Nitrogen mineralization in high-elevation forests of the appalachians. II. Patterns with stand development in fir waves

Wave-like patterns of mortality and regeneration of balsam fir (Abies balsamea {L.} Mill.) and Fraser fir (A. fraseri {Pursh.} Poir.) forests at high elevations in the Appalachian Mountains offer a unique opportunity to study the effects of stand development on nitrogen cycling. We sampled two fir waves, one with Fraser fir on Mt. LeConte in the Great Smoky Mountains National Park, and one with balsam fir on Whiteface Mountain in New York. Net nitrogen mineralization for 3 summer months at Mt. LeConte was high in the dead fir zone (47 kg-N/ha), lower in the regeneration and juvenile zones (24 and 21 kg-N/ha), and highest in the mature zone (61 kg-N/ha). This sampling period probably accounted for about 60% of the annual total. The pattern was similar in the balsam fir wave on Whiteface Mountain, with N mineralization rates of 39 and 33 kg-N/ha over 2 months for the regenerating and juvenile zones, and 43 and 54 kg-N/ha for the mature and dead zones. Throughfall nitrogen followed a fairly similar pattern, with rates ranging from 4.5 to 10 kg-N/ha for 2 or 3 months across all zones at both sites. Tension-free lysimeters indicated very little leaching of nitrogen below 30 cm depth (the maximum was 2.6 kg-N/ha), but these estimates may be low. We conclude that nitrogen mineralization is high at all stages of stand development, perhaps exceeding the uptake capacity of the trees. Rates of nitrogen leaching may be high in these ecosystems and should receive further attention. Request for offprints     

Potential of Margosan-O, an azadirachtin-containing formulation from neem seed extract, as a control agent for spruce budworm, Choristoneura fumiferana

The effects on spruce budworm larvae, Choristoneura fumiferana (Clem.), produced by ingestion of Margosan-O, a commercially available neem seed extract formulation containing 0.3% azadirachtin, were investigated. Bioassays with the test material were conducted using various instars of spruce budworm larvae, with either artificial diet, cut branches of balsam fir, Abies balsamea (L.) Mill., or small growing balsam fir trees as substrates. The dose-response data on feeding reduction, developmental retardation, and mortality (LC50, LC95, and LD50) suggest that Margosan-O has promise as a control agent for spruce budworm in an integrated pest management program.     

Production,application, and field performance of Abietiv™, the balsam fir sawfly nucleopolyhedrovirus

Beginning in the early 1990s, the balsam fir sawfly (Neodiprion abietis) became a significant defoliating insect of precommercially thinned balsam fir (Abies balsamea (L.) Mill.) stands in western Newfoundland, Canada. In 1997, a nucleopolyhedrovirus (NeabNPV) was isolated from the balsam fir sawfly and, as no control measures were then available, NeabNPV was developed for the biological control of balsam fir sawfly. In order to register NeabNPV for operational use under the Canadian Pest Control Products Act, research was carried out in a number of areas including NeabNPV field efficacy, non-target organism toxicology, balsam fir sawfly ecology and impact on balsam fir trees, and NeabNPV genome sequencing and analysis. As part of the field efficacy trials, approximately 22 500 hectares of balsam fir sawfly-infested forest were aerially treated with NeabNPV between 2000 and 2005. NeabNPV was found to be safe, efficacious, and economical for the suppression of balsam fir sawfly outbreak populations. Conditional registration for the NeabNPV-based product, Abietiv™, was received from the Pest Management Regulatory Agency (Health Canada) in April 2006. In July 2006, Abietiv was applied by spray airplanes to 15 000 ha of balsam fir sawfly-infested forest in western Newfoundland in an operational control program.      

