FIELD NOTE: IRRIGATION OF TREE STANDS WITH GROUNDWATER CONTAINING 1,4-DIOXANE

Coniferous and deciduous tree stands totaling 14 ha were recently planted on a closed landfill, and when mature, the stands are expected to be part of a natural treatment system for recovered groundwater. The trees would be irrigated at the rate of 189 L/min year-round with water containing 1,4-dioxane (< 10 mg/L), a compound that would be taken up and phytovolatilized by the trees. The water is moderately saline and contains elevated levels of manganese. This paper describes a concurrent series of preliminary studies, performed prior to the full-scale planting, to assess the feasibility of the phytoremediation system. Greenhouse experiments were carried out to identify tree species that can take up 1,4-dioxane and are tolerant of the water. Estimates were made of the area of the tree stand necessary to transpire the irrigation water plus precipitation. The landfill matrix was characterized in terms of its percolation rate and water holding capacity and based on those results salinity-modeling studies were carried out to estimate the fate and leaching potential of the various inorganic species that would accumulate in the root-zone of the trees. A pilot study, currently in progress on the landfill, suggested that the landfill cap is a suitable matrix for the establishment of large trees, and that the stands could be irrigated without the production of excess drainage.     

Clonal Variation in Morphology of Populus Root Systems Following Irrigation with Landfill Leachate or Water during 2 Years of Establishment

Increased municipal solid waste generation in North America has prompted the use of Populus for phytoremediation of waste waters including landfill leachate. Populus species and hybrids are ideal for such applications because of their high water usage rates, fast growth, and extensive root systems. Adventitious rooting (i.e., lateral rooting from primordia and basal rooting from callus) of Populus is important for phytotechnologies to ensure successful plantation establishment with genotypes that thrive when irrigated with highly variable or specific contaminants. We evaluated differences in root system morphology following establishment with high-salinity municipal solid waste landfill leachate or uncontaminated well water (control). Populus clones (NC13460, NC14018, NC14104, NC14106, DM115, DN5, NM2, and NM6) were irrigated during 2005 and 2006 in northern Wisconsin, USA and tested for differences in morphology of lateral and basal root types, as well as fine (0–2 mm diameter), small (2–5 mm), and coarse (>5 mm) roots. Across treatments and clones, trees averaged five roots per root type. Leachate-irrigated trees had 87% (lateral) and 105% (basal) as many roots as those irrigated with water. Leachate-irrigated trees had 96% as many fine roots as watering with irrigation water, whereas trees with leachate had 112% (small) and 88% (coarse) as many roots versus water. Despite root necrosis and regrowth in 23% of the trees, leachate irrigation did not negatively affect root diameter or dry mass. Given that adequate rooting is necessary for plantation establishment, leachate and similar waste waters are viable irrigation and fertilization sources of Populus crops used as feedstocks for biofuels, bioenergy, and bioproducts.     

Groundwater Capture Using Hybrid Poplar Trees: Evaluation of a System in Ogden,Utah

A phytoremediation system was installed in 1996 in Ogden, Utah, with the objective of controlling groundwater containing petroleum hydrocarbons. Hybrid poplar trees were deeply and densely planted in rows oriented perpendicular to the direction of groundwater flow, and the stand was never irrigated. Piezometers were installed to measure water table elevation and contaminant levels upgradient, within, and down-gradient of the trees. In 1998, an analysis of the root structure of a representative tree indicated that roots had extended down to the saturated zone, approximately 6 ft below ground surface. The rate of water use by the stand during 1998 was estimated from reference evapotranspiration (ET_o), leaf area, and a water use multiplication factor ($tH) specific to poplar trees. Sap velocity data were collected to measure actual water use by the stand in late summer of 1998. Estimated and measured values compared favorably, with measured values averaging 2.8 gallons day^-1 tree^-1 (1.7 mm day^-1 tree^-1). Water use by the stand in 1999 averaged an estimated 445 gallons day^-1 (6.9 mm day^-1 for the stand). Although the trees transpired a volume of water equivalent to a 10-ft thickness of the saturated zone, water table elevation data collected in 1999 did not indicate a depression in the water table.     

