Accurately predicting biomass allocation to below- versus above-ground tree parts is crucial in estimating carbon stocks in forest ecosystems. A 9-year outdoor experiment was conducted to analyze the variations in biomass allocation in below- and above-ground parts in black spruce trees growing at the edge or at the center of a raised garden bed. Dry biomass, length, and radial growth of stems, branches, and roots were measured as well as the annual above-ground growth for the five most recent years prior to harvesting. We found that more than 90% of dry biomass was allocated to the above-ground tree parts whereas less than 10% was allocated to the root system. Strong correlations were found between the different tree parts regardless of the tree’s position in the delimited growth area. Annual growth variables declined from 2018 to 2022, likely due to increased competition for resources. The dry biomass of the woody root and the root surface close to the stem were well correlated to the above-ground tree parts (varying between 45% and 95%). Thus, the strong link between the root system and the above tree parts is confirmed.
Controlled experiments suggest that the seasonal build-up of nitrogen (N) limitation constrains the responses of forest autumn phenology to elevated temperatures. Therefore, rising soil N is expected to increase the delaying effects of elevated temperature on the end of the season, i.e., leaf senescence. However, the interactive effects of temperature, soil N, and aridity on xylem autumn phenology remain unknown. We conducted a wide spatial analysis from 75 conifer sites in the Northern Hemisphere and found that rising soil N increases the delaying effects of elevated temperature on the end of xylem cell wall thickening but reduced the delaying effects on the cessation of cell enlargement, especially in humid regions. The contrasting effects of elevated soil N on cell enlargement versus cell wall thickening could affect xylem cell anatomy, thereby induce changes in wood density, and induce a decoupling of stem size growth from photosynthate production. These analyses extend previous findings on forest autumn phenology by systematically investigating the spatial variation in the interactive effects of temperature and soil N on xylem autumn phenology at the cellular scale.
Wood growth is key to understanding the feedback of forest ecosystems to the ongoing climate warming. An increase in spatial synchrony (i.e., coincident changes in distant populations) of spring phenology is one of the most prominent climate responses of forest trees. However, whether temperature variability contributes to an increase in the spatial synchrony of spring phenology and its underlying mechanisms remains largely unknown. Here, we analyzed an extensive dataset of xylem phenology observations of 20 conifer species from 75 sites over the Northern Hemisphere. Along the gradient of increase in temperature variability in the 75 sites, we observed a convergence in the onset of cell enlargement roughly toward the 5th of June, with a convergence in the onset of cell wall thickening toward the summer solstice. The increase in rainfall since the 5th of June is favorable for cell division and expansion, and as the most hours of sunlight are received around the summer solstice, it allows the optimization of carbon assimilation for cell wall thickening. Hence, the convergences can be considered as the result of matching xylem phenological activities to favorable conditions in regions with high temperature variability. Yet, forest trees relying on such consistent seasonal cues for xylem growth could constrain their ability to respond to climate warming, with consequences for the potential growing season length and, ultimately, forest productivity and survival in the future.
Large areas in boreal forests are classified as peatlands, characterized by organic soils with a high water table. Black spruce (Picea mariana (Mill.) B.S.P.) is one of the species capable of growing in this inauspicious environment, where an adaptation of the root system can be expected. We studied young black spruces growing in peat moss with two different hydrological conditions over a 19 years timespan: saturated and well-drained peat. We identified the initial and adventitious roots of the trees and compiled radial growth measurements of each root. The general growth pattern of the roots was identified and compared to the annual radial growth within the stem. We observed growth reductions during the first years after the planting shock, followed by a growth increase in the roots and stems for both hydrological conditions. The continuous formation of adventitious roots in trees growing in the saturated environment was the main adaptation noticed. The largest radial growth values were registered in the younger adventitious roots growing in saturated conditions. The lowest radial growth in the adventitious roots were registered in the well-drained condition. PCA analyses revealed the influence of root depth with regard to stem height and diameter at soil level. The black spruce trees displayed the required plasticity to form adventitious roots outside the range of the water table, allowing them a better access to oxygen and nutrients. This survival strategy implies to allocate a higher amount of energy to the root system instead of the aerial part of the tree in which overall productivity is low.
