PhenoCaB: a new phenological model based on carbon balance in boreal conifers.

Traditional phenological models use chilling and thermal forcing (temperature sum or degree-days) to predict budbreak. Because of the heightening impact of climate and other related biotic or abiotic stressors, a model with greater biological support is needed to better predict budbreak. Here, we present an original mechanistic model based on the physiological processes taking place before and during budbreak of conifers. As a general principle, we assume that phenology is driven by the carbon status of the plant, which is closely related to environmental variables and the annual cycle of dormancy-activity. The carbon balance of a branch was modelled from autumn to winter with cold acclimation and dormancy and from winter to spring when deacclimation and growth resumption occur. After being calibrated in a field experiment, the model was validated across a large area (> 34 000 km2 ), covering multiple conifers stands in Québec (Canada) and across heated plots for the SPRUCE experiment in Minnesota (USA). The model accurately predicted the observed dates of budbreak in both Québec (±3.98 d) and Minnesota (±7.98 d). The site-independent calibration provides interesting insights on the physiological mechanisms underlying the dynamics of dormancy break and the resumption of vegetative growth in spring.

Photosynthetic parameters and stomatal conductance in attached and detached balsam fir foliage.

Leaf level gas-exchange measurements can be made on detached foliage to address the challenge of access to the crown of tall trees. However, detachment may impact leaf gas exchange. This necessitates the study of gas-exchange characteristics of foliage on detached branches to assess the feasibility of using detached branches for gas-exchange analysis. We compared photosynthetic parameters and stomatal conductance in foliage of attached and detached branches of balsam fir [Abies balsamea (L.) Mill.] during the growing season. Data were analyzed using a linear mixed-effect model, with fixed and random effects (branch status and measurement month, and tree number, respectively). Branch detachment had no significant effects on: (i) photosynthesis at the current ambient CO2 concentration (400 µmol mol-1, A 400); (ii) maximum rates of Ribulose-1,5-bisphosphate (RuBP) carboxylation (V cmax) and regeneration (J max); (iii) the ratio of J max to V cmax (i.e., J max:V cmax), and (iv) stomatal conductance (g s) during the study period (p = 0.120-0.335). There was a strong seasonal effect on all gas-exchange variables (p ≤ 0.001-0.015). Gas-exchange measurements made on detached foliage during the warm summer months should be performed with care. Reliable gas-exchange measurements can be obtained using balsam fir foliage on detached branches 50-80 cm in length, in cooler growing-season months, up to 30 min after detachment.

Host Tree Species Affects Spruce Budworm Winter Survival.

With current trends in global warming, it has been suggested that spruce budworm outbreaks may spread to northern parts of the boreal forest. However, the major constraints for a northward expansion are the availability of suitable host trees and the insect winter survival capacity. This study aimed to determine the effect of larval feeding on balsam fir, white spruce and black spruce on various spruce budworm life history traits of both the parental and the progeny generations. Results indicated that the weight of the overwintering larval progeny and their winter survival were influenced by host tree species on which larvae of the parental generation fed. White spruce was the most suitable host for the spruce budworm, producing the heaviest pupae and the heaviest overwintering larvae while black spruce was the least suitable, producing the smallest pupae and the smallest overwintering progeny. Overwintering larvae produced by parents that fed on black spruce also suffered higher winter mortality than individuals coming from parents that fed on balsam fir or white spruce. With current trends in global warming, spruce budworm is expected to expand its range to northern boreal forests where black spruce is the dominant tree species. Such northern range expansion might not result in outbreaks if low offspring winter survival on black spruce persist.

Responses of bud-break phenology to daily-asymmetric warming: daytime warming intensifies the advancement of bud break.

There is evidence that the ongoing climate change is happening through nighttime rather than daytime warming. How such a daily-asymmetric warming modifies plant phenology is still unclear. We investigated the effects of asymmetric warming on bud break by daily monitoring seedlings belonging to 26 black spruce [Picea mariana (Mill.) BSP.] and 15 balsam fir [Abies balsamea (L.) Mill.] provenances from the native range in Canada. Seedlings were subjected to either daytime or nighttime warming in three growth chambers at temperatures ranging between 10 and 24 °C. On average, a warming of 4 °C advanced the timings of bud break in both species by 2.4 days, with the later phases being more sensitive to the treatment. Bud break of both species responded more strongly to daytime warming, with the bud break occurred 1.2 and 3.2 days earlier under daytime than nighttime warming in black spruce and balsam fir, respectively. A marked ecotypic differentiation was only observed in black spruce that originated from provenances distributed broadly across Canada, with seedlings from the warmest provenance completing bud break 8.3 days later than those from the coldest one. However, no significant effect of provenance was observed for balsam fir, the narrowly distributed species. Overall, the above results suggest that a higher temporal resolution such as temperatures during daytime and nighttime, and higher spatial resolution should be taken into account to improve the accuracy of phenological model predictions under global change scenarios. Phenological models based on daily average temperature should take into account the diverging impacts of asymmetric warming on plant phenology. Our findings may indicate that the influence of warming on plant phenology may be less dramatic than expected.

