The ecological function of boreal forests is challenged by drastically changing climate conditions. Although an increasing number of studies are investigating how climate change is influencing growth ...and distribution of boreal tree species, there is a lack of studies examining the potential of these species to genetically adapt or phenotypically adjust. Here, we sampled clonally and non-clonally growing white spruce trees (
Moench Voss) to investigate spatial and genetic effects on tree ring width and on six xylem anatomical traits representing growth, water transport, mechanical support, and wood density. We compared different methods for estimating broad sense heritability (H
) of each trait and we evaluated the effects of spatial grouping and genetic grouping on the xylem anatomical traits with linear models. We found that the three different methods used to estimate H
were quite robust, showing overall consistent patterns, while our analyses were unsuccessful at fully separating genetic from spatial effects. By evaluating the effect size, we found a significant effect of genetic grouping in latewood density and earlywood hydraulic diameter. However, evaluating model performances showed that spatial grouping was a better predictor than genetic grouping for variance in earlywood density, earlywood hydraulic diameter and growth. For cell wall thickness neither spatial nor genetic grouping was significant. Our findings imply that (1) the variance in the investigated xylem anatomical traits and growth is mainly influenced by spatial clustering (most probably caused by microhabitat conditions), which (2) makes it rather difficult to estimate the heritability of these traits in naturally grown trees
. Yet, (3) latewood density and earlywood hydraulic diameter qualified for further analysis on the genetic background of xylem traits and (4) cell wall thickness seems a useful trait to investigate large-scale climatic effects, decoupled from microclimatic, edaphic and genetic influences.
Treeline ecosystems are of great scientific interest to study the effects of limiting environmental conditions on tree growth. However, tree growth is multidimensional, with complex interactions ...between height and radial growth. In this study, we aimed to disentangle effects of height and climate on xylem anatomy of white spruce
Picea glauca
(Moench) Voss at three treeline sites in Alaska; i.e., one warm and drought-limited, and two cold, temperature-limited. To analyze general growth differences between trees from different sites, we used data on annual ring width, diameter at breast height (DBH), and tree height. A representative subset of the samples was used to investigate xylem anatomical traits. We then used linear mixed-effects models to estimate the effects of height and climatic variables on our study traits. Our study showed that xylem anatomical traits in white spruce can be directly and indirectly controlled by environmental conditions: hydraulic-related traits seem to be mainly influenced by tree height, especially in the earlywood. Thus, they are indirectly driven by environmental conditions, through the environment’s effects on tree height. Traits related to mechanical support show a direct response to environmental conditions, mainly temperature, especially in the latewood. These results highlight the importance of assessing tree growth in a multidimensional way by considering both direct and indirect effects of environmental forcing to better understand the complexity of tree growth responses to the environment.
Human-driven peatland drainage has occurred in Europe for centuries, causing habitat degradation and leading to the emission of greenhouse gases. As such, in the last decades, there has been an ...increase in policies aiming at restoring these habitats through rewetting. Alder (
L.) is a widespread species in temperate forest peatlands with a seemingly high waterlogging tolerance. Yet, little is known about its specific response in growth and wood traits relevant for tree functioning when dealing with changing water table levels. In this study, we investigated the effects of rewetting and extreme flooding on alder growth and wood traits in a peatland forest in northern Germany. We took increment cores from several trees at a drained and a rewetted stand and analyzed changes in ring width, wood density, and xylem anatomical traits related to the hydraulic functioning, growth, and mechanical support for the period 1994-2018. This period included both the rewetting action and an extreme flooding event. We additionally used climate-growth and climate-density correlations to identify the stand-specific responses to climatic conditions. Our results showed that alder growth declined after an extreme flooding in the rewetted stand, whereas the opposite occurred in the drained stand. These changes were accompanied by changes in wood traits related to growth (i.e., number of vessels), but not in wood density and hydraulic-related traits. We found poor climate-growth and climate-density correlations, indicating that water table fluctuations have a stronger effect than climate on alder growth. Our results show detrimental effects on the growth of sudden water table changes leading to permanent waterlogging, but little implications for its wood density and hydraulic architecture. Rewetting actions should thus account for the loss of carbon allocation into wood and ensure suitable conditions for alder growth in temperate peatland forests.
The carbon pool of peatlands has been considered as potentially unstable in a changing climate. This study is the first presenting carbon dioxide (CO₂) net ecosystem exchange, CO₂ efflux due to ...ecosystem respiration and CO₂ uptake by gross primary production over a complete growing season for different microforms of a boreal peatland in Russia (61°56′N, 50°13′E). CO₂ fluxes were measured using the closed chamber technique from the 25th April in the period of snow melt until the end of the vegetation period and the first frost on the 20th October 2008 at seven different microform types: minerogenous and ombrogenous hollows, lawns and hummocks, respectively, and Carex lawns situated in a transition zone between minerogenous and ombrogenous mire parts. The total number of chamber flux measurements was 5,517. Ombrogenous hummocks and lawns were sources of CO₂ over the investigation period whereas hollows and minerogenous lawns were CO₂ sinks. Some plots of Carex lawns and minerogenous hummocks were sinks while other plots of these microform types were sources. The CO₂ fluxes were characterised by large variability not only between the microform types but also within the respective microform types. Of all microform types, the Carex, ombrogenous, and minerogenous lawns showed the highest variability in CO₂ fluxes, which is probably related to a stronger within-microform heterogeneity in vegetation composition and coverage as well as in the water table level. Air temperature was one of the dominant controls on the CO₂ flux dynamics. Water table and green area index were found to have strong influence on CO₂ fluxes both within different patches of the same microform type as well as between different microforms.