Despite growing interest in predicting plant phenological shifts, advanced spring phenology by global climate change remains debated. Evidence documenting either small or large advancement of spring phenology to rising temperature over the spatio‐temporal scales implies a potential existence of a thermal threshold in the responses of forests to global warming. We collected a unique data set of xylem cell‐wall‐thickening onset dates in 20 coniferous species covering a broad mean annual temperature (MAT) gradient (−3.05 to 22.9°C) across the Northern Hemisphere (latitudes 23°–66° N). Along the MAT gradient, we identified a threshold temperature (using segmented regression) of 4.9 ± 1.1°C, above which the response of xylem phenology to rising temperatures significantly decline. This threshold separates the Northern Hemisphere conifers into cold and warm thermal niches, with MAT and spring forcing being the primary drivers for the onset dates (estimated by linear and Bayesian mixed‐effect models), respectively. The identified thermal threshold should be integrated into the Earth‐System‐Models for a better understanding of spring phenology in response to global warming and an improved prediction of global climate‐carbon feedbacks.
Research on endangered species is important for species conservation. Some lines of conservation research require tissue or fluids so that molecular data can be obtained. Thus, there is an increasing demand for methods of DNA sampling in endangered species that minimize harm to live animals. This especially concerns such highly threatened and protected freshwater mussels as the thick-shelled river mussel Unio crassus. With two different field experiments in Germany under natural conditions, the short- and long-term survival of Unio crassus after hemolymph extraction from the foot tissue was evaluated. We detected no negative effects in individuals of over 40 mm shell length and older than five years. We also followed some individuals for up to three years after treatment and detected no harm to these individuals. DNA quality was high and proved fully suitable for single-gene sequencing (COI) and microsatellite analyses.
Eastern spruce budworm (Choristoneura fumiferana Clem.) is considered the most important disturbing insect in coniferous stands in eastern North America. During an outbreak, spruce budworm can cause severe defoliation in balsam fir (Abies balsamea (L.) Mill.), which can affect wood properties such as moisture content and mechanical properties. This project aimed to assess the influence of the duration of spruce budworm defoliation on the wood quality of mature balsam fir trees. To do this, we studied sapwood proportion, decay, moisture content, mechanical properties and tracheid dimensions in stands that had suffered three, four or five years of defoliation. We also compared living and dead balsam firs and evaluated the change in wood properties with time. Our results showed that dead balsam firs suffered from a loss of wood quality rapidly after their death, particularly in terms of moisture content and decay in the sapwood. Sapwood proportion was similar between living and dead trees, but the sapwood of dead trees contained more decay and had a lower moisture content than living trees. Mechanical properties and tracheid dimensions were 10% and 4% lower in dead trees than in living trees. We did not observe any major differences in wood properties between the three durations of defoliation, suggesting that wood degradation occurs before that. The study did not make it possible to determine the optimal duration of defoliation to harvest the stands.
a dozen the Tree-Ring Research University of Arizona for an International Workshop on Dendroclimatology. Organized by Harold C. Fritts, the Workshop was designed, among other things, to acquaint important members of the dendrochronological community with one another and to develop common goals, procedures, and standards for the burgeoning practice of dendroclimatology. Dieter was an enthusiastic participant in these efforts and was instrumental in furthering the goals of the Workshop both during and after its deliberations. Notable accomplishments of the meeting include establishing a collec-tive basis for future dendroclimatic research, laying the founda- tions for what would become the International Tree-Ring Data Base, and transforming the somewhat parochial Tree-Ring So- ciety into an international organization that represents dendrochronology around the globe. My contact with Dieter did not end with the Dendroclimatic Workshop. As long-term officers of the reformulated Tree-Ring Society we frequently consulted on Society policies and operations as well as current developments in the fields. As an archaeologist, I am particularly interested in and appreciative of Dieter’s interests in and manifest contributions to the many aspects of dendroarchaeology. I remain in awe of his pioneering work at Haithabu (Hedeby) and his continuing work along these lines. We had many fruitful discussions of is-sues concerning the relationships between tree-ring dates and other kinds of archaeological data.
Black spruce (Picea mariana (Mill.) BSP) is the main tree species growing within peatlands. The difficult growing conditions within peatlands are associated with low individual wood productivity, and only tree species tolerant to high ground water level can survive. We examine the effect of water-saturated soil on the growth potential of black spruce trees and specific adaptation of the root system. Experimental mesocosms were constructed with two drainage regimes (saturated and well-drained soil conditions). We measured biomass, height, and diameter at stem base of 55 black spruce saplings and noted the location of each horizontal root. Black spruce exhibited a very shallow root system located above saturated commercial peat 19 years after the experiment was initiated. Most roots were adventitious in the water-saturated mesocosms. Overall aerial and root biomass accumulation of the black spruce trees growing under saturated soil conditions was significantly lower than that of the well-drained mesocosms. Interestingly, root–shoot ratios were similar across the two drainage regimes. Soil conditions induced adaptation of the root system in black spruce trees, and physiological stress affected the entire individual with lower biomass productivity in all components (stem, branches, root system). However, biomass distribution remained similar to that of trees growing in well-drained mesocosms.
