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Agricultural bioinoculants containing arbuscular mycorrhizal fungi represent a potential opportunity to reduce the dependence of grapevines (Vitis) on agrochemicals. This field study assessed the ability of four commercial bioinoculants to colonize grapevine roots and their effects on petiole nutrient concentration, berry composition, and root morphology of ‘Pinot noir’ (Vitis vinifera) grafted onto rootstock ‘Couderc 3309’ (Vitis riparia × Vitis rupestris) and ‘Riesling’ (V. vinifera) grafted onto ‘Couderc 3309’ and Selection Oppenheim four (Vitis berlandieri × V. riparia). Three bioinoculants increased root mycorrhizal colonization; however, regardless of the treatment, mycorrhizal fungal structures were enhanced. Grapevine petiole nutrient concentration was improved by bioinoculants. Root diameter, root length density, and specific root length increased with greater mycorrhizal root colonization. Using bioinoculants to reduce chemical fertilizers may be a good strategy to improve grapevine productivity and health in cool climates; however, the impact of mycorrhizal bioinoculants in the vineyard may differ among scion–rootstocks, edaphoclimatic conditions, and vineyard soil microbiomes.

In recent years, important processes controlling ecosystem carbon dynamics have been connected to fine-root exudation of soluble carbon compounds. Root exudation patterns may change depending on plant interactions and plant susceptibility to and recovery from drought. Recent investigations suggest that root exudation tends to increase with stress events, but quantification of the amount of carbon released from roots across soil depths with differing water availability and species interactions are missing.We tested if root exudation rates were negatively correlated with soil water content across soil depths during and after drought. We further tested if species in mixture, often considered to be less stressed under drought, exuded less carbon than species in monospecific environments. Exudates were sampled in a mature Fagus sylvatica L. and Picea abies (L.) Karst. forest at the end of a five-year throughfall exclusion period and again one year after the drought ended. We quantified root exudates and their variation with soil depth for both tree species in monospecific and mixed species zones.Carbon exudation significantly increased in fine roots exposed to dry soils (

Forest soils are crucial for many ecosystem services that rely on soil organic matter (SOM) stability. Carbon allocated to roots and released as exudates to the rhizosphere plays a key role in SOM stabilization. Under periodic drought, elevated root exudation and SOM accumulation have been reported. Yet, whether root exudates control SOM formation and stability in mature forests once the drought ends is largely unknown. We examined whether root exudates from P. abies and F. sylvatica trees relate to SOM formation and stability in soil depth profiles one year following five years of experimental drought (Kroof experiment, Germany). We collected root exudates throughout the rooting zone and combined the data with thermogravimetric analysis of SOM in the rhizosphere and non-rooted soil. We found that the rhizosphere of both species was characterized by stable SOM fractions that did not decrease post-drought, suggesting potential protection of SOM due to rhizodeposition and root exudates. In contrast, stable SOM fractions decreased relative to controls in non-rooted topsoil below P. abies, indicating a loss of stabilized SOM from drought-affected and re-wetted soil. Our measurements provide valuable insights into post-drought SOM formation and mechanisms of SOM stabilization in forest ecosystems under climate change.

Drought tolerance is a highly complex trait controlled by numerous interconnected pathways with substantial variation within and across plant species. This complexity makes it difficult to distill individual genetic loci underlying tolerance, and to identify core or conserved drought responsive pathways. Here, we collected drought physiology and gene expression datasets across diverse genotypes of the C4 cereals sorghum and maize and searched for signatures defining water deficit responses. Differential gene expression identified few overlapping drought associated genes across sorghum genotypes, but using a predictive modeling approach, we found a shared core drought response across development, genotype, and stress severity. Our model had similar robustness when applied to datasets in maize, reflecting a conserved drought response between sorghum and maize. The top predictors are enriched in functions associated with various abiotic stress responsive pathways as well as core cellular functions. These conserved drought response genes were less likely to contain deleterious mutations than other gene sets, suggesting that core drought responsive genes are under evolutionary and functional constraints. Our findings support a broad evolutionary conservation of drought responses in C4 grasses regardless of innate stress tolerance, which could have important implications for developing climate resilient cereals.Significance StatementDrought is a complex and variable stress that is difficult to quantify and link to underlying mechanisms both within and across species. Here, we developed a predictive model to classify drought stress responses in sorghum and identify important features that are responsive to water deficit. Our model has high predictive accuracy across development, genotype, and stress severity, and the top features are enriched in genes related to classical stress responses and have functional and evolutionary conservation. We applied this sorghum trained model to maize, and observed similar predictive accuracy of drought responses, supporting transfer learning across plant species. Our findings suggest there are deeply conserved drought responses across C4 grasses that are unrelated to tolerance.

Successful control and prevention of biological invasions depend on identifying traits of non-native species that promote fitness advantages in competition with native species. Here, we show that, among 76 native and non-native woody plants of deciduous forests of North America, invaders express a unique functional syndrome that combines high metabolic rate with robust leaves of longer lifespan and a greater duration of annual carbon gain, behaviours enabled by seasonally plastic xylem structure and rapid production of thin roots. This trait combination was absent in all native species examined and suggests the success of forest invaders is driven by a novel resource-use strategy. Furthermore, two traits alone-annual leaf duration and nuclear DNA content-separated native and invasive species with 93% accuracy, supporting the use of functional traits in invader risk assessments. A trait syndrome reflecting both fast growth capacity and understorey persistence may be a key driver of forest invasions.

