Genetics and Genomics Commons^™

Genetic Markers And Tree Properties Predicting Wood Biorefining Potential In Aspen (Populus Tremula) Bioenergy Feedstock, Sacha Escamez, Kathryn M. Robinson, Mikko Luomaranta, Madhavi Latha Gandla, Niklas Mähler, Zakiya Yassin, Thomas Grahn, Gerhard Scheepers, Lars-Göran Stener, Stefan Jansson, Leif J. Jönsson, Nathaniel R. Street, Hannele Tuominen

Aspen Bibliography

Background Wood represents the majority of the biomass on land and constitutes a renewable source of biofuels and other bioproducts. However, wood is recalcitrant to bioconversion, raising a need for feedstock improvement in production of, for instance, biofuels. We investigated the properties of wood that affect bioconversion, as well as the underlying genetics, to help identify superior tree feedstocks for biorefining.

Results We recorded 65 wood-related and growth traits in a population of 113 natural aspen genotypes from Sweden (https://doi.org/10.5061/dryad.gtht76hrd). These traits included three growth and field performance traits, 20 traits for wood chemical composition, 17 traits for wood anatomy …

Less Fuel For The Next Fire? Short-Interval Fire Delays Forest Recovery And Interacting Drivers Amplify Effects, Kristin H. Braziunas, Nathan G. Kiel, Monica G. Turner

Aspen Bibliography

As 21st-century climate and disturbance dynamics depart from historic baselines, ecosystem resilience is uncertain. Multiple drivers are changing simultaneously, and interactions among drivers could amplify ecosystem vulnerability to change. Subalpine forests in Greater Yellowstone (Northern Rocky Mountains, USA) were historically resilient to infrequent (100–300 year), severe fire. We sampled paired short-interval (<30-year) and long-interval (>125-year) post-fire plots most recently burned between 1988 and 2018 to address two questions: (1) How do short-interval fire, climate, topography, and distance to unburned live forest edge interact to affect post-fire forest regeneration? (2) How do forest biomass and fuels vary following short-interval versus long-interval severe fires? …

Regeneration Strategies And Forest Resilience To Changing Fire Regimes: Insights From A Goldilocks Model, Tanjona Ramiadantsoa, Zak Ratajczak, Monica G. Turner

Aspen Bibliography

Disturbances are ubiquitous in ecological systems, and species have evolved a range of strategies to resist or rebound following disturbance. Understanding how the presence and complementarity of regeneration traits will affect community responses to disturbance is increasingly urgent as disturbance regimes shift beyond their historical ranges of variability. We define "disturbance niche" as a species' fitness across a range of disturbance sizes and frequencies that can reflect the fundamental or realized niche, that is, whether the species occurs alone or with other species. We developed a model of intermediate complexity (i.e., a Goldilocks model) to infer the disturbance niche. We …

Revisiting Trophic Cascades And Aspen Recovery In Northern Yellowstone, Robert L. Beschta, Luke E. Painter, William J. Ripple

Aspen Bibliography

We revisit the nature and extent of trophic cascades and quaking aspen (Populus tremuloides) recovery in the northern range of Yellowstone National Park (YNP), where studies have reported on Rocky Mountain elk (Cervus canadensis) browsing and young aspen heights following the St. John, 1995-96 reintroduction of gray wolves (Canis lupus). A recent study by Brice et al. (2021) expressed concerns about methodologies employed in earlier aspen studies and that results from those studies exaggerated the extent to which a trophic cascade has benefitted aspen, concerns such as: (a) the selection of aspen stands, (b) …

Bison Influences On Composition And Diversity Of Riparian Plant Communities In Yellowstone National Park, J. Boone Kauffman, Dian Lyn Cummings, Cimarron Kauffman, Robert L. Beschta, Jeremy Brooks, Keeley Macneill, William J. Ripple

Aspen Bibliography

Riparian zones are among the most biologically diverse ecosystems in the Intermountain West, USA, and provide valuable ecosystem services, including high rates of biotic productivity, nutrient processing, and carbon storage. Thus, their sustainability is a high priority for land managers. Large ungulates affect composition and structure of riparian/stream ecosystems through herbivory and physical effects, via trailing and trampling. Bison (Bison bison) in Yellowstone National Park (YNP) have been characterized as "ecosystem engineers" because of their demonstrated effects on phenology, aboveground productivity of grasses, and woody vegetation structure. Bison have greatly increased in numbers during the last two decades …