Persistence of balsam fir and black spruce populations in the mixedwood and coniferous bioclimatic domain of eastern North America.

The boreal ecocline (ca 49°N) between the southern mixedwood (dominated by balsam fir) and the northern coniferous bioclimatic domain (dominated by black spruce) may be explained by a northward decrease of balsam fir regeneration, explaining the gradual shift to black spruce dominance. 7,010 sample plots, with absence of major disturbances, were provided by the Quebec Ministry of Forest, Fauna, and Parks. The regeneration (sapling abundance) of balsam fir and black spruce were compared within and between the two bioclimatic domains, accounting for parental trees, main soil type (clay and till) and climate conditions, reflected by summer growing degree-days above 5°C (GDD_5), total summer precipitation (May-August; PP_MA). Parental trees and soil type determined balsam fir and black spruce regeneration. Balsam fir and black spruce, respectively, showed higher regeneration in the mixedwood and the coniferous bioclimatic domains. Overall, higher regeneration was obtained on till for balsam fir, and on clay soils for black spruce. GDD_5 and PP_MA were beneficial for balsam fir regeneration on clay and till soils, respectively, while they were detrimental for black spruce regeneration. At a population level, balsam fir required at least 28% of parental tree basal area in the mixedwood, and 38% in the coniferous bioclimatic domains to maintain a regeneration at least equal to the mean regeneration of the whole study area. However, black spruce required 82% and 79% of parental trees basal area in the mixedwood and the coniferous domains, respectively. The northern limit of the mixedwood bioclimatic domain was attributed to a gradual decrease toward the north of balsam fir regeneration most likely due to cooler temperatures, shorter growing seasons, and decrease of the parental trees further north of this northern limit. However, balsam fir still persists above this northern limit, owing to a patchy occurrence of small parental trees populations, and good establishment substrates.

Conifer Regeneration After Experimental Shelterwood and Seed-Tree Treatments in Boreal Forests: Finding Silvicultural Alternatives.

Forest regeneration is a key element in achieving sustainable forest management. Partial harvest methods have been used extensively in temperate broadleaf and mixedwood ecosystems to promote regeneration on poorly stocked sites and to maintain forest composition and productivity. However, their effectiveness in promoting conifer establishment has yet to be demonstrated in unmanaged boreal forests, especially those dominated by black spruce (Picea mariana (Mill.) BSP) where constraints for regeneration differ from those found in more meridional regions. We aimed to evaluate conifer seedling density and dimensions, 10 years after the onset of a gradient of silvicultural treatments varying in harvesting intensities, and to identify the critical factors driving the regeneration process. Study blocks of even-aged black spruce stands in the eastern Canadian boreal forest were submitted to three variants of shelterwood harvesting: a seed-tree harvest, a clear-cut and an untreated control. Shelterwood and seed-tree harvesting were combined with spot scarification to promote regeneration. Shelterwood and seed-tree harvesting produced a density of conifer regeneration sufficient to maintain forest productivity, but they did not promote seedling growth. Black spruce was the predominant species in terms of regeneration density, with proportions 3-5× higher than that for balsam fir (Abies balsamea (L.) Mill.). Ten years after treatment, seed-origin black spruce seedlings were abundant in skidding trails, while layers dominated the residual strips. Balsam fir density was not influenced by treatment nor by tree position relative to skidding trails. Balsam fir and black spruce had different responses to treatment in terms of height and diameter, the former exhibiting a better growth performance and larger diameter in the residual strips. Spot scarification created micro-sites that had a significant impact on the regeneration process. Overall, our results support that shelterwood and seed-tree harvesting combined with scarification enable adequate regeneration in black spruce stands, confirming these treatments as viable silvicultural alternatives to clear-cutting when required by sustainable forest management objectives.