Rhizosphere bacterial communities (‘rhizobacteria’) can mediate plant-soil feedbacks that enhance plant drought tolerance, but the composition and ecological characteristics of these populations are poorly understood. We investigated the impact of drought on rhizobacteria communities and the effect of propagating drought-impacted communities on the drought tolerance of naive plants in a two-phase experiment. We tested whether propagating rhizosphere soils from drought-stressed shrub willow (Salix purpurea), or continuously watered controls, affected the performance of a subsequent generation of droughted plants. The impacts of drought on rhizobacteria and plant fitness were evaluated using 16S rRNA gene sequencing and measures of plant growth (root mass and stem length etc.) and stress (abscisic acid, osmolality). Uninoculated reference plants were used to isolate the effects of inoculation. Drought had a significant and lasting impact on the structure of rhizobacterial communities, characterized by increased populations of Actinobacteria (Acidimicrobiia, Thermoleophilia and Micrococcaceae). Foliar osmolality was significantly higher in plants receiving drought-selected communities relative to uninoculated controls, but no significant differences in plant growth were observed. Inoculation with rhizosphere soil from watered controls significantly reduced plant growth. The rhizosphere of watered control plants was dominated by Proteobacteria and Bacteroidetes. Our findings demonstrate that plants are affected by the legacy effects of drought on the rhizosphere microbiome. This drought legacy was propagated and persisted throughout nine weeks of plant growth, independent of prevailing water stress. Drought-impacted rhizospheres had larger populations of desiccation-tolerant (ex. Arthrobacter) and putatively endophytic taxa (ex. Rhizobium) with established plant growth promoting capabilities.

The xylem network, the water conduction system in wood determines the ability of trees to avoid hydraulic failure during drought stress. The capability to withstand embolisms, disruptions of the water column by gas bubbles that contribute to hydraulic failure, is mainly determined by the anatomical arrangement and connectedness (topology) of xylem vessels. However, the quantification of xylem network characteristics has been difficult, so that relating network properties and topology to hydraulic vulnerability and predicting xylem function remains challenging. We studied the xylem vessel networks of three diffuse- (Fagus sylvatica, Liriodendron tulipifera, Populus x canadensis) and three ring-porous (Carya ovata, Fraxinus pennsylvatica, Quercus montana) tree species using volumetric images of xylem from laser ablation tomography (LATscan). Using convolutional neural networks for image segmentation, we generated three-dimensional, high-resolution maps of xylem vessels, with detailed measurements of morphology and topology. We studied the network topologies by incorporating multiple network metrics into a multidimensional analysis and simulated the robustness of these networks against the loss of individual vessel elements that mimic the obstruction of water flow from embolisms. This analysis suggested that networks inPopulus x canadensisandCarya ovataare quite similar despite being different wood types. Similar networks had comparable experimental measurements of P50 values (pressure inducing 50% hydraulic conductivity loss) obtained from hydraulic vulnerability curves, a common tool to quantify the cavitation resistance of xylem networks. This work produced novel data on plant xylem vessel networks and introduces new methods for analyzing the biological impact of these network structures.Significance statementThe resilience of fluid transport networks such as xylem vessels that conduct water in trees depends on both the structure of the network and features of the individual network elements. High-resolution reconstruction of xylem networks from six tree species provided novel, three-dimensional, structural data which enabled the xylem networks to be described using graph theory. Using an array of network metrics as multidimensional descriptors, we compared the xylem networks between species and showed relationships to simulated and experimental measures of drought resistance. In addition to providing insight on drought resistance, these approaches offer new ways for comparative analysis of networks applicable to many systems.

There is urgency in optimizing agricultural production in light of climate uncertainty, human population growth and resource limitations. The rhizosphere microbiome is a promising genetic resource, yet it is poorly understood. Additionally, research into rhizosphere microbial community assembly (MCA) lacks a consistent ecological framework that incorporates the dynamics of the occurring interactions. We explore the ecological principles that guide community establishment as they pertain to the rhizosphere while reviewing relevant research and highlighting the gaps in knowledge. We propose a conceptual model for studying the rhizosphere, and argue for higher resolution characterizations of the rhizosphere, under a framework of root and microbial traits that will determine its resulting composition. For this, we borrow concepts from ecological theory to chronologically describe MCA as a function of colonization, distribution and succession under changing plant-imposed filters. We argue that there is a need to consider the temporal and spatial scale of rhizosphere processes in a manner that is relevant to its microbial components. Finally, we discuss strategies for managing the rhizosphere microbiome for agriculture and the remaining gaps in knowledge that impede their application. Synthesis and applications. We aim to initiate a comprehensive conversation on ecological processes and plant and microbial traits that affect rhizosphere assembly. Rhizosphere microbiome research is relatively new and highly multidisciplinary. The use of a shared vocabulary and ecological theory will facilitate the development and advances in this discipline.

