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Plant roots influence many ecological and biogeochemical processes, such as carbon, water and nutrient cycling. Because of difficult accessibility, knowledge on plant root growth dynamics in field conditions, however, is fragmentary at best. Minirhizotrons, i.e. transparent tubes placed in the substrate into which specialized cameras or circular scanners are inserted, facilitate the capture of high-resolution images of root dynamics at the soil-tube interface with little to no disturbance after the initial installation. Their use, especially in field studies with multiple species and heterogeneous substrates, though, is limited by the amount of work that subsequent manual tracing of roots in the images requires. Furthermore, the reproducibility and objectivity of manual root detection is questionable. Here, we use a Convolutional Neural Network (CNN) for the automatic detection of roots in minirhizotron images and compare the performance of our RootDetector with human analysts with different levels of expertise. Our minirhizotron data come from various wetlands on organic soils, i.e. highly heterogeneous substrates consisting of dead plant material, often times mainly roots, in various degrees of decomposition. This may be seen as one of the most challenging soil types for root segmentation in minirhizotron images. RootDetector showed a high capability to correctly segment root pixels in minirhizotron images from field observations (F1 = 0.6044; r2 compared to a human expert = 0.99). Reproducibility among humans, however, depended strongly on expertise level, with novices showing drastic variation among individual analysts and annotating on average more than 13-times higher root length/cm2 per image compared to expert analysts. CNNs such as RootDetector provide a reliable and efficient method for the detection of roots and root length in minirhizotron images even from challenging field conditions. Analyses with RootDetector thus save resources, are reproducible and objective, and are as accurate as manual analyses performed by human experts.
Biological invasions pose global threats to biodiversity and ecosystem functions. Invasive species often display a high degree of phenotypic plasticity, enabling them to adapt to new environments. This study examines plasticity to water stress in native and invasive Opuntia ficus-indica populations, a prevalent invader in arid and semi-arid ecosystems. Through controlled greenhouse experiments, we evaluated three native and nine invasive populations. While all plants survived the dry treatment, natives exhibited lower plasticity to high water availability with only a 36% aboveground biomass increase compared to the invasives with a greater increase of 94%. In terms of belowground biomass, there was no significant response to increased water availability for native populations, but plants from the invasive populations showed a 75% increase from the dry to the wet treatment. Enhanced phenotypic plasticity observed in invasive populations of O. ficus-indica is likely a significant driver of their success and invasiveness across different regions, particularly with a clear environmental preference towards less arid conditions. Climate change is expected to amplify the invasion success due to the expansion of arid areas and desertification. Opuntia ficus-indica adapts to diverse environments, survives dry spells, and grows rapidly in times of high-water supply, making it a candidate for increased invasion potential with climate change.
Understanding the effects of temperature and moisture on radial growth is vital for assessing the impacts of climate change on carbon and water cycles. However, studies observing growth at sub-daily temporal scales remain scarce.
We analysed sub-daily growth dynamics and its climatic drivers recorded by point dendrometers for 35 trees of three temperate broadleaved species during the years 2015–2020. We isolated irreversible growth driven by cambial activity from the dendrometer records. Next, we compared the intra-annual growth patterns among species and delimited their climatic optima.
The growth of all species peaked at air temperatures between 12 and 16°C and vapour pressure deficit (VPD) below 0.1 kPa. Acer pseudoplatanus and Fagus sylvatica, both diffuse-porous, sustained growth under suboptimal VPD. Ring-porous Quercus robur experienced a steep decline of growth rates with reduced air humidity. This resulted in multiple irregular growth peaks of Q. robur during the year. By contrast, the growth patterns of the diffuse-porous species were always right-skewed unimodal with a peak in June between day of the year 150–170.
Intra-annual growth patterns are shaped more by VPD than temperature. The different sensitivity of radial growth to VPD is responsible for unimodal growth patterns in both diffuse-porous species and multimodal growth pattern in Q. robur.