Assessment Of Carbon Productivity Dynamics In Aspen Stands Under Climate Change Based On Forest Inventories In Central Siberia, Andrey Andreevich Vais, Valentina Valerievna Popova, Alina Andreevna Andronova, Viktor Nikolaevich Nemich, Artem Gennadievich Nepovinnykh, Pavel Vladimirovich Mikhaylov

Aspen Bibliography

The aim of the present research was to study the dynamics of growth and conditions of aspen stands under climate change, according to different periods of forest inventory. The study was conducted in modal aspen forests growing in the subtaiga/forest steppe region of Central Siberia. Aspen forests grow intensively at young age, which allows them to realize maximum carbon sequestration potential. The research was based on forest inventory data from 1972, 1982, 2002, and 2021 (the study was conducted on a limited territory). There was a steady increase in temperatures in the growing season from 1982 to 2002. The amount …

Big Meadows Mountain Meadow & Aspen Restoration Project 2017 – 2023 Summary Report, Scott River Watershed Council

Aspen Bibliography

Big Meadows Mountain Meadow and Aspen Restoration Project is designed to enhance aspen and mountain meadow habitats at Big Meadows, a property that is privately owned by Ecotrust Forest Management (EFM). EFM owns approximately 40,000 acres in the Klamath Mountains of Siskiyou County, known as the Scott River Headwaters Property. The Scott River Headwaters Property Management Plan, developed by EFM, specifically identifies and prioritizes the restoration of aspen woodlands. As such, EFM and Scott River Watershed Council (SRWC) partnered with the goal of improving these critical habitats. The Big Meadows complex is one of the largest mountain meadow systems within …

Tree Biomass – A Fragile Carbon Storage In Old-Growth Birch And Aspen Stands In Hemiboreal Latvia, Laura Ķēniņa, Didzis Elferts, Ieva Jaunslaviete, Endijs Bāders, Guntars Šņepsts, Āris Jansons

Aspen Bibliography

Birch (Betula pendula Roth, Betula pubescens Ehrh.) and European aspen (Populus tremula L.) stands dominate the deciduous forests of Northern Europe. Due to increasing forest protections, more deciduous stands will reach the old-growth stage. Thus, data on the carbon storage potential in such areas are essential. We aimed to establish a benchmark for carbon stocks of the main carbon pools in old-growth deciduous hemiboreal stands. Carbon pools were calculated from measurements in forty old-growth (104–148 years-old) deciduous stands in forests on fertile mineral soil. The carbon stock in these stands is distributed across tree biomass (~ 60%), mineral soil (~ …

Aspen Leaves As A "Chemical Landscape" For Fungal Endophyte Diversity - Can Nitrogen And Herbivory Shape The Community Composition In Controlled Conditions?, Johanna Witzell, Vicki Huizu Guo Decker, Marta Agostinelli, Carmen Romeralo Tapia, Michelle Cleary, Benedicte Riber Albrectsen

Aspen Bibliography

The endophytic microbiome may influence the ecological performance of plants, including forest trees. Various abiotic and biotic factors may shape the endophyte communities directly but also indirectly, by modifying the quality of host plants as a substrate. We hypothesized that potentially antifungal or fungistatic condensed tannins (CTs) would determine the quality of aspen (Populus tremula) leaves as a substrate for endophytic fungi. By subjecting the plants to nitrogen fertilization (N) or herbivory (H; leaf beetles) we aimed to change the internal “chemical landscape”, especially the CT levels, in aspen leaves. We expected that this would lead to changes in the …

Ungulate Herbivores As Drivers Of Aspen Recruitment And Understory Composition Throughout Arid Montane Landscapes, Elizabeth S. Reikowski, Tyler Refsland, J. Hall Cushman

Aspen Bibliography

Herbivory by wild and domestic ungulates can influence tree recruitment and understory forest communities throughout the world. Herbivore-driven declines in tree recruitment have been observed for quaking aspen (Populus tremuloides), a foundation species whose health and management is recognized as a critical priority throughout much of its range. Livestock fencing is commonly used to promote aspen regeneration, but its effectiveness is rarely assessed, especially across large spatial scales. Using a livestock-reduction experiment, we evaluated the effects of ungulate herbivory on aspen in the Great Basin and southern Cascades, an expansive and environmentally heterogeneous region where aspen faces the …

