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Northern temperate coniferous forests serve as crucial connectors between boreal and temperate forests, yet they are vulnerable to various stressors such as climate change and human activities. Severe drought poses a significant threat to plant species within these forests, prompting recent research into its impacts. However, many studies lack explicit definitions of post-disturbance vegetation processes and fail to identify potential interactions with disturbance factors, necessitating comprehensive discussions. This study examines the effects of drought on tree growth patterns of the main dominant species in northern temperate regions: Abies nephrolepis and Picea jezoensis, along with two commonly associated Betula ermanii, and Quercus mongolica. Additionally, new disturbance factors in forests inhabited by these species (A. nephrolepis and P. jezoensis) were evaluated based on community classification. The study sites were located in the Mt. Baekdu (Changbai) and South Korea regions, which are positioned at the southern limit of the phytogeographical patterns of target species. Results indicate that A. nephrolepis and P. jezoensis exhibit high levels of recovery and resilience, while B. ermanii and Q. mongolica demonstrate high resistance. Species-specific responses align with drought intensity, with resistance, recovery, and resilience decreasing notably with increasing pre-drought radial growth. South Korean forests, the invasion of the vine species Tripterygium regelii after the death of A. nephrolepis in the overstory vegetation threatens the regeneration of new trees. However, certain environmental factors, such as high rock exposure and dense overstory canopy, limit vine invasion. Based on the results, pre-drought radial growth emerges as a key determinant in how trees respond to drought. Additionally, the results suggest the potential for new disturbances to emerge in forest gaps due to overstory vegetation mortality induced by global warming. These findings contribute to a deeper understanding of increasing drought stress, aid in identifying climate refugia, and inform conservation priorities based on habitat characteristics., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 The Authors. Published by Elsevier B.V. All rights reserved.)

Lowland northern white-cedar (Thuja occidentalis L.) forests are increasingly exposed to extreme droughts and floods that cause tree mortality. However, it is not clear the extent to which these events may differentially affect regeneration of cedar and its increasingly common associate, balsam fir (Abies balsamea (L.) Mill.). To test this, we measured how seedlings of cedar and fir were able to avoid, resist and recover from experimental drought and flood treatments of different lengths (8 to 66 days). Overall, we found that cedar exhibited a strategy of stress resistance and growth recovery (resilience) from moderate drought and flood stress. Fir, on the other hand, appears to be adapted to avoid drought and flood stress and exhibited overall lower growth resilience. In drought treatments, we found evidence of different stomatal behaviors. Cedar used available water quickly and therefore experienced more drought stress than fir, but cedar was able to survive at water potentials > 3 MPa below key hydraulic thresholds. On the other hand, fir employed a more conservative water-use strategy and therefore avoided extremely low water potential. In response to flood treatments, cedar survival was higher and only reached 50% if exposed to 23.1 days of flooding in contrast to only 7.4 days to reach 50% mortality for fir. In both droughts and floods, many stressed cedar were able to maintain partially brown canopies and often survived the stress, albeit with reduced growth, suggesting a strategy of resistance and resilience. In contrast, fir that experienced drought or flood stress had a threshold-type responses and they either had full live canopies with little effect on growth or they died suggesting reliance on a strategy of drought avoidance. Combined with increasingly variable precipitation regimes, seasonal flooding and complex microtopography that can provide safe sites in these forests, these results inform conservation and management of lowland cedar stands., (Published by Oxford University Press 2024.)

Mistletoes are xylem-tapping hemiparasites that rely on their hosts for water and nutrient uptake. Thus, they impair tree performance in the face of environmental stress via altering the carbon and water relations and nutritional status of trees. To improve our understanding of physiological responses to mistletoe and ongoing climate change, we investigated radial growth, stable carbon and oxygen isotopic signals, and elemental composition of tree rings in silver fir (Abies alba Mill.) and Scots pine (Pinus sylvestris L.) forests infested with Viscum album L. We compared temporal series (1990-2020) of basal area increment (BAI), intrinsic water-use efficiency (iWUE), oxygen isotope composition (δ18O), nutrient concentrations and stoichiometric ratios between non-infested (NI) and severely infested (SI) fir and pine trees from populations located close to the xeric distribution limit of the species in north-eastern Spain. The SI trees showed historically higher growth, but the BAI trend was negative for more than three decades before 2020 and their growth rates became significantly lower than those of NI trees by the mid-2010s. Mistletoe infestation was related to an enhanced sensitivity of radial growth to vapour pressure deficit (atmospheric drought). The SI trees showed less pronounced iWUE increases (fir) and lower iWUE values (pine) than NI trees. The lower tree-ring δ18O values of SI trees may be the result of several superimposed effects operating simultaneously, including leaf-level evaporative enrichment, source water isotopic signals, and anatomical and phenological differences. We observed a deterioration of potassium (K) nutrition in tree-ring wood of both species in SI trees, along with accumulation of manganese (Mn). We suggest that such nutritional patterns are driven by the indirect effect of mistletoe-induced drought stress, particularly in pine. The combined analyses of different physiological indicators imprinted on tree rings provided evidence of the progressive onset of carbon, water and nutrient imbalances in mistletoe-infested conifers inhabiting seasonally dry regions., (© The Author(s) 2024. Published by Oxford University Press.)

Disentangling the factors influencing the climate sensitivity of trees is crucial to understanding the susceptibility of forests to climate change. Reducing tree-to-tree competition and mixing tree species are two strategies often promoted to reduce the drought sensitivity of trees, but it is unclear how effective these measures are in different ecosystems. Here, we studied the growth and physiological responses to climate and severe droughts of silver fir and Douglas-fir growing in pure and mixed conditions at three sites in Switzerland. We used tree-ring width data and carbon (δ13C), oxygen (δ18O) and hydrogen (δ2H) stable isotope ratios from tree-ring cellulose to gain novel information on water relations and the physiology of trees in response to drought and how tree species mixture and competition modulate these responses. We found significant differences in isotope ratios between trees growing in pure and mixed conditions for the two species, although these differences varied between sites, e.g. trees growing in mixed conditions had higher δ13C values and tree-ring width than trees growing in pure conditions for two of the sites. For both species, differences between trees in pure and mixed conditions regarding their sensitivity to temperature, precipitation, climatic water balance and vapor pressure deficit were minor. Furthermore, trees growing in pure and mixed conditions showed similar responses of tree-ring width and isotope ratios to the past severe droughts of 2003, 2015 and 2018. Competition had only a significantly negative effect on δ13C of silver fir, which may suggest a decrease in photosynthesis due to higher competition for light and nutrients. Our study highlights that tree species mixture may have only moderate effects on the radial growth and physiological responses of silver fir and Douglas-fir to climatic conditions and that site condition effects may dominate over mixture effects., (© The Author(s) 2024. Published by Oxford University Press.)

The resilience of forests to drought events has become a major natural resource sustainability concern, especially in response to climate change. Yet, little is known about the legacy effects of repeated droughts, and tree species ability to respond across environmental gradients. In this study, we used a tree-ring database (121 sites) to evaluate the overall resilience of tree species to drought events in the last century. We investigated how climate and geography affected the response at the species level. We evaluated temporal trends of resilience using a predictive mixed linear modeling approach. We found that pointer years (e.g., tree growth reduction) occurred during 11.3% of the 20th century, with an average decrease in tree growth of 66% compared to the previous period. The occurrence of pointer years was associated with negative values of the Standardized Precipitation Index (SPI, 81.6%) and Palmer Drought Severity Index (PDSI, 77.3%). Tree species differed in their resilience capacity, however, species inhabiting xeric conditions were less resistant but with higher recovery rates (e.g., Abies concolor, Pinus lambertiana, and Pinus jeffreyi). On average, tree species needed 2.7 years to recover from drought events, with extreme cases requiring more than a decade to reach pre-drought tree growth rates. The main abiotic factor related to resilience was precipitation, confirming that some tree species are better adapted to resist the effects of droughts. We found a temporal variation for all tree resilience indices (scaled to 100), with a decreasing resistance (-0.56 by decade) and resilience (-0.22 by decade), but with a higher recovery (+1.72 by decade) and relative resilience rate (+0.33 by decade). Our results emphasize the importance of time series of forest resilience, particularly by distinguishing the species-level response in the context of legacy of droughts, which are likely to become more frequent and intense under a changing climate., (© 2023 John Wiley & Sons Ltd.)