Water holdup capacity and residence time of red spruce and balsam fir branches

Summary This study examined the throughfall dynamics of high-elevation red spruce (Picea rubens Sarg.) and balsam fir [Abies balsamea (L.) Mill.] branches. A site was established at an elevation of 1160 m on Mt. Washington, New Hampshire, USA, and branches were collected from the canopies of mature trees. Throughfall water dynamics of branches collected in September 1988 and March 1989 were determined using a fluorescent tracer in an artificial precipitation apparatus. Water holdup capacities of spruce and fir branches from different canopy positions were similar. Spruce retained more water per unit area than fir. For rain, a three-compartment model that was used to analyze water dynamics showed that the bulk of water initially on the branch experienced very slow turnover, while intercepted water left the branch quickly. There did not appear to be any difference in throughfall dynamics between spruce and fir. Our results suggest that the initial composition of rain and mobilized dry deposition will influence the composition of water in contact with the branch for a relatively long time, as branch water composition will respond slowly to changes in precipitation composition. These predictions require field testing where sequential sampling of throughfall and precipitation occurs on a time scale equivalent to 0.1–0.2 mm of precipitation.     

Effect of climatic variation on the morphological characteristics of 37‐year‐old balsam fir provenances planted in a common garden in New Brunswick,Canada

The extent of the effect of projected changes in climate on trees remains unclear. This study investigated the effect of climatic variation on morphological traits of balsam fir [Abies balsamea (L.) Mill.] provenances sourced from locations spanning latitudes from 44° to 51°N and longitudes from 53° to 102°W across North America, growing in a common garden in eastern Canada. Lower latitude provenances performed significantly better than higher latitude provenances (p < .05) with regard to diameter at breast height (DBH), height (H), and crown width (CW), a distinction indicative of genotypic control of these traits. There was, however, no significant difference among provenances in terms of survival (p > .05), an indication of a resource allocation strategy directed at survival relative to productivity in higher latitude provenances as seen in their lower DBH, H, and CW compared to the lower latitude provenances. Temperature had a stronger relationship with DBH, H, and CW than precipitation, a reflection of adaptation to local conditions in populations of the species along latitudinal gradients. Both climatic variables had some effect on tree survival. These results suggest that the response of balsam fir to climatic variation will likely not be uniform in the species, but differ based on genetic characteristics between populations located in the northern and southern parts of the species’ range. Population differences in response to climatic variation may be evident earlier in growth traits, compared to survival in balsam fir. The findings of this study will facilitate modeling in the species that is reflective of genetic variation in response to climatic conditions, and guide provenance selection for utilization in terms of productivity or resilience as well as breeding programs directed at obtaining species that possibly combine both traits.     

Does climate affect host-plant quality? Annual variation in the quality of balsam fir as food for spruce budworm

Summary The hypothesis that annual climatic variation can affect the nutritive value of host-plant tissue for a herbivore was tested. Larvae of a single generation of western spruce budworm, Choristoneura occidentalis Free., were fed foliages that had been collected on five consecutive springs from a stand of balsam fir Abies balsamea (L.) Mill. Budworm of both sexes were about 20 percent heavier at pupation when fed folages that had been produced in early rather than later spring seasons. This could result in an estimated 25 percent difference in the reproductive rate of females. It is concluded that annual variations in spring climate can affect the nutritive value of forage.     

Evidence of a direct chemical plant defense role for maltol against spruce budworm

This study examines the direct chemical defensive role of maltol, a previously identified secondary metabolite found in balsam fir, Abies balsamea (L.) Mill. (Pinaceae), that was detected during herbivory of spruce budworm, Choristoneura fumiferana (Clemens) (Lepidoptera: Tortricidae). Although used extensively in many industries, in addition to being found in multiple plant species, its functional role in plants remains unknown. The objectives of this study were to quantify the amount of free maltol and its potential conjugated form, maltol glucoside, in various foliage age classes and to evaluate whether constitutive foliage levels of maltol have an impact on spruce budworm fitness in maltol supplementation assays. Gas chromatography–mass spectrometry (GC‐MS) analysis of balsam fir foliage showed that maltol is produced in all foliage age classes tested; however, concentrations were significantly higher in older foliage. Liquid chromatography–mass spectrometry–mass spectrometry (LC‐MS‐MS) analysis showed that maltol also exists in balsam fir in its glucosylated form, a unique discovery in conifers. Similar to maltol, maltol glucoside is also present in current and 1‐year‐old balsam fir foliage and in significantly higher concentration in older foliage. We investigated the impact of maltol‐treated diet on spruce budworm fitness. Maltol additions that reflected constitutive foliage concentrations caused a significant reduction in larval development rate and pupal mass, whereas higher concentrations were required to cause significant mortality. These results suggest that maltol may be an important component of a direct defense strategy in balsam fir against spruce budworm herbivory.     