Coping With Environmental Constraints: Geographically Divergent Adaptive Evolution And Germination Plasticity In The Transcontinental Populus Tremuloides, Roos Goessen, Nathalie Isabel, Christian Wehenkel, Nathalie Pavy, Lisa Tischenko, Lyne Touchette, Isabelle Giguère, Marie-Claude Gros-Louis, Jérôme Laroche, Brian Boyle, Raju Soolanayakanahally, Karen E. Mock, Javier Hernández-Velasco, Sergio Leonel Simental-Rodriguez, Jean Bousquet, Ilga M. Porth

Aspen Bibliography

Societal Impact Statement

Syntheses clearly show that global warming is affecting ecosystems and biodiversity around the world. New methods and measures are needed to predict the climate resilience of plant species critical to ecosystem stability, to improve ecological management and to support habitat restoration and human well-being. Widespread keystone species such as aspen are important targets in the study of resilience to future climate conditions because they play a crucial role in maintaining various ecosystem functions and may contain genetic material with untapped adaptive potential. Here, we present a new framework in support of climate-resilient revegetation based on comprehensively understood …

Growing Space Management In Boreal Mixedwood Forests: 22 Year Results, Richard Kabzems, George Harper, Che Elkin

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Boreal mixed forests of trembling aspen (Populus tremuloides) and white spruce (Picea glauca) can provide higher outputs of many ecosystem goods and services, combined with resilience for changing environmental conditions. In this study, we examine the growth of white spruce and aspen over a range of stand compositions created by spot and broadcast treatments of broadleaves using manual and chemical means, aspen spacing, and untreated control. Twenty-two growing seasons postharvest, treatment responses created a range of broadleaf densities and spatial arrangements, reflected in significant differences in heights and diameters of the spruce mixedwood component. At this …

Recovery Of Trembling Aspen, Tamarack, And White Spruce Seedlings From Nacl Stress Following Winter Dormancy: Implications For Increased Foliar Potassium, Necrosis, And Sodium Management As Stress Resistance Mechanisms, Nathan T. Lauer

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Revegetation of boreal forest lands disturbed by surface mining for bitumen can be challenging due to fluctuating levels of soil NaCl and harsh winter temperatures. These stressors may hinder the growth and survival of planted tree seedlings. Two experiments were carried out to examine the processes of recovery from NaCl stress and overwintering in trembling aspen, tamarack, and white spruce seedlings. In the recovery experiment, seedlings were treated with 0, 50, or 100 mM NaCl for 60 days and then allowed to recover for 60 days. Most of the examined physiological variables (total dry weight, chlorophyll concentration, photosynthesis, and transpiration) …

Compensatory Phenolic Induction Dynamics In Aspen After Aphid Infestation, Rajarshi Kumar Gaur, Ilka Nacif De Abreu, Benedicte Riber Albrectsen

Aspen Bibliography

Condensed tannins (CTs) are polyphenolics and part of the total phenolic (TP) pool that shape resistance in aspen (Populus tremula). CTs are negatively associated with pathogens, but their resistance properties against herbivores are less understood. CTs shape resistance to pathogens and chewing herbivores and could also shape resistance to aphids. Being chemical pools that are highly variable it can further be questioned whether CT-shaped resistance is better described by constitutive levels, by the induced response potential, or by both. Here, aspen genotypes were propagated and selected to represent a range of inherent abilities to produce and store foliar …

Young Forests And Fire: Using Lidar–Imagery Fusion To Explore Fuels And Burn Severity In A Subalpine Forest Reburn, Kristin H. Braziunas, Diane C. Abendroth, Monica G. Turner

Aspen Bibliography

Anticipating fire behavior as climate change and fire activity accelerate is an increasingly pressing management challenge in fire-prone landscapes. In subalpine forests adapted to infrequent, stand-replacing fire, self-limitation of burn severity in short-interval fire is incompletely understood. Spatially explicit fuels data can support assessments of landscape-scale fire risk and fuel feedbacks on burn severity. For a ~1450-km² largely forested landscape in the US Northern Rocky Mountains, we used airborne lidar and imagery to predict and map canopy and surface fuels. In a fire that burned mature (>125-year-old) and also reburned young (~30-year-old) subalpine forest, we then asked: (1) …

