Severe drought may increase physiological stress on long-lived woody vegetation, occasionally leading to mortality of overstory trees. Little is known about the factors determining tree survival and ...subsequent recovery after drought. We used structural equation modeling to analyse the recovery of Scots pine (Pinus sylvestris) trees 4 yr after an extreme drought episode occurred in 2004-2005 in north-east Spain. Measured variables included the amount of green foliage, carbon reserves in the stem, mistletoe (Viscum album) infection, needle physiological performance and stem radial growth before, during and after the drought event. The amount of green leaves and the levels of carbon reserves were related to the impact of drought on radial growth, and mutually correlated. However, our most likely path model indicated that current depletion of carbon reserves was a result of reduced photosynthetic tissue. This relationship potentially constitutes a feedback limiting tree recovery. In addition, mistletoe infection reduced leaf nitrogen content, negatively affecting growth. Finally, successive surveys in 2009-2010 showed a direct association between carbon reserves depletion and drought-induced mortality. Severe drought events may induce long-term physiological disorders associated with canopy defoliation and depletion of carbon reserves, leading to prolonged recovery of surviving individuals and, eventually, to delayed tree death.
Patterns of plant biomass allocation and functional adjustments along climatic gradients are poorly understood, particularly belowground. Generally, low temperatures suppress nutrient release and ...uptake, and forests under such conditions have a greater proportion of their biomass in roots. However, it is not clear whether ‘more roots’ means better capacity to acquire soil resources.
Herein we quantified patterns of fine-root anatomy and their biomass distribution across Scots pine (Pinus sylvestris) populations both along a 2000-km latitudinal gradient and within a common garden experiment with a similar range of populations.
We found that with decreasing mean temperature, a greater percentage of Scots pine root biomass was allocated to roots with higher potential absorptive capacity. Similar results were seen in the common experimental site, where cold-adapted populations produced roots with greater absorptive capacity than populations originating from warmer climates.
These results demonstrate that plants growing in or originated from colder climates have more acquisitive roots, a trait that is likely adaptive in the face of the low resource availability typical of cold soils.
Radioactive contamination of the natural areas is one of the most long-lasting anthropogenic impacts on the environment. Scots pine (Pinus sylvestris L.) is a promising organism for radiation-related ...research because of its high radiosensitivity, but the genome size of Pinacea species has imposed obstacles for high-throughput studies so far. In this work, we conducted the analysis of the de novo assembled transcriptome of Scots pine populations growing in the Chernobyl-affected zone, which is still today contaminated with radionuclides because of the accident at the nuclear power plant in 1986. The transcriptome profiles indicate a clear pattern of adaptive stress response, which seems to be dose-dependent. The transcriptional response indicates a continuous modulation of the cellular redox system, enhanced expression of chaperones and histones, along with the control of ions balance. Interestingly, the activity of transposable element families is inversely correlated to the exposure levels to radiation. These adaptive responses, which are triggered by radiation doses 30 times lower than the one accepted as a safe for biota species by international regulations, suggest that the environmental management in radiation protection should be reviewed.
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•De novo transcriptome assembly of Scots pine trees from Chernobyl affected zone.•The profiles indicate stress adaptation to doses considered as safe for biota species.•The response pattern suggests a sustained control of the redox system and ion balance.•Transposons activity essentially inversely correlated to the levels of irradiation.
De novo transcriptome analysis of chronically irradiated Scots pine trees from Chernobyl affected zone reveals a unique stress response profile.
Forestry reshapes ecosystems with respect to tree age structure, soil properties and vegetation composition. These changes are likely to be paralleled by shifts in microbial community composition ...with potential feedbacks on ecosystem functioning. Here, we assessed fungal communities across a chronosequence of managed Pinus sylvestris stands and investigated correlations between taxonomic composition and extracellular enzyme activities. Not surprisingly, clear-cutting had a negative effect on ectomycorrhizal fungal abundance and diversity. In contrast, clear-cutting favoured proliferation of saprotrophic fungi correlated with enzymes involved in holocellulose decomposition. During stand development, the re-establishing ectomycorrhizal fungal community shifted in composition from dominance by Atheliaceae in younger stands to Cortinarius and Russula species in older stands. Late successional ectomycorrhizal taxa correlated with enzymes involved in mobilisation of nutrients from organic matter, indicating intensified nutrient limitation. Our results suggest that maintenance of functional diversity in the ectomycorrhizal fungal community may sustain long-term forest production by retaining a capacity for symbiosis-driven recycling of organic nutrient pools.
