Plant ecophysiology by Peter Petrík

European spruce bark beetle Ips typographus is an important driver of ecological processes in spr... more European spruce bark beetle Ips typographus is an important driver of ecological processes in spruce stands, with severe effects on forestry economies. To prevent bark beetle outbreaks, early detection of infestations is a crucial step in forest management. It is expected that bark beetle infestation modifies biochemical composition of wood and needles, alters physiological responses in the early stage of infestation, which results in the reduction of tree growth and ultimately a tree death. Therefore, we studied the differences in biochemical composition of wood, content of photosynthesis-related pigments, shoot morphology, and growth between the healthy Siberian spruce trees (Picea obovata) and trees which were infested by I. typographus. The study was performed in five experimental plots established in the south of the Udmurt Republic in the European part of the Russian Federation. Three infested and three non-infested trees were chosen on each plot. Our results showed no significant effect of bark beetle infestation on the content of main structural components of wood-holocellulose (cellulose and hemicellulose) and lignin. On the other hand, we found differences in the content of extractive substances in the wood. Specifically, we found a higher tannin content in the non-infested trees. The content of photosynthesis-related pigments differed between the non-infested and infested trees as well. Unexpectedly, bark beetle infestation caused the overproduction of both pigment types rather than their degradation. Moreover, we observed that a higher amount of total extractive substances positively affected the incremental growth, whereas tannins supported the growth of shoots and needles.

The stability of monocultural, even-aged spruce forests at lower altitudes in Central Europe is s... more The stability of monocultural, even-aged spruce forests at lower altitudes in Central Europe is seriously threatened by the prospects of global climate change. The thermostability and water use efficiency of their photosynthetic apparatus might play a vital role in their successful acclimation. In this study, photosystem II (PSII) performance (OJIP transient, rapid light curves) and thermostability were analyzed in Norway spruce (Picea abies (L.) Karst.) throughout the growing season of the exceptionally warm year 2018 (May–September) in the Western Carpathians, Slovakia. These measurements were accompanied by analysis of pigment concentrations in the needles. In addition, gas-exchange temperature curves were produced weekly from June until September to obtain intrinsic water use efficiencies. At the beginning of the growing season, needles exposed to heat stress showed significantly higher basal fluorescence and lower quantum yield, performance index, critical temperature thresholds of PSII inactivation and non-photochemical yield in comparison to other months. The overall thermostability (heat-resistance) of PSII peaked in July and August, reflected in the lowest basal fluorescence and the highest quantum yield of PSII, critical temperature thresholds and yield of non-photochemical quenching under heat stress. Additionally, the ratio between chlorophyll and carotenoids was the highest in August and had a positive impact on PSII thermostability. Moreover, the high-temperature intrinsic water use efficiency was significantly higher during July and August than in June. Results show that 15-year-old trees of Picea abies at 840 m a.s.l. exhibited acclimative seasonal responses of PSII thermostability and intrinsic water use efficiency during an exceptionally warm year. Our results suggest that mountainous P. abies at lower altitudes can acclimate their photosynthetic apparatus to higher temperatures during summer.

Nowadays, a large area of Norway spruce forest stands in Europe is disturbed by windstorm and, su... more Nowadays, a large area of Norway spruce forest stands in Europe is disturbed by windstorm and, subsequently, bark beetle outbreaks. We investigated the state of three disturbed spruce stands along an altitudinal gradient in Tatra National Park (Slovakia) through various physiological processes. Tree-growth characteristics, the mineral nutrition in the needles, and photosynthetic efficiency were assessed. Two techniques of chlorophyll a fluorescence and analyses of assimilatory pigments were used to detect the changes in photosynthesis functioning. Also, the heat sensitivity of photosystem II was tested. Our results showed that these stands are located in nutrient-poor environments. We recorded similar contents of nitrogen, phosphorus, potassium, sodium, zinc, and iron in all stands. Down the vertical transect, the contents of calcium, magnesium, and manganese significantly decreased and the non-essential aluminium increased. Based on stem circumference measurements, water deficit occurred during the vegetation season in all stands, but with the smallest magnitude highest U-stand. We found some photosynthetic constraints: slightly lower chlorophyll contents in all stands were recorded; however, seasonal dynamics with increasing chlorophyll concentration in the highest U-stand were observed. Moreover, the photochemistry of the lowest D-stand was the most negatively influenced by simulated heat, as the photosynthetic performance index, and the density of the active reactions centres significantly decreased and the values of the K-step and basal fluorescence increased. Therefore, we can conclude the different levels of physiological vitality in these naturally damaged spruce stands, with the best physiological performance of the trees in the highest stand.