We studied a pristine, prominently patterned raised bog in Tierra del Fuego, Argentina, to disentangle the complex interactions among plants and water and peat. The studied bog lacks complicating features often posed by other bogs. It is completely dominated by Sphagnum magellanicum, which covers all niches and growth forms, and is joined by only a dozen higher plant species; it is entirely ombrotrophic with very sharp borders to the surrounding fen; it has only one type of peat that shows an only limited range in degree of decomposition; and it is situated in a very even climate with minimal differences in rainfall and temperature over the year. We present detailed measurements along a 498-m-long transect crossing the bog, including water table measurements (n = 498), contiguous vegetation relevés (n = 248), hydraulic conductivity just below the water table (n = 246), and hydraulic conductivity in 11 depth profiles (n = 291); degree of humification of the corresponding peat was assessed in conjunction with the hydraulic conductivity measurements (n = 537). Sphagnum magellanicum moss samples were collected every 2 m along this transect as well and genotyped (n = 242). In addition, along short, 26-m-long transects crossing strings and flarks water table and hydraulic conductivity just below the water table were measured every meter. Sphagnum growth forms were assessed, and the vegetation of the entire bog was mapped in 10 × 10-m relevés (n = 3322). A simulation model was applied to a generalized shape of the bog and produced surface patterns that well matched those seen in the field. The results were integrated with information from the literature and discussed in the framework of a self-regulating and self-organizing raised bog. We identified 19 hydrological feedback mechanisms. We found that the various mechanisms overlap in both space and time, which means there is redundancy in the self-regulation of the system. Raised bogs, when in a natural state, are among the most resilient ecosystems known; resilience that is provided by feedbacks and backup systems to these feedbacks.
Forests influence the climate of our Earth and provide habitat and food for many species and resources for human use. These valuable ecosystems are threatened by fast environmental changes caused by human-induced climte change. Negative growth responses and higher tree mortality rates were associated with increasing physiological stress induced by global warming. Especially boreal forests at high latitudes in the arctic region are threatened, a region predicted to undergo the highest increase in temperature during the next decades. Therefore, it is important to assess the adaptation potential in trees. For this purpose, I studied natural populations of white spruce (Picea glauca (Moench) Voss) in Alaska. In this thesis, I present three scientific papers in which my co-authors and I studied the phenotypic plasticity and genetic basis of tree growth, wood anatomy and drought tolerance as well as the genetic structure of white spruce populations in contrasting environments. We established three sites representing two cold-limited treelines and one drought-limited treeline with a paired plot design including one plot located at the treeline and one plot located in a closed-canopy forest, respectively. Additionally, the study design included one forest plot as reference. Within the entire project, in total 3,000 trees were measured, genotyped and dendrochronological data was obtained. I used several approaches to estimate the neutral and adaptive genetic diversity and phenotypic plasticity of white spruce as a model organism to explore the adaptation potential of trees to climate change.
In the first chapter, I combined neutral genetic markers with dendrochronological and climatic data to investigate population structure and individual growth of white spruce. Several individual-based dendrochronological approaches were applied to test the influence of genetic similarity and microenvironment on growth performance. The white spruce populations of the different sites showed high gene flow and high genetic diversity within and low genetic differentiation among populations, rather explained by geographic distance. The individual growth performances showed a high plasticity rather influenced by microenvironment than genetic similarity.
In the second chapter, I investigated the populations of the drought and cold-limited treeline sites to decipher the underlying genetic structure of drought tolerance using different genotype-phenotype association analyses. Based on tree-ring series and climatic data, growth declines caused by drought stress were identified and the individual reaction to the drought stress event was determined. A subset of 458 trees was genotyped, using SNPs in candidate genes and associated with the individual drought response. Most of the associations were revealed by an approach which took into account small-effect size SNPs and their interactions. Populations of the contrasting treelines responded differently to drought stress events. Populations further showed divergent genetic structures associated with drought responsive traits, most of them in the drought-limited site, indicating divergent selection pressure.