AbstractThe taxonomy of the Mediterranean Aristolochia pallida complex has been under debate since several decades with the following species currently recognized: A. pallida, A. lutea, A. nardiana, A. microstoma, A. merxmuelleri, A. croatica, and A. castellana. These taxa are distributed from Iberia to Turkey. To reconstruct phylogenetic and biogeographic patterns, we employed cpDNA sequence variation using both noncoding (intron and spacer) and protein‐coding regions (i.e., trnK intron, matK gene, and trnK‐psbA spacer). Our results show that the morphology‐based traditional taxonomy was not corroborated by our phylogenetic analyses. Aristolochia pallida, A. lutea, A. nardiana, and A. microstoma were not monophyletic. Instead, strong geographic signals were detected. Two major clades, one exclusively occurring in Greece and a second one of pan‐Mediterranean distribution, were found. Several subclades distributed in Greece, NW Turkey, Italy, as well as amphi‐Adriatic subclades, and a subgroup of southern France and Spain, were revealed. The distribution areas of these groups are in close vicinity to hypothesized glacial refugia areas in the Mediterranean. According to molecular clock analyses the diversification of this complex started around 3–3.3 my, before the onset of glaciation cycles, and the further evolution of and within major lineages falls into the Pleistocene. Based on these data, we conclude that the Aristolochia pallida alliance survived in different Mediterranean refugia rarely with low, but often with a high potential for range extension, and a high degree of morphological diversity.
During spruce budworm (Choristoneura fumiferana (Clem.)) outbreaks, stands dominated by balsam fir (Abies balsamea (L.) Mill.) suffer marked levels of mortality after consecutive years of defoliation. The balsam fir mortality rate is also influenced by such factors as stand age and forest composition. Nonetheless, the relative importance of these factors when predicting balsam fir mortality during an outbreak remains uncertain. Here we present models for predicting balsam fir mortality rates on the basis of defoliation duration and intensity, stand age, and forest composition. We found a significant increase in balsam fir mortality rates with an increased duration and intensity of defoliation. Balsam fir experienced >22% mortality after five years of continuous severe defoliation. The model identified cumulative defoliation-a commonly used measure in Canada of the sum of annual defoliation intensities experienced by a stand over a given defoliation period-as an effective index for predicting mortality. Detailed cumulative defoliation, a factor specifying the frequency of different levels of annual defoliation intensities experienced by a stand over a given defoliation period, provided even more reliable results. Mature stands experienced 5% greater mortality than immature stands, and mortality was 3% and 8% greater in pure balsam fir stands than in mixed balsam fir-spruce and mixed balsam fir-hardwood stands, respectively. The proportion of other tree species also affected the mortality prediction model. We attained 70% accuracy for our best model for predicting balsam fir mortality rate in heterogeneous mixed stands; this model included detailed cumulative defoliation, stand age, and the proportion of other species in the stand. Therefore, these factors must be included in decisions related to recovery cutting in forests affected by spruce budworm outbreaks.
Abstract In boreal landscapes, emphasis is currently placed on close‐to‐nature management strategies, which aim to maintain the biodiversity and ecosystem services related to old‐growth forests. The success of these strategies, however, depends on an accurate understanding of the dynamics within these forests. While moderate‐severity disturbances have recently been recognized as important drivers of boreal forests, little is known about their effects on stand structure and growth. This study therefore aimed to reconstruct the disturbance and postdisturbance dynamics in boreal old‐growth forests that are driven by recurrent moderate‐severity disturbances. We studied eight primary old‐growth forests in Québec, Canada, that have recorded recurrent and moderately severe spruce budworm (Choristoneura fumiferana [Clem.]) outbreaks over the 20th century. We applied an innovative dendrochronological approach based on the combined study of growth patterns and releases to reconstruct stand disturbance and postdisturbance dynamics. We identified nine growth patterns; they represented trees differing in age, size, and canopy layer. These patterns highlighted the ability of suppressed trees to rapidly fill gaps created by moderate‐severity disturbances through a single and significant increase in radial growth and height. Trees that are unable to attain the canopy following the disturbance tend to remain in the lower canopy layers, even if subsequent disturbances create new gaps. This combination of a low stand height typical of boreal forests, periodic disturbances, and rapid canopy closure often resulted in stands constituted mainly of dominant and codominant trees, similar to even‐aged forests. Overall, this study underscored the resistance of boreal old‐growth forests owing to their capacity to withstand repeated moderate‐severity disturbances. Moreover, the combined study of growth patterns and growth release demonstrated the efficacy of such an approach for improving the understanding of the fine‐scale dynamics of natural forests. The results of this research will thus help develop silvicultural practices that approximate the moderate‐severity disturbance dynamics observed in primary and old‐growth boreal forests.