Recent changes in the floodplain forests of Central Europe, caused mainly by changes in hydrologi... more Recent changes in the floodplain forests of Central Europe, caused mainly by changes in hydrological management and the increased frequency of droughts due to climate change, have led to severe degradation of floodplain ecosystems. Our main objective was to determine the sensitivity of trees to drought by observing the response of the tree phenology, stem radial growth, and physiology (sap flow) of three predominant tree species, namely English oak, narrow-leaved ash, and common hornbeam, to the environmental variables (climate). Stem radial growth began before bud break in ring-porous oak and ash, whereas in diffuse-porous hornbeam, growth onset occurred after leaf formation. The early onset with intense growth during favorable months (April-May) observed in ring-porous species was a major prerequisite for the successful growth of oak and ash at this site. Tree water deficit (TWD), an indicator of stem hydration, was triggered by decreasing soil moisture in all species, and was most prominent in ash, followed by oak. Intriguingly, sap flow was decoupled from TWD in all species and was driven primarily by evaporative demand from the atmosphere. Oak was the least conservative in regulating sap flow under atmospheric drought followed by hornbeam, whereas ash was most restricted and reduced its transpiration during dry periods. In contrast, ash was characterized by the highest radial growth and growthbased water-use efficiency. The lower water storage capacity of oak and ash is likely compensated by deep rooting and drought avoidance strategies, respectively. Tree species that tend to use surface soil water could be severely limited by more extractive species such as hornbeam. Despite the contrasting leaf and wood phenology, stomatal control, and rooting depth among the studied floodplain tree species, they exhibited analogous sap flow and water storage dynamics responses to drier conditions that enabled them to co-exist in the South Moravian Region. Nevertheless, our results suggest that the severe droughts and human-induced alterations in groundwater pose serious threats to floodplain forests in Central Europe, with certain tree species being unable to adapt to these altered conditions.

Partitioning of evapotranspiration (ET) into transpiration (T) and residual evaporation (E) is a ... more Partitioning of evapotranspiration (ET) into transpiration (T) and residual evaporation (E) is a challenging but important task in order to assess the dynamics of increasingly scarce water resources in forest ecosystems. The T/ET ratio has been linked to the ecosystem water use efficiency of temperate forests, and thus is an important index for understanding utilization of water resources under global climate change. We used concurrent sap flow and eddy-covariance measurements to quantify the ET partitioning in pure European beech forest during the 2019-2020 period. The sap flow data were upscaled to stand level T and combined with stand level ET to calculate the T/ET ratio. We analysed intra-annual dynamics, the effect of seasonality and the impact of meteorological conditions on T, ET and T/ET. Annual T/ET of a pure European beech ecosystem was 0.48, falling at the lower end of reported global T/ET values for forest ecosystems. T/ET showed significant seasonal differences throughout spring (T/ET = 0.28), summer (T/ET = 0.62) and autumn (T/ET = 0.35). Air temperature (R 2 = 0.45-0.63), VPD (R 2 = 0.47-0.6) and PAR (R 2 = 0.32-0.63) affected the daily dynamics of T, ET and T/ET; however, soil water content (SWC) had no significant effect. Mature European beech trees showed more anisohydric behaviour and relatively stable T/ET, even under decreasing SWC. The results improve the understanding of ecosystem scale T, ET and T/ET intraannual dynamics and environmental constraints in anisohydric mature European beech.

• The current projections of climate change might exceed the ability of European forest trees to ... more • The current projections of climate change might exceed the ability of European forest trees to adapt to upcoming environmental conditions. However, stomatal and leaf morphological traits could greatly influence the acclimation potential of forest tree species subjected to global warming, including the single most important forestry species in Europe, European beech. • We analysed stomatal (guard cell length, stomatal density and potential conductance index) and leaf (leaf area, leaf dry weight and leaf mass per area) morphological traits of ten provenances from two provenance trials with contrasting climates between 2016 and 2020. The impact of meteorological conditions of the current and preceding year on stomatal and leaf traits was tested by linear and quadratic regressions. Ecodistance was used to capture the impact of adaptation after the transfer of provenances to new environments. • Interactions of trial-provenance and trial-year factors were significant for all measured traits. Guard cell length was lowest and stomatal density was highest across beech provenances in the driest year, 2018. Adaptation was also reflected in a significant relationship between aridity ecodistance and measured traits. Moreover, the meteorological conditions of the preceding year affected the interannual variability of stomatal and leaf traits more than the meteorological conditions of the spring of the current year, suggesting the existence of plant stress memory. • High intraspecific variability of stomatal and leaf traits controlled by the interaction of adaptation, acclimation and plant memory suggests a high acclimation potential of European beech provenances under future conditions of global climate change.