Climate change challenges the adaptive capacity of several forest tree species in the face of increasing drought and rising temperatures. Therefore, understanding the mechanistic connections between genetic diversity and drought resilience is highly valuable for conserving drought-sensitive forests. Nonetheless, the post-drought recovery in trees from a transcriptomic perspective has not yet been studied by comparing contrasting phenotypes. Here, experimental drought treatments, gas-exchange dynamics and transcriptomic analysis (RNA-seq) were performed in the relict and drought-sensitive fir Abies pinsapo Boiss. to identify gene expression differences over immediate (24 h) and extended drought (20 days). Post-drought responses were investigated to define resilient and sensitive phenotypes. Single nucleotide polymorphisms (SNPs) were also studied to characterize the genomic basis of A. pinsapo drought resilience. Weighted gene co-expression network analysis showed an activation of stomatal closing and an inhibition of plant growth-related genes during the immediate drought, consistent with an isohydric dynamic. During the extended drought, transcription factors, as well as cellular damage and homeostasis protection-related genes prevailed. Resilient individuals activate photosynthesis-related genes and inhibit aerial growth-related genes, suggesting a shifting shoot/root biomass allocation to improve water uptake and whole-plant carbon balance. About, 152 fixed SNPs were found between resilient and sensitive seedlings, which were mostly located in RNA-activity-related genes, including epigenetic regulation. Contrasting gene expression and SNPs were found between different post-drought resilience phenotypes for the first time in a forest tree, suggesting a transcriptomic and genomic basis for drought resilience. The obtained drought-related transcriptomic profile and drought-resilience candidate genes may guide conservation programs for this threatened tree species., (© The Author(s) 2022. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permission@oup.com.)

(1) Background: The genetic basis of local adaptation in conifers remains poorly understood because of limited research evidence and the lack of suitable genetic materials. Sakhalin fir ( Abies sachalinensis ) is an ideal organism for elucidating the genetic basis of local adaptation because its altitudinal adaptation has been demonstrated, and suitable materials for its linkage mapping are available. (2) Method: We constructed P336 and P236 linkage maps based on 486 and 516 single nucleotide polymorphisms, respectively, that were derived from double digest restriction site-associated DNA sequences. We measured the growth and eco-physiological traits associated with morphology, phenology, and photosynthesis, which are considered important drivers of altitudinal adaptation. (3) Results: The quantitative trait loci (QTLs) for growth traits, phenology, needle morphology, and photosynthetic traits were subsequently detected. Similar to previous studies on conifers, most traits were controlled by multiple QTLs with small or moderate effects. Notably, we detected that one QTL for the crown area might be a type-A response regulator, a nuclear protein responsible for the cytokinin-induced shoot elongation. (4) Conclusion: The QTLs detected in this study include potentially important genomic regions linked to altitudinal adaptation in Sakhalin fir.

Old-growth forests play a major role in conserving biodiversity, protecting water resources, and sequestrating carbon, as well as serving as indispensable resources for indigenous societies. Novel silvicultural practices must be developed to emulate the natural dynamics and structural attributes of old-growth forests and preserve the ecosystem services provided by these boreal ecosystems. The success of these forest management strategies depends on developing an accurate understanding of natural regeneration dynamics. Our goal was therefore to identify the main patterns and drivers involved in the regeneration dynamics of old-growth forests with a focus on boreal stands dominated by black spruce (Picea mariana (L.) Mill.) and balsam fir (Abies balsamea (L.) Mill.) in eastern Canada. We sampled 71 stands in a 2 200 km2 study area located within Quebec's boreal region. For each stand, we noted tree regeneration (seedlings and saplings), structural attributes (diameter distribution, deadwood volume, etc.), and abiotic (slope and soil) factors. The presence of seed-trees located nearby and slopes having moderate to high angles most influenced balsam fir regeneration. In contrast, the indirect indices of recent secondary disturbances (e.g., insect outbreaks or windthrows) and topographic constraints (slope and drainage) most influenced black spruce regeneration. We propose that black spruce regeneration dynamics can be separated into distinct phases: (i) layering within the understory, (ii) seedling growth when gaps open in the canopy, (iii) gradual canopy closure, and (iv) production of new layers once the canopy is closed. These dynamics are not observed in paludified stands or stands where balsam fir is more competitive than black spruce. Overall, this research helps explain the complexity of old-growth forest dynamics, where many ecological factors interact at multiple temporal and spatial scales. This study also improves our understanding of ecological processes within primary old-growth forests and identifies the key factors to consider when ensuring the sustainable management of old-growth boreal stands., Competing Interests: The authors have declared that no competing interests exist.

Synchrony between host budburst and insect emergence greatly influences the time window for insect development and survival. A few alterations of bud phenology have been reported under defoliation without clear consensus regarding the direction of effects, i.e., advance or delay. Here, we compared budburst phenology between conifers in defoliation and control treatments, and measured carbon allocation as a potential mechanistic explanation of changes in phenology. In a 2-year greenhouse experiment, saplings of balsam fir, black spruce and white spruce of two different provenances (north and south) were subjected to either control (no larvae) or natural defoliation treatment (larvae added) by spruce budworm. Bud and instar phenology, primary and secondary growth, defoliation and non-structural carbohydrates were studied during the growing season. No differences were observed in bud phenology during the first year of defoliation. After 1 year of defoliation, bud phenology advanced by 6-7 days in black spruce and balsam fir and by 3.5 days in white spruce compared with the control. Because of this earlier bud break, apical and shoot growth exceeded 50% of its final length before mature instar defoliation occurred, which decreased the overall level of damage. A sugar-mediated response, via earlier starch breakdown, and higher sugar availability to buds explains the advanced budburst in defoliated saplings. The advanced phenological response to defoliation was consistent across the conifer species and provenances except for one species × provenance combination. Allocation of carbon to buds and shoots growth at the expense of wood growth in the stem and reserve accumulation represents a shift in the physiological resources priorities to ensure tree survival. This advancement in bud phenology could be considered as a physiological response to defoliation based on carbohydrate needs for primary growth, rather than a resistance trait to spruce budworm., (© The Author(s) 2018. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Treeline advance has occurred throughout the twentieth century in mountainous regions around the world; however, local variation and temporal lags in responses to climate warming indicate that the upper limits of some treelines are not necessarily in climatic equilibrium. These observations suggest that factors other than climate are constraining tree establishment beyond existing treelines. Using a seed addition experiment, we tested the effects of seed availability, predation and microsite limitation on the establishment of two subalpine tree species (Picea engelmannii and Abies lasiocarpa) across four treelines in the Canadian Rocky Mountains. The effect of vegetation removal on seedling growth was also determined, and microclimate conditions were monitored. Establishment limitations observed in the field were placed in context with the effects of soil properties observed in a parallel experiment. The seed addition experiment revealed reduced establishment with increasing elevation, suggesting that although establishment within the treeline ecotone is at least partially seed limited, other constraints are more important beyond the current treeline. The effects of herbivory and microsite availability significantly reduced seedling establishment but were less influential beyond the treeline. Microclimate monitoring revealed that establishment was negatively related to growing season temperatures and positively related to the duration of winter snow cover, counter to the conventional expectation that establishment is limited by low temperatures. Overall, it appears that seedling establishment beyond treeline is predominantly constrained by a combination of high soil surface temperatures during the growing season, reduced winter snowpack and unfavourable soil properties. Our study supports the assertion that seedling establishment in alpine treeline ecotones is simultaneously limited by various climatic and nonclimatic drivers. Together, these factors may limit future treeline advance in the Canadian Rocky Mountains and should be considered when assessing the potential for treeline advance in alpine systems elsewhere., (© 2018 John Wiley & Sons Ltd.)