Research Highlights: Radial growth patterns of trees growing in old-growth boreal forests in eastern Canada can be grouped into a small number of simple patterns that are specific to different old-growth forest types or successional stages. Background and Objectives: Identifying the main radial growth trends in old-growth forests could help to develop silvicultural treatments that mimic the complex dynamics of old-growth forests. Therefore, this study aimed to identify the main radial growth patterns and determine how their frequencies change during forest succession in old-growth forests, focusing on boreal landscapes in eastern Canada. Materials and Methods: We used dendrochronological data sampled from 21 old-growth stands in the province of Quebec, Canada. Tree-ring chronologies were simplified into chronologies of equal length to retain only primary growth trends. We used k-means clustering to identify individual growth patterns and the difference in growth-pattern frequency within the studied stands. We then used non-parametric analyses of variance to compare tree or stand characteristics among the clusters. Results: We identified six different growth patterns corresponding to four old-growth forest types, from stands at the canopy breakup stage to true old-growth stands (i.e., when all the pioneer cohort had disappeared). Secondary disturbances of low or moderate severity drove these growth patterns. Overall, the growth patterns were relatively simple and could be generally separated into two main phases (e.g., a phase of limited radial increment size due to juvenile suppression and a phase of increased radial increment size following a growth release). Conclusions: The complexity of old-growth forest dynamics was observed mainly at the stand level, not at the tree level. The growth patterns observed in true old-growth forests were similar to those observed following partial or stem-selection cuts in boreal stands; thus, these silvicultural treatments may be effective in mimicking old-growth dynamics.
Significance Forest trees can live for hundreds to thousands of years, and they play a critical role in mitigating global warming by fixing approximately 15% of anthropogenic CO 2 emissions annually by wood formation. However, the environmental factors triggering wood formation onset in springtime and the cellular mechanisms underlying this onset remain poorly understood, since wood forms beneath the bark and is difficult to monitor. We report that the onset of wood formation in Northern Hemisphere conifers is driven primarily by photoperiod and mean annual temperature. Understanding the unique relationships between exogenous factors and wood formation could aid in predicting how forest ecosystems respond and adapt to climate warming, while improving the assessment of long-term and high-resolution observations of global biogeochemical cycles.
Understanding the biological basis of social anxiety disorder (SAD), one of the most disabling of the anxiety disorders, will allow for novel treatment strategies to be developed. Here, we show that gut microbiota may be such a target. Mice ...Social anxiety disorder (SAD) is a crippling psychiatric disorder characterized by intense fear or anxiety in social situations and their avoidance. However, the underlying biology of SAD is unclear and better treatments are needed. Recently, the gut ...
In this work, we examine the effects of commercial thinning on stand volume and individual stem form in nine naturally regenerated black spruce (Picea mariana (Mill.) B.S.P.) stands. We compared these study sites with controls in the commercial boreal forest of northern Quebec, Canada. At stand level, dendrochronological data provided insight into changes in stand volume ten years after thinning. Analysis of a subsample of six individual trees from each commercially thinned stand and three individual trees from the controls illustrated the role of thinning on stem shape development. Although average volume increased for residual stems in thinned stands slightly more than in the controls (110% versus 106%), the treatment effect stand-level volume increment or stand-level total volume harvested (ten years after treatment) was not statistically significant. Moreover, at tree level, thinning did not significantly affect stem volume increment. However, radial growth increment significantly increased after treatment. Trees from commercially thinned stands showed a significantly higher growth increment along the lower first two-thirds of the stem. Response to thinning at tree level correlated strongly with the size and number of harvested competitors around a residual stem. We conclude that commercial thinning modified wood allocation rather than wood volume and did not affect taper and stem shape. These patterns of post-cutting growth are explained by wood allocation following thinning. After commercial thinning, growth increment is favored at the expense of height growth. As the treatment effect was found at the stem scale rather than at the stand scale, further research is needed in regard to commercial thinning treatments to investigate how to increase productivity at the stand scale.
