Search

Water and nutrient are two critical factors that limit plant growth to spatial-temporal extents. Tree root foraging behavior has not received adequate attention in heterogeneous soil environments in temperate forest under drought pressure. In this study, birch (Betula platyphylla) and larch (Larix olgensis) seedlings were raised in pots in a split-root system with artificially heterogeneous soil environments to study the root foraging response to drought. Potted space was split into two halves where substrates were mixed with fertilizers in 67.5 mg nitrogen (N) plant-1 (N-P2O5-K2O, 14-13-13) to both halves as to create a homogeneous condition. Otherwise, a rate of 135 mg N plant-1 of fertilizers was delivered to a random half to create a heterogeneous condition. Half of seedlings were fully sub-irrigated every three days with the other half received the drought treatment by being watered every six days. Both birch and larch seedlings showed greater net shoot growth and biomass increment in well-watered condition, while root morphology was promoted by drought. Both species placed more fine roots with higher root N concentration in nutrient-enriched patches. In the heterogeneous pattern, birch showed a higher foraging precision assessed by biomass and greater foraging plasticity assessed in morphology and physiology. In contrast, larch seedlings had higher root N concentration in the well-watered condition. Neither species showed a significant response of N utilization to the heterogeneous pattern, but both used more N when water supply was improved. Overall, birch is better at acclimating to heterogeneous soil conditions, but its ability to seize N was lower than larch when drought was alleviated., Competing Interests: The authors declare no conflict of interest. The commercial affiliation of Harbin Gloria Pharmaceuticals Ltd. (Harbin city, postal code 150040, China) does not alter authors’ adherence to PLOS ONE policies on sharing data and materials.

Numerous long-term, free-air plant growth facilities currently explore vegetation responses to the ongoing climate change in northern latitudes. Open top chamber (OTC) experiments as well as the experimental set-ups with active warming focus on many facets of plant growth and performance, but information on morphological alterations of plant cells is still scarce. Here we compare the effects of in-situ warming on leaf epidermal cell expansion in dwarf birch, Betula nana in Finland, Greenland, and Poland. The localities of the three in-situ warming experiments represent contrasting regions of B. nana distribution, with the sites in Finland and Greenland representing the current main distribution in low and high Arctic, respectively, and the continental site in Poland as a B. nana relict Holocene microrefugium. We quantified the epidermal cell lateral expansion by microscopic analysis of B. nana leaf cuticles. The leaves were produced in paired experimental treatment plots with either artificial warming or ambient temperature. At all localities, the leaves were collected in two years at the end of the growing season to facilitate between-site and within-site comparison. The measured parameters included the epidermal cell area and circumference, and using these, the degree of cell wall undulation was calculated as an Undulation Index (UI). We found enhanced leaf epidermal cell expansion under experimental warming, except for the extremely low temperature Greenland site where no significant difference occurred between the treatments. These results demonstrate a strong response of leaf growth at individual cell level to growing season temperature, but also suggest that in harsh conditions other environmental factors may limit this response. Our results provide evidence of the relevance of climate warming for plant leaf maturation and underpin the importance of studies covering large geographical scales., Competing Interests: The authors have declared that no competing interests exist.

Birches are generally known for their high genetic and morphological variability, which has resulted in the description of many species. Ojców birch was described in 1809 by Willibald Suibert Joseph Gottlieb Besser in Poland. Since then, several studies assessing its taxonomy were conducted. Today, various authors present Ojców birch at different taxonomic ranks. In Czechia, the Ojców birch is classified a critically endangered taxon and confirmed at one locality consisting of several tens of individuals. However, before a strategy for its conservation can be applied, we consider it necessary to assess the taxonomic position of the endangered Czech population and to evaluate its relationship to the original Polish population. This study aimed to evaluate the morphometric and genetic variability between populations of B. ×oycoviensis in Poland and the Czechia and their relationship to regional populations of B. pendula, one of the putative parental species of the Ojców birch. Altogether, 106 individuals were sampled, including the holotype of B. szaferi, the second putative parental species of B. ×oycoviensis, received from the herbarium of W. Szafer, which is deposited at the Institute of Botany in Kraków. Morphological analyses identified differences in leaves between B. ×oycoviensis and B. pendula. However, no significant differences were found in genome size between selected taxa/working units except for B. pendula sampled in Czechia. The identified difference of the Czech population of B. pendula is probably caused by population variability. Genetic variability between all the taxa under comparison, regardless of their origin, was also very low; only the benchmark taxa (B. nana and B. humilis) clearly differed from all samples analyzed. The results indicate minute morphological and negligible genetic variability between the Czech and Polish populations of B. ×oycoviensis. In light of our results, the classification of B. ×oycoviensis as B. pendula var. oycoviensis seems more accurate than all hitherto presented alternatives (e.g. B. ×oycoviensis as a separate species)., Competing Interests: The authors have declared that no competing interests exist.