Opposing effects of mortality factors on progeny operational sex ratio may thwart adaptive manipulation of primary sex ratio.

Despite extensive research on mechanisms generating biases in sex ratios, the capacity of natural enemies to shift or further skew operational sex ratios following sex allocation and parental care remains largely unstudied in natural populations. Male cocoons of the sawfly Neodiprion abietis (Hymenoptera: Diprionidae) are consistently smaller than those of females, with very little overlap, and thus, we were able to use cocoon size to sex cocoons. We studied three consecutive cohorts of N. abietis in six forest stands to detect cocoon volume-associated biases in the attack of predators, pathogens, and parasitoids and examine how the combined effect of natural enemies shapes the realized operational sex ratio. Neodiprion abietis mortality during the cocoon stage was sex-biased, being 1.6 times greater for males than females. Greater net mortality in males occurred because male-biased mortality caused by a pteromalid parasitic wasp and a baculovirus was greater and more skewed than female-biased mortality caused by ichneumonid parasitic wasps. Variation in the susceptibility of each sex to each family of parasitoids was associated with differences in size and life histories of male and female hosts. A simulation based on the data indicated that shifts in the nature of differential mortality have different effects on the sex ratio and fitness of survivors. Because previous work has indicated that reduced host plant foliage quality induces female-biased mortality in this species, bottom-up and top-down factors acting on populations can affect operational sex ratios in similar or opposite ways. Shifts in ecological conditions therefore have the potential to alter progeny fitness and produce extreme sex ratio skews, even in the absence of unbalanced sex allocation. This would limit the capacity of females to anticipate the operational sex ratio and reliably predict the reproductive success of each gender at sex allocation.

Terrestrial acidification and ecosystem services: effects of acid rain on bunnies, baseball, and Christmas trees.

Often termed "acid rain," combined nitrogen and sulfur deposition can directly and indirectly impact the condition and health of forest ecosystems. Researchers use critical loads (CLs) to describe response thresholds, and recent studies on acid-sensitive biological indicators show that forests continue to be at risk from terrestrial acidification. However, rarely are impacts translated into changes in "ecosystem services" that impact human well-being. Further, the relevance of this research to the general public is seldom communicated in terms that can motivate action to protect valuable resources. To understand how changes in biological indicators affect human well-being, we used the STEPS (Stressor-Ecological Production function-final ecosystem Services) Framework to quantitatively and qualitatively link CL exceedances to ecosystem service impacts. We specified the cause-and-effect ecological processes linking changes in biological indicators to final ecosystem services. The Final Ecosystem Goods and Services Classification System (FEGS-CS) was used within the STEPS Framework to classify the ecosystem component and the beneficiary class that uses or values the component. We analyzed two acid-sensitive tree species, balsam fir (Abies balsamea) and white ash (Fraxinus americana), that are common in northeastern USA. These well-known species provide habitat for animals and popular forest products that are relatable to a broad audience. We identified 160 chains with 10 classes of human beneficiaries for balsam fir and white ash combined, concluding that there are resources at risk that the public may value. Two stories resulting from these explorations into the cascading effects of acid rain on terrestrial resources are ideal for effective science communication: the relationship between (1) balsam fir as a popular Christmas tree and habitat for the snowshoe hare, a favorite of wildlife viewers, and (2) white ash because it is used for half of all baseball bats, fine wood products, and musical instruments. Thus, rather than focusing on biological indicators that may only be understood or appreciated by specific stakeholders or experts, this approach extends the analysis to include impacts on FEGS and humans. It also lays the foundation for developing stakeholder-specific narratives, quantitative measures of endpoints, and for conducting demand-based valuations of affected ecosystem services.

New antibacterial hydrophobic assay reveals Abies balsamea oleoresin activity against Staphylococcus aureus and MRSA.