Oystershell Scale: An Invasive Threat To Aspen Conservation, Amanda M. Grady, Connor D. Crouch, Nicholas P. Wilhelmi, Richard W. Hofstetter, Kristen M. Waring

Aspen Bibliography

Aspen decline is an acute and chronic problem in Arizona, where high levels of overstory mortality and a lack of recruitment continue to be observed. Oystershell scale (Lepidosaphes ulmi; OSS), an invasive sapsucking insect, has recently become widespread in native aspen stands in the southwestern U.S., further contributing to aspen mortality. Damage is severe in lower elevation stands and within ungulate exclosures created to conserve aspen. Young recruiting aspen that are rare on the landscape incur high levels of OSS-caused mortality when infested (Fig. 1). OSS has only recently become a pest of concern in the Southwest and …

Emulating Succession Of Boreal Mixedwood Forests In Alberta Using Understory Protection Harvesting, Ivan Bjelnanovic, Philip G. Comeau, Sharon Meredith, Brian Roth

Aspen Bibliography

Understory protection harvesting is a form of partial cutting that can be used in aspen (Populus tremuloides Michx.)-dominated stands that have understories of white spruce (Picea glauca (Moench) Voss). This practice involves removing 75% to 85% of the merchantable aspen while minimizing damage to the advance spruce regeneration, in addition to leaving 15% to 25% of the aspen standing to reduce potential windthrow of the spruce understory. In this paper, we summarize results from 18 stands measured 10 to 12 years after understory protection harvest. Diameter growth of spruce increased during the first five years after harvest while …

Climate Effects On Black Spruce And Trembling Aspen Productivity In Natural Origin Mixed Stands, Mahadev Sharma

Aspen Bibliography

Forest managers need site productivity estimates for tree species growing in mixed stands. Models developed in the past are generally for pure stands and don’t factor in the effects of climate change on site productivity. Therefore, site index (SI) models were developed for black spruce (Picea mariana Mill. B.S.P.) and trembling aspen (Populus tremuloides Michx.) trees grown in natural origin mixed stands. For this, 186 trees (93 black spruce and trembling aspen each) were sampled from 31 even-aged natural mixed stands (sites) (3 trees/species/site) across Ontario, Canada. Stand height growth models were developed by incorporating climate variables during …

Facilitation Differentially Affects Competitive Responses Of Aspen And Subalpine Fir Through Stages Of Stand Development, Rebecca Lee Molinari, Matthew F. Bekker, Benjamin D. St. Clair, Jason Bartholomew, R. Justin Derose, Stanley G. Kitchen, Samuel B. St. Clair

Aspen Bibliography

Spatial interactions between trees influence forest community succession. The objective of this study was to investigate how shifts in forest composition and proximity between tree species affect stand development over time in mixed forest systems. At six locations across the Fishlake National Forest, Utah, USA, in stands where facilitation has been documented previously, tree-ring samples were collected from aspen and subalpine fir trees. Basal area increment was calculated to characterize the effects of the proximity of overstory trees on multidecadal growth responses of aspen and subalpine fir in aspen-dominant and mixed aspen–conifer stands. Subalpine fir seedlings were established next to …

Bottom-Up Drivers Of Future Fire Regimes In Western Boreal North America, Adrianna C. Foster, Jacquelyn K. Shuman, Brendan M. Rogers, Xanthe J. Walker, Michelle C. Mack, Laura L. Bourgeau-Chavez, Sander Veraverbeke, Scott J. Goetz

Aspen Bibliography

Forest characteristics, structure, and dynamics within the North American boreal region are heavily influenced by wildfire intensity, severity, and frequency. Increasing temperatures are likely to result in drier conditions and longer fire seasons, potentially leading to more intense and frequent fires. However, an increase in deciduous forest cover is also predicted across the region, potentially decreasing flammability. In this study, we use an individual tree-based forest model to test bottom-up (i.e. fuels) vs top-down (i.e. climate) controls on fire activity and project future forest and wildfire dynamics. The University of Virginia Forest Model Enhanced is an individual tree-based forest model …

Identifying Conifer Tree Vs. Deciduous Shrub And Tree Regeneration Trajectories In A Space-For-Time Boreal Peatland Fire Chronosequence Using Multispectral Lidar, Humaira Enayetullah, Laura Chasmer, Christopher Hopkinson, Dan Thompson, Danielle Cobbaert

