Saline and sodic soils that cannot be used for agriculture occur worldwide. Cultivating stress‐tolerant trees to obtain biomass from salinized areas has been suggested. Various tree species of ...economic importance for fruit, fibre and timber production exhibit high salinity tolerance. Little is known about the mechanisms enabling tree crops to cope with high salinity for extended periods. Here, the molecular, physiological and anatomical adjustments underlying salt tolerance in glycophytic and halophytic model tree species, such as Populus euphratica in terrestrial habitats, and mangrove species along coastlines are reviewed. Key mechanisms that have been identified as mediating salt tolerance are discussed at scales from the genetic to the morphological level, including leaf succulence and structural adjustments of wood anatomy. The genetic and transcriptomic bases for physiological salt acclimation are salt sensing and signalling networks that activate target genes; the target genes keep reactive oxygen species under control, maintain the ion balance and restore water status. Evolutionary adaptation includes gene duplication in these pathways. Strategies for and limitations to tree improvement, particularly transgenic approaches for increasing salt tolerance by transforming trees with single and multiple candidate genes, are discussed.
The molecular, physiological and anatomical adjustments underlying salt tolerance in glycophytic and halophytic tree species are reviewed.
Tolerance mechanisms include salt sensing and signalling, activation of transport processes to maintain the ion balance, restoration of the water status and removal of excess reactive oxygen species.
Forest tree breeding to increased salinity tolerance can be accelerated by use of novel genomic resources and gene stacking.
Cultivation of salt tolerant trees may generate economic revenues on degraded, saline farmland and contribute to climate mitigation.
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BFBNIB, FZAB, GIS, IJS, KILJ, NLZOH, NUK, OILJ, SBCE, SBMB, UL, UM, UPUK
Heavy metal (HM)-accumulating herbaceous and woody plants are employed for phytoremediation. To develop improved strategies for enhancing phytoremediation efficiency, knowledge of the ...microstructural, physiological and molecular responses underlying HM-accumulation is required. Here we review the progress in understanding the structural, physiological and molecular mechanisms underlying HM uptake, transport, sequestration and detoxification, as well as the regulation of these processes by signal transduction in response to HM exposure. The significance of genetic engineering for enhancing phytoremediation efficiency is also discussed. In herbaceous plants, HMs are taken up by roots and transported into the root cells via transmembrane carriers for nutritional ions. The HMs absorbed by root cells can be further translocated to the xylem vessels and unloaded into the xylem sap, thereby reaching the aerial parts of plants. HMs can be sequestered in the cell walls, vacuoles and the Golgi apparatuses. Plant roots initially perceive HM stress and trigger the signal transduction, thereby mediating changes at the molecular, physiological, and microstructural level. Signaling molecules such as phytohormones, reactive oxygen species (ROS) and nitric oxide (NO), modulate plant responses to HMs via differentially expressed genes, activation of the antioxidative system and coordinated cross talk among different signaling molecules. A number of genes participated in HM uptake, transport, sequestration and detoxification have been functionally characterized and transformed to target plants for enhancing phytoremediation efficiency. Fast growing woody plants hold an advantage over herbaceous plants for phytoremediation in terms of accumulation of high HM-amounts in their large biomass. Presumably, woody plants accumulate HMs using similar mechanisms as herbaceous counterparts, but the processes of HM accumulation and signal transduction can be more complex in woody plants.
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GEOZS, IJS, IMTLJ, KILJ, KISLJ, NUK, OILJ, PNG, SAZU, SBCE, SBJE, UL, UM, UPCLJ, UPUK, ZRSKP
We reviewed the impact of fungal volatile organic compounds (VOCs) on soil-inhabiting organisms and their physiological and molecular consequences for their targets. Because fungi can only move by ...growth to distinct directions, a main mechanism to protect themselves from enemies or to manipulate their surroundings is the secretion of exudates or VOCs. The importance of VOCs in this regard has been significantly underestimated. VOCs not only can be means of communication, but also signals that are able to specifically manipulate the recipient. VOCs can reprogram root architecture of symbiotic partner plants or increase plant growth leading to enlarged colonization surfaces. VOCs are also able to enhance plant resistance against pathogens by activating phytohormone-dependent signaling pathways. In some cases, they were phytotoxic. Because the response was specific to distinct species, fungal VOCs may contribute to regulate the competition of plant communities. Additionally, VOCs are used by the producing fungus to attack rivaling fungi or bacteria, thereby protecting the emitter or its nutrient sources. In addition, animals, like springtails, nematodes, and earthworms, which are important components of the soil food web, respond to fungal VOCs. Some VOCs are effective repellents for nematodes and, therefore, have applications as biocontrol agents. In conclusion, this review shows that fungal VOCs have a huge impact on soil fauna and flora, but the underlying mechanisms, how VOCs are perceived by the recipients, how they manipulate their targets and the resulting ecological consequences of VOCs in inter-kingdom signaling is only partly understood. These knowledge gaps are left to be filled by future studies.