Plant module size influences the intra‐tree distribution and abundance of a shoot‐boring sawfly in young balsam fir

Field surveys were carried out to assess the effects of intra‐tree variation in developing shoot length within and among crown levels on the density and abundance of the balsam shoot‐boring sawfly, Pleroneura brunneicornis Rohwer (Hymenoptera: Xyelidae), in young balsam fir, Abies balsamea (L.) Mill. (Pinaceae). Overall, cardinal direction had no influence on shoot‐borer density or abundance; however, the highest percentage and abundance of bored shoots occurred on intermediate‐sized shoots within the crown (i.e., in the mid‐crown and on the distal‐lateral and medial‐lateral shoots). Comparatively, few shoot borers occurred in the upper or lower crown levels, or on the relatively large terminal shoots within branches. This distribution appears indicative of the higher suitability of intermediate‐sized shoots within hosts for either egg lay or larval performance. Results of this study are most consistent with predictions of the ‘optimal module size’ hypothesis, which posits that herbivore responses to plant module size should reflect the balance of tradeoffs between utilizing relatively large, nutritious shoots vs. small, more easily exploited shoots.     

Relating mechanical strength at the stem level to values obtained from defect-free wood samples

Stem or branch failure is a recurrent problem in silviculture and arboriculture. The risk of rupture varies with species in relation to the inherent mechanical properties of the species and the presence of defects. In general, calculations of critical loads for breakage are based on mechanical properties determined from defect-free samples and adjustment factors that try to scale up to full trees that include defects. This study aims at developing an objective method to scale up mechanical resistance to breakage from defect-free samples to full trees, including different types of defects. It combines two approaches. In the first one, a correction factor is determined from a meta-analysis of various tree-pulling studies involving balsam fir [Abies balsamea (L.) Mill.], white spruce [Picea glauca (Moench) Voss], jack pine [Pinus banksiana Lamb.], and black spruce [Picea mariana (Mill.) B.S.P.]. The second approach consists in obtaining empirical data from three-point bending tests using 8-foot (2.44 m) logs with various amounts of decay. Results show that the correction required varies according to the species and the presence of some defects. For balsam fir, which was the species showing the most important difference between whole log and small sample values, differences in correction factors were found between tree-pulling tests and three-point bending tests. Data from winching tests tend to underestimate the stem’s resistance to breakage since they likely represent the weakest trees among those tested. No relationship was found between the adjustment factors and different indices used in arboriculture to account for decay, showing the complexity of mechanical resistance at the stem level.     

Effects of soil calcium and aluminum on the physiology of balsam fir and red spruce saplings in northern New England

We examined the influence of calcium (Ca) and aluminum (Al) nutrition on the foliar physiology of red spruce (Picea rubens Sarg.) and balsam fir [Abies balsamea (L.) Mill.] in northern New England, USA. At the Hubbard Brook Experimental Forest (NH, USA), spruce and fir saplings were sampled from control, Al-, and Ca-supplemented plots at a long-established nutrient perturbation (NuPert) study in fall 2008. Measurements included cation concentrations (roots and foliage), dark-adapted chlorophyll fluorescence (F _v/F _m), soluble sugar concentrations, and ascorbate peroxidase (APX) and glutathione reductase (GR) activity in current-year foliage. Additional untreated saplings were sampled from base-rich Sleepers River (VT) and base-poor Jeffers Brook (NH) for F _v/F _m and foliar nutrient concentrations. At NuPert, there were significantly greater Ca concentrations and Ca:Al ratios in roots from the Ca end vs. the Al end of the Al-control-Ca addition gradient. There were also trends toward greater foliar Ca and Ca:Al ratios and lower Al concentrations across the treatment gradient at NuPert and for foliage at Sleepers River vs. Jeffers Brook. At NuPert, F _v/F _m and APX activity increased across the treatment gradient, and red spruce was higher in these measures than balsam fir. These patterns were also observed when Jeffers Brook and Sleepers River were compared. Increased Ca availability appeared to enhance the ability of red spruce and balsam fir to repair oxidative stress damage, including photooxidation. Our findings support work indicating a greater contemporary level of stress for balsam fir relative to red spruce, which is surprising considering the well-documented regional decline of spruce.     