We tested two methods for non-destructive assessment of wood density of Scots pine standing trees: one based on penetration depth of a steel pin (Pilodyn) and the other on micro-drilling resistance ...(Resistograph). As a benchmark we used wood density data from x-ray analysis (SilviScan). We assessed in total 622 trees of 175 full-sib families growing in a single progeny test. Pilodyn was applied with bark (PIL) and without bark (PILB). Raw Resistograph drilling profiles (RES) were adjusted (RESTB) in order to eliminate increasing trend caused by needle friction. Individual narrow-sense heritability of benchmark SilviScan density (DEN; 0.46) was most closely approached by that of adjusted RESTB (0.43). Heritabilities were lower for unadjusted RES (0.35) as well as for PIL and PILB (both 0.32). Additive genetic correlations of the benchmark DEN with RES, RESTB, PIL and PILB were 0.89, 0.96, 0.59 and 0.71, respectively. Our results suggest that Resistograph is a more reliable tool than Pilodyn for wood density assessment of Scots pine; however, we highly recommend adjusting Resistograph drilling profiles prior to further analyses.
• The extent to which water availability can be used to predict the enlargement and final dimensions of xylem conduits remains an open issue.
• We reconstructed the time course of tracheid ...enlargement in Pinus sylvestris trees in central Spain by repeated measurements of tracheid diameter on microcores sampled weekly during a 2 yr period. We analyzed the role of water availability in these dynamics empirically through time-series correlation analysis and mechanistically by building a model that simulates daily tracheid enlargement rate and duration based on Lockhart’s equation and water potential as the sole input.
• Tracheid enlargement followed a sigmoid-like time course, which varied intra- and interannually. Our empirical analysis showed that final tracheid diameter was strongly related to water availability during tracheid enlargement. The mechanistic model was calibrated and successfully validated (R² = 0.92) against the observed tracheid enlargement time course. The model was also able to reproduce the seasonal variations of tracheid enlargement rate, duration and final diameter (R² = 0.84–0.99).
• Our results support the hypothesis that tracheid enlargement and final dimensions can be modeled based on the direct effect of water potential on turgor-driven cell expansion. We argue that such a mechanism is consistent with other reported patterns of tracheid dimension variation.
Polyamines are small metabolites present in all living cells and play fundamental roles in numerous physiological events in plants. The aminopropyltransferases (APTs), spermidine synthase (SPDS), ...spermine synthase (SPMS) and thermospermine synthase (ACL5), are essential enzymes in the polyamine biosynthesis pathway. In angiosperms, SPMS has evolved from SPDS via gene duplication, whereas in gymnosperms APTs are mostly unexplored and no SPMS gene has been reported. The present study aimed to investigate the functional properties of the SPDS and ACL5 proteins of Scots pine (Pinus sylvestris L.) in order to elucidate the role and evolution of APTs in higher plants.
Germinating Scots pine seeds and seedlings were analysed for polyamines by high-performance liquid chromatography (HPLC) and the expression of PsSPDS and PsACL5 genes by in situ hybridization. Recombinant proteins of PsSPDS and PsACL5 were produced and investigated for functional properties. Also gene structures, promoter regions and phylogenetic relationships of PsSPDS and PsACL5 genes were analysed.
Scots pine tissues were found to contain spermidine, spermine and thermospermine. PsSPDS enzyme catalysed synthesis of both spermidine and spermine. PsACL5 was found to produce thermospermine, and PsACL5 gene expression was localized in the developing procambium in embryos and tracheary elements in seedlings.
Contrary to previous views, our results demonstrate that SPMS activity is not a novel feature developed solely in the angiosperm lineage of seed plants but also exists as a secondary property in the Scots pine SPDS enzyme. The discovery of bifunctional SPDS from an evolutionarily old conifer reveals the missing link in the evolution of the polyamine biosynthesis pathway. The finding emphasizes the importance of pre-existing secondary functions in the evolution of new enzyme activities via gene duplication. Our results also associate PsACL5 with the development of vascular structures in Scots pine.