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CEKLJ, DOBA, EMUNI, FZAB, GEOZS, IJS, IMTLJ, IZUM, KILJ, KISLJ, MFDPS, NUK, OILJ, PILJ, PNG, SAZU, SBCE, SBJE, SBMB, SBNM, UILJ, UKNU, UL, UM, UPUK, VKSCE, ZAGLJ
Climatic stresses limit plant growth and productivity. In the past decade, tree improvement programs were mainly focused on yield but it is obvious that enhanced stress resistance is also required. ...In this review we highlight important drought avoidance and tolerance mechanisms in forest trees. Genomes of economically important trees species with divergent resistance mechanisms can now be exploited to uncover the mechanistic basis of long-term drought adaptation at the whole plant level. Molecular tree physiology indicates that osmotic adjustment, antioxidative defense and increased water use efficiency are important targets for enhanced drought tolerance at the cellular and tissue level. Recent biotechnological approaches focused on overexpression of genes involved in stress sensing and signaling, such as the abscisic acid core pathway, and down-stream transcription factors. By this strategy, a suite of defense systems was recruited, generally enhancing drought and salt stress tolerance under laboratory conditions. However, field studies are still scarce. Under field conditions trees are exposed to combinations of stresses that vary in duration and magnitude. Variable stresses may overrule the positive effect achieved by engineering an individual defense pathway. To assess the usability of distinct modifications, large-scale experimental field studies in different environments are necessary. To optimize the balance between growth and defense, the use of stress-inducible promoters may be useful. Future improvement programs for drought resistance will benefit from a better understanding of the intricate networks that ameliorate molecular and ecological traits of forest trees.
Phosphorus (P) is a major plant nutrient. It is transported into and allocated inside plants by four families of phosphate transporters (PHT1 to PHT4) with high or low affinity to phosphate. Here, we ...studied whole-plant P uptake kinetics and expression profiles of members of the PHT families under high, intermediate and low P availability in the woody crop poplar (Populus × canescens) in relation to plant performance.
Poplars exhibited strong growth reduction and increased P use efficiency in response to lower P availabilities. The relative P uptake rate increased with intermediate and decreased with low P availability. This decrease was not energy-limited because glucose addition could not rescue the uptake. The maximum P uptake rate was more than 13-times higher in P-starved than in well-supplied poplars. The K
for whole-root uptake ranged between 26 μM and 20 μM in poplars with intermediate and low P availability, respectively. In well-supplied plants, only low uptake rate was found. The minimum concentration for net P uptake from the nutrient solution was 1.1 μM. All PHT1 members studied showed significant up-regulation upon P starvation and were higher expressed in roots than leaves, with the exception of PtPHT1;3. PtPHT1;1 and PtPHT1;2 showed root- and P starvation-specific expression. Various members of the PHT2, PHT3 and PHT4 families showed higher expression in leaves than in roots, but were unresponsive to P deprivation. Other members (PtPHT3;1, PtPHT3;2, PtPHT3;6, PtPHT4;6 to PtPHT4;8) exhibited higher expression in roots than in leaves and were in most cases up-regulated in response to P deficiency.
Expression profiles of distinct members of the PHT families, especially those of PHT1 were linked with changes in P uptake and allocation at whole-plant level. The regulation was tissue-specific with lower P responsiveness in leaves than in roots. Uptake efficiency for P increased with decreasing P availability, but could not overcome a threshold of about 1 μM P in the nutrient solution. Because the P concentrations in soil solutions are generally in the lower micro-molar range, even below the apparent K
-values, our findings suggest that bare-rooted poplars are prone to suffer from P limitations in most environments.