Abies religiosa forests in central Mexico are the only overwinter refuge of the monarch butterfly and provide important ecosystem services. These forests have lost 55% of their original area and as a consequence, diversity and biotic interactions in these ecosystems are in risk. The aim of this study was to compare the soil fungal diversity and community structure in the Abies religiosa forests and surrounding Pinus montezumae, Pinus hartwegii, and coniferous mixed forest plant communities to provide data on ecology of mycorrhizal interactions for the assisted migration of A. religiosa. We sampled soil from five coniferous forests, extracted total soil DNA, and sequenced the ITS2 region by Illumina MiSeq. The soil fungi community was integrated by 1746 taxa with a species turnover ranging from 0.280 to 0.461 between sampling sites. In the whole community, the more abundant and frequent species were Russula sp. (aff. olivobrunnea), Mortierella sp.1, and Piloderma sp. (aff. olivacearum). The ectomycorrhizal fungi were the more frequent and abundant functional group. A total of 298 species (84 ectomycorrhizal) was shared in the five conifer forests; these widely distributed species were dominated by Russulaceae and Clavulinaceae. The fungal community composition was significantly influenced by altitude and the lowest species turnover happened between the two A. religiosa forests even though they have different soil types. As Pinus montezumae forests have a higher altitudinal distribution adjacent to A. religiosa and share the largest number of ectomycorrhizal fungi with it, we suggest these forests as a potential habitat for new A. religiosa populations.

Morphology, anatomy and physiology of sun and shade leaves of Abies alba were investigated and major differences were identified, such as sun leaves being larger, containing a hypodermis and palisade parenchyma as well as possessing more stomata, while shade leaves exhibit a distinct leaf dimorphism. The large size of sun leaves and their arrangement crowded on the upper side of a plagiotropic shoot leads to self-shading which is explainable as protection from high solar radiation and to reduce the transpiration via the lamina. Sun leaves furthermore contain a higher xanthophyll cycle pigment amount and Non-Photochemical Quenching (NPQ) capacity, a lower amount of chlorophyll b and a total lower chlorophyll amount per leaf, as well as an increased electron transport rate and an increased photosynthesis light saturation intensity. However, sun leaves switch on their NPQ capacity at rather low light intensities, as exemplified by several parameters newly measured for conifers. Our holistic approach extends previous findings about sun and shade leaves in conifers and demonstrates that both leaf types of A. alba show structural and physiological remarkable similarities to their respective counterparts in angiosperms, but also possess unique characteristics allowing them to cope efficiently with their environmental constraints., (© 2018 John Wiley & Sons Ltd.)

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

Improving our understanding of the potential of forest adaptation is an urgent task in the light of predicted climate change. Long-term alternatives for susceptible yet economically important tree species such as Norway spruce (Picea abies) are required, if the frequency and intensity of summer droughts will continue to increase. Although Silver fir (Abies alba) and Douglas fir (Pseudotsuga menziesii) have both been described as drought-tolerant species, our understanding of their growth responses to drought extremes is still limited. Here, we use a dendroecological approach to assess the resistance, resilience, and recovery of these important central Europe to conifer species the exceptional droughts in 1976 and 2003. A total of 270 trees per species were sampled in 18 managed mixed-species stands along an altitudinal gradient (400-1200 m a.s.l.) at the western slopes of the southern and central Black Forest in southwest Germany. While radial growth in all species responded similarly to the 1976 drought, Norway spruce was least resistant and resilient to the 2003 summer drought. Silver fir showed the overall highest resistance to drought, similarly to Douglas fir, which exhibited the widest growth rings. Silver fir trees from lower elevations were more drought prone than trees at higher elevations. Douglas fir and Norway spruce, however, revealed lower drought resilience at higher altitudes. Although the 1976 and 2003 drought extremes were quite different, Douglas fir maintained consistently the highest radial growth. Although our study did not examine population-level responses, it clearly indicates that Silver fir and Douglas fir are generally more resistant and resilient to previous drought extremes and are therefore suitable alternatives to Norway spruce; Silver fir more so at higher altitudes. Cultivating these species instead of Norway spruce will contribute to maintaining a high level of productivity across many Central European mountain forests under future climate change., (© 2017 John Wiley & Sons Ltd.)

Tree populations usually show adaptations to their local environments as a result of natural selection. As climates change, populations can become locally maladapted and decline in fitness. Evaluating the expected degree of genetic maladaptation due to climate change will allow forest managers to assess forest vulnerability, and develop strategies to preserve forest health and productivity. We studied potential genetic maladaptation to future climates in three major European tree species, Norway spruce (Picea abies), silver fir (Abies alba), and European beech (Fagus sylvatica). A common garden experiment was conducted to evaluate the quantitative genetic variation in growth and phenology of seedlings from 77 to 92 native populations of each species from across Switzerland. We used multivariate genecological models to associate population variation with past seed source climates, and to estimate relative risk of maladaptation to current and future climates based on key phenotypic traits and three regional climate projections within the A1B scenario. Current risks from climate change were similar to average risks from current seed transfer practices. For all three climate models, future risks increased in spruce and beech until the end of the century, but remained low in fir. Largest average risks associated with climate projections for the period 2061-2090 were found for spruce seedling height (0.64), and for beech bud break and leaf senescence (0.52 and 0.46). Future risks for spruce were high across Switzerland. However, areas of high risk were also found in drought-prone regions for beech and in the southern Alps for fir. Genetic maladaptation to future climates is likely to become a problem for spruce and beech by the end of this century, but probably not for fir. Consequently, forest management strategies should be adjusted in the study area for spruce and beech to maintain productive and healthy forests in the future., (© 2017 John Wiley & Sons Ltd.)

Warmer and drier climatic conditions are projected for the 21st century; however, the role played by extreme climatic events on forest vulnerability is still little understood. For example, more severe droughts and heat waves could threaten quaternary relict tree refugia such as Circum-Mediterranean fir forests (CMFF). Using tree-ring data and a process-based model, we characterized the major climate constraints of recent (1950-2010) CMFF growth to project their vulnerability to 21st-century climate. Simulations predict a 30% growth reduction in some fir species with the 2050s business-as-usual emission scenario, whereas growth would increase in moist refugia due to a longer and warmer growing season. Fir populations currently subjected to warm and dry conditions will be the most vulnerable in the late 21st century when climatic conditions will be analogous to the most severe dry/heat spells causing dieback in the late 20th century. Quantification of growth trends based on climate scenarios could allow defining vulnerability thresholds in tree populations. The presented predictions call for conservation strategies to safeguard relict tree populations and anticipate how many refugia could be threatened by 21st-century dry spells., Competing Interests: The authors declare no conflict of interest.