In the third chapter, my co-authors and I studied xylem anatomical traits at one of the cold-limited treeline sites to investigate whether genetic or spatial grouping affected the anatomy and growth of white spruce. Annual growth and xylem anatomy were compared between spatial groups and between genetic groups and individuals. Overall, wood traits were rather influenced by spatial than genetic grouping. Genetic effects were only found in earlywood hydraulic diameter and latewood density. Environmental conditions indirectly influenced traits related to water transport.
In conclusion, white spruce showed a high genetic diversity within and a low genetic differentiation among populations influenced by high gene flow rates. Genetic differences among populations are rather caused by geographical distance and therefore genetic drift. Differing selection pressure at the treeline ecotones presumably lead to divergent genetic structures underlying drought-tolerant phenotypes among the populations. Thus, adaptation to drought most likely acts on a local scale and involves small frequency shifts in several interacting genes. The identified genes with adaptive growth traits can be used to further exlore local adaptation in white spruce. Tree growth and wood anatomical traits are rather influenced by the environment than genetics and showed a high phentoypic plasticity. The high genetic diverstiy and phenotypic plasticity of white spruce may help the species to cope with rapid environmental changes. Still, additional work is needed to further explore adaptation processes to estimate how tree species reacted to rapid climate change. The presented thesis shed some light on the adaptation potential of trees by the example of white spruce using several approaches.
Late to bed, late to rise—Warmer autumn temperatures delay spring phenology by delaying dormancy
(2021)
Abstract
Spring phenology of temperate forest trees has advanced substantially over the last decades due to climate warming, but this advancement is slowing down despite continuous temperature rise. The decline in spring advancement is often attributed to winter warming, which could reduce chilling and thus delay dormancy release. However, mechanistic evidence of a phenological response to warmer winter temperatures is missing. We aimed to understand the contrasting effects of warming on plants leaf phenology and to disentangle temperature effects during different seasons. With a series of monthly experimental warming by ca. 2.4°C from late summer until spring, we quantified phenological responses of forest tree to warming for each month separately, using seedlings of four common European tree species. To reveal the underlying mechanism, we tracked the development of dormancy depth under ambient conditions as well as directly after each experimental warming. In addition, we quantified the temperature response of leaf senescence. As expected, warmer spring temperatures led to earlier leaf‐out. The advancing effect of warming started already in January and increased towards the time of flushing, reaching 2.5 days/°C. Most interestingly, however, warming in October had the opposite effect and delayed spring phenology by 2.4 days/°C on average; despite six months between the warming and the flushing. The switch between the delaying and advancing effect occurred already in December. We conclude that not warmer winters but rather the shortening of winter, i.e., warming in autumn, is a major reason for the decline in spring phenology.
Abstract
Myxomycetes are terrestrial protists with many presumably cosmopolitan species dispersing via airborne spores. A truly cosmopolitan species would suffer from outbreeding depression hampering local adaptation, while locally adapted species with limited distribution would be at a higher risk of extinction in changing environments. Here, we investigate intraspecific genetic diversity and phylogeography of Physarum albescens over the entire Northern Hemisphere. We sequenced 324 field collections of fruit bodies for 1–3 genetic markers (SSU, EF1A, COI) and analysed 98 specimens with genotyping by sequencing. The structure of the three‐gene phylogeny, SNP‐based phylogeny, phylogenetic networks, and the observed recombination pattern of three independently inherited gene markers can be best explained by the presence of at least 18 reproductively isolated groups, which can be seen as cryptic species. In all intensively sampled regions and in many localities, members of several phylogroups coexisted. Some phylogroups were found to be abundant in only one region and completely absent in other well‐studied regions, and thus may represent regional endemics. Our results demonstrate that the widely distributed myxomycete species Ph. albescens represents a complex of at least 18 cryptic species, and some of these seem to have a limited geographical distribution. In addition, the presence of groups of presumably clonal specimens suggests that sexual and asexual reproduction coexist in natural populations of myxomycetes.