Heat negatively affects photosynthesis, thus tree growth and eventually survivability. The light-... more Heat negatively affects photosynthesis, thus tree growth and eventually survivability. The light-dependent photosynthesis reactions are considered highly heat sensitive; therefore, we studied the impact of supra-optimal temperatures on photosystem II (PSII) in five important tree species (Abies alba, Fagus sylvatica, Picea abies, Pinus sylvestris and Quercus petraea agg.). We simulated short-term heat stress at different levels and assessed PSII thermotolerance via fluorescence parameters derived from the OKJIP transient. To capture the seasonal changes in PSII thermostability and the stress tolerance index, the measurements were performed twice: at the beginning of the vegetation season before the natural heat events occurred and during the peak of the vegetation season. Moreover, the content of photosynthesis-related pigments was analysed. We confirmed the relationship between the appearance of the K-step at the OKJIP curve and the concentrations of photosynthesis-related pigments. We found that broadleaved tree species showed overall higher PSII thermostability and had a higher content of pigments than coniferous species. The most heat-tolerant species was Q. petraea agg., which presented enhanced PSII thermotolerance after heat events. F. sylvatica responded by a decrease in PSII thermostability during the vegetation season, though its thermostability was still higher than that of the coniferous species and was comparable to that of Q. petraea agg. The PSII thermotolerance of P. abies was relatively high compared to that of the other coniferous species on both dates, but its response was inconsistent across the assessed parameters. Contrary, the PSII thermostability of A. alba and P. sylvestris was generally low on both measurement dates.

Climate change-induced elevated temperatures and drought are considered to be serious threats to ... more Climate change-induced elevated temperatures and drought are considered to be serious threats to forest ecosystems worldwide, negatively affecting tree growth and viability. We studied nine European beech (Fagus sylvatica L.) provenances located in two provenance trial plots with contrasting climates in Central Europe. Stomata play a vital role in the water balance of plants by regulating gaseous exchanges between plants and the atmosphere. Therefore, to explain the possible adaptation and acclimation of provenances to climate conditions, stomatal (stomatal density, the length of guard cells, and the potential conductance index) and leaf morphological traits (leaf size, leaf dry weight and specific leaf area) were assessed. The phenotypic plasticity index was calculated from the variability of provenances' stomatal and leaf traits between the provenance plots. We assessed the impact of various climatic characteristics and derived indices (e.g., ecodistance) on intraspecific differences in stomatal and leaf traits. Provenances transferred to drier and warmer conditions acclimated through a decrease in stomatal density, the length of guard cells, potential conductance index, leaf size and leaf dry weight. The reduction in stomatal density and the potential conductance index was proportional to the degree of aridity difference between the climate of origin and conditions of the new site. Moreover, we found that the climate heterogeneity and latitude of the original provenance sites influence the phenotypic plasticity of provenances. Provenances from lower latitudes and less heterogeneous climates showed higher values of phenotypic plasticity. Furthermore, we observed a positive correlation between phenotypic plasticity and mortality in the arid plot but not in the more humid plot. Based on these impacts of the climate on stomatal and leaf traits of transferred provenances, we can improve the predictions of provenance reactions for future scenarios of global climate change.

Understanding of the intraspecific variability in the physiological stress response of trees may ... more Understanding of the intraspecific variability in the physiological stress response of trees may enable to mitigate the impact of climate change on forest ecosystems in the future. We studied the photosynthetic performance of five silver fir (Abies alba Mill.) provenances originating from climatically distinct localities. The study was performed in the trial plot of the silver fir provenance experiment IUFRO 2005 on two dates: in the early summer and in the late summer. Heat waves and a decrease in water availability occurred between the two measurement dates, allowing us to study the response of the provenances to suboptimal growing conditions. The provenances were evaluated at the level of PSII photochemistry and CO 2 assimilation by measuring photosynthesis-related pigment content, chlorophyll a fluorescence, and gas exchange parameters. Significant climatic clines were confirmed: the photosynthetic performance before and after the stress period increased with the increasing altitude and precipitation at the site of origin. In contrast, photosynthetic performance declined with the increasing temperature and Ellenberg's quotient of the origin site. We concluded that provenances originating from high altitudes, corresponding well with more humid and colder conditions in Central Europe, showed the greatest photosynthetic performance and were less responsive to moderate heat and drought. This documents inter-population variation in physiological traits, which needs to be considered in setting rules and recommendations for the transfer of forest reproductive materials.