Toxicity Assessment of Volatile Organic Compounds in Poplar Trees

A method is described to assess the toxicity of selected volatile organic compounds (VOCs) in poplar trees. The method is illustrated for a specific aqueous mixture of VOCs that contaminates the groundwater at a site for which phytoremediation was being considered. The VOC mixture contained a variety of aromatic compounds, chlorinated aliphatics, and alcohols. Poplar tree cuttings planted in 50-gallon barrels in the greenhouse were watered via subirrigation. The subirrigant contained either a low (42 mg/L), medium (85 mg/L), or high (169 mg/L) dose of the VOC mixture, or water only (experimental control). Phytotoxicity was evaluated by measuring the physiological parameters of stomatal conductance, shoot elongation, and biomass production. Two experiments are briefly described: (1) The poplar tree cuttings were allowed to become established in the barrels and then treated to gradually increasing concentrations of the VOC mixture until the final dose was reached. The objective was to establish a detailed dose-response relationship. (2) The poplar tree cuttings were given the low, medium, or high dose of the VOC mixture immediately after planting. The aim of this experiment was to determine if the VOC mixture would be inhibitory to root development. Phytotoxic effects were not observed in either experiment.     

Volatile organic compound fate in phytoremediation applications: natural and engineered systems

Unique sampling techniques have generated a new understanding regarding the fate of volatile organic compounds (VOCs) in phytoremediation systems. Tissue sampling and diffusion traps were used to determine how VOCs are transported in and diffuse from vegetation, particularly woody species. These techniques were then utilized to observe how plants interact with different contaminated media, showing transport of contaminants occurs from the vadose zone (vapor phase) as well as the saturated zone (aqueous phase). Data was gathered in laboratory studies, in native vegetation, and in engineered phytoremediation systems. The findings reveal that diffusion from the xylem tissues to the atmosphere is a major fate for VOCs in phytoremediation applications. Linking VOCs' fate with groundwater hydraulics, mass removal rates from contaminant plumes can be estimated. These techniques were also utilized to observe the impact of engineered plant/microbe systems, which utilize recombinant, root-colonizing organisms to selectively degrade compounds and subsequently alter the fate of VOCs and other organic compounds. The genetically enhanced rhizoremediation methods pose a novel approach that may allow for biodegradation of compounds that formerly were considered recalcitrant.     

Competitive uptake and phytomonitoring of chlorinated contaminant mixtures by Redosier dogwood (Cornus sericea)

Plant uptake is an important process in phytoremediation. The robust uptake of volatile organic compounds (VOCs) by plants offers opportunities to establish quantitative relationships between VOCs in plant tissues and in groundwater for the purpose of phytoscreening or phytomonitoring. Most previous research pertaining to phytoremediation neglected the competitive effects of co-contaminants on the uptake of VOCs by plants, yet recent studies appeared to indicate high competitive effects of co-contamination. This study investigated the competitive uptake of three chlorinated compounds in the presence and absence of other co-contaminants by Redosier dogwood in a greenhouse and examined the implications of this competitive phenomenon for phytomonitoring of contaminant mixtures in groundwater. Concentrations of VOCs in stems decreased along the height in both single and bi-solute systems, in agreement with previous observations in the literature. Examination of the VOCs in single and bi-solute systems showed that concentrations of individual compounds are comparable in single and bi-solute systems, yet the ratios of contaminants along the height in bi-solute systems revealed interesting trends. TCE/PCE ratio increased along height while TCE/1,1,2-TCA ratio was roughly constant. The result indicated that sampling point as well as the physicochemical properties of co-contaminants is highly important in phytomonitoring of contaminant mixtures.     

Free and bound fatty acid oxidation products in archaeological ceramic vessels

While oxidation products of unsaturated fatty acids, for example dicarboxylic acids (hereafter diacids), must form during the use of unglazed ceramic vessels for the processing of animal and plant products, such components have never been observed during studies of absorbed lipids. Their absence from the extractable lipid fraction is presumed to be the result of their loss from potsherds through groundwater leaching. Lipid oxidation products including short-chain dicarboxylic acids, ω-hydroxy acids and longer-chain hydroxy and dihydroxy acids have now been observed as components probably covalently bound into solvent insoluble residues of potsherds recovered from waterlogged deposits. These components were only revealed following alkaline treatment of the insoluble residues. A similar mixture of diacids was observed in high abundance in the free lipid fraction of vessels recovered from an exceptionally arid deposit where groundwater leaching would never have occurred. These results confirm the formation of oxidation and probable polymerization products of unsaturated fatty acids during vessel use and burial.     