In many woody dicot plant species, colder temperatures correlate with a greater degree of leaf dissection and with larger and more abundant leaf teeth (the serrated edges along margins). The measurement of site-mean characteristics of leaf size and shape (physiognomy), including leaf dissection and tooth morphology, has been an important paleoclimate tool for over a century. These physiognomic-based climate proxies require that all woody dicot plants at a site, regardless of species, change their leaf shape rapidly and predictably in response to temperature. Here we experimentally test these assumptions by growing five woody species in growth cabinets under two temperatures (17 and 25°C). In keeping with global site-based patterns, plants tend to develop more dissected leaves with more abundant and larger leaf teeth in the cool treatment. Overall, this upholds the assumption that leaf shape responds in a particular direction to temperature change. The assumption that leaf shape variables respond to temperature in the same way regardless of species did not hold because the responses varied by species. Leaf physiognomic models for inferring paleoclimate should take into account these species-specific responses., Competing Interests: The authors have declared that no competing interests exist.

The alternative prey hypothesis predicts that the interaction between generalist predators and their main prey is a major driver of population dynamics of alternative prey species. In Fennoscandia, changes in climate and human land use are assumed to alter the dynamics of cyclic small rodents (main prey) and lead to increased densities and range expansion of an important generalist predator, the red fox Vulpes vulpes. In order to better understand the role of these potential changes in community structure on an alternative prey species, willow ptarmigan Lagopus lagopus, we analyzed nine years of population census data from SE Norway to investigate how community interactions affected their population dynamics. The ptarmigan populations showed no declining trend during the study period, and annual variations corresponded with marked periodic small rodent peaks and declines. Population growth and breeding success were highly correlated, and both demographic variables were influenced by an interaction between red fox and small rodents. Red foxes affected ptarmigan negatively only when small rodent abundance was low, which is in accordance with the alternative prey hypothesis. Our results confirm the important role of red fox predation in ptarmigan dynamics, and indicate that if small rodent cycles are disrupted, this may lead to decline in ptarmigan and other alternative prey species due to elevated predation pressure., Competing Interests: The authors have declared that no competing interests exist.

Worldwide tree mortality as induced by climate change presents a challenge to forest managers. To successfully manage vulnerable forests requires the capacity of regeneration to compensate for losses from tree mortality. We observed rapid regeneration and the growth release of young trees after warming-induced mortality in a David aspen-dominated (Populus davidiana) broad-leaved forest in Inner Mongolia, China, as based on individual tree measurements taken in 2012 and 2015 from a 6-ha permanent plot. Warming and drought stress killed large trees 10-15 m tall with a total number of 2881 trees during 2011-2012, and also thinned the upper crowns. David aspen recruitment increased 2 times during 2012-2015 and resulted in a high transition probability of David aspen replacing the same or other species, whereas the recruitment of Mongolian oak (Quercus mongolica) was much lower: it decreased from 2012 to 2015, indicating that rapid regeneration represented a regrowth phase for David aspen, and not succession to Mongolian oak. Further, we found that the recruitment density increased with canopy openness, thus implying that warming-induced mortality enhanced regeneration. Our results suggest that David aspen has a high regrowth ability to offset individual losses from warming-induced mortality. This important insight has implications for managing this vulnerable forest in the semi-arid region of northern China.

We sampled shrub canopy volume (height times area) and environmental factors (soil wetness, soil depth of thaw, soil pH, mean July air temperature, and typical date of spring snow loss) on 471 plots across five National Park Service units in northern Alaska. Our goal was to determine the environments where tall shrubs thrive and use this information to predict the location of future shrub expansion. The study area covers over 80,000 km2 and has mostly tundra vegetation. Large canopy volumes were uncommon, with volumes over 0.5 m3/m2 present on just 8% of plots. Shrub canopy volumes were highest where mean July temperatures were above 10.5°C and on weakly acid to neutral soils (pH of 6 to 7) with deep summer thaw (>80 cm) and good drainage. On many sites, flooding helped maintain favorable soil conditions for shrub growth. Canopy volumes were highest where the typical snow loss date was near 20 May; these represent sites that are neither strongly wind-scoured in the winter nor late to melt from deep snowdrifts. Individual species varied widely in the canopy volumes they attained and their response to the environmental factors. Betula sp. shrubs were the most common and quite tolerant of soil acidity, cold July temperatures, and shallow thaw depths, but they did not form high-volume canopies under these conditions. Alnus viridis formed the largest canopies and was tolerant of soil acidity down to about pH 5, but required more summer warmth (over 12°C) than the other species. The Salix species varied widely from S. pulchra, tolerant of wet and moderately acid soils, to S. alaxensis, requiring well-drained soils with near neutral pH. Nearly half of the land area in ARCN has mean July temperatures of 10.5 to 12.5°C, where 2°C of warming would bring temperatures into the range needed for all of the potential tall shrub species to form large canopies. However, limitations in the other environmental factors would probably prevent the formation of large shrub canopies on at least half of the land area with newly favorable temperatures after 2°C of warming.