ETHNOPHARMACOLOGICAL RELEVANCE: Oleoresin of Abies balsamea (L.) Mill. was used by Native Americans of the boreal forest of Canada and French Canadians to treat various infections, suggesting that oleoresin has antibacterial properties. AIM OF THE STUDY: In this study, the antibacterial activity of whole oleoresin from A. balsamea was investigated against E. coli, S. aureus and two methicillin-resistant S. aureus (MRSA) strains using a new sensitive assay developed to evaluate hydrophobic matrix and compounds. MATERIALS AND METHODS: Antibacterial activity of oleoresin was first investigated using dilution and disk diffusion methods against E. coli and S. aureus, and compared to a new sensitive assay for hydrophobic matrix. Moreover, whole oleoresin was analyzed by GC-MS to characterize the composition and to identify the compounds responsible of the antibacterial activity. RESULTS: The results showed that whole oleoresin was inactive against Gram-negative E. coli (MIC90 >90µg/ml) but active against Gram-positive S. aureus and MRSA with MIC90 ranging from 18.2 to 30µg/ml. The oleoresin is mainly composed of monoterpene (28%), sesquiterpenes (2%), and diterpenes (45%). Resin acids were found, in part, responsible for the antibacterial activity of whole oleoresin. Isopimaric acid and levopimaric acid are the most active with a MIC90 of respectively 9.7µg/ml and 10µg/ml. CONCLUSION: This study supports the use of oleoresin of A. balsamea by the Native Americans and French Canadians to treat bacterial infections due to S. aureus.

Synchronisms and correlations of spring phenology between apical and lateral meristems in two boreal conifers.

Phenological synchronisms between apical and lateral meristems could clarify some aspects related to the physiological relationships among the different organs of trees. This study correlated the phenological phases of bud development and xylem differentiation during spring 2010-14 in balsam fir (Abies balsamea Mill.) and black spruce [(Picea mariana Mill. (BSP)] of the Monts-Valin National Park (Quebec, Canada) by testing the hypothesis that bud development occurs after the reactivation of xylem growth. From May to September, we conducted weekly monitoring of xylem differentiation using microcores and bud development with direct observations on terminal branches. Synchronism between the beginning of bud development and xylem differentiation was found in both species with significant correlations between the phases of bud and xylem phenology. Degree-day sum was more appropriate in assessing the date of bud growth resumption, while thermal thresholds were more suitable for cambium phenology. Our results provide new knowledge on the dynamics of spring phenology and novel information on the synchronisms between two meristems in coniferous trees. The study demonstrates the importance of precisely defining the phases of bud development in order to correctly analyse the relationships with xylem phenology.

Regulation of liver cell glucose homeostasis by dehydroabietic acid, abietic acid and squalene isolated from balsam fir (Abies balsamea (L.) Mill.) a plant of the Eastern James Bay Cree traditional pharmacopeia.

In our previous study, Abies balsamea (L.) Mill., a plant used in Cree traditional medicine, had a strong effect on the regulation of glucose homeostasis in liver cells. This study aimed to isolate and identify its active constituents using a bioassay-guided fractionation approach as well as to elucidate their mechanism(s) of action. The effect of the crude extract and its constituents was evaluated on the activity of Glucose-6-Phosphatase (G6Pase) and Glycogen Synthase (GS) and phosphorylation of three kinases, AMP-activated protein kinase (AMPK), Akt and Glycogen Synthase Kinase-3 (GSK-3). Three compounds, abietic acid, dehydroabietic acid and squalene, were isolated from the most active fraction in the bioassays (hexane). The compounds were able to decrease the activity of G6Pase and to stimulate GS. Their effect on G6Pase activity involved both Akt and AMPK phosphorylation with significant correlations between insulin-dependent and -independent pathways and the bioassay. In addition, the compounds were able to stimulate GS through GSK-3 phosphorylation with a significant correlation between the signaling pathway and the bioassay. Dehydroabietic acid stood out for its strongest effect in all the experiments close to that of the crude extract. These compounds may have potential applications in the treatment of type 2 diabetes and insulin resistance.

Integrating phylogeography and paleoecology to investigate the origin and dynamics of hybrid zones: insights from two widespread North American firs.

Secondary contact between closely related taxa routinely occurs during postglacial migrations. After initial contact, the location of hybrid zones may shift geographically or remain spatially stable over time in response to various selective pressures or neutral processes. Studying the extent and direction of introgression using markers having contrasted levels of gene flow can help unravel the historical dynamics of hybrid zones. Thanks to their contrasted maternal and paternal inheritance, resulting in different levels of gene flow for mitochondrial and chloroplast DNA (mtDNA and cpDNA), the Pinaceae stand out as a relevant biological model for this purpose. The objective of the study was to assess whether the hybrid zone between Abies balsamea and Abies lasiocarpa (two largely distributed Pinaceae) has moved or remained stable over time by analysing the distribution of cytoplasmic DNA variation as well as published palaeobotanical data. Interspecific gene flow was higher for cpDNA than mtDNA markers; hence, the geographic distribution of mitotypes was more congruent with species distributions than chlorotypes. This genetic signature was contrary to expectations under a moving hybrid zone scenario, as well as empirical observations in other conifers. Genetic evidence for this rare instance of stable hybrid zone was corroborated by the colonization chronology derived from published fossil data, indicating that the two fir species initially came into contact in the area corresponding to the current sympatric zone 11 kyr ago. While an explanatory analysis suggested the putative influence of various environmental factors on the relative abundance of cytoplasmic genome combinations, further research appears necessary to assess the role of both demographic history and selective factors in driving the dynamics of hybrid zones.