Mass balance studies of volatile chlorinated hydrocarbon phytoremediation

In field-scale mass balance studies of poplar remediation of carbon tetrachloride (CT), more than 95% of the mass of CT was degraded with all of the CT chlorine accountable as chloride ion accumulation in the soil. Atmospheric loss of CT through leaf transpiration and trunk diffusion was insignificant. These findings are consistent with previously reported uptake and degradation of trichloroethylene (TCE) by poplar. Poplar phytoremediation of CT and TCE results in little decrease in aqueous concentration, since water is taken up at about the same rate as the chlorinated compounds. From this result we predict that phytoirrigation--the application of pumped contaminated groundwater to planted systems--will result in concentrations of the pollutants at the bottom of the root zone that are higher than permitted regulatory levels. Such plantations will be susceptible to loss of contaminants during rainfall events, possibly resulting in pollution of uncontaminated soil. Greenhouse studies of pollutant profiles in the media beneath poplar trees that were surface irrigated with TCE and CT confirmed that regulatory concentrations of these pollutants were not achieved in the root zone of the poplar; rather concentrations fell by less than 50%.     

Nutrient content of throughfall and stemflow in fertilized and irrigatedPinus resinosa ait. Stands

Summary The P, K, Ca, and Mg contents of throughfall and stemflow from K-fertilized and/or irrigated plots from adjacent sites differing in productivity in 39-year-oldPinus resinosa plantations were determined. The amounts of these elements leached from the tree canopies (throughfall plus stemflow) during April through October varied significantly according to site and treatment. These amounts ranged in kg per ha from −0.03 to 0.23 for P, 2.05 to 8.78 for K, 1.55 to 3.63 for Ca, and 0.02 to 0.44 for Mg. Leaching of P, K, and Ca from the trees was usually greater on the more productive site than on the poorer one. In general, the fertilization treatment was correlated with increasing amounts of P, K, and Ca leached, whereas the irrigation treatment was correlated with decreasing amounts of P, K, and Ca leached. For Mg, leaching was consistently greater on the poorer site than on the more productive one; further, it was greatest on the control plots and least on plots both fertilized and irrigated. Gross precipitation of 61.26 cm contained P, K, Ca, and Mg in amounts of 0.33, 0.84, 2.96, and 0.83 kg per ha, respectively. Volumes of both throughfall and stemflow were significantly affected by site conditions. Although throughfall was not affected by treatment, stemflow varied significantly according to plot treatment and was also highly and positively correlated with tree diameter. Stemflow accounted for about 2 per cent of the total water volume collected beneath the canopies, and contained, on a relative basis, considerably less P, an equivalent amount of K, twice as much Mg, and four times as much Ca as throughfall. Contribution of the Northeastern Forest Experiment Station, Forest Service, USDA, and Department of Silviculture and Forest Influences, State University of New York College of Environmental Science and Forestry, Syracuse, New York, 13210.     

我国北方干旱半干旱地区人工造林对地下水位变化影响的模拟研究

水资源短缺已成为阻碍我国社会可持续发展的至关因素,而其中地下水资源短缺更甚。自1952年我国于干旱半干旱地区实施大规模人工造林以来,大量研究资料显示人工林的蒸散量要普遍高于当地自然植被的蒸散量,这可能会打破当地的地下水平衡。而以往我国有关人工造林对地下水位影响的研究较欠缺,因此在基于两种假设条件下,运用7种蒸散发模型测算了人工造林活动对北方干旱半干旱地区的地下水位影响。结果表明大规模的人工造林活动会降低地下水位高度,基于假设条件一,甘肃、宁夏和新疆地下水位下降严重;基于假设条件二,北京地下水位下降明显。提出我国在未来生态修复实践中要考虑对地下水供给的影响,确定合理造林规模并选择栽培乡土树种,真正提高地下水资源利用率。     

Phytoremediation of groundwater contaminated with pesticides using short-rotation willow crops: A case study of an apple orchard