Aspen Bibliography

Wildland fires and anthropogenic disturbances can cause changes in vegetation species composition and structure in boreal peatlands. These could potentially alter regeneration trajectories following severe fire or through cumulative impacts of climate-mediated drying, fire, and/or anthropogenic disturbance. We used lidar-derived point cloud metrics, and site-specific locational attributes to assess trajectories of post-disturbance vegetation regeneration in boreal peatlands south of Fort McMurray, Alberta, Canada using a space-for-time-chronosequence. The objectives were to (a) develop methods to identify conifer trees vs. deciduous shrubs and trees using multi-spectral lidar data, (b) quantify the proportional coverage of shrubs and trees to determine environmental conditions driving …

Effects Of Buried Wood On The Development Of Populus Tremuloides On Various Oil Sands Reclamation Soils, Kaitlyn E. Trepanier, Laura Manchola-Rojas, Bradley D. Pinno

Aspen Bibliography

Buried wood is an important but understudied component of reclamation soils. We examined the impacts of buried wood amounts and species on the growth of the common reclamation tree species trembling aspen (Populus tremuloides). In a greenhouse study, aspen seedlings were planted into four soil types, upland derived fine forest floor-mineral mix (fFFMM), coarse forest floor-mineral mix (cFFMM), and lowland derived peat and peat-mineral mix (PMM), that were mixed with either aspen or pine wood shavings at four concentrations (0%, 10%, 20% and 50% of total volume). Height and diameter growth, chlorophyll concentration, and leaf and stem biomass were measured. …

Sampling Bias Exaggerates A Textbook Example Of A Trophic Cascade, Elaine M. Brice, Eric J. Larsen, Daniel R. Macnulty

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Understanding trophic cascades in terrestrial wildlife communities is a major challenge because these systems are difficult to sample properly. We show how a tradition of non-random sampling has confounded this understanding in a textbook system (Yellowstone National Park) where carnivore [Canis lupus (wolf)] recovery is associated with a trophic cascade involving changes in herbivore [Cervus canadensis (elk)] behaviour and density that promote plant regeneration. Long-term data indicate a practice of sampling only the tallest young plants overestimated regeneration of overstory aspen (Populus tremuloides) by a factor of 4–7 compared to random sampling because it favoured plants taller than the preferred …

Growth–Defense Trade-Offs Shape Population Genetic Composition In An Iconic Forest Tree Species, Olivia L. Cope, Ken Keefover-Ring, Eric L. Kruger, Richard L. Lindroth

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All organisms experience fundamental conflicts between divergent metabolic processes. In plants, a pivotal conflict occurs between allocation to growth, which accelerates resource acquisition, and to defense, which protects existing tissue against herbivory. Trade-offs between growth and defense traits are not universally observed, and a central prediction of plant evolutionary ecology is that context-dependence of these trade-offs contributes to the maintenance of intraspecific variation in defense [Züst and Agrawal, Annu. Rev. Plant Biol., 68, 513–534 (2017)]. This prediction has rarely been tested, however, and the evolutionary consequences of growth–defense trade-offs in different environments are poorly understood, especially in long-lived species …

Determining The Novel Pathogen Neodothiora Populina As The Causal Agent Of The Aspen Running Canker Disease In Alaska, Loretta M. Winton, Gerard C. Adams, Roger W. Ruess

Aspen Bibliography

Neodothiora populina Crous, G.C. Adams & Winton was determined to be a new pathogen of trembling aspen (Populus tremuloides) growing in Alaska, based on completion of Koch’s Postulates in replicated forest and growth chamber inoculation trials. The pathogen is responsible for severe damage and widespread rapid mortality of sapling to mature aspen (≥ 80 years) in the boreal forests of interior Alaska, due to large diffuse annual (1–2 years) cankers. Isolation of the pathogen was challenging, and identification based on cultural characters was difficult. Fruiting bodies were not found on wild diseased trees, but erumpent pycnidia were found …

Availability And Structure Of Coarse Woody Debris In Hemiboreal Mature To Old-Growth Aspen Stands And Its Implications For Forest Carbon Pool, Silva Šēnhofa, Guntars Šnepsts, Kārlis Bičkovskis, Ieva Jaunslaviete, Līga Liepa, Inga Straupe, Āris Jansons