Temperate forest ecosystems are exposed to a higher frequency, duration and severity of drought. To promote forest longevity in a changing climate, we require a better understanding of the long-term impacts of repetitive drought events on fine-root dynamics in mature forests. Using minirhizotron methods, we investigated the effect of seasonal drought on fine-root dynamics in single-species and mixed-species arrangements of Fagus sylvatica (European beech) and Picea abies (Norway spruce) by means of a 4-yr-long throughfall-exclusion experiment. Fine-root production of both species decreased under drought. However, this reduction was not evident for P. abies when grown intermixed with F. sylvatica. Throughfall-exclusion prolonged the lifespan of P. abies roots but did not change the lifespan of F. sylvatica roots, except in 2016. Fagus sylvatica responded to drought by reducing fine-root production at specific depths and during roof closure. This is the first study to examine long-term trends in mature forest fine-root dynamics under repetitive drought events. Species-specific fine-root responses to drought have implications for the rate and depth of root-derived organic matter supply to soil. From a root dynamics perspective, intermixing tree species is not beneficial to all species but dampens drought impacts on the belowground productivity of P. abies.

After drought events, tree recovery depends on sufficient carbon (C) allocation to the sink organs. The present study aimed to elucidate dynamics of tree-level C sink activity and allocation of recent photoassimilates (C$_{new}$) and stored C in c. 70-year-old Norway spruce (Picea abies) trees during a 4-week period after drought release. We conducted a continuous, whole-tree $^{13}$C labeling in parallel with controlled watering after 5 years of experimental summer drought. The fate of C$_{new}$ to growth and CO$_{2}$ efflux was tracked along branches, stems, coarse- and fine roots, ectomycorrhizae and root exudates to soil CO$_{2}$ efflux after drought release. Compared with control trees, drought recovering trees showed an overall 6% lower C sink activity and 19% less allocation of C$_{new}$ to aboveground sinks, indicating a low priority for aboveground sinks during recovery. In contrast, fine-root growth in recovering trees was seven times greater than that of controls. However, only half of the C used for new fine-root growth was comprised of C$_{new}$ while the other half was supplied by stored C. For drought recovery of mature spruce trees, in addition to C$_{new}$, stored C appears to be critical for the regeneration of the fine-root system and the associated water uptake capacity.

Carbon (C) exuded via roots is proposed to increase under drought and facilitate important ecosystem functions. However, it is unknown how exudate quantities relate to the total C budget of a drought-stressed tree, that is, how much of net-C assimilation is allocated to exudation at the tree level. - We calculated the proportion of daily C assimilation allocated to root exudation during early summer by collecting root exudates from mature Fagus sylvatica and Picea abies exposed to experimental drought, and combining above- and belowground C fluxes with leaf, stem and fine-root surface area. - Exudation from individual roots increased exponentially with decreasing soil moisture, with the highest increase at the wilting point. Despite c. 50% reduced C assimilation under drought, exudation from fine-root systems was maintained and trees exuded 1.0% (F. sylvatica) to 2.5% (P. abies) of net C into the rhizosphere, increasing the proportion of C allocation to exudates two- to three-fold. Water-limited P. abies released two-thirds of its exudate C into the surface soil, whereas in droughted F. sylvatica it was only one-third. - Across the entire root system, droughted trees maintained exudation similar to controls, suggesting drought-imposed belowground C investment, which could be beneficial for ecosystem resilience.

Drought is a severe natural risk that increases drying-rewetting frequencies of soils. Yet, it remains largely unknown how forest ecosystems respond to dry-wet cycles, hampering our ability to evaluate the overall sink and source functionality for this large carbon pool. Recent investigations suggest that the release of soluble carbon via root exudation increases under drought, influencing soil carbon stabilization and mineralization. However, an integration of root exudation into the carbon allocation dynamics of drought stressed trees is missing. We hypothesized that roots in dry soil layers have a higher exudation rate than roots in more moist layers across different soil depths. Further, we tested if higher exudation rates under drought are attenuated by reduced root abundance in dry soils and if the fraction of root exudation from total carbon allocation increases with decreasing photosynthesis rates under drought. At the KROOF experimental site in southern Germany, where mature beech (Fagus sylvatica L.) and spruce (Picea abies (L.) Karst.) trees were exposed to artificial drought stress for five consecutive growing seasons, we show that at the root level root exudation rate increases in drier soils. Especially roots in the upper soil profile and roots of spruce trees increased root exudation under drought. When scaled to whole tree level, we did not find differences in total exudation between drought stressed and control trees, indicating sustained root exudation at the tree level under drought. As photosynthesis rates and therefore total carbon assimilation was substantially reduced under drought (by 50 % in beech and almost 70 % in spruce), the fraction of root exudation from total assimilation slightly increased for drought stressed trees. Our results demonstrate that stimulation of root exudation rates with drought exists in natural temperate forest ecosystems but might be mitigated by reduced fine root abundance under drought. Nevertheless, increased exudation per root surface area will have localized impacts on rhizosphere microbial composition and activity especially in the topsoil exposed to more extreme dry-wet cycles. Finally, also the exudate composition can help to determine how priming of soil organic matter relates to belowground carbon allocation dynamics and to disclose processes of complementary species interaction and should be emphasised in future studies.