Significant increases in tree mortality due to drought-induced physiological stress have been documented worldwide. This trend is likely to continue with increased frequency and severity of extreme drought events in the future. Therefore, understanding the factors that influence variability in drought responses among trees will be critical to predicting ecosystem responses to climate change and developing effective management actions. In this study, we used hierarchical mixed-effects models to analyze drought responses of Pseudotsuga menziesii in 20 unmanaged forests stands across a broad range of environmental conditions in northeastern Washington, USA. We aimed to 1) identify the biotic and abiotic attributes most closely associated with the responses of individual trees to drought and 2) quantify the variability in drought responses at different spatial scales. We found that growth rates and competition for resources significantly affected resistance to a severe drought event in 2001: slow-growing trees and trees growing in subordinate canopy positions and/or with more neighbors suffered greater declines in radial growth during the drought event. In contrast, the ability of a tree to return to normal growth when climatic conditions improved (resilience) was unaffected by competition or relative growth rates. Drought responses were significantly influenced by tree age: older trees were more resistant but less resilient than younger trees. Finally, we found differences between resistance and resilience in spatial scale: a significant proportion (approximately 50%) of the variability in drought resistance across the study area was at broad spatial scales (i.e. among different forest types), most likely due to differences in the total amount of precipitation received at different elevations; in contrast, variation in resilience was overwhelmingly (82%) at the level of individual trees within stands and there was no difference in drought resilience among forest types. Our results suggest that for Pseudotsuga menziesii resistance and resilience to drought are driven by different factors and vary at different spatial scales.

Productivity of conifers in seasonally snow-covered forests is high before and during snowmelt when environmental conditions are optimal for photosynthesis. Climate change is altering the timing of spring in many locations, and changes in the date of transition from winter dormancy can have large impacts on annual productivity. Sap flow methods provide a promising approach to monitor tree activity during the cold season and the winter-spring and fall-winter transitions. Although sap flow techniques have been widely used, cold season results are generally not reported. Here we examine the feasibility of using the Granier thermal dissipation (TD) sap flux density method to monitor transpiration and dormancy of evergreen conifers during the cold season. We conducted a laboratory experiment which demonstrated that the TD method reliably detects xylem water transport (when it occurs) both at near freezing temperature and at low flow rate, and that the sensors can withstand repeated freeze-thaw events. However, the dependence between sensor output and water transport rate in these experiments differed from the established TD relation. In field experiments, sensors installed in two Abies forests lasted through two winters and a summer with low failure. The baseline (no-flow) sensor output varied considerably with temperature during the cold season, and a new baseline algorithm was developed to accommodate this variation. The Abies forests differed in elevation (2070 and 2620 m), and there was a clear difference in timing of initiation and cessation of transpiration between them. We conclude that the TD method can be reliably used to examine water transport during cold periods with associated low flow conditions., (© The Author 2017. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Forests are being clearcut over extensive areas of western North America, but plant community response to harvesting and slashburning under varying climatic conditions in central British Columbia, Canada is still largely unknown. Evaluation of resilience is hampered by the short history of logging, lack of long-term experiments and methodological limitations. To test the effect of clearcut logging, prescribed burning and reforestation on forest resilience, we recorded vascular plant cover repeatedly after treatment between 1981 and 2008 in 16 permanent research installations in three biogeoclimatic zones: Engelmann Spruce-Subalpine Fir, Interior Cedar-Hemlock and Sub-Boreal Spruce. We created a plant-trait dataset for the 181 recorded species to define plant functional types representing groups of plants that behave in similar ways and/or produce similar ecological outcomes. These plant functional types, along with taxonomic analysis of diagnostic and indicator species, were then used to evaluate plant community response to disturbance. Twenty years post-treatment, species diversity increased in all zones and plant abundance was greatest in the Interior Cedar-Hemlock. Cover of understory plant functional types associated with mature conifer forests increased in all zones, constituting a significant proportion (> 40%) of the vegetation community by year 20. Response patterns varied by zone and with time. Understory species diagnostic of mature forests were present in all zones by year 20, but we identified indicator species sensitive to slashburning or requiring more time for recovery, including white-flowered rhododendron (Rhododendron albiflorum) and devil's club (Oplopanax horridus). Overall, loss of compositional or functional diversity following harvest and site remediation was not detected, suggesting that montane and subalpine forests in British Columbia are resilient to this treatment. However, because these forests can be slow to recover from disturbance, the post-disturbance assessment window of this study may not have been long enough to detect diminishment of ecosystem resilience.

Changes in climate can alter the distribution and population dynamics of tree species by altering their recruitment patterns, especially at range edges. However, geographical patterns of genetic diversity could buffer the negative consequences of changing climate at rear range edges where populations might also harbour individuals with drought-adapted genotypes. Silver fir (Abies alba Mill.) reaches its south-western distribution limit in the Spanish Pyrenees, where recent climatic dieback events have disproportionately affected westernmost populations. We hypothesized that silver fir populations from the eastern Pyrenees are less vulnerable to the expected changing climate due to the inclusion of drought-resistant genotypes. We performed an experiment under strictly controlled conditions simulating projected warming and drought compared with current conditions and analysed physiology, growth and survival of silver fir seedlings collected from eastern and western Pyrenean populations. Genetic analyses separated eastern and western provenances in two different lineages. Climate treatments affected seedling morphology and survival of both lineages in an overall similar way: elevated drought diminished survival and induced a higher biomass allocation to roots. Increased temperature and drought provoked more negative stem water potentials and increased δ 13 C ratios in leaves. Warming reduced nitrogen concentration and increased soluble sugar content in leaves, whereas drought increased nitrogen concentration. Lineage affected these physiological parameters, with western seedlings being more sensitive to warming and drought increase in terms of δ 13 C, nitrogen and content of soluble sugars. Our results demonstrate that, in A. alba, differences in the physiological response of this species to drought are also associated with differences in biogeographical history., (© The Author 2016. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Background: Local adaptation and phenotypic plasticity are important components of plant responses to variations in environmental conditions. While local adaptation has been widely studied in trees, little is known about plasticity of gene expression in adult trees in response to ever changing environmental conditions in natural habitats. Here we investigate plasticity of gene expression in needle tissue between two Douglas-fir provenances represented by 25 adult trees using deep RNA sequencing (RNA-Seq)., Results: Using linear mixed models we investigated the effect of temperature, soil water availability and photoperiod on the abundance of 59189 detected transcripts. Expression of more than 80 % of all identified transcripts revealed a response to variations in environmental conditions in the field. GO term overrepresentation analysis revealed gene expression responses to temperature, soil water availability and photoperiod that are highly conserved among many plant taxa. However, expression differences between the two Douglas-fir provenances were rather small compared to the expression differences observed between individual trees. Although the effect of environment on global transcript expression was high, the observed genotype by environment (GxE) interaction of gene expression was surprisingly low, since only 21 of all detected transcripts showed a GxE interaction., Conclusions: The majority of the transcriptome responses in plant leaf tissue is driven by variations in environmental conditions. The small variation between individuals and populations suggests strong conservation of this response within Douglas-fir. Therefore we conclude that plastic transcriptome responses to variations in environmental conditions are only weakly affected by local adaptation in Douglas-fir.

The most widespread response to global warming among alpine treeline ecotones is not an upward shift, but an increase in tree density. However, the impact of increasing density on interactions among trees at treeline is not well understood. Here, we test if treeline densification induced by climatic warming leads to increasing intraspecific competition. We mapped and measured the size and age of Smith fir trees growing in two treelines located in the southeastern Tibetan Plateau. We used spatial point-pattern and codispersion analyses to describe the spatial association and covariation among seedlings, juveniles, and adults grouped in 30-yr age classes from the 1860s to the present. Effects of competition on tree height and regeneration were inferred from bivariate mark-correlations. Since the 1950s, a rapid densification occurred at both sites in response to climatic warming. Competition between adults and juveniles or seedlings at small scales intensified as density increased. Encroachment negatively affected height growth and further reduced recruitment around mature trees. We infer that tree recruitment at the studied treelines was more cold-limited prior to 1950 and shifted to a less temperature-constrained regime in response to climatic warming. Therefore, the ongoing densification and encroachment of alpine treelines could alter the way climate drives their transitions toward subalpine forests., (© 2016 by the Ecological Society of America.)