Aspen Bibliography

European aspen deadwood is extensively studied as a habitat for saproxylic species, while less is known of its dynamics and role in carbon sequestration. We studied unmanaged mature (41–60 years), moderately overmature (61–80 years), overmature (81–100 years), and old-growth (101–140 years) and managed mature and moderately overmature aspen stands on fertile mineral soils. In unmanaged stands, marginal mean CWD volume was from 67.3 ± 12.1 m³ ha⁻¹ in moderately overmature to 92.4 ± 5.1 m³ ha⁻¹ in old-growth stands, with corresponding marginal mean CWD carbon pool 8.2 ± 1.6 t ha⁻¹ and 12.5 ± 0.7 …

Coarse Woody Debris Decomposition Assessment Tool: Model Validation And Application, Zhaohua Dai, Carl C. Trettin, Andrew J. Burton, Martin F. Jurgensen, Deborah S. Page-Dumroese, Brian T. Forschler, Jonathan S. Schilling, Daniel L. Lindner

Aspen Bibliography

Coarse woody debris (CWD) is a significant component of the forest biomass pool; hence a model is warranted to predict CWD decomposition and its role in forest carbon (C) and nutrient cycling under varying management and climatic conditions. A process-based model, CWDDAT (Coarse Woody Debris Decomposition Assessment Tool) was calibrated and validated using data from the FACE (Free Air Carbon Dioxide Enrichment) Wood Decomposition Experiment utilizing pine (Pinus taeda), aspen (Populous tremuloides) and birch (Betula papyrifera) on nine Experimental Forests (EF) covering a range of climate, hydrology, and soil conditions across the continental USA. …

Tree Establishment On Post-Mining Waste Soils: Species, Density, And Mixture Effects, Degi Harja Asmara, Suzanne Allaire, Meine Van Noordwijk, Damase P. Khasa

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Tree establishment to restore degraded boreal post-mining lands is challenged by low soil productivity, a harsh microclimate, and potentially high contaminant levels. The use of mixed vegetation can facilitate the microclimate but increase competition for soil resources. A statistical accounting of plant–plant interactions and adaptation to multispecies conditions is hard to achieve in field experiments; trials under controlled conditions can distinguish effects of planting density and species interactions in the early stages of plant establishment. A greenhouse trial was established in containers (“mesocosms”) with waste rock or fine tailings from gold mines. Pregerminated (1-week-old) seedlings (Alnus viridis subsp. crispa …

Spatial And Temporal Variation Of Epigaeic Beetle Assemblages (Coleoptera, Carabidae, Staphylinidae) In Aspen-Dominated Mixedwood Forests Across North-Central Alberta, H. E. James Hammond, Sergio García-Tejero, Greg R. Pohl, David W. Langor, John R. Spence

Aspen Bibliography

Epigaeic beetle assemblages were surveyed using continuous pitfall trapping during the summers of 1992 and 1993 in six widely geographically distributed locations in Alberta’s aspen-mixedwood forests prior to initial forest harvest. Species composition and turnover (β-diversity) were evaluated on several spatial scales ranging from Natural Regions (distance between samples 120–420 km) to pitfall traps (40–60 m). A total of 19,885 ground beetles (Carabidae) representing 40 species and 12,669 rove beetles (non-Aleocharinae Staphylinidae) representing 78 species was collected. Beetle catch, species richness, and diversity differed significantly among the six locations, as did the identity of dominant species. Beetle species composition differed …

Splitting The Difference: Heterogeneous Soil Moisture Availability Affects Aboveground And Belowground Reserve And Mass Allocation In Trembling Aspen, Ashley T. Hart, Morgane Merlin, Erin Wiley, Simon M. Landhäusser

Aspen Bibliography

When exploring the impact of resource availability on perennial plants, artificial treatments often apply conditions homogeneously across space and time, even though this rarely reflects conditions in natural systems. To investigate the effects of spatially heterogeneous soil moisture on morphological and physiological responses, trembling aspen (Populus tremuloides) saplings were used in a split-pot experiment. Following the division of the root systems, saplings were established for a full year and then subjected to either heterogeneous (portion of the root system exposed to non-lethal drought) or homogeneous (whole root system exposed to non-lethal drought or well-watered) treatments. Above- and belowground …