Forests are critical in the terrestrial carbon cycle, and the knowledge of their response to ongoing climate change will be crucial for determining future carbon fluxes and climate trajectories. In ...areas with contrasting seasons, trees form discrete annual rings that can be assigned to calendar years, allowing to extract valuable information about how trees respond to the environment. The anatomical structure of wood provides highly-resolved information about the reaction and adaptation of trees to climate. Quantitative wood anatomy helps to retrieve this information by measuring wood at the cellular level using high-resolution images of wood micro-sections. However, whereas large advances have been made in identifying cellular structures, obtaining meaningful cellular information is still hampered by the correct annual tree ring delimitation on the images. This is a time-consuming task that requires experienced operators to manually delimit ring boundaries. Classic methods of automatic segmentation based on pixel values are being replaced by new approaches using neural networks which are capable of distinguishing structures, even when demarcations require a high level of expertise. Although neural networks have been used for tree ring segmentation on macroscopic images of wood, the complexity of cell patterns in stained microsections of broadleaved species requires adaptive models to accurately accomplish this task. We present an automatic tree ring boundary delineation using neural networks on stained cross-sectional microsection images from beech cores. We trained a UNETR, a combined neural network of UNET and the attention mechanisms of Visual Transformers, to automatically segment annual ring boundaries. Its accuracy was evaluated considering discrepancies with manual segmentation and the consequences of disparity for the goals of quantitative wood anatomy analyses. In most cases (91.8%), automatic segmentation matched or improved manual segmentation, and the rate of vessels assignment to annual rings was similar between the two categories, even when manual segmentation was considered better. The application of convolutional neural networks-based models outperforms human operator segmentations when confronting ring boundary delimitation using specific parameters for quantitative wood anatomy analysis. Current advances on segmentation models may reduce the cost of massive and accurate data collection for quantitative wood anatomy.
Hydrological conditions play a key role in the carbon cycle of northern peatlands. This study examines the evapotranspiration (ET) dynamics and its impact on the water and energy balance in response ...to differing meteorological conditions during the exceptionally dry year 2006 and the normal wet year 2007 at a boreal peatland in Finland. Energy and water vapor fluxes were determined continuously using the eddy covariance approach. Daily ET rates varied considerably during the growing season and averaged 2.23 ± 0.15 mm d−1 and 1.59 ± 0.07 mm d−1 in the dry and wet year, respectively. Synoptic weather conditions as reflected by incoming radiation and water vapor pressure deficit (VPD) were the key factors controlling ET. Differences in the precipitation patterns and summer temperature also accounted for some of the observed differences in ET between the 2 years. No evidence was found for a relationship between ET rates and water table level, probably due to the relatively high water table level even in the dry year. Latent heat flux dominated the energy balance, particularly in the dry year 2006 with 60% of cumulative precipitation returned to the atmosphere through ET. In the wet year 2007, runoff dominated the water loss, and only 36% of the cumulative precipitation was returned to the atmosphere through ET. While the annual water balance regime of the peatland was mainly regulated by the precipitation pattern, daily measured ET was closely related to potential evaporation, and latent heat flux could be well modeled by the Penman‐Monteith approach, suggesting two feasible schemes for ET prediction in peatlands under well watered conditions.
The ranges of black and white spruce are largely sympatric, suggesting both species have similar climate requirements. The two species, however, are highly segregated across the landscape with black ...spruce most common on nutrient‐poor sites with cold, poorly drained soils and white spruce more common on productive sites with warmer, well‐drained soils. Because site conditions influence tree climate–growth responses, it is difficult to compare white and black spruce climate–growth responses as these responses are confounded by the differences in site conditions in which the two species naturally occur. As the climate warms dramatically in northern latitudes, it is critical to understand how a changing climate and associated changes in permafrost and fire regimes will interact to shape future species composition and ecosystem functioning in the boreal forest. In this study, we examined the climate–growth responses of black and white spruce growing in the same sites. This approach eliminates the confounding factor of site conditions and facilitates our understanding of how these two species respond to climate. We included standardized thaw depth of the active layer in our analysis as a representation of permafrost, which is a key factor delineating these two species' habitat preferences and is actively warming and thawing as the climate warms. Our most important finding was that the climate–growth responses of the two species, but especially white spruce, hinged on the thaw depth of the active layer. Specifically, with increasing June–July temperatures white spruce radial growth increased in areas with deep thaw or no near‐surface permafrost, but strongly decreased when growing in areas with near‐surface permafrost. Black spruce radial growth was less sensitive to June–July temperature than white spruce but had a consistent and more positive response to summer precipitation. These findings point to a primary mechanism potentially driving the positioning of these two tree species within the landscapes of boreal interior Alaska and imply widespread thawing of permafrost may foster expansion of white spruce in this region at the expense of black spruce, but that in a wetter climate, black spruce may gain competitive advantage over white spruce in some landscape positions.