Hydraulic impairment due to xylem embolism and carbon starvation are the two proposed mechanisms explaining drought-induced forest dieback and tree death. Here, we evaluate the relative role played by these two mechanisms in the long-term by quantifying wood-anatomical traits (tracheid size and area of parenchyma rays) and estimating the intrinsic water-use efficiency (iWUE) from carbon isotopic discrimination. We selected silver fir and Scots pine stands in NE Spain with ongoing dieback processes and compared trees showing contrasting vigour (declining vs nondeclining trees). In both species earlywood tracheids in declining trees showed smaller lumen area with thicker cell wall, inducing a lower theoretical hydraulic conductivity. Parenchyma ray area was similar between the two vigour classes. Wet spring and summer conditions promoted the formation of larger lumen areas, particularly in the case of nondeclining trees. Declining silver firs presented a lower iWUE than conspecific nondeclining trees, but the reverse pattern was observed in Scots pine. The described patterns in wood anatomical traits and iWUE are coherent with a long-lasting deterioration of the hydraulic system in declining trees prior to their dieback. Retrospective quantifications of lumen area permit to forecast dieback in declining trees 2-5 decades before growth decline started. Wood anatomical traits provide a robust tool to reconstruct the long-term capacity of trees to withstand drought-induced dieback., (© 2016 John Wiley & Sons Ltd.)

The complex inner mechanisms that create typical conifer tree-ring structure (i.e. the transition from large, thin-walled earlywood cells to narrow, thick-walled latewood cells) were recently unraveled. However, what physiological or environmental factors drive xylogenesis key processes remain unclear. Here, we aim to quantify the influence of seasonal variations in climatic factors on the spectacular changes in the kinetics of wood cell differentiation and in the resulting tree-ring structure. Wood formation was monitored in three sites over 3 years for three coniferous species (Norway spruce [Picea abies], Scots pine [Pinus sylvestris], and silver fir [Abies alba]). Cell differentiation rates and durations were calculated and related to tracheid final dimensions and corresponding climatic conditions. On the one hand, we found that the kinetics of cell enlargement and the final size of the tracheids were not explained by the seasonal changes in climatic factors. On the other hand, decreasing temperatures strongly constrained cell wall deposition rates during latewood formation. However, the influence of temperature was permanently written into tree-ring structure only for the very last latewood cells, when the collapse of the rate of wall deposition was no longer counterbalanced by the increase of its duration. Our results show that the formation of the typical conifer tree-ring structure, in normal climatic conditions, is only marginally driven by climate, suggesting strong developmental control of xylogenesis. The late breakage of the compensatory mechanism at work in the wall deposition process appears as a clue to understand the capacity of the maximum latewood density to record past temperature conditions., (© 2016 American Society of Plant Biologists. All Rights Reserved.)

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

Recent large and severe outbreaks of native bark beetles have raised concern among the general public and land managers about potential for amplified fire activity in western North America. To date, the majority of studies examining bark beetle outbreaks and subsequent fire severity in the U.S. Rocky Mountains have focused on outbreaks of mountain pine beetle (MPB; Dendroctonus ponderosae) in lodgepole pine (Pinus contorta) forests, but few studies, particularly field studies, have addressed the effects of the severity of spruce beetle (Dendroctonus rufipennis Kirby) infestation on subsequent fire severity in subalpine Engelmann spruce (Picea engelmannii) and subalpine fir (Abies lasiocarpa) forests. In Colorado, the annual area infested by spruce beetle outbreaks is rapidly rising, while MPB outbreaks are subsiding; therefore understanding this relationship is of growing importance. We collected extensive field data in subalpine forests in the eastern San Juan Mountains, southwestern Colorado, USA, to investigate whether a gray-stage (< 5 yr from outbreak to time of fire) spruce beetle infestation affected fire severity. Contrary to the expectation that bark beetle infestation alters subsequent fire severity, correlation and multivariate generalized linear regression analysis revealed no influence of pre-fire spruce beetle severity on nearly all field or remotely sensed measurements of fire severity. Findings were consistent across moderate and extreme burning conditions. In comparison to severity of the pre-fire beetle outbreak, we found that topography, pre-outbreak basal area, and weather conditions exerted a stronger effect on fire severity. Our finding that beetle infestation did not alter fire severity is consistent with previous retrospective studies examining fire activity following other bark beetle outbreaks and reiterates the overriding influence of climate that creates conditions conducive to large, high-severity fires in the subalpine zone of Colorado. Both bark beetle outbreaks and wildfires have increased autonomously due to recent climate variability, but this study does not support the expectation that post-beetle outbreak forests will alter fire severity, a result that has important implications for management and policy decisions.

Leaves growing in the forest understory usually present a decreased mesophyll conductance (gm) and photosynthetic capacity. The role of leaf anatomy in determining the variability in gm among species is known, but there is a lack of information on how the acclimation of gm to shade conditions is driven by changes in leaf anatomy. Within this context, we demonstrated that Abies pinsapo Boiss. experienced profound modifications in needle anatomy to drastic changes in light availability that ultimately led to differential photosynthetic performance between trees grown in the open field and in the forest understory. In contrast to A. pinsapo, its congeneric Abies alba Mill. did not show differences either in needle anatomy or in photosynthetic parameters between trees grown in the open field and in the forest understory. The increased gm values found in trees of A. pinsapo grown in the open field can be explained by occurrence of stomata at both needle sides (amphistomatous needles), increased chloroplast surface area exposed to intercellular airspace, decreased cell wall thickness and, especially, decreased chloroplast thickness. To the best of our knowledge, the role of such drastic changes in ultrastructural needle anatomy in explaining the response of gm to the light environment has not been demonstrated in field conditions., (© The Author 2015. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

The total phenol content in decomposing litter not only acts as a crucial litter quality indicator, but is also closely related to litter humification due to its tight absorption to clay particles. However, limited attention has been focused on the total phenol dynamics in foliar litter in relation to forest gaps. Here, the foliar litter of six representative tree species was incubated on the forest floor from the gap center to the closed canopy of an alpine Minjiang fir (Abies faxoniana) forest in the upper reaches of the Yangtze River and eastern Tibetan Plateau. The dynamics of total phenol concentration in the incubated litter was measured from November 2012 to October 2014. Over two-year incubation, 78.22% to 94.06% of total phenols were lost from the foliar litter, but 52.08% to 86.41% of this occurred in the first year. Forest gaps accelerated the loss of total phenols in the foliar litter in the winter, although they inhibited the loss of total phenols during the growing season in the first year. In comparison with the effects of forest gaps, the variations of litter quality among different species were much stronger on the dynamics of total phenols in the second year. Overall, the loss of total phenols in the foliar litter was slightly higher in both the canopy gap and the expanded gap than in the gap center and under the closed canopy. The results suggest that the predicted decline in snow cover resulting from winter warming or vanishing gaps caused by forest regeneration will retard the loss of total phenol content in the foliar litter of alpine forest ecosystems, especially in the first decomposition year.