Spruce budworm (SBW - Choristoneura fumifercma Clem.) is one of the most damaging defoliating insects in the coniferous forests of eastern North America. In Canada, the widely distributed balsam fir (Abies balsamea L. (Mill)) and black spruce (Picea mariana B.S.P. (Mill)) are its most important hosts. Defoliation by SBW reduces growth in the host trees and can lead to host mortality. Although SBW impacts on growth are well documented, much less is known about changes in wood properties resulting from defoliation. To address this knowledge gap, we sampled 36 SBW-infested stands located in the boreal forest of Quebec (Canada) to determine whether defoliation modifies the wood quality of affected trees. The selected stands had been subjected to one to four years of SSW defoliation. For both species, we assessed ring growth, wood density, and the anatomical characteristics of stem wood formed during the outbreak years. We determined that rings formed during the SBW outbreak had a significant and progressive loss of biomass production with a longer duration of defoliation. SBW significantly reduced latewood density in the second and third year of defoliation for black spruce and the third and fourth year for balsam fir. Average ring density was reduced only in black spruce and only after four years of defoliation. The observed changes in growth and density were associated with changes in anatomical features. While the cellular characteristics of the earlywood remained fairly constant, significant reductions occurred in latewood cell-wall thickness after three years of defoliation. Our study shows that not only do SBW outbreaks reduce annual radial growth, but the cellular characteristics in latewood cells are also modified momentarily. Thus, SBW outbreaks affect wood density and quality in both black spruce and balsam fir.
Branch diameter is an important aspect of wood quality, as lumber grades can be determined based on the maximum diameter of branches. Crown and branch development can be influenced by the environment surrounding the trees, and silvicultural interventions, which reduce stand density and increase the growth of residual trees, could therefore alter branch properties. We evaluated maximum branch diameter within the crown of residual black spruce (Picea mariana (Mill.) B.S.P.) subjected to five types of silvicultural intervention—three partial-cutting and two clearcutting treatments—as well as trees within unmanaged control stands. We sampled a total of 41 stands and 223 trees. We collected 15 whorls from the live crown of each tree and measured the diameters of the largest branches. For all treatments, we observed a curvilinear relationship between maximum branch diameter and distance from the stem apex, and the largest branches were located in the lower third of the live crown. DBH before treatment and treatment were the variables that best explained maximum branch diameter in the lowest portion of the crown. A generalized additive model showed that maximum branch diameter in black spruce following silvicultural treatment will not differ significantly from trees of unmanaged control stands. Therefore, the studied partial cutting and clearcutting treatments do not have adverse effects on maximum branch diameter when compared to unmanaged control stands. However, DBH prior to treatment must be considered before any treatment is applied in forest management operations if maximum branch diameter is an important wood quality factor at the time of the final harvest of the stands.
Wood properties were generally similar in unmanaged even-aged and uneven-aged black spruce stands. In contrast, in managed forests, wood density was higher in even-aged than in uneven-aged stands. As ring width is the discriminant factor, forest management should regulate the intensity of partial cutting to maintain a high wood density. North American boreal forests experience recurrent disturbances, such as fire and insect outbreaks, that shape the structure of even-aged and uneven-aged stands. Ecosystem-based forest management practices, such as partial cutting, aim to increase timber yields, and these forest harvesting approaches are being applied more frequently. Although the influence of these practices on tree growth is well known, few studies have examined their impact on anatomy and density of the resulting wood. This research studied the relationships between wood properties and ring width and tree heights in even- and uneven-aged black spruce (Picea mariana (Mill.) BSP) stands and how these relationships are modified following partial cutting. Wood density (ring, earlywood, latewood, and maximum density) and anatomical traits (lumen area, cell-wall thickness, and tracheid length) were measured at breast height on the stem of black spruce trees. The measured wood properties were then correlated with ring width and stem height. The model demonstrated significant correlations between most measured wood parameters and ring width with only small differences between even- and uneven-aged unmanaged stands. Five wood properties were influenced by tree height in even-aged and uneven-aged unmanaged plots. Partial cutting increased the number of significant relationships between wood properties (ring, latewood, maximum density, early- and latewood lumen area, tracheid length) and tree height. We detected differences in wood properties between even- and uneven-aged plots. Although we detected no change in average ring density, the variation of intra-annual density increased, due to changes in earlywood and latewood density that resulted in more heterogeneous rings. Partial cutting should take into consideration the age structure of the stand when the density of wood products matters.