The occurrence of pesticides in groundwater represents an important health issue, notably for population whose drinking water supply source is located in agricultural areas. However, few solutions have been considered with regard to this issue. We tested the efficacy of a vegetal filtering system made of shrub willows planted at a high density (16,000 plants ha^?1) to filter or degrade pesticides found in the groundwater flowing out of an apple orchard. Ethylene urea (EU), ethylene thiourea (ETU), tetrahydrophthalimide (THPI), atrazine, and desethylatrazine were monitored in the soil solution in willow and control plots over one growing season. ETU and atrazine concentrations were lower in the willow plots relative to the control plots, whereas desethylatrazine concentration was higher in the willow plots. No significant difference was detected for EU and THPI. Furthermore, pesticide concentrations displayed complex temporal patterns. These results suggest that willow filter systems can filter or degrade pesticides, notably ETU and atrazine, and could be used for phytoremediation purposes. Yet, this potential remains to be quantified with further studies using experimental settings allowing more estimation in time and space.     

PHYTOREMEDIATION OF BTEX HYDROCARBONS: POTENTIAL IMPACTS OF DIURNAL GROUNDWATER FLUCTUATION ON MICROBIAL DEGRADATION

Volatile hydrocarbons have multiple potential fates in phytoremediation. This research investigated the relationship between biodegradation and plant uptake of BTEX compounds in laboratory and field settings. At a phytoremediation site, preliminary studies revealed minimal uptake into trees and enhanced degradation potential in the rhizosphere and in the bulk soil. Increased oxygen transport to the vadose zone caused by diurnal rise and fall of the water table was hypothesized to enhance degradation in the bulk soil. A detailed greenhouse study was then conducted to investigate potential bioremediation impacts using field-site soil and DN34 hybrid poplar trees.

In rhizosphere soils, the contaminated-planted reactor had significantly higher BTEX degrader populations versus the uncontaminated-planted reactor, as was anticipated. The bulk soil in the planted-contaminated reactor had increased degrader populations than the unplanted-contaminated soil or planted-uncontaminated soil, and planting increased degradation throughout the soil profile, not just in the limited volume of rhizosphere soils. Oxygen diffusive and advective transport into reactors was modeled and calculated. Oxygen input in planted reactors was at least 3 to 5 times higher than in unplanted reactors, and increasing oxygen input lead to increased degrader populations in a linear manner. These results combined with the knowledge that high-transpiration trees draw the contaminated groundwater to the capillary fringe and the rhizosphere indicate that phytoremediation can aid microbial degradation via multiple mechanisms: increasing degrader populations, increasing oxygen input via groundwater diurnal fluctuations, and transporting contaminants to the biologically-enriched soil profile.     

Predicting the Effect of Deep-Rooted Hybrid Poplars on the Groundwater Flow System at a Large-Scale Phytoremediation Site

Estimating the effect of phreatophytes on the groundwater flow field is critical in the design or evaluation of a phytoremediation system. Complex hydrogeological conditions and the transient water use rates of trees require the application of numerical modeling to address such issues as hydraulic containment, seasonality, and system design.

In 1999, 809 hybrid poplars and willows were planted to phytoremediate the 317 and 319 Areas of Argonne National Laboratory near Chicago, Illinois. Contaminants of concern are volatile organic compounds and tritium. The site hydrogeology is a complex framework of glacial tills interlaced with sands, gravels, and silts of varying character, thickness, and lateral extent. A total of 420 poplars were installed using a technology to direct the roots through a 25--ft (8--m)-thick till to a contaminated aquifer.

Numerical modeling was used to simulate the effect of the deep-rooted poplars on this aquifer of concern. Initially, the best estimates of input parameters and boundary conditions were determined to provide a suitable match to historical transient ground-water flow conditions. The model was applied to calculate the future effect of the developing deep-rooted poplars over a 6 year period. The first 3 years represent the development period of the trees. In the fourth year, canopy closure is expected to occur; modeling continues through the first 3 years of the mature plantation. Monthly estimates of water use by the trees are incorporated. The modeling suggested that the mature trees in the plantation design will provide a large degree of containment of groundwater from the upgradient source areas, despite the seasonal nature of the trees' water consumption. The results indicate the likely areas where seasonal dewatering of the aquifer may limit the availability of water for the trees. The modeling also provided estimates of the residence time of groundwater in the geochemically altered rhizosphere of the plantation.     