The frequently observed forest decline in water‐limited regions may be associated with impaired tree hydraulics, but the precise physiological mechanisms remain poorly understood. We compared ...hydraulic architecture of Mongolian pine (Pinus sylvestris var. mongolica) trees of different size classes from a plantation and a natural forest site to test whether greater hydraulic limitation with increasing size plays an important role in tree decline observed in the more water‐limited plantation site. We found that trees from plantations overall showed significantly lower stem hydraulic efficiency. More importantly, plantation‐grown trees showed significant declines in stem hydraulic conductivity and hydraulic safety margins as well as syndromes of stronger drought stress with increasing size, whereas no such trends were observed at the natural forest site. Most notably, the leaf to sapwood area ratio (LA/SA) showed a strong linear decline with increasing tree size at the plantation site. Although compensatory adjustments in LA/SA may mitigate the effect of increased water stress in larger trees, they may result in greater risk of carbon imbalance, eventually limiting tree growth at the plantation site. Our results provide a potential mechanistic explanation for the widespread decline of Mongolian pine trees in plantations of Northern China.
Hydraulic failure has been proposed as an important reason causing tree die‐off in vast areas of water‐limited land across the globe, especially under the influence of climate change, but the underlying mechanisms remain poorly understood. Decline has been more frequently observed in larger trees, and significant changes in hydraulic architecture are also commonly found with increasing tree size, suggesting a mechanistic linkage between tree decline and hydraulic limitations mediated by variation in tree size. Here, we compared hydraulic architecture of Pinus sylvestris var. mongolica trees between a plantation and a natural forest site across different tree size classes to test whether greater hydraulic limitation in larger trees is responsible for tree decline in the more water‐limited plantation site. We observed clear contrasts in hydraulic‐related characteristics between trees of the 2 sites and found clear trends of change in these traits with increasing tree size at the plantation site but not at the natural forest site, which provide a potential mechanistic explanation for the widespread decline of this species in plantations of Northern China.
Aggressive reactive oxygen species (ROS) play an important role in plant defence against biotic stressors, including herbivorous insects. Plants may even generate ROS in response to insect eggs, thus ...effectively fighting against future larval herbivory. However, so far nothing is known on how ROS-mediated plant defence against insect eggs is enzymatically regulated. Neither do we know how insects cope with egg-induced plant ROS. We addressed these gaps of knowledge by studying the activities of ROS-related enzymes in Pinus sylvestris deposited with eggs of the herbivorous sawfly Diprion pini. This species cuts a slit into pine needles and inserts its eggs into the needle tissue. About a quarter of egg-deposited needles show chlorotic tissue at the oviposition sites, indicating hypersensitive response-like direct defence responses resulting in reduced larval hatching from eggs. Hydrogen peroxide and peroxidase sensitive staining of sections of egg-deposited pine needles revealed the presence of hydrogen peroxide and peroxidase activity in needle tissue close to the eggs. Activity of ROS-producing NADPH-oxidase did not increase after egg deposition. However, the activity of the ROS-detoxifying enzyme catalase decreased after egg deposition and ovipositional wounding of needles. These results show that local ROS accumulation at the oviposition site is not caused by increased NADPH-oxidase activity, but reduced activity of pine needle catalase may contribute to it. However, our data suggest that pine sawflies can counteract the egg deposition-induced hydrogen peroxide accumulation in pine needles by high catalase activity in their oviduct secretion which is released with the eggs into pine tissue.
The purpose of this study was to determine the concentrations of heavy metals (cadmium, iron, manganese, lead and zinc) in current-year, 1-year old and 2-year old needles of
Pinus sylvestris
L. Trees ...were from three heavily polluted (immediate vicinity of zinc smelter, iron smelter and power plant) and three relatively clean sites (nature reserve, ecologically clean site and unprotected natural forest community) in southern Poland. Analysis also concerned the antioxidant response and contents of protein, proline, total glutathione, non-protein thiols and activity of guaiacol peroxidase (GPX) in the needles. Generally, in pine needles from the polluted sites, the concentrations of the metals were higher and increased with the age of needles, and in most cases, antioxidant responses also were elevated. The highest levels of Cd, Pb and Zn were found in 2-year old pine needles collected near the polluted zinc smelter (respectively: 6.15, 256.49, 393.5 mg kg
−1
), Fe in 2-year old pine needles in the vicinity of the iron smelter (206.82 mg kg
−1
) and Mn in 2-year old needles at the ecologically clean site (180.32 mg kg
−1
). Positive correlations were found between Fe, Mn and Pb and the content of proteins and NPTs, between Cd and non-protein –SH groups, and between Zn and proline levels. The activity of GPX increased under the influence of Mn, while glutathione levels tended to decrease as Mn levels rose. The data obtained show that the levels of protein and non-protein –SH groups may be useful in biological monitoring, and that these ecophysiological parameters seem to be good evidence of elevated oxidative stress caused by heavy metals.
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