Understanding local adaptation in forest trees is currently a key research and societal priority. Geographically and ecologically marginal populations provide ideal case studies, because environmental stress along with reduced gene flow can facilitate the establishment of locally adapted populations. We sampled European silver fir (Abies alba Mill.) trees in the French Mediterranean Alps, along the margin of its distribution range, from pairs of high- and low-elevation plots on four different mountains situated along a 170-km east-west transect. The analysis of 267 SNP loci from 175 candidate genes suggested a neutral pattern of east-west isolation by distance among mountain sites. F(ST) outlier tests revealed 16 SNPs that showed patterns of divergent selection. Plot climate was characterized using both in situ measurements and gridded data that revealed marked differences between and within mountains with different trends depending on the season. Association between allelic frequencies and bioclimatic variables revealed eight genes that contained candidate SNPs, of which two were also detected using F(ST) outlier methods. All SNPs were associated with winter drought, and one of them showed strong evidence of selection with respect to elevation. Q(ST)-F(ST) tests for fitness-related traits measured in a common garden suggested adaptive divergence for the date of bud flush and for growth rate. Overall, our results suggest a complex adaptive picture for A. alba in the southern French Alps where, during the east-to-west Holocene recolonization, locally advantageous genetic variants established at both the landscape and local scales., (© 2015 John Wiley & Sons Ltd.)

As sessile organisms, plants have evolved different methods to defend against attacks and have adapted their defense measures to discriminate between mechanical damage and herbivory by insects. One of the ways that plant defenses are triggered is via elicitors from insect oral secretions (OS). In this study, we investigated the ability of second-instar (L2) spruce budworm [SBW; Choristoneura fumiferana (Clemens)] to alter the volatile organic compounds (VOCs) of four conifer species [Abies balsamea (L.) Mill., Picea mariana (Miller) B.S.P., Picea glauca (Moench) Voss, Picea rubens (Sargent)] and found that the emission profiles from all host trees were drastically changed after herbivory. We then investigated whether some of the main elicitors (fatty acid conjugates [FACs], β-glucosidase, and glucose oxidase) studied were present in SBW OS. FACs (glutamine and glutamic acid) based on linolenic, linoleic, oleic, and stearic acids were all observed in varying relative quantities. Hydroxylated FACs, such as volicitin, were not observed. Enzyme activity for β-glucosidase was also measured and found present in SBW OS, whereas glucose oxidase activity was not found in the SBW labial glands. These results demonstrate that SBW L2 larvae have the ability to induce VOC emissions upon herbivory and that SBW OS contain potential elicitors to induce these defensive responses. These data will be useful to further evaluate whether these elicitors can separately induce the production of specific VOCs and to investigate whether and how these emissions benefit the plant., (© Crown copyright 2015.)

Rates and spatial patterns of tree mortality are predicted to change during forest structural development. In young forests, mortality should be primarily density dependent due to competition for light, leading to an increasingly spatially uniform pattern of surviving trees. In contrast, mortality in old-growth forests should be primarily caused by contagious and spatially autocorrelated agents (e.g., insects, wind), causing spatial aggregation of surviving trees to increase through time. We tested these predictions by contrasting a three-decade record of tree mortality from replicated mapped permanent plots located in young (< 60-year-old) and old-growth (> 300-year-old) Abies amabilis forests. Trees in young forests died at a rate of 4.42% per year, whereas trees in old-growth forests died at 0.60% per year. Tree mortality in young forests was significantly aggregated, strongly density dependent, and caused live tree patterns to become more uniform through time. Mortality in old-growth forests was spatially aggregated, but was density independent and did not change the spatial pattern of surviving trees. These results extend current theory by demonstrating that density-dependent competitive mortality leading to increasingly uniform tree spacing in young forests ultimately transitions late in succession to a more diverse tree mortality regime that maintains spatial heterogeneity through time.

Greater demand for wood material has converged with greater demand for biodiversity conservation to make balancing forest ecosystem services a key societal issue. Forest managers, owners, or policymakers need new approaches and methods to evaluate their ability to adapt to this dual objective. We analyze the ability of forest owners to define sustainable forest management options based on viability theory and a new flexibility index. This new indicator gauges the adaptive capacity of forest owners based on the number of sustainable actions available to them at a given time. Here we study a public forest owner who regulates harvest intensity and frequency in order to meet demand for timber wood at forest scale and to meet a biodiversity recommendation via a minimum permanently maintained volume of deadwood per hectare at stand scale. Dynamical systems theory was used to model uneven-aged forest dynamics-including deadwood dynamics-and the dynamics of timber wood demand and tree removals. Uneven-aged silver fir forest management in the "Quatre Montagnes region" (Vercors, France) is used as an illustrative example. The results explain situations where a joint increase in wood production and deadwood retention does not reduce the flexibility index more than increasing either one dimension alone, thus opening up ecological intensification options. To conclude, we discuss the value of the new flexibility index for addressing environmental management and ecological intensification issues.

Many studies have demonstrated linkages between the occurrence of fog and ecophysiological functioning in cloud forests, but few have investigated hydraulic functioning as a determining factor that explains sharp changes in vegetation. The objective of this study was to compare the plant water status during cloud-immersed and non-immersed conditions and hydraulic vulnerability in branches and roots of species across a temperate, mountain fog ecotone. Because cloud forests are often dark, cool and very moist, we expected cloud forest species to have less drought-tolerant characteristics (i.e., lower Pe and P50-the pressures required to induce a 12 and 50% loss in hydraulic conductivity, respectively) relative to non-cloud forest species in adjacent (lower elevation) forests. Additionally, due to the ability of cloud forest species to absorb cloud-fog water, we predicted greater improvements in hydraulic functioning during fog in cloud forest species relative to non-cloud forest species. Across the cloud forest ecotone, most species measured were very resistant to losses in conductivity with branch P50 values from -4.5 to -6.0 MPa, hydraulic safety margins (Ψmin - P50) >1.5 MPa and low calculated hydraulic conductivity losses. Roots had greater vulnerabilities, with P50 values ranging from -1.4 to -2.5 MPa, leading to greater predicted losses in conductivity (∼20%). Calculated values suggested strong losses of midday leaf hydraulic conductance in three of the four species, supporting the hydraulic segmentation hypothesis. In both cloud forest and hardwood species, Ψs were greater on foggy days than sunny days, demonstrating the importance of fog periods to plant water balance across fog regimes. Thus, frequent fog did not result in systemic changes in hydraulic functioning or vulnerability to embolism across our temperate cloud forest ecotone. Finally, roots functioned with lower hydraulic conductivity than branches, suggesting that they may serve as more sensitive indicators of hydraulic functioning in these mesic, foggy ecosystems., (© The Author 2015. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Photosynthetic light-use efficiency (LUE) has gained wide interest as an input to modeling forest gross primary productivity (GPP). The photochemical reflectance index (PRI) has been identified as a principle means to inform LUE-based models, using airborne and satellite-based observations of canopy reflectance. More recently, low-cost electronics have become available with the potential to provide for dense in situ time-series measurements of PRI. A recent design makes use of interference filters to record light transmission within narrow wavebands. Uncertainty remains as to the dynamic range of these sensors and performance under low light conditions, the placement of the reference band, and methodology for reflectance calibration. This paper presents a low-cost sensor design and is tested in a laboratory set-up, as well in the field. The results demonstrate an excellent performance against a calibration standard (R2 = 0.9999) and at low light conditions. Radiance measurements over vegetation demonstrate a reversible reduction in green reflectance that was, however, seen in both the reference and signal wavebands. Time-series field measurements of PRI in a Douglas-fir canopy showed a weak correlation with eddy-covariance-derived LUE and a significant decline in PRI over the season. Effects of light quality, bidirectional scattering effects, and possible sensor artifacts on PRI are discussed.