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DOBA, IZUM, KILJ, NUK, PILJ, PNG, SAZU, SIK, UILJ, UKNU, UL, UM, UPUK
Increasing salinity is one of the major drawbacks for plant growth. Besides the ion itself being toxic to plant cells, it greatly interferes with the supply of other macronutrients like potassium, ...calcium and magnesium. However, little is known about how sodium affects the translocation of these nutrients from the root to the shoot. The major driving force of this translocation process is thought to be the water flow through the xylem driven by transpiration. To dissect the effects of transpiration from those of salinity we compared salt stressed, ABA treated and combined salt- and ABA treated poplars with untreated controls. Salinity reduced the root content of major nutrients like K.sup.+, Ca.sup.2+ and Mg.sup.2+ . Less Ca.sup.2+ and Mg.sup.2+ in the roots resulted in reduced leaf Ca.sup.2+ and leaf Mg.sup.2+ levels due to reduced stomatal conductance and reduced transpiration. Interestingly, leaf K.sup.+ levels were positively affected in leaves under salt stress although there was less K.sup.+ in the roots under salt. In response to ABA, transpiration was also decreased and Mg.sup.2+ and Ca.sup.2+ levels decreased comparably to the salt stress treatment, while K.sup.+ levels were not affected. Thus, our results suggest that loading and retention of leaf K.sup.+ is enhanced under salt stress compared to merely transpiration driven cation supply.
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DOBA, IZUM, KILJ, NUK, PILJ, PNG, SAZU, SIK, UILJ, UKNU, UL, UM, UPUK
Abstract
To investigate how long-lived forest trees cope with low soil phosphorus (P) availabilities, we characterized P nutrition of beech (Fagus sylvatica, L.) in soils from P-rich and P-poor beech ...forests throughout an annual growth cycle. Young trees were excavated with intact soil cores in mono-specific beech forests, kept under common garden conditions, and used for 33P labeling, analyses of P uptake, P content and biomass during five phenological stages (dormancy in winter, bud swelling in early spring, mature leaves in early and late summer, and senescent leaves in fall). Seasonal allocation patterns showed that young, emerging leaves were preferred sinks for P under P-poor conditions, thereby keeping foliar P concentrations at levels similar to those of trees grown in P-rich soil. Phosphorus concentrations in stems and roots of trees from the P-poor conditions were lower than those from P-rich conditions. Coarse roots were the main P storage tissue, supplying inorganic P to newly formed leaves, originating from the inorganic and organic P pools under low and high P conditions, respectively. Beech trees in P-poor soil exhibited net biomass increment early in the annual growth along with a strong P deficit, which was replenished by enhanced uptake in late summer and fall. Trees in P-rich soil grew until late summer, and showed a moderate P decline in organic pools and recovery late in fall, which coincided with elevated P uptake from soil. Beech in P-poor soil produced more biomass per unit of P but at a slower growth rate than those in P-rich soil, thereby exhibiting similar P-use efficiencies. Temporal decoupling of growth and P acquisition in combination with internal P trade-off between storage tissues and leaves facilitated flexible acclimation of beech to a wide range of soil P availabilities.
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IZUM, KILJ, NUK, PILJ, PNG, SAZU, UL, UM, UPUK
Phosphorus (P) is an essential plant nutrient, but its availability is often limited in soil. Here, we studied changes in the transcriptome and in nutrient element concentrations in leaves and roots ...of poplars (Populus × canescens) in response to P deficiency. P starvation resulted in decreased concentrations of S and major cations (K, Mg, Ca), in increased concentrations of N, Zn and Al, while C, Fe and Mn were only little affected. In roots and leaves >4,000 and >9,000 genes were differently expressed upon P starvation. These genes clustered in eleven co-expression modules of which seven were correlated with distinct elements in the plant tissues. One module (4.7% of all differentially expressed genes) was strongly correlated with changes in the P concentration in the plant. In this module the GO term "response to P starvation" was enriched with phosphoenolpyruvate carboxylase kinases, phosphatases and pyrophosphatases as well as regulatory domains such as SPX, but no phosphate transporters. The P-related module was also enriched in genes of the functional category "galactolipid synthesis". Galactolipids substitute phospholipids in membranes under P limitation. Two modules, one correlated with C and N and the other with biomass, S and Mg, were connected with the P-related module by co-expression. In these modules GO terms indicating "DNA modification" and "cell division" as well as "defense" and "RNA modification" and "signaling" were enriched; they contained phosphate transporters. Bark storage proteins were among the most strongly upregulated genes in the growth-related module suggesting that N, which could not be used for growth, accumulated in typical storage compounds. In conclusion, weighted gene coexpression network analysis revealed a hierarchical structure of gene clusters, which separated phosphate starvation responses correlated with P tissue concentrations from other gene modules, which most likely represented transcriptional adjustments related to down-stream nutritional changes and stress.