Factors affecting oral regurgitation by larval spruce budworm

This study evaluated factors that influence the regurgitation behaviour of sixth instar spruce budworm, Choristoneura fumiferana Clemens (Lepidoptera: Tortricidae), reared on balsam fir, Abies balsamea (L.) Mill. (Pinaceae), under various experimental conditions in the laboratory. Upon physical disturbance, larvae discharged a median volume of regurgitant of 0.4 μl when fed and 1.6 μl when food‐deprived. Larvae deprived of food for 24 or 48 h disgorged more regurgitant than larvae feeding on balsam fir foliage, and the effect was consistent for laboratory‐reared and field‐collected larvae. The water content of the foliage fed upon by larvae had no immediate impact on the volume of regurgitant; following a 24‐h period of food deprivation, however, larvae that previously fed on fresh foliage discharged >2.5 times more regurgitant than larvae that previously fed on dry foliage. Self‐regulated regurgitation by larvae, measured using the number of regurgitant stains on filter paper, was >10 times higher when larvae had access to balsam fir foliage than when they were starved. The number of larvae confined inside the Petri dish (one or four individuals) had a relatively small effect on regurgitation. Larvae were deterred from feeding when balsam fir needles were entirely covered with regurgitant, but not when only a portion of the foliage was treated. These results suggest that the regurgitant does not serve as resource marking or spacing pheromone. The high level of regurgitation by larvae after contact with ants suggests that the regurgitant has evolved in part as a defence mechanism against natural enemies.     

Acceptance and suitability of novel trees for <Emphasis Type="Italic">Orthotomicus erosus</Emphasis>, an exotic bark beetle in North America

To predict whether an herbivorous pest insect will establish in a new area, the potential host plants must be known. For invading bark beetles, adults must recognize and accept trees suitable for larval development. The preference-performance hypothesis predicts that adults will select host species that maximize the fitness of their offspring. We tested five species of North American conifers and one angiosperm for adult acceptance and suitability for reproduction of the Mediterranean pine engraver, Orthotomicus erosus (Wollaston). Red pine, Pinus resinosa Aiton, and white spruce, Picea glauca (Moench) Voss, were accepted by adult beetles and suitable for reproduction to the extent of adult replacement. Others, such as balsam fir, Abies balsamea (L.) Mill., eastern hemlock, Tsuga canagensis (L.) Carrière, and tamarack, Larix laricina (Du Roi) Koch, were acceptable but unsuitable. The presence of tree species that are acceptable to adults but unsuitable for reproduction may affect the ability of O. erosus to establish across North America.     

Genetic discontinuity revealed by chloroplast microsatellites in eastern North American Abies (Pinaceae)

Development of conservation strategies for Fraser fir (Abies fraseri) in the southern Appalachian Mountains depends in part on recognition of the extent to which Fraser fir is genetically distinct from the closely related balsam (A. balsamea) and intermediate (A. balsamea var. phanerolepis) fir. These sibling species have exhibited intergrading, clinal variation in morphological, chemical, and genetic characteristics in prior research. Chloroplast microsatellite markers were polymerase chain reaction amplified from genomic DNA samples of 78 individuals representing the geographic ranges of Fraser, balsam, and intermediate fir. Gene diversity levels at two loci ranged among taxa from 0.65 to 0.84. Allele frequencies demonstrated significant differentiation among taxa, with R(ST) values of 0.36 and 0.10. Haplotype diversity and D(SH) were highest for balsam fir and lowest for intermediate fir. A haplotype network analysis based on allele size distribution for the two loci revealed two distinct clusters of haplotypes and population-specific haplotypes. Ninety-two percent of the haplotypes in one cluster were from balsam fir and intermediate fir, and 84% of the haplotypes in the other cluster were from Fraser fir and intermediate fir. The genetic differentiation of chloroplast DNA markers provides justification for the recognition of Fraser fir as a distinct Management Unit (MU) for conservation purposes, regardless of its taxonomic classification.     