Plants allocate an estimated 11% of the C that they fixate as root exudates, a complex mixture of compounds that helps engineer the plant’s subterranean habitat. Root exudates stabilize soil aggregates, improve water retention, and shape rhizosphere microbial community composition. Exudates are also thought to contribute to the formation of stable mineral-associated organic matter. However, the function and fate of exudates along the soil profile may differ. We hypothesize that in topsoils with highly active microbial populations and mineral surfaces saturated with organic matter, root exudates may be rapidly intercepted and assimilated by soil microbes, and later adsorbed to surfaces as microbial necromass. But in subsoils with low microbial activity, exudates may directly adsorb on unsaturated mineral surfaces. The magnitude of these divergent pathways can shape the role of root exudates in rhizosphere C cycling. However, little is known about how adsorption vs. decomposition processes at the root-soil interface control i) the chemical transformations of C occurring along the root-microbe-mineral pathway, and ii) the spatial distribution and heterogeneity of exuded and processed exudate C. Our objective was to investigate the effect of microbial activity, and reactive mineral surfaces on the spatial distribution and functional group chemistry of root exudates at the root-microbe-mineral interface.We packed samples from O, A, B, and C horizons collected from a grassland Mollisol, into individual microcosms, and installed a porous microdialysis membrane which served as an artificial root. Through this root, we injected either root exudates collected from maize plants, or dissolved organic carbon extracted from plant litter collected at the site. This comparison allowed us to study the dynamics of organic inputs entering the soil profile from the litter layer vs. directly from the roots. We destructively sampled the microcosms and obtained intact cross sections containing the artificial root and surrounding mineral/pore structures at different time points throughout the experiment. Here, we will present results obtained using synchrotron-radiation FTIR-microscopy of the temporal evolution, and spatial distribution of organic matter functional group chemistry in an artificial rhizosphere.

Tracers provide a way to determine, follow and quantify biochemical cycles and energy fluxes within the soil-plant-atmosphere continuum (SPAC). Thereby, different tracers, such as dyes, carbonyl sulfite or stable isotopes are employed. One major disadvantage of many tracers is, that very often, plants have to be destructively harvested to analyze the tracer concentration, making it difficult to measure continuous fluxes. Additionally, for stable isotope studies, fractionation or exchange effects can make interpretation and quantification of biogeochemical fluxes difficult. Novel tracers that are already frequently used in animal systems, are fluorescent C-dots. These nanoparticles (5-50 nm diameter) provide a non-destructive imaging option using “in vivo imaging systems” (IVIS). We examined a first approach to apply and measure C-dots as possible tracers in tree saplings of three species (Picea glauca, Pinus strobus, Tsuga canadensis). Roots were excavated from soils and exposed to 20 µmol/l liquid silica-based nanoparticles (diameter of 5.1 nm) labeled with a near-infrared fluorophore, cyanine 5.5 (excitation maximum 646 nm, emission maximum 662 nm). Subsequent continuous IVIS imaging showed real-time uptake and transport of the C-dots by the trees. Respective fluorescence intensity revealed the concentration of C-dots in each of the tissues at measured time steps. Subsequent cross-sectioning of roots, stems and leaves elucidated the internal transport pathway of the C-dots inside the saplings’ tissues. Finally, measurements of stomatal conductance, photosynthesis, transpiration rate, stem hydraulic conductivity and pre-dawn leaf water potentials in comparison to control saplings revealed no phytotoxic effect by the C-dots on plant functioning. In conclusion, C-dots provide a non-toxic new technique for measuring biochemical cycles within the SPAC. Future applications include high resolution tracing of water flow cycles and turnover times within the SPAC, compound specific analyses of root exudation or determining mechanisms of pest influences on plant metabolism.

Drought is considered a severe natural risk that increases drying-rewetting frequencies of soil. Yet, it remains largely unknown how forest ecosystems respond, hampering our ability to evaluate the overall sink and source functionality for this large carbon pool. Recent investigations present that the loss of soluble carbon via root exudation increases under drought, facilitating fundamental carbon stabilization and mineralization dynamics. However, information on the vertical variation of root exudation from interacting tree species is missing. Here we show that drought increases root exudation rates only in the upper soil profile, while exudation rates decrease in the deeper profile under drought. These trends occurred in both, monocultures and species mixtures. Surprisingly, beech (Fagus sylvatica) and spruce (Picea abies) trees showed opposing results depending on species mixture. While root exudation rates increased in beech growing together with spruce, drought-susceptible spruce had higher exudation rates when grown in monoculture, suggesting the benefit of spruce in mixed cultures via reduced belowground carbon loss. Our results demonstrate that stimulation of root exudation rates with drought exists in natural temperate forest ecosystems, but only in shallow soil depths. In contrast, decreased exudation rates in deeper soil during drought suggest carbon stabilization. The exudate composition can help to determine how priming of soil organic matter relates to microbial respiration and to disclose belowground processes of complementary species interaction.

Forest cover exerts a significant control on the partitioning of precipitation between evapotranspiration and surface runoff. Thus, understanding how plants take up and transpire water in forested catchments is essential to predict flooding potential and hydrologic cycling. A growing literature underscores the importance of integrating whole-plant hydraulics, including such processes as the spatial variability of root distribution and the temporally dynamic nature of root water uptake by depth in understanding the relationship between changes in vegetation and hydrology. The analysis of stable isotopes of water (18O and 2H) sourced from soils and plant tissue has enabled the estimation of tree root water uptake depths and water use strategies. Despite the general acceptance of stable water isotopic data to estimate plant hydraulic dynamics, this methodology imposes assumptions that may produce spurious results. For example, end member mixing analysis neglects time-delays during tree-water storage. Also, it is likely that hydraulic redistribution processes of plants, which transport water across soil depths and both into and out of plant tissue, modify δ18O and δ2H; the isotopic signature of a collected sample may thus reflect a history of transport and exposure to fractionating processes not accounted for in analysis. We tested the feasibility of C-dots, core-shell silica polyethylene-glycol coated fluorescent nano-particles (5.1 nm diameter) in 20 µmol/l solution with H2O labeled with a near-infrared fluorophore, cyanine 5.5 (excitation maximum of 646 nm, emission maximum of 662 nm), as an alternative to stable water isotopes in the investigation of plant hydraulics. We examined the absorption and transport of C-dots through soil, as well as roots and aerial structures of Eastern hemlock (Tsuga canadensis), Eastern white pine (Pinus strobus), and white spruce (Picea glauca) saplings (n = 12 each) via an IVIS-200 luminescence in-situ imaging system. We compared the fluid mechanics, residence times and mixing schemes of C-dots with 2H-labeled water during transport within these plant species to establish the nanoparticles as a viable alternative through a split-root hydraulic redistribution experiment under moderate and severe drought conditions. We present a residence-time distribution to elucidate the mixing scheme of C-dot solution and calibration curves to aid future studies. This research is the premier assessment of this nanoparticle as an alternative tracer to stable water isotopes, and as such may yield insights for broader applications.

Phenolic compounds perform various functions in soil ranging from microbial substrate to toxin and form the basis of several plant-mediated processes. The aim of this study was to investigate how phenolics commonly exuded by tree roots influence soil organic matter (SOM) decomposition and interact with other labile forms of carbon (C) abundant in root exudates. Therefore, we performed a 38-day incubation experiment and assessed whether phenolic compounds (benzoic acid, caffeic acid and catechin) facilitated or inhibited SOM decomposition in a glucose-amended forest soil. Changes in decomposition, substrate use, fungal and bacterial community composition, and microbial abundance and activity were measured over time using 13C-stable-isotope tracing, DNA-based molecular methods and enzyme assays. Our findings showed that phenolics inhibited microbial activity and abundance to varying degrees. Yet, benzoic acid was the only compound producing a substantial priming effect leading to a 21% increase in SOM decomposition, which was amplified in glucose-amended soils. This stimulation in microbial activity was associated with an increase in β-1,4-glucosidase activity and the bacterial genera Paraburkholderia and Caballeronia of the Burkholderiaceae family. Phenolics drove microbial community shifts in glucose-amended soils with negligible interactive effects. In conclusion, phenolic priming of SOM decomposition is associated with microbial community shifts and amplified in the presence of glucose. This evidence emphasizes the need for considering phenolics and interactions among root exudates as priming mechanisms in the rhizosphere and other soil environments where aromatics and phenolic compounds are abundant.

Key message Under severe drought, growth of Norway spruce suffered much more than European beech. Norway spruce benefited from growing in the environment of beech, and both species acclimated slightly to 5 years of experimentally extended drought. Abstract Recent studies show that the detrimental effects of drought on stand growth are mitigated when the stand contains mixed tree species. We analysed the growth responses of Norway spruce and European beech to episodic and experimentally extended drought in intra- and inter-specific mature stands. We used annual diameter growth records dating back to 1998 to determine the impact of the natural episodic drought in 2003 and 2015. To analyse extended drought, spruce and beech trees were exposed to extreme drought under automatic throughfall exclusion roofs from 2014 to 2018. The growth of spruce in an inter-specific environment with beech was 20–50% less affected by natural episodic drought compared with an intra-specific constellation. When beech grew in an inter-specific environment, it was by 23% more affected by drought compared to intra-specific conditions, but seemed to recover faster. The induced drought from 2014 to 2018 resulted in a strong growth reduction in the first year particularly for spruce, followed by a slight acclimation to the dry conditions. Beech acclimated and recovered faster than spruce across all growing conditions, while spruce only acclimatized faster in the environment of beech. Both species showed a higher mortality under induced drought compared with the controls; for spruce, the mortality rate was fivefold higher compared to the long-term mortality. The long-term moderate-growth stabilization and the growth increase after the 5-year exposure to drought suggest a gradual acclimation to drought by beech. The resistance and acclimation to drought of spruce when growing in mixture should be considered when designing resource efficient and productive mixed conifer-broadleaved stands for future climates.

Hydraulic redistribution (HR) of soil water through plant roots is widely described; however its extent, especially in temperate trees, remains unclear. Here, we quantified HR of five temperate tree species. We hypothesized that both, HR within a plant and into the soil increase with higher water‐potential gradients, larger root conduit diameters and root‐xylem hydraulic conductivities as HR driving factors. Saplings of conifer (Picea abies, Pseudotsuga menziesii), diffuse‐porous (Acer pseudoplatanus) and ring‐porous species (Castanea sativa, Quercus robur) were planted in split‐root systems, where one plant had its roots split between two pots with different water‐potential gradients (0.23–4.20 MPa). We quantified HR via deuterium labelling. Species redistributed 0.39 ± 0.14 ml of water overnight (0.08 ± 0.01 ml/g root mass). Higher pre‐dawn water‐potential gradients, hydraulic conductivities and larger conduits significantly increased HR quantity. Hydraulic conductivity was the most important driving factor on HR amounts, within the plants (0.03 ± 0.01 ml/g) and into the soil (0.06 ± 0.01 ml/g). Additional factors as soil‐root contact should be considered, especially when calculating water transfer into the soil. Nevertheless, trees maintaining high‐xylem hydraulic conductivity showed higher HR amounts, potentially making them valuable ‘silvicultural tools’ to improve plant water status. A free Plain Language Summary can be found within the Supporting Information of this article.

Mixed stands of European beech (Fagus sylvatica L.) and Norway spruce (Picea abies (L.) Karst.) frequently over-yield, when compared to respective monospecific stands. Over-yielding is attributed to enhanced resource uptake efficiency through niche complementarity alleviating species competition, for example through enhanced root stratification in mixture. Under severe and frequent summer drought, however, water limitation may become crucial in modifying the prevailing competitive interaction in mixed beech-spruce forests. We hypothesize, therefore, that under drought (H I) inter-specific interaction with beech reduces water accessibility for spruce more than intra-specific conditions, thus (H II) exacerbating drought susceptibility of spruce in terms of reduced photosynthesis and stem growth. Reactions at the organ (leaf, fine root), tree and stand scale were analysed in a mature forest with beech-spruce group mixture. Under inter-specific conditions spruce’s fine-root production and depth of water uptake (assessed via δ18O of xylem water) shifted to shallow, drought-prone soil horizons, in agreement with H I. Overall, lowered fine root production and ramification along with a reduction in long-distance explorative ectomycorrhizal types resulted in decreased soil exploitation in spruce when growing together with beech. Spruce’s drought sensitivity was exemplified by a distinct decrease in stomatal conductance, net CO2 uptake rate and stem growth during periods of water limitation. Notwithstanding, species interaction effects were absent in leaf gas exchange and stem diameter growth, during a six-week summer drought period in 2013 as well as in the extremely dry year of 2003, hence rejecting H II. Based on results from soil moisture measurements and water uptake depth, we interpret the conflicting findings for H I and H II to result from: (i) seasonal shifts between positive (during spring drought) and negative (during summer drought) effects of beech neighbourhood on soil water availability for spruce, possibly overriding each other in their effect on annual stem diameter growth and (ii) the group-wise mixture pattern, where spruce is exposed to competition with beech only along group edges, i.e. laterally only, so that the putatively adverse beech effect on water accessibility stays limited. Our results suggest, compared to single tree mixture, group-wise mixture of beech and spruce to be a favourable silvicultural option in the face of climate change.

The (13/12) C ratio in plant roots is likely dynamic depending on root function (storage versus uptake), but to date, little is known about the effect of season and root order (an indicator of root function) on the isotopic composition of C-rich fractions in roots. To address this, we monitored the stable isotopic composition of one evergreen (Picea abies) and one deciduous (Fagus sylvatica), tree species' roots by measuring δ(13) C of bulk, respired and labile C, and starch from first/second and third/fourth order roots during spring and fall root production periods. In both species, root order differences in δ(13) C were observed in bulk organic matter, labile, and respired C fractions. Beech exhibited distinct seasonal trends in δ(13) C of respired C, while spruce did not. In fall, first/second order beech roots were significantly depleted in (13) C, whereas spruce roots were enriched compared to higher order roots. Species variation in δ (13) C of respired C may be partially explained by seasonal shifts from enriched to depleted C substrates in deciduous beech roots. Regardless of species identity, differences in stable C isotopic composition of at least two root order groupings (first/second, third/fourth) were apparent, and should hereafter be separated in belowground C-supply-chain inquiry.

Root trenching of Norway spruce and European beech caused no long-term decrease of stem growth and only temporal growth reductions in European beech. Root volume reduction is a common occurrence in silvicultural, urban, and scientific environments, often the direct result of trenching applications. Although this interference is widespread near forest roads and in rural and urban areas, trees’ growth reactions to these limitations have rarely been analysed. Here, we compare the growth of 81 mature Norway spruce (Picea abies [L.] Karst) and European beech (Fagus sylvatica [L.]) trees, trenched for an experiment down to 1 m depth in a Luvisol soil (Parabraunerde) in 2010, and compared with 69 untrenched trees. The trenching was conducted prior to a long-term precipitation-exclusion treatment in a temperate mixed forest in Southern Germany. Stem and crown sizes, tree positions, distance to the trench, and annual stem growth of the 150 trees were measured from 2006 through 2013. Statistical analysis revealed that (1) stem growth reduction does not correlate with the loss of rooting area by trenching, (2) trenching does not significantly decrease the stem growth in the long-term post-trenching application, and (3) European beech showed a greater reaction in the first years after trenching as indicated by a greater decrease in stem growth. The relationship between stem growth reduction and the loss of rooting area arises during the first year post-trenching, but diminishes by the third year. The trenching effect on stem growth was strongest in trees at the plot edge and diminished towards the centre of the plots, where the actual study trees of the precipitation-exclusion experiment were located. The relevance of trenching for growth reduction is discussed, including potential underlying mechanisms and conclusions about experimental interferences.

Plants require the capacity for quick and precise recognition of external stimuli within their environment for survival. Upon exposure to biotic (herbivores and pathogens) or abiotic stressors (environmental conditions), plants can activate hydraulic, chemical, or electrical long-distance signals to initiate systemic stress responses. A plant's stress reactions can be highly precise and orchestrated in response to different stressors or stress combinations. To date, an array of information is available on plant responses to single stressors. However, information on simultaneously occurring stresses that represent either multiple, within, or across abiotic and biotic stress types is nascent. Likewise, the crosstalk between hydraulic, chemical, and electrical signaling pathways and the importance of each individual signaling type requires further investigation in order to be fully understood. The overlapping presence and speed of the signals upon plant exposure to various stressors makes it challenging to identify the signal initiating plant systemic stress/defense responses. Furthermore, it is thought that systemic plant responses are not transmitted by a single pathway, but rather by a combination of signals enabling the transmission of information on the prevailing stressor(s) and its intensity. In this review, we summarize the mode of action of hydraulic, chemical, and electrical long-distance signals, discuss their importance in information transmission to biotic and abiotic stressors, and suggest future research directions.

Absorptive fine roots are an important driver of soil biogeochemical cycles. Yet, the spatio-temporal dynamics of those roots in the presence of neighboring species remain poorly understood. The aim of this study was to analyze shifts in absorptive fine-root traits in monoculture or mixtures of Fagus sylvatica [L.] and Picea abies [L.] Karst. We hypothesized that root competition would be higher under single-species than mixed-species interactions, leading to changes in (i) root survivorship, diameter and respiration and (ii) spatio-temporal patterns of root growth and death. Using minirhizotron methods, we monitored the timing and location of absorptive fine-root growth and death at an experimental forest in southern Germany from 2011 to 2013. We also measured root respiration in the spring and fall seasons of 2012 and 2013. Our findings show that the absorptive fine roots of F. sylvatica had a 50% higher risk of root mortality and higher respiration rates in the single-species compared to mixed-species zones. These results support our hypothesis that root competition is less intense for F. sylvatica in mixture versus monoculture. We were unable to find confirmation for the same hypothesis for P. abies. To analyze spatio-temporal patterns of absorptive fine-root production and mortality, we used a mixed-effects model considering root depth (space) and seasons (time) simultaneously. This analysis showed that F. sylvatica shifts root production towards shallower soil layers in mixed-species stands, besides significant seasonal fluctuations in root production depths for both species. Ultimately, the impact of neighbor species identity on root traits observed in this study has important implications for where, when and how fast root-facilitated carbon cycling takes place in single-species versus mixed-species forests. In addition, our study highlights the need for inclusion of absorptive fine-root spatio-temporal dynamics when examining belowground plant interactions and biogeochemical cycles.

1. Context. Precipitation regimes are changing in response to climate change, yet understanding of how forest ecosystems respond to extreme droughts and pluvials remains incomplete. As future precipitation extremes will likely fall outside the range of historical variability, precipitation manipulation experiments (PMEs) are critical to advancing knowledge about potential ecosystem responses. However, few PMEs have been conducted in forests compared to short‐statured ecosystems, and forest PMEs have unique design requirements and constraints. Moreover, past forest PMEs have lacked coordination, limiting cross‐site comparisons. Here, we review and synthesize approaches, challenges, and opportunities for conducting PMEs in forests, with the goal of guiding design decisions, while maximizing the potential for coordination.2. Approach. We reviewed 63 forest PMEs at 70 sites worldwide. Workshops, meetings, and communications with experimentalists were used to generate and build consensus around approaches for addressing the key challenges and enhancing coordination.3. Results. Past forest PMEs employed a variety of study designs related to treatment level, replication, plot and infrastructure characteristics, and measurement approaches. Important considerations for establishing new forest PMEs include: selecting appropriate treatment levels to reach ecological thresholds; balancing cost, logistical complexity, and effectiveness in infrastructure design; and preventing unintended water subsidies. Response variables in forest PMEs were organized into three broad tiers reflecting increasing complexity and resource intensiveness, with the first tier representing a recommended core set of common measurements.4. Conclusions. Differences in site conditions combined with unique research questions of experimentalists necessitate careful adaptation of guidelines for forest PMEs to balance local objectives with coordination among experiments. We advocate adoption of a common framework for coordinating forest PME design to enhance cross‐site comparability and advance fundamental knowledge about the response and sensitivity of diverse forest ecosystems to precipitation extremes.

Background and Aims The objective of this study was to assess the effect of root pruning (RP) and annual under-trellis cover crops (UTCC) on vegetative growth and physiological processes in Cabernet Franc (Vitis vinifera L.) grapevines grafted on 3309C rootstock. Methods and Results Five root-zone management treatments, namely under-trellis vegetation-free strip control (C), RP and three distinct annual UTCC [annual ryegrass (AR), buckwheat, (BW) and turnip (TP)] were evaluated and compared over three seasons in a vigorous vineyard in the Northeastern USA. Root pruning had the greatest impact in reducing grapevine size; AR reduced the duration of shoot growth 2 out of 3 years. Competition between grapevine and AR and BW fine roots resulted in a lower grapevine fine root production in shallow soil layers and in a decrease in fine root lifespan. Conclusions Root pruning was effective in reducing grapevine size for 3 years; RP effectiveness, however, may vary greatly depending on site conditions. Annual UTCC hold potential for reducing grapevine size in vigorous plantings in regions with ample precipitation. Appropriate choice of cover crop species, however, remains unclear. Significance of the Study This work demonstrates that root-zone management practices impact grapevine vegetative growth and induce changes in root morphological traits.

Summary Non-native understorey woody species have been shown to extend leaf display and inhabit vacant phenological niches in early spring and late autumn when growing with native counterparts in temperate deciduous forests across the world. Despite the potential competitive advantages, extended leaf duration also subjects non-native species to possible hydraulic risks associated with maintaining leaves during periods of increased frost probability. It remains unclear how non-native species are able to maintain xylem function within this context. Leaf phenology in temperate deciduous trees has been shown to be a function of xylem anatomy, with earlier bud break associated with smaller xylem vessels due to the presumed resistance of smaller vessels to freezing-induced cavitation. We examined relationships between leaf phenology and xylem vessel traits across 82 native and non-native understorey deciduous woody species common to eastern U.S. deciduous forests. We hypothesized that non-native species possess xylem vessel traits associated with maximum hydraulic safety during frost-prone spring and autumn leaf display without compromising rapid growth rate. Larger metaxylem vessels in non-native species were associated with both faster spring growth and delayed autumn leaf fall compared to native species. Non-native species also had smaller latewood vessel diameter, latewood vessel area percentage and a higher proportion of solitary vessels in the entire secondary xylem cross section compared to natives, potentially increasing their resistance to freezing- and/or drought-induced cavitation in autumn, thus allowing for delayed autumn leaf fall. Native and non-native species exhibited similar dates of spring bud break and leaf emergence, consistent with similar xylem vessel size and vessel area percentage within metaxylem and earlywood. Within both groups, species with earlier bud and leaf emergence had a higher total percentage of vessel area within metaxylem and earlywood. This suggests understorey species need sufficient water to support their early spring growth at the risk of freezing-induced cavitation. Our study suggests xylem vessel properties, along with cross-sectional spatial xylem vessel distribution, reflect the capacity of non-native plants to thrive in a new environment and deepen our understanding of the physiological mechanisms of successful invasions of non-native understorey woody plant species.

Root–soil interactions fundamentally affect the terrestrial carbon (C) cycle and thereby ecosystem feedbacks to climate change. This study addressed the question of whether the secondary metabolism of different temperate forest tree species can affect soil microbial respiration. We hypothesized that phenolics can both increase and decrease respiration depending on their function as food source, mobilizer of other soil resources, signaling compound, or toxin. We analyzed the phenolic compounds from root exudates and root tissue extracts of six tree species grown in a glasshouse using high-performance liquid chromatography. We then tested the effect of individual phenolic compounds, representing the major identified phenylpropanoid compound classes, on microbial respiration through a 5-d soil incubation. Phenolic root profiles were highly species-specific. Of the eight classes identified, flavonoids were the most abundant, with flavanols being the predominating sub-class. Phenolic effects on microbial respiration ranged from a 26% decrease to a 46% increase, with reduced respiration occurring in the presence of compounds possessing a catechol ring. Tree species variation in root phenolic composition influences the magnitude and direction of root effects on microbial respiration. Our data support the hypothesis that functional group rather than biosynthetic class determines the root phenolic effect on soil C cycling.

In our study, we investigated whether root hydraulic conductance is related to post-transplant recovery. We used two Quercus species that differ in their transplant ability, Q. bicolor and Q. macrocarpa. Q. bicolor easily survives transplanting, whereas Q. macrocarpa often does not. We compared root hydraulic conductance after transplanting between control (without root pruning) and root-pruned, 1-year-old, small-caliper trees. We also examined the effects of transplant timing on post-transplant recovery of large-caliper trees. Hydraulic conductance in fine roots was correlated with recovery of the two Quercus species after transplanting. Six months after transplanting, small-caliper Q. bicolor trees had similar specific hydraulic conductance (KS) in fine roots compared with the KS before root-pruning, whereas fine root KS in small-caliper Q. macrocarpa trees decreased. Lower pre-dawn and midday xylem water potential in root-pruned Q. macrocarpa 6 weeks after transplanting indicates that root-pruned Q. macrocarpa experienced transplanting-induced water stress. For large-caliper trees, all Q. macrocarpa trees exhibited typical symptoms of transplant shock regardless of transplant timing, which was the result of higher vulnerability to mild water stress compared with Q. bicolor, resulting in a large reduction in fine root KS. Fine root KS in spring-transplanted Q. bicolor trees was much higher than that in fall-transplanted trees, implying spring transplanting is optimal for Q. bicolor. Other intrinsic characteristics of the species should be considered in the future when making better decisions on transplant timing such as xylem anatomy, carbon storage, rhizosphere conditions, and plant growth.

Tree root systems are inherently dynamic in their distribution within a soil volume. Analysis of tree root system space occupation through time can improve not only our implicit understanding of a virtually hidden portion of a plant, but influence future management decisions through a more thorough understanding of root placement within a soil volume. We compared root standing crop populations of four ornamental tree species including Acer rubrum L. ‘Franksred’ (Acer), Carpinus betula L. ‘Columnaris’ (Carpinus), Gleditsia tricanthos L. var. inermis ‘Skycole’ (Gleditsia), and Quercus rubra L. ‘Rubrum’