Evaluation of Populus and Salix continuously irrigated with landfill leachate II. soils and early tree development

Soil contaminant levels and early tree growth data are helpful for assessing phytoremediation systems. Populus (DN17, DN182, DN34, NM2, and NM6) and Salix (94003, 94012, S287, S566, and SX61) genotypes were irrigated with landfill leachate or municipal water and tested for differences in (1) element concentrations (P, K, Ca, Mg, S, Zn, B, Mn, Fe, Cu, Al, and Na) of a topsoil layer and a layer of sand in tanks with a cover crop of trees or no trees and (2) height, diameter, volume, and dry mass of leaves, stems, and roots. Trees were irrigated with leachate or water during the final 12 wk of the 18-wk study. Differences in most soil element concentrations were negligible (P > 0.05) for irrigation treatments and cover main effects. Phosphorous, K, Mg, S, Zn, Mn, Fe, and Al concentrations were greater in topsoil than sand (P = 0.0011 for Mg; P < 0.0001 for others). There was broad variation between genera and among clones for all growth traits. The treatment x clone interaction governed height, volume, and root dry mass, with (94012, SX61), (NM2, S566, SX61), and (S287, S566) exhibiting the greatest levels, respectively,following leachate application. Given the broad amount of variability among and within these genera, there is great potential for the identification and selection of specific genotypes with a combination of elevated phytoremediation capabilities and tree yield. From a practical standpoint, these results may be used as a baseline for the development of future remediation systems.     

Monitoring the Effect of Poplar Trees on Petroleum-Hydrocarbon and Chlorinated-Solvent Contaminated Ground Water

At contaminated groundwater sites, poplar trees can be used to affect ground-water levels, flow directions, and ultimately total groundwater and contaminant flux to areas downgradient of the trees. The magnitude of the hydrologic changes can be monitored using fundamental concepts of groundwater hydrology, in addition to plant physiology-based approaches, and can be viewed as being almost independent of the contaminant released. The affect of poplar trees on the fate of groundwater contaminants, however, is contaminant dependent. Some petroleum hydrocarbons or chlorinated solvents may be mineralized or transformed to innocuous compounds by rhizospheric bacteria associated with the tree roots, mineralized or transformed by plant tissues in the transpiration stream or leaves after uptake, or passively volatilized and rapidly dispersed or oxidized in the atmosphere. These processes also can be monitored using a combination of physiological- or geochemical-based field or laboratory approaches. When combined, such hydrologic and contaminant monitoring approaches can result in a more accurate assessment of the use of poplar trees to meet regulatory goals at contaminated groundwater sites, verify that these goals continue to be met in the future, and ultimately lead to a consensus on how the performance of plant-based remedial strategies (phytoremediation) is to be assessed.     

汽油添加剂甲基叔丁基醚(MTBE)污染的植物修复

甲基叔丁基醚(MTBE)是北美燃料市场最常用的汽油添加剂。由于其在土壤中的不吸附性和极高的水溶性,MTBE已成为一种蔓延性的地下水污染物。本实验用一自行设计的植物反应器来观察和测定在不同温度条件下柳树(Salix alba)对MTBE污染水溶液的修复潜力。长出新根须和嫩叶的柳树枝条在一容积500mL的锥型瓶中生长12d(其中MTBE溶液450mL)来观察MTBE对柳树生长的影响,同时测定柳树对MTBE的吸收和降解。MTBE及其主要降解产物叔丁基醇(TBA)用气相色谱来检测。本实验结果表明在为期12d的时间内,水溶液中24．84～53．27％MTBE可以通过柳树的蒸腾作用去除。同时在15℃～25℃的温度范围内,：MTBE的去除率(％)和柳树的蒸腾量(g)之间存在着明显的线性关系。由于没有发现TBA和其它可能的降解产物,植物挥发是MTBE植物修复技术中主要的作用机理。尽管植物在MTBE污染的修复过程中只是起着中间传输媒介的作用,大量的MTBE通过植物的蒸腾作用以气态的形式释放到大气中,但由于MTBE在气态下的光氧化非常快,MTBE不会造成空气污染。我们认为植物修复技术对MTBE污染的土壤和地下水仍不失为一个有效的修复手段。     

Long-term changes in above ground biomass after disturbance in a neotropical dry forest,Hellshire Hills,Jamaica