Spread of a Gammabaculovirus within Larval Populations of Its Natural Balsam Fir Sawfly (Neodiprion abietis) Host Following Its Aerial Application.

Field trials and assessments of the balsam fir sawfly (Neodiprion abietis) nucleopolyhedrovirus (NeabNPV: Baculoviridae, Gammabaculovirus) against its natural host were conducted in July and August 2002 near Corner Brook, Newfoundland and Labrador, Canada, in naturally regenerated, precommercially thinned stands dominated by balsam fir (Abies balsamea). Two experimental blocks, each with its own untreated control, were established. The purpose of the Island Pond block was to examine the spread of NeabNPV from a 313-ha aerial treatment block out into adjacent populations of balsam fir sawflies. The purpose of the Old Man's Pond block (2,093 ha) was to determine whether NeabNPV could disperse into populations of balsam fir sawflies within a 200-m zone between spray swaths. NeabNPV was applied to treatment blocks by a Cessna 188B AgTruck aircraft equipped with MicronAir AU4000 rotary atomizers at an application rate equivalent to 1 × 108 NeabNPV occlusion bodies/ha in 2.5 L of 20% aqueous molasses. At Island Pond, NeabNPV infection increased with time following the spray, especially for individuals close to the treatment block, and infection rate decreased to a measured distance of 400 m from the treatment block. At Old Man's Pond, NeabNPV infection rose higher (80% vs. 15%) and sawfly densities declined more (84% vs. 60%) in the area between spray swaths than in the control block.

Transmission of a Gammabaculovirus within Cohorts of Balsam Fir Sawfly (Neodiprion abietis) Larvae.

Nucleopolyhedroviruses (NPV: Gammabaculovirus: Baculoviridae) of diprionid sawflies (Diprionidae: Hymenoptera) are highly host specific and only infect the midgut epithelium. While still alive, infected sawfly larvae excrete NPV-laden diarrhea that contaminates food sources. The diarrhea can then be consumed by conspecific larvae, resulting in rapid horizontal transmission of the virus. To better understand the efficacy of Gammabaculovirus-based biological control products, the horizontal spread of such a virus (NeabNPV) within cohorts of balsam fir sawfly (Neodiprion abietis) larvae was studied by introducing NeabNPV-treated larvae into single-cohort groups at densities similar to those observed during the increasing (field study) and peak (laboratory study) phases of an outbreak. In field studies (~200 N. abietis larvae/m² of balsam fir (Abies balsamea) foliage), NeabNPV-induced mortality increased positively in a density-dependent manner, from 23% (in control groups) to 51% with the addition of one first-instar NeabNPV-treated larva, to 84% with 10 first-instar-treated larvae. Mortality was 60% and 63% when one or 10 NeabNPV-treated third-instar larva(e), respectively, were introduced into groups. Slightly higher levels of NeabNPV-induced mortality occurring when NeabNPV-treated larvae were introduced into first- rather than third-instar cohorts suggests that early instars are more susceptible to the virus. In the laboratory (~1330 N. abietis larvae/ m² of foliage), NeabNPV-caused mortality increased from 20% in control groups to over 80% with the introduction of one, five or 10 NeabNPV-treated larvae into treatment groups of first-instar larvae.

Production, Application, and Field Performance of AbietivTM, the Balsam Fir Sawfly Nucleopolyhedrovirus / 中国病毒学

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.

Testing the ecophysiological basis for the control of monoterpene concentrations in thinned and unthinned balsam fir stands across different drainage classes.