The phylogeographic structure and postglacial history of balsam fir (Abies balsamea), a transcontinental North American boreal conifer, was inferred using mitochondrial DNA (mtDNA) and chloroplast DNA (cpDNA) markers. Genetic structure among 107 populations (mtDNA data) and 75 populations (cpDNA data) was analyzed using Bayesian and genetic distance approaches. Population differentiation was high for mtDNA (dispersed by seeds only), but also for cpDNA (dispersed by seeds and pollen), indicating that pollen gene flow is more restricted in balsam fir than in other boreal conifers. Low cpDNA gene flow in balsam fir may relate to low pollen production due to the inherent biology of the species and populations being decimated by recurrent spruce budworm epidemics, and/or to low dispersal of pollen grains due to their peculiar structural properties. Accordingly, a phylogeographic structure was detected using both mtDNA and cpDNA markers and population structure analyses supported the existence of at least five genetically distinct glacial lineages in central and eastern North America. Four of these would originate from glacial refugia located south of the Laurentide ice sheet, while the last one would have persisted in the northern Labrador region. As expected due to reduced pollen-mediated gene flow, congruence between the geographic distribution of mtDNA and cpDNA lineages was higher than in other North American conifers. However, concordance was not complete, reflecting that restricted but nonetheless detectable cpDNA gene flow among glacial lineages occurred during the Holocene. As a result, new cpDNA and mtDNA genome combinations indicative of cytoplasmic genome capture were observed.

To avoid winter frost damage, evergreen coniferous species develop cold hardiness with suitable phenology for the local climate regime. Along the elevational gradient, a genetic cline in autumn phenology is often recognised among coniferous populations, but further quantification of evolutionary adaptation related to the local environment and its responsible signals generating the phenological variation are poorly understood. We evaluated the timing of cold hardening among populations of Abies sachalinensis, based on time series freezing tests using trees derived from four seed source populations × three planting sites. Furthermore, we constructed a model to estimate the development of hardening from field temperatures and the intraspecific variations occurring during this process. An elevational cline was detected such that high-elevation populations developed cold hardiness earlier than low-elevation populations, representing significant genetic control. Because development occurred earlier at high-elevation planting sites, the genetic trend across elevation overlapped with the environmental trend. Based on the trade-off between later hardening to lengthen the active growth period and earlier hardening to avoid frost damage, this genetic cline would be adaptive to the local climate. Our modelling approach estimated intraspecific variation in two model components: the threshold temperature, which was the criterion for determining whether the trees accumulated the thermal value, and the chilling requirement for trees to achieve adequate cold hardiness. A higher threshold temperature and a lower chilling requirement could be responsible for the earlier phenology of the high-elevation population. These thermal responses may be one of the important factors driving the elevation-dependent adaptation of A. sachalinensis., (© 2014 German Botanical Society and The Royal Botanical Society of the Netherlands.)

Low temperatures are associated high-altitude treelines, but the functional mechanism of treeline formation remains controversial. The relative contributions of carbon limitation (source activity) and growth limitation (sink activity) require more tests across taxa and regions. We examined temporal variations of mobile carbon supply in different tissues of Abies fargesii across treeline ecotones on north- and south-facing slopes of the Qinling Mountains, China. Non-structural carbohydrate (NSC) concentrations in tissues along the altitudinal gradient on both slopes changed significantly in the early and late growing season, but not in the mid-growing season, indicating the season-dependent carbon supply status. Late in the growing season on both slopes, trees at the upper limits had the highest NSC concentrations and total soluble sugars and lowest starch concentrations compared to trees at the lower elevations. NSC concentrations tended to increase in needles and branches throughout the growing season with increasing elevation on both slopes, but declined in roots and stems. NSC concentrations across sampling dates also indicated increases in needles and branches, and decreases in roots and stem with increasing elevation. Overall altitudinal trends of NSC in A. fargesii revealed no depletion of mobile carbon reserves at upper elevation limits, suggesting limitation of sink activity dominates tree life across treeline ecotones in both north- and south-facing slopes. Carbon reserves in storage tissues (especially roots) in the late growing season might also play an important role in winter survival and early growth in spring at upper elevations on both slopes, which define the uppermost limit of A. fargesii., (© 2014 German Botanical Society and The Royal Botanical Society of the Netherlands.)

Plant species living in the understory increase carbon (C) allocation toward leaf production for maximizing light capture at the expense of roots and stems, with negative consequences for the whole-plant hydraulic conductance. Moreover, under some conditions, the high atmospheric evaporative demand occurring in Mediterranean areas may be not well buffered by the canopy, which might be the case for relict conifer Abies pinsapo Boiss. growing in the forest understory. We hypothesized that acclimation to combined understory shade and high atmospheric dryness can be achieved through the adjustment of water losses to cope with the restriction in water transport. The results reveal high structural plasticity in A. pinsapo that allows light harvesting of this species to maximize light capture in the forest understory, and maintain a positive C balance under low light conditions. However, growth in the understory resulted in reduced leaf-specific conductivity, up to approximately four to five times, implying decreased plant capacity to supply water to the leaves. In order to cope with the high atmospheric evaporative demand in the understory, there is an adjustment of the stomatal conductance to the hydraulic conductivity by means of a reduction in the stomatal density in understory individuals, which is due to the almost complete lack of stomata in the adaxial side of the needles. To the extent of our knowledge, such a drastic phenotypic response found in a conifer when growing under shaded conditions had not been previously reported., (© The Author 2014. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

We developed models to describe the responses of four commonly examined leaf traits (mass per area, weight, area and nitrogen (N) concentration) to gradients of light, soil nutrients and tree height in three conifer species of contrasting shade tolerance. Our observational dataset from the sub-boreal spruce forests of British Columbia included subalpine fir (Abies lasioscarpa [Hook.] Nutt; high shade tolerance), interior spruce (Picea glauca × Picea engelmannii [Moench] Voss; intermediate shade tolerance) and lodgepole pine (Pinus contorta Dougl. ex Loud. var. latifolia; low shade tolerance) saplings from 0.18 to 4.87 m tall, in 8-98% of total incident light, from field sites with <17.6 kg ha(-1) to >46.8 kg ha(-1) total dissolved N. Leaf weights and areas showed strong positive responses to light and height, but little or no response to soil nutrients. Parameter estimates indicated that the shape of leaf weight and area responses to light corresponded with shade tolerance ranking for the three species; pine had the most linear response whereas spruce and fir had asymptotic responses. Leaf N concentration responded positively to soil nutrients, negatively to light and idiosyncratically to height. The negative effect of light was only apparent on sites of high soil nutrient availability, and parameter estimates for the shape of the negative response also corresponded to shade tolerance ranking (apine = -0.79, aspruce = -0.15, afir = -0.07). Of the traits we measured, leaf mass per area showed the least response to light, soil nutrient and height gradients. Although it is a common practice in comparisons across many species, characterizing these conifers by mean values of their leaf traits would miss important intraspecific variation across environmental and size gradients. In these forests, parameter estimates representing the intraspecific variability of leaf trait responses can be used to understand relative shade tolerances., (© The Author 2014. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Leaf life span is an important plant trait associated with interspecific variation in leaf, organismal, and ecosystem processes. We hypothesized that intraspecific variation in gymnosperm needle traits with latitude reflects both selection and acclimation for traits adaptive to the associated temperature and moisture gradient. This hypothesis was supported, because across 127 sites along a 2,160-km gradient in North America individuals of Picea glauca, Picea mariana, Pinus banksiana, and Abies balsamea had longer needle life span and lower tissue nitrogen concentration with decreasing mean annual temperature. Similar patterns were noted for Pinus sylvestris across a north-south gradient in Europe. These differences highlight needle longevity as an adaptive feature important to ecological success of boreal conifers across broad climatic ranges. Additionally, differences in leaf life span directly affect annual foliage turnover rate, which along with needle physiology partially regulates carbon cycling through effects on gross primary production and net canopy carbon export. However, most, if not all, global land surface models parameterize needle longevity of boreal evergreen forests as if it were a constant. We incorporated temperature-dependent needle longevity and %nitrogen, and biomass allocation, into a land surface model, Community Atmosphere Biosphere Land Exchange, to assess their impacts on carbon cycling processes. Incorporating realistic parameterization of these variables improved predictions of canopy leaf area index and gross primary production compared with observations from flux sites. Finally, increasingly low foliage turnover and biomass fraction toward the cold far north indicate that a surprisingly small fraction of new biomass is allocated to foliage under such conditions.