We used data from experimental plots (control, partially cut and clear-cut) established in 1998, in a tropical dry forest (TDF) in Jamaica, to assess changes in above ground biomass (AGB) 10 years after disturbance. The treatments reduced AGB significantly in 1999 (partially cut: 37.6 %, clear-cut: 94.4 %) and after 10 years, AGB did not recover overall, nor did it recover in the clear-cut plots. Partially cut plots, however, recovered the lost AGB in 10 years via growth of uncut trees, which contributed significantly to biomass recovery, with only minor contributions from recruited trees and coppice shoots. For clear-cut plots, coppice shoots contributed significantly to the recovered AGB when compared with recruited biomass. Together, they recovered 26 % of AGB lost, despite recovering 78 % of the density and height of the control plots. The probability of survivorship decreased for trees with higher pre-treatment AGB values, but was higher for trees with multiple stems in 1998, regardless of treatment. The magnitude of biomass reduction varied among the species assessed and this had a differential effect on their ability to recover AGB. We estimate that it will take approximately 45.4 years for the clear-cut plots to recover pre-treatment AGB; this is significantly longer than AGB recovery time for some successional rainforests on abandoned pastures/farmland. Consequently, this TDF may not be as resilient as tropical rainforests.     

Phytoremediation removal rates of benzene,toluene, and chlorobenzene

Phytoremediation is a sustainable remedial approach, although performance efficacy is rarely reported. In this study, we assessed a phytoremediation plot treating benzene, toluene, and chlorobenzene. A comparison of the calculated phytoremediation removal rate with estimates of onsite contaminant mass was used to forecast cleanup periods. The investigation demonstrated that substantial microbial degradation was occurring in the subsurface. Estimates of transpiration indicated that the trees planted were removing approximately 240,000 L of water per year. This large quantity of water removal implies substantial removal of contaminant due to large amounts of contaminants in the groundwater; however, these contaminants extensively sorb to the soil, resulting in large quantities of contaminant mass in the subsurface. The total estimate of subsurface contaminant mass was also complicated by the presence of non-aqueous phase liquids (NAPL), additional contaminant masses that were difficult to quantify. These uncertainties of initial contaminant mass at the site result in large uncertainty in the cleanup period, although mean estimates are on the order of decades. Collectively, the model indicates contaminant removal rates on the order of 10^?2–10⁰ kg/tree/year. The benefit of the phytoremediation system is relatively sustainable cleanup over the long periods necessary due to the presence of NAPL.     

Root distribution following spatial separation of water and nitrogen supply in furrow irrigated corn

Proper management of water and fertilizer placement in irrigated corn (Zea mays L.) has the potential to reduce nitrate leaching into the groundwater. Potential management practices tested in a two year field experiment included row or furrow fertilizer placement combined with every or alternate furrow irrigation. To understand how fertilizer availability to plants could be affected by these management practices, root growth and distribution in a Ulm clay loam soil were examined. Spring rains were greater than normal in both years providing adequate moisture for early root growth in both irrigated and non-irrigated furrows. As the non-irrigated furrow began to dry, root biomass increased as much as 126% compared with the irrigated furrow. The greatest increase was at lower depths, however, where moisture was still plentiful. When early season moisture was available, roots proliferated throughout the soil profile and quickly became available to take up fertilizer N in both irrigated and non-irrigated furrows. Root growth responded positively to fertilizer placement in the furrow in 1996 but not in 1995. Excessive N leaching in 1995 may have limited the response to fertilizer N.     

Memory of environmental conditions across generations affects the acclimation potential of scots pine

Long generation times have been suggested to hamper rapid genetic adaptation of organisms to changing environmental conditions. We examined if environmental memory of the parental Scots pines (Pinus sylvestris L.) drive offspring survival and growth. We used seeds from trees growing under naturally dry conditions (control), irrigated trees (irrigated from 2003 to 2016), and formerly irrigated trees (“irrigation stop”; irrigated from 2003–2013; control condition since 2014). We performed two experiments, one under controlled greenhouse conditions and one at the experimental field site. In the greenhouse, the offspring from control trees exposed regularly to drought were more tolerant to hot–drought conditions than the offspring from irrigated trees and showed lower mortality even though there was no genetic difference. However, under optimal conditions (high water supply and full sunlight), these offspring showed lower growth and were outperformed by the offspring of the irrigated trees. This different offspring growth, with the offspring of the “irrigation-stop” trees showing intermediate responses, points to the important role of transgenerational memory for the long-term acclimation of trees. Such memory effects, however, may be overridden by climatic extremes during germination and early growth stages such as the European 2018 mega-drought that impacted our field experiment.