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DOBA, IZUM, KILJ, NUK, PILJ, PNG, SAZU, SIK, UILJ, UKNU, UL, UM, UPUK
The present study introduces metabolic modeling as a new tool to analyze the network of redox reactions composing the superoxide dismutase-ascorbate (Asc)-glutathione (GSH) cycle. Based on previously ...determined concentrations of antioxidants and defense enzymes in chloroplasts, kinetic properties of antioxidative enzymes, and nonenzymatic rate constants of antioxidants with reactive oxygen, models were constructed to simulate oxidative stress and calculate changes in concentrations and fluxes of oxidants and antioxidants. Simulated oxidative stress in chloroplasts did not result in a significant accumulation of $\text{O}_{2}{}^{.-}$ and H2O2 when the supply with reductant was sufficient. Model results suggest that the coupling between Asc- and GSH-related redox systems was weak because monodehydroascorbate radical reductase prevented dehydroascorbate (DHA) formation efficiently. DHA reductase activity was dispensable. Glutathione reductase was mainly required for the recycling of GSH oxidized in nonenzymatic reactions. In the absence of monodehydroascorbate radical reductase and DHA reductase, glutathione reductase and GSH were capable to maintain the Asc pool more than 99% reduced. This suggests that measured DHA/Asc ratios do not reflect a redox balance related to the Asc-GSH-cycle. Decreases in Asc peroxidase resulted in marked H2O2 accumulation without significant effects on the redox balance of Asc/DHA or GSH/GSSG. Simulated loss of SOD resulted in higher H2O2 production rates, thereby affecting all subsequent steps of the Asc-GSH-cycle. In conclusion, modeling approaches contribute to the theoretical understanding of the functioning of antioxidant systems by pointing out questions that need to be validated and provide additional information that is useful to develop breeding strategies for higher stress resistance in plants.
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BFBNIB, NMLJ, NUK, PNG, SAZU, UL, UM, UPUK
To investigate N metabolism of two contrasting Populus species in acclimation to low N availability, saplings of slow-growing species (Populus popularis, Pp) and a fast-growing species (Populus alba ...× Populus glandulosa, Pg) were exposed to 10, 100, or 1000 μM NH4NO3. Despite greater root biomass and fine root surface area in Pp, lower net influxes of NH4 + and NO3 – at the root surface were detected in Pp compared to those in Pg, corresponding well to lower NH4 + and NO3 – content and total N concentration in Pp roots. Meanwhile, higher stable N isotope composition (δ15N) in roots and stronger responsiveness of transcriptional regulation of 18 genes involved in N metabolism were found in roots and leaves of Pp compared to those of Pg. These results indicate that the N metabolism of Pp is more sensitive to decreasing N availability than that of Pg. In both species, low N treatments decreased net influxes of NH4 + and NO3 –, root NH4 + and foliar NO3 – content, root NR activities, total N concentration in roots and leaves, and transcript levels of most ammonium (AMTs) and nitrate (NRTs) transporter genes in leaves and genes involved in N assimilation in roots and leaves. Low N availability increased fine root surface area, foliar starch concentration, δ15N in roots and leaves, and transcript abundance of several AMTs (e.g. AMT1;2) and NRTs (e.g. NRT1;2 and NRT2;4B) in roots of both species. These data indicate that poplar species slow down processes of N acquisition and assimilation in acclimation to limiting N supply.
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