Stand thinning across different soil drainage types was used to test the ecophysiological basis for the control of foliar monoterpenes in current-year foliage of balsam trees [Abies balsamea (L.) P. Mill.] the year following treatment. Photosynthetic capacity (A max) was greater on mesic sites than on sub-hygric and hydric sites (1.81, 1.33 and 0.88 µmol m-2 s-1, respectively) and greater in mid-July than at the end of June (1.86 vs 0.77 µmol m-2 s-1). Foliar N was greater on thinned plots than control plots (2.6 vs 2.2 g m-2); starch was greater on mesic control plots than mesic thinned plots (16.3 vs 10.9 g m-2); and total soluble sugars (TSS) were higher in mid-July than at the end of June (16.5 vs 12.8 g m-2). Total monoterpenes, as well as five individual monoterpenes, showed a complex interaction between site drainage class, thinning treatment and time of sampling (P<0.05). For all sites and dates combined, total monoterpenes were negatively correlated with specific leaf area (r 2=0.46), and positively but weakly correlated with A max (r 2=0.35), foliar N (r 2=0.29) and TSS (r 2=0.24). Total monoterpenes were not correlated with the different measures of carbon to nitrogen ratio as predicted by the carbon-nutrient balance hypothesis. Our results offer partial support to the different hypotheses proposing that foliage having greater resource availability, enzymatic machinery and potential for building storage structures will have greater monoterpene content. However, the strength of the relationships with monoterpenes often varied with sampling date, suggesting that phenological development may play a strong role in determining which factors control monoterpenes at a given time.

Testing the ecophysiological basis for the control of monoterpene concentrations along canopy profiles in thinned and unthinned balsam fir stands.

To determine which ecophysiological factors appear to control monoterpene concentrations in balsam fir foliage [Abies balsamea (L.) P. Mill.], the percentage of photosynthetically active radiation (%PAR), specific leaf area (SLA), light-saturated photosynthesis (A max), and concentrations per unit leaf area of foliar nitrogen (N), total soluble sugars (TSS), starch and monoterpenes were measured on current-year needles from three canopy levels (upper, middle and lower) the year following a pre-commercial thinning. The thinning only modestly changed the light profile within the canopy. %PAR was negatively correlated with SLA (r 2=0.62 in June, r 2=0.53 in July and August) and positively correlated with foliar nitrogen concentrations (r 2=0.51) within the crown profile. The positive relationship between N and A max was quite weak (r 2=0.15), suggesting significant variations in non-photosynthetic N within the canopies. Total monoterpenes were positively correlated with both %PAR (r 2=0.29) and A max (r 2=0.27), and negatively correlated with SLA (r 2=0.30). Contrary to that predicted by the carbon-nutrient balance hypothesis, total monoterpenes were negatively and only very weakly correlated with the starch/N ratio (r 2=0.06) and were not significantly correlated with either the TSS/N or the [TSS+starch]/N ratios. Monoterpenes were positively correlated with both N and TSS, although the relationship varied with the phenological state of the foliage, i.e., monoterpenes were more highly correlated with TSS (r 2=0.67) (immature foliage) in June, and in July and August with N (r 2=0.63) (mature foliage). Thus, it appears that monoterpene concentrations may be controlled primarily by carbohydrate supply in the early growing season and later by enzymatic capacity. Data expressed on a dry weight basis showed a similar pattern.

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

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.

Factors influencing choice of balsam fir twigs from thinned and unthinned stands by moose.

Moose were observed to browse preferentially on balsam fir (Abies balsamea) trees in stands where the stem density had been mechanically reduced to about 2000 stems/ha from over 30,000 stems/ha. Twigs from trees in thinned and unthinned stands were analyzed to test the hypotheses that moose were choosing thinned stands to maximize intake of a nutrient, or avoiding plant secondary compounds deliterious to digestion. Analyses included: twig length, weight, diameter, acid detergent fiber, neutral detergent fiber, dry matter digestibility, macroelements, crude fat (resins), crude protein, ash, and volatile secondary metabolites. Significantly higher concentrations of crude protein, P, Ca, Na, and crude fats occurred in trees from thinned compared to unthinned stands. Twigs from thinned stands were more digestible, longer, heavier, and had a greater diameter than those from unthinned stands. Several secondary metabolites were found in highest concentration in thinned stands. We suggest that moose chose thinned stands over unthinned stands for feeding because of high protein levels and large twigs. Trees in unthinned stands had protein levels below reported maintenance levels required by moose. Secondary metabolite concentrations were opposite to the direction predicted and the data do not support plant defence hypotheses for the chemicals analyzed.