Widespread fire suppression and thinning have altered the structure and composition of many forests in the western United States, making them more susceptible to the synergy of large-scale drought and fire events. We examine how these changes affect carbon storage and stability compared to historic fire-adapted conditions. We modeled carbon dynamics under possible drought and fire conditions over a 300-year simulation period in two mixed-conifer conditions common in the western United States: (1) pine-dominated with an active fire regime and (2) fir-dominated, fire suppressed forests. Fir-dominated stands, with higher live- and dead-wood density, had much lower carbon stability as drought and fire frequency increased compared to pine-dominated forest. Carbon instability resulted from species (i.e., fir's greater susceptibility to drought and fire) and stand (i.e., high density of smaller trees) conditions that develop in the absence of active management. Our modeling suggests restoring historic species composition and active fire regimes can significantly increase carbon stability in fire-suppressed, mixed-conifer forests. Long-term management of forest carbon should consider the relative resilience of stand structure and composition to possible increases in disturbance frequency and intensity under changing climate.

Silvicultural treatments such as thinning have been suggested as management tools against the spruce budworm, Choristoneura fumiferana (Clemens) (Lepidoptera: Tortricidae). Among other things, parasitoids are also proposed to be influenced by silvicultural procedures, but the effect of thinning on spruce budworm's natural enemies has not been tested yet. In this study, the influence of partial cutting on parasitism of endemic spruce budworm populations has been investigated in mature balsam fir-white birch forests. Two intensities of partial cutting (25 and 40% stand basal area reduced) were conducted in 2009 and parasitism of introduced spruce budworm larvae and pupae was determined during the 3 yr after these treatments. Pupal parasitism was too low for comparison between treatments. However, 2 yr after treatments, parasitism of the fourth- and fifth-instar larvae was significantly reduced in plots with both intensities of partial cutting, which was attributed to the parasitoid Tranosema rostrale (Brischke). Three years after treatments, no significant influence of partial cutting on parasitism of spruce budworm larvae was found. This study suggests that the influence of partial cutting on parasitism of endemic spruce budworm populations is not consistent, but that under certain circumstances parasitism is reduced by partial cutting.

In cloud forests, foliar uptake (FU) of water has been reported for numerous species, possibly acting to relieve daily water and carbon stress. While the prevalence of FU seems common, how daily variation in fog timing may affect this process has not been studied. We examined the quantity of FU, water potentials, gas exchange and abiotic variation at the beginning and end of a 9-day exposure to fog in a glasshouse setting. Saplings of Abies fraseri (Pursh) Poir. and Picea rubens Sarg. were exposed to morning (MF), afternoon (AF) or evening fog (EF) regimes to assess the ability to utilize fog water at different times of day and after sustained exposure to simulated fog. The greatest amount of FU occurred during MF (up to 50%), followed by AF (up to 23%) and then EF, which surprisingly had no FU. There was also a positive relationship between leaf conductance and FU, suggesting a role of stomata in FU. Moreover, MF and AF lead to the greatest improvements in daily water balance and carbon gain, respectively. Foliar uptake was important for improving plant ecophysiology but was influenced by diurnal variation in fog. With climate change scenarios predicting changes to cloud patterns and frequency that will likely alter diurnal patterns, cloud forests that rely on this water subsidy could be affected., (© The Author 2014. Published by Oxford University Press. All rights reserved.)

We present a novel measurement setup for monitoring changes in leaf water status using nondestructive terahertz time-domain spectroscopy (THz-TDS). Previous studies on a variety of plants showed the principal applicability of THz-TDS. In such setups, decreasing leaf water content directly correlates with increasing THz transmission. Our new system allows for continuous, nondestructive monitoring of the water status of multiple individual plants each at the same constant leaf position. It overcomes previous drawbacks, which were mainly due to the necessity of relocating the plants. Using needles of silver fir (Abies alba) seedlings as test subjects, we show that the transmission varies along the main axis of a single needle due to a variation in thickness. Therefore, the relocation of plants during the measuring period, which was necessary in the previous THz-TDS setups, should be avoided. Furthermore, we show a highly significant correlation between gravimetric water content and respective THz transmission. By monitoring the relative change in transmission, we were able to narrow down the permanent wilting point of the seedlings. Thus, we established groups of plants with well-defined levels of water stress that could not be detected visually. This opens up the possibility for a broad range of genetic and physiological experiments.

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

We examined the ecophysiology and growth of 0.3-1.3 m tall advance red spruce (Picea rubens Sarg.) and balsam fir (Abies balsamea [L.] Mill.) regeneration during a 5-year period following the application of different harvest types producing three sizes of canopy openings: (i) small gaps (<100 m(2) in area; SMA) created by partial uniform single-tree harvest; (ii) irregular gaps of intermediate size (100-300 m(2); INT) created by group-selection harvest (removal of groups of trees, mainly balsam fir, with uniform partial removal between groups); and (iii) large circular gaps (700 m(2); LAR) created by patch-selection harvest (removal of trees in 30-m diameter circular areas with uniform partial removal between gaps). An unharvested control (CON) was monitored for comparison. At the ecophysiological level, we mainly found differences in light-saturated photosynthesis of red spruce and specific leaf area of balsam fir among treatments. Consequently, we observed good height growth of both species in CON and INT, but fir surpassed spruce in SMA and LAR. Results suggest that intermediate 100-300 m(2) irregular openings create microenvironmental conditions that may promote short-term ecophysiology and growth of red spruce, allowing the species to compete with balsam fir advance regeneration. Finally, results observed for spruce in large 700-m(2) openings confirm its inability to grow as rapidly as fir in comparable open conditions.

Cloud immersion can provide a potentially important moisture subsidy to plants in areas of frequent fog including the threatened spruce-fir communities of the southern Appalachian Mountains (USA). These mountaintop communities grow only above ~1,500 m elevation, harbor the endemic Abies fraseri, and have been proposed to exist because of frequent cloud immersion. While several studies have demonstrated the importance of cloud immersion to plant water balance, no study has evaluated the proportion of plant water derived from cloud moisture in this ecosystem. Using the isotopic mixing model, IsoSource, we analyzed the isotopic composition of hydrogen and oxygen for water extracted from ground water, deep soil, shallow soil, fog, and plant xylem at the upper and lower elevational limits both in May (beginning of the growing season) and October (end of the growing season). Cloud-immersion water contributed up to 31% of plant water at the upper elevation sites in May. High-elevation plants of both species also experienced greater cloud immersion and had greater cloud water absorption (14-31%) compared to low-elevation plants (4-17%). Greater cloud water uptake occurred in May compared to October, despite similar rainfall and cloud-immersion frequencies. These results demonstrate the important water subsidy that cloud-immersion water can provide. With a warming climate leading potentially to increases in the ceiling of the cloud base and, thus, less frequent cloud immersion, persistence of these relic mountaintop forests may depend on the magnitude of these changes and the compensating capabilities of other water sources.

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