The recent developments in artificial intelligence have the potential to facilitate new research methods in ecology. Especially Deep Convolutional Neural Networks (DCNNs) have been shown to ...outperform other approaches in automatic image analyses. Here we apply a DCNN to facilitate quantitative wood anatomical (QWA) analyses, where the main challenges reside in the detection of a high number of cells, in the intrinsic variability of wood anatomical features, and in the sample quality. To properly classify and interpret features within the images, DCNNs need to undergo a training stage. We performed the training with images from transversal wood anatomical sections, together with manually created optimal outputs of the target cell areas. The target species included an example for the most common wood anatomical structures: four conifer species; a diffuse-porous species, black alder (
L.); a diffuse to semi-diffuse-porous species, European beech (
L.); and a ring-porous species, sessile oak (
Liebl.). The DCNN was created in Python with Pytorch, and relies on a Mask-RCNN architecture. The developed algorithm detects and segments cells, and provides information on the measurement accuracy. To evaluate the performance of this tool we compared our Mask-RCNN outputs with U-Net, a model architecture employed in a similar study, and with ROXAS, a program based on traditional image analysis techniques. First, we evaluated how many target cells were correctly recognized. Next, we assessed the cell measurement accuracy by evaluating the number of pixels that were correctly assigned to each target cell. Overall, the "learning process" defining artificial intelligence plays a key role in overcoming the issues that are usually manually solved in QWA analyses. Mask-RCNN is the model that better detects which are the features characterizing a target cell when these issues occur. In general, U-Net did not attain the other algorithms' performance, while ROXAS performed best for conifers, and Mask-RCNN showed the highest accuracy in detecting target cells and segmenting lumen areas of angiosperms. Our research demonstrates that future software tools for QWA analyses would greatly benefit from using DCNNs, saving time during the analysis phase, and providing a flexible approach that allows model retraining.
Coastal dunes near the Baltic Sea are often stabilized by Scots pine forests and are characterized by a mild climate. These ecosystems are affected by water shortages and might be influenced by ...climate extremes. Considering future climate change, utilizing tree rings could help assess the role of climate extremes on coastal forest growth. We used superposed epoch analysis to study Scots pine responses to droughts and cold winters, with focus on frequency, timing, and duration. We measured ring widths (RW) and latewood blue intensity (LBI) on samples extracted from trees growing at dune ridge and bottom microsites at the south Baltic Sea. At the regional scale, we observed some similarities in tree responses to both extremes between RW and LBI within the same microsite type and region. At the local scale, RW and LBI were more frequently influenced by cold winters than droughts. RW and LBI from dune ridges were more frequently influenced by droughts than RW and LBI from dune bottoms. LBI from both microsites was more often influenced by droughts than RW. RW and LBI from both microsites were similarly often influenced by cold winters. At both scales, the response time of RW and LBI after droughts predominantly lagged by one year, while cold winters were recorded in the same year. The typical duration of growth reductions after both extremes was one year for both RW and LBI. Our study indicates that Scots pine from the Baltic Sea region is sensitive to climate extremes, especially cold winters.
Local adaptations to environmental conditions are of high ecological importance as they determine distribution ranges and likely affect species responses to climate change. Increased environmental ...stress (warming, extreme drought) due to climate change in combination with decreased genetic mixing due to isolation may lead to stronger local adaptations of geographically marginal than central populations. We experimentally observed local adaptations of three marginal and four central populations of Fagus sylvatica L., the dominant native forest tree, to frost over winter and in spring (late frost). We determined frost hardiness of buds and roots by the relative electrolyte leakage in two common garden experiments. The experiment at the cold site included a continuous warming treatment; the experiment at the warm site included a preceding summer drought manipulation. In both experiments, we found evidence for local adaptation to frost, with stronger signs of local adaptation in marginal populations. Winter frost killed many of the potted individuals at the cold site, with higher survival in the warming treatment and in those populations originating from colder environments. However, we found no difference in winter frost tolerance of buds among populations, implying that bud survival was not the main cue for mortality. Bud late frost tolerance in April differed between populations at the warm site, mainly because of phenological differences in bud break. Increased spring frost tolerance of plants which had experienced drought stress in the preceding summer could also be explained by shifts in phenology. Stronger local adaptations to climate in geographically marginal than central populations imply the potential for adaptation to climate at range edges. In times of climate change, however, it needs to be tested whether locally adapted populations at range margins can successfully adapt further to changing conditions.
Seedlings of F. sylvatica showed evidence for local adaptations to winter and spring frosts which were stronger developed in three marginal than in four central populations referring to the range of the species. We confirmed that preceding summer drought can improve frost tolerance and found no difference in the sensitivity of populations in this drought effect. The response of bud frost tolerance in winter to continuous warming, however, differed between central and marginal populations, thereby raising the question how marginal populations will be able to adapt to ongoing climate change.
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