Interspecific variation in resistance of two host tree species to spruce budworm

Woody plants regularly sustain biomass losses to herbivorous insects. Consequently, they have developed various resistance mechanisms to cope with insect attack. However, these mechanisms of defense and how they are affected by resource availability are not well understood. The present study aimed at evaluating and comparing the natural resistance (antibiosis and tolerance) of balsam fir (Abies balsamea [L.] Mill.) and white spruce (Picea glauca [Moench) Voss] to spruce budworm, Choristoneura fumiferana (Clem.), and how drainage site quality as a component of resource availability affects the expression of resistance over time (6 years). Our results showed that there are differences in natural resistance between the two tree species to spruce budworm, but it was not significantly affected by drainage quality. Balsam fir exhibited higher foliar toxic secondary compounds concentrations than white spruce in all drainage classes, resulting in lower male pupal mass, survival and longer male developmental time. This, however, did not prevent spruce budworm from consuming more foliage in balsam fir than in white spruce. This response suggests that either natural levels of measured secondary compounds do not provide sufficient toxicity to reduce defoliation, or spruce budworm has developed compensatory mechanisms, which allow it to utilize food resources more efficiently or minimize the toxic effects that are produced by its host's defensive compounds. Larvae exhibited lower pupal mass and higher mortality in rapidly drained and subhygric sites. Drainage class also affected the amount of foliage destroyed but its impact varied over the years and was probably influenced by climatic variables. These results demonstrate the complexity of predicting the effect of resource availability on tree defenses, especially when other confounding environmental factors can affect tree resource allocation and utilization.     

Daily weather response of balsam fir (<Emphasis Type="Italic">Abies balsamea</Emphasis> (L.) Mill.) stem radius increment from dendrometer analysis in the boreal forests of Québec (Canada)

Daily stem radial growth of balsam fir [ Abies balsamea (L.) Mill.] was studied between 1998 and 2001 using automated point dendrometers to investigate meteorological influence. By dividing the dendrometer day-night variation, the diurnal growth pattern was resolved into the three phases of (1) contraction, (2) expansion and (3) stem radius increment (SRI). The entire circadian cycle (4) defined by the three previous phases was considered as a fourth phase. The mean weather conditions of each phase were compared with the SRI using simple correlation and response function analysis. It was found that the weather conditions prevailing from 1600/1700 hours to 0800/0900 hours corresponding with the expansion-SRI phases had greater impact on SRI. Response function results confirmed most of the correlation analyses and explained up to 95% of the variance of the SRI series. Total rainfall in phases 2, 3 and 4 was correlated positively with SRI, and hence verifies the importance of daily water balance. The importance of water was also demonstrated by the negative effect of high vapour pressure deficit of phase 2, decreasing the possibility of cell radial expansion. The maximum temperature of phase 3 was the only temperature variable having a positive impact on SRI suggesting that night temperature was more important than day temperature in controlling radial growth. These results may influence the process of cell enlargement and reflect only the mechanical aspect of growth.     

IN VITRO ALLELOPATHIC INHIBITION OF NITRIFICATION BY BALSAM POPLAR AND BALSAM FIR

A soil percolation apparatus was used to study the effect of plant extracts on nitrification. Foliar leachates, leaf extracts, and bud extracts of balsam fir (Abies balsamea) and balsam poplar (Populus balsamifera) were added to the ammonium percolation solution. Color tests and quantitative measurement of residual ammonium and formed nitrate were applied. Foliar leachates of balsam fir strongly inhibited nitrification; leachates of balsam poplar were slightly less inhibitory. Balsam fir needle and balsam poplar dormant bud extracts (5% w/v) completely prevented the oxidation of ammonium. The same estracts (2% w/v) still prevented the oxidation of ammonium to nitrate to a large extent. Once again balsam fir extracts inhibited the process more strongly than balsam poplar. No evidence was obtained for the presence of a specific inhibitor of nitrifier microorganisms.     

Design and evaluation of an aerial spray trial with true replicates to test the efficacy of Bacillus thuringiensis insecticide in a boreal forest

A field trial using true replicates was conducted successfully in a boreal forest in 1996 to evaluate the efficacy of two aerially applied Bacillus thuringiensis formulations, ABG 6429 and ABG 6430. A complete randomized design with four replicates per treatment was chosen. Twelve to 15 balsam fir (Abies balsamea [L.] Mill.) per plot were randomly selected as sample trees. Interplot buffer zones, > or = 200 m wide, adequately prevented cross contamination from sprays that were atomized with four rotary atomizers (volume median diameters ranging from 64.6 to 139.4 microm) and released approximately 30 m above the ground. The B. thuringiensis formulations were not significantly different (P > 0.05) from each other in reducing spruce budworm (Choristoneura fumiferana [Clem.]) populations and protecting balsam trees from defoliation but both formulations were significantly more efficacious than the controls. The results suggest that true replicates are a feasible alternative to pseudoreplication in experimental forest aerial applications.     

A Matrix model for Predicting Foliage Weight of Trees by Age Classes

A Leslie-type matrix has been developed to model the annual growth of foliage on individual trees for any species that retains its foliage for more than one year. In order to apply the model to any given species, one must know: (1) age-specific foliage photosynthetic efficiencies; (2) the proportion of foliage of each age class that remains to become one year older; and (3) the diameter of the tree at any two points in time, several years apart. These diameters are used to estimate the maximum eigenvalue of the matrix, which is needed in the determination of the specific numerical matrix which corresponds to a particular individual tree. Although the model is a general one, applicable to any species that has age classes of foliage, it is developed with occasional references to balsam fir, Abies balsamea (L.) Mill. For a data set of 66 dominant and codominant balsam fir trees, the model predicted the weights of the first three age classes of foliage accurately (less than 10% error) over a 6-year period. There was a 23% error for all age classes combined.     

Transgenic Christmas Trees Regenerated from Agrobacterium tumefaciens-mediated Transformation of Zygotic Embryos Using the Green Fluorescence Protein as a Reporter

Mature zygotic embryos of recalcitrant Christmas tree species Fraser fir [Abies fraseri (Pursh) Poir], and Nordmann fir (Abies nordmanniana L.k.), and Virginia pine (Pinus virginiana Mill.) were used as explants for Agrobacterium tumefaciens strain GV3850-mediated transformation using the gfp (green fluorescent protein) gene as a reporter. Factors including media used for inoculation and co-cultivation, concentrations of acetosyringone, and antibiotics in tissue culture media have been evaluated. A high transformation frequency was obtained on TE medium containing 50μM acetosyringone and using 500 mg/l timentin to eliminate bacteria. Transient gene expression was observed in all three Christmas tree species, but transgenic plants were only produced from Virginia pine. Stable integration and expression of transgenes in the plant genome of Virginia pine was confirmed by polymerase chain reaction (PCR), Southern and northern blot analyses. These results demonstrated that a stable transformation system has been established in Virginia pine and this system would provide an opportunity to transfer economically important genes into Christmas tree species.     

Foliage‐age mixing within balsam fir increases the fitness of a generalist caterpillar

1. Manipulative field studies were carried out to evaluate the foliage age preference–performance relationship for an extreme generalist herbivore, the whitemarked tussock moth (Orygia leucostigma Smith) (Lepidoptera: Lymantriidae), within balsam fir [Abies balsamea (L.) Mill]. 2. Field surveys indicated that early instar caterpillars fed almost exclusively on young (i.e. current‐year) foliage, whereas late instars caterpillars fed on both young and mature (i.e. 1‐ and 2‐year‐old) foliage. 3. Survival of early instar caterpillars was highest in treatments where current‐year and/or 1‐year old foliage were available, but decreased significantly on older foliage. In contrast, late instar caterpillars had the highest survival when allowed to feed on all age classes of foliage, whereas potential fecundity was highest for late instars that fed on young foliage. 4. Overall, caterpillars had 32–65% higher fitness when able to feed on all rather than just one age class of foliage. 5. These results support both the ‘complementary diet' hypothesis, which states that dietary mixing of different‐aged foliage can increase nutrient uptake and/or dilute harmful secondary plant chemicals, and the ‘ontogeny’ hypothesis, which attributes changes in diet to changes in the nutritional needs and/or tolerance to plant defences of juvenile insects as they develop.