Abstract
Drainage has turned 650,000 km2 of peatlands worldwide into greenhouse gas sources. To counteract climate change, large‐scale rewetting is necessary while agricultural use of rewetted areas, termed paludiculture, is still possible. However, more information is required on the performance of suitable species, such as cattail, in the range of environmental conditions after rewetting. We investigated productivity and biomass quality (morphological traits and tissue chemical composition) of Typha angustifolia and Typha latifolia along gradients of water table depth (−45 to +40 cm) and nutrient addition (3.6–400 kg N ha−1 a−1) in a six‐month mesocosm experiment with an emphasis on their high‐value utilization, e.g., as building material, paper, or biodegradable packaging. Over a wide range of investigated conditions, T. latifolia was more productive than T. angustifolia. Productivity was remarkably tolerant of low nutrient addition, suggesting that long‐term productive paludiculture is possible. Low water tables were beneficial for T. latifolia productivity and high water tables for T. angustifolia biomass quality. Rewetting will likely create a mosaic of different water table depths. Our findings that the yield of T. angustifolia and tissue chemical composition of T. latifolia were largely unaffected by water table depth are therefore promising. Depending on intended utilization, optimal cultivation conditions and preferable species differ. Considering yield or diameter, e.g., for building materials, T. latifolia is generally preferable over T. angustifolia. A low N, P, K content, high Si content and high C/N‐ratio can be beneficial for processing into disposable tableware, charcoal, or building material. For these utilizations, T. angustifolia is preferable at high water tables, and both species should be cultivated at a low nutrient supply. When cellulose and lignin contents are relevant, e.g., for paper and biodegradable packaging, T. angustifolia is preferable at high water tables and both species should be cultivated at nutrient additions of about 20 kg N ha−1 a−1.
Abstract
Climate change is increasing the frequency and intensity of drought events in many boreal forests. Trees are sessile organisms with a long generation time, which makes them vulnerable to fast climate change and hinders fast adaptations. Therefore, it is important to know how forests cope with drought stress and to explore the genetic basis of these reactions. We investigated three natural populations of white spruce (Picea glauca) in Alaska, located at one drought‐limited and two cold‐limited treelines with a paired plot design of one forest and one treeline plot. We obtained individual increment cores from 458 trees and climate data to assess dendrophenotypes, in particular the growth reaction to drought stress. To explore the genetic basis of these dendrophenotypes, we genotyped the individual trees at 3000 single nucleotide polymorphisms in candidate genes and performed genotype–phenotype association analysis using linear mixed models and Bayesian sparse linear mixed models. Growth reaction to drought stress differed in contrasting treeline populations. Therefore, the populations are likely to be unevenly affected by climate change. We identified 40 genes associated with dendrophenotypic traits that differed among the treeline populations. Most genes were identified in the drought‐limited site, indicating comparatively strong selection pressure of drought‐tolerant phenotypes. Contrasting patterns of drought‐associated genes among sampled sites and in comparison to Canadian populations in a previous study suggest that drought adaptation acts on a local scale. Our results highlight genes that are associated with wood traits which in turn are critical for the establishment and persistence of future forests under climate change.
Abstract
River estuaries are characterized by mixing processes between freshwater discharge and marine water masses. Since the first are depleted in heavier stable isotopes compared with the marine realm, estuaries often show a linear correlation between salinity and water stable isotopes (δ18O and δ2H values). In this study, we evaluated spatial and seasonal isotope dynamics along three estuarine lagoon transects, located at the northern German Baltic Sea coast. The data show strong seasonality of isotope values, even at locations located furthest from the river mouths. They further reveal a positive and linear salinity‐isotope correlation in spring, but ‐in two of the three studied transects‐ hyperbolic and partially reverse correlations in summers. We conclude that additional hydrological processes partially overprint the two‐phase mixing correlation during summers: aside from the isotope seasonality of the riverine inflows, the shallow inner lagoons in the studied estuaries are influenced by evaporation processes. In contrast the estuarine outflow regions are under impact of significant salinity and isotope fluctuations of the Baltic Sea. Deciphering those processes is crucial for the understanding of water isotope and salinity dynamics. This is also of relevance in context of ecological studies, for example, when interpreting oxygen and hydrogen isotope data in aquatic organisms that depend on ambient estuarine waters.