Betula platyphylla Suk (birch) is a fast-growing woody species that is important in pulp industries and the biofuels. However, as an important pulp species, few studies had been performed on its wood formation. In the present study, we investigated the molecular responses of birch xylem to artificial bending and gravitational stimuli. After trunks of birch trees were subjected to bending for 8 weeks, the cellulose content was significantly greater in tension wood (TW) than in opposite wood (OW) or normal wood (NW), whereas the lignin content in TW was significantly lower than that in OW and NW. In addition, TW grew more rapidly than OW and generated TW-specific fibers with an additional G-layer. Three transcriptome libraries were constructed from TW, OW and NW of B. platyphylla, respectively, after the plants were subjected to artificial bending. Overall, 80,909 nonredundant unigenes with a mean size of 768 nt were assembled. Expression profiles were generated, and 9,684 genes were found to be significantly differentially expressed among the TW, OW and NW libraries. These included genes involved in secondary cell wall structure, wood composition, and cellulose or lignin biosynthesis. Our study showed that during TW formation, genes involved in cellulose synthesis were induced, while the expression of lignin synthesis-related genes decreased, resulting in increased cellulose content and decreased lignin levels in TW. In addition, fasciclin-like arabinogalactan proteins play important role in TW formation. These findings may provide important insights into wood formation at the molecular level.

To contribute to a better understanding of the regeneration strategy of Betula albosinensis forests and the likely reasons behind either the successful recovery or failure after strip clearcutting, we compared the population structures and spatial patterns of B. albosinensis in eight B. albosinensis stands in Qinling Mountains, China. Four cut and four uncut stands were selected, and each sampled using a single large plot (0.25 ha). Results indicated that, on the one hand, B. albosinensis recruitment was scarce (average of 48 stems ha(-1)) in the uncut stands, relative to the mature population (average of 259 stems ha(-1)), suggesting a failure of recruitment. On the other hand, the subsequent regeneration approximately 50 years after the strip clearcutting showed that the density of the target species seedlings and saplings has increased significantly, and the current average density of seedlings and saplings was 156 stems ha(-1). The clumped spatial pattern of B. albosinensis suggested that their regeneration was highly dependent on canopy disturbance. However, recruitment remained poor in the uncut stands because most gaps were small in scale. The successful regeneration of sunlight-loving B. albosinensis after strip clearcutting was attributed to the exposed land and availability of more sunlight. Bamboo density did not influence B. albosinensis recruitment in the uncut stands. However, stand regeneration was impeded after strip clearcutting; thus, removing bamboo is essential in improving the competitive status of B. albosinensis at the later stage of forest regeneration after clearcutting. The moderate severity of disturbance resulting from strip clearcutting reversed the degeneration trend of primary B. albosinensis stands. This outcome can help strike a balance between forest conservation and the demand for wood products by releasing space and exposing the forested land for recruitment. Life history traits and spatiotemporal disturbance magnitude are important factors to consider in implementing effective B. albosinensis regeneration strategies.

Immediate phenotypic variation and the lagged effect of evolutionary adaptation to climate change appear to be two key processes in tree responses to climate warming. This study examines these components in two types of growth models for predicting the 2010-2099 diameter growth change of four major boreal species Betula papyrifera, Pinus banksiana, Picea mariana, and Populus tremuloides along a broad latitudinal gradient in eastern Canada under future climate projections. Climate-growth response models for 34 stands over nine latitudes were calibrated and cross-validated. An adaptive response model (A-model), in which the climate-growth relationship varies over time, and a fixed response model (F-model), in which the relationship is constant over time, were constructed to predict future growth. For the former, we examined how future growth of stands in northern latitudes could be forecasted using growth-climate equations derived from stands currently growing in southern latitudes assuming that current climate in southern locations provide an analogue for future conditions in the north. For the latter, we tested if future growth of stands would be maximally predicted using the growth-climate equation obtained from the given local stand assuming a lagged response to climate due to genetic constraints. Both models predicted a large growth increase in northern stands due to more benign temperatures, whereas there was a minimal growth change in southern stands due to potentially warm-temperature induced drought-stress. The A-model demonstrates a changing environment whereas the F-model highlights a constant growth response to future warming. As time elapses we can predict a gradual transition between a response to climate associated with the current conditions (F-model) to a more adapted response to future climate (A-model). Our modeling approach provides a template to predict tree growth response to climate warming at mid-high latitudes of the Northern Hemisphere.

