During periods (a) and (b), heterotrophic respiration (R (h)) from decomposition of fine and coarse litter greatly exceeded net primary productivity (NPP = GPP - R (a)) so that Douglas-fir stands were large sources of CO(2). During period (c), NPP exceeded R (h) so that these stands became large sinks for CO(2). Hypothesis 2 explained how declines in NPP during later growth in period (c) could be caused by lower hydraulic conductances in taller trees that would force lower canopy water potentials and hence greater sensitivity of stomatal conductances and CO(2) uptake to vapor pressure deficits. Enhanced sensitivity to vapor pressure deficits was also apparent in the EC measurements over the post-clearcut chronosequence. Hypothesis 3 did not contribute to the explanation of forest age effects on NEP.

Abies↗