Silicon is absorbed as uncharged mono-silicic acid by plant roots through passive absorption of L... more Silicon is absorbed as uncharged mono-silicic acid by plant roots through passive absorption of Lsi1, an influx transporter belonging to the aquaporin protein family. Lsi2 then actively effluxes silicon from root cells towards the xylem from where it is exported by Lsi6 for silicon distribution and accumulation to other parts. Recently, it was proposed that silicon nanoparticles (SiNPs) might share a similar route for their uptake and transport. SiNPs then initiate a cascade of morphophysiological adjustments that improve the plant physiology through regulating the expression of many photosynthetic genes and proteins along with photosystem I (PSI) and PSII assemblies. Subsequent improvement in photosynthetic performance and stomatal behaviour correspond to higher growth, development, and productivity. On many occasions, SiNPs have demonstrated a protective role during stressful environments by improving plant-water status, source-sink potential, reactive oxygen species (ROS) metabolism, and enzymatic profile. The present review comprehensively discusses the crop improvement potential of SiNPs stretching their role during optimal and abiotic stress conditions including salinity, drought, temperature, heavy metals, and ultraviolet (UV) radiation. Moreover, in the later section of this review, we offered the understanding that most of these upgrades can be explained by SiNPs intricate correspondence with phytohormones, antioxidants, and signalling molecules. SiNPs can modulate the endogenous phytohormones level such as abscisic acid (ABA), auxins (IAAs), cytokinins (CKs), ethylene (ET), gibberellins (GAs), and jasmonic acid (JA). Altered phytohormones level affects plant growth, development, and productivity at various organ and tissue levels. Similarly, SiNPs regulate the activities of catalase (CAT), ascorbate peroxidase (APX), superoxide dismutase (SOD), and ascorbate-glutathione (AsA-GSH) cycle leading to an upgraded defence system. At the cellular and subcellular levels, SiNPs crosstalk with various signalling molecules such as Ca 2+ , K + , Na + , nitric oxide (NO), ROS, soluble sugars, and transcription factors (TFs) was also explained.

The cultivation of lemongrass (Cymbopogon flexuosus) crop is dominated by its medicinal, food pre... more The cultivation of lemongrass (Cymbopogon flexuosus) crop is dominated by its medicinal, food preservative, and cosmetic demands. The growing economy of the lemongrass market suggests the immense commercial potential of lemongrass and its essential oil. Nevertheless, the continuous increase of the saline regime threatens the growth and productivity of most of the plant life worldwide. In this regard, the present experiment explores the salt sensitiveness of the lemongrass crop against five different levels of salt stress. Metabolomic analyses suggest that lemongrass plants can effectively tolerate a salt concentration of up to 80 mM and retain most of their growth and productivity. However, extreme NaCl concentrations (≥160 mM) inflicted significant (α = 0.05) damage to the plant physiology and exhausted the lemongrass antioxidative defence system. Therefore, the highest NaCl concentration (240 mM) minimised plant height, chlorophyll fluorescence, and essential oil production by up to 50, 27, and 45%. The overall data along with the salt implications on photosynthetic machinery and ROS metabolism suggest that lemongrass can be considered a moderately sensitive crop to salt stress. The study, sensu lato, can be used in reclaiming moderately saline lands with lemongrass cultivation converting such lands from economic liability to economic asset.
Papers by Peter Petrík

AoB PLANTS
The increasing evaporative demand due to climate change will significantly affect the balance of ... more The increasing evaporative demand due to climate change will significantly affect the balance of carbon assimilation and water losses of plants worldwide. The development of crop varieties with improved water-use efficiency (WUE) will be critical for adapting agricultural strategies under predicted future climates. This review aims to summarize the most important leaf morpho-physiological constraints of WUE in C3 plants and identify gaps in knowledge. From the carbon gain side of the WUE, the discussed parameters are mesophyll conductance, carboxylation efficiency and respiratory losses. The traits and parameters affecting the waterside of WUE balance discussed in this review are stomatal size and density, stomatal control and residual water losses (cuticular and bark conductance), nocturnal conductance and leaf hydraulic conductance. In addition, we discussed the impact of leaf anatomy and crown architecture on both the carbon gain and water loss components of WUE. There are multip...
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Plant ecophysiology by Peter Petrík
Papers by Peter Petrík