Vessels of broad-leaved trees have been analyzed to study how trees deal with various environmental factors. Cambial injury, in particular, has been reported to induce the formation of narrower conduits. Yet, little or no effort has been devoted to the elaboration of vessel sampling strategies for retrospective injury detection based on vessel lumen size reduction. To fill this methodological gap, four wounded individuals each of grey alder (Alnus incana (L.) Moench) and downy birch (Betula pubescens Ehrh.) were harvested in an avalanche path. Earlywood vessel lumina were measured and compared for each tree between the injury ring built during the growing season following wounding and the control ring laid down the previous year. Measurements were performed along a 10 mm wide radial strip, located directly next to the injury. Specifically, this study aimed at (i) investigating the intra-annual duration and local extension of vessel narrowing close to the wound margin and (ii) identifying an adequate sample of earlywood vessels (number and intra-ring location of cells) attesting to cambial injury. Based on the results of this study, we recommend analyzing at least 30 vessels in each ring. Within the 10 mm wide segment of the injury ring, wound-induced reduction in vessel lumen size did not fade with increasing radial and tangential distances, but we nevertheless advise favoring early earlywood vessels located closest to the injury. These findings, derived from two species widespread across subarctic, mountainous, and temperate regions, will assist retrospective injury detection in Alnus, Betula, and other diffuse-porous species as well as future related research on hydraulic implications after wounding.

Densification of the shrub layer has been reported in many subarctic regions, raising questions about the implication for large herbivores and their resources. Shrubs can tolerate browsing and their level of tolerance could be affected by browsing and soils productivity, eventually modifying resource availability for the caribou. Our objective was to assess the compensatory growth potential of a subarctic shrub, Betula glandulosa Michx., in relation with caribou browsing and nutriment availability for the plants. We used a simulated browsing (0, 25 and 75% of available shoots) and nitrogen-fertilisation (0 and 10 g m(-2)) experiment to test two main hypotheses linking tolerance to resource availability, the Compensatory Continuum Hypothesis and the Growth Rate Hypothesis as well as the predictions from the Limiting Resource Model. We seek to explicitly integrate the relative browsing pressure in our predictions since the amount of tissues removed could affect the capacity of long-lived plants to compensate. Birches fully compensated for moderate browsing with an overall leaf biomass similar to unbrowsed birches but undercompensated under heavy browsing pressure. The main mechanism explaining compensation appears to be the conversion of short shoots into long shoots. The leaf area increased under heavy browsing pressure but only led to undercompensation. Fertilisation for two consecutive years did not influence the response of birch, thus we conclude that our results support the LRM hypothesis of equal tolerance under both high and low nitrogen availability. Our results highlight that the potential for compensatory growth in dwarf birch is surpassed under heavy browsing pressure independently of the fertilisation regime. In the context of the worldwide decline in caribou herds, the reduction in browsing pressure could act synergistically with global climate change to promote the current shrub expansion reported in subarctic regions.

Most manipulation experiments simulating global change in tundra were short-term or did not measure plant growth directly. Here, we assessed the growth of three shrubs (Cassiope tetragona, Empetrum hermaphroditum and Betula nana) at a subarctic heath in Abisko (Northern Sweden) after 22 years of warming (passive greenhouses), fertilisation (nutrients addition) and shading (hessian fabric), and compare this to observations from the first decade of treatment. We assessed the growth rate of current-year leaves and apical stem (primary growth) and cambial growth (secondary growth), and integrated growth rates with morphological measurements and species coverage. Primary- and total growth of Cassiope and Empetrum were unaffected by manipulations, whereas growth was substantially reduced under fertilisation and shading (but not warming) for Betula. Overall, shrub height and length tended to increase under fertilisation and warming, whereas branching increased mostly in shaded Cassiope. Morphological changes were coupled to increased secondary growth under fertilisation. The species coverage showed a remarkable increase in graminoids in fertilised plots. Shrub response to fertilisation was positive in the short-term but changed over time, likely because of an increased competition with graminoids. More erected postures and large, canopies (requiring enhanced secondary growth for stem reinforcement) likely compensated for the increased light competition in Empetrum and Cassiope but did not avoid growth reduction in the shade intolerant Betula. The impact of warming and shading on shrub growth was more conservative. The lack of growth enhancement under warming suggests the absence of long-term acclimation for processes limiting biomass production. The lack of negative effects of shading on Cassiope was linked to morphological changes increasing the photosynthetic surface. Overall, tundra shrubs showed developmental plasticity over the longer term. However, such plasticity was associated clearly with growth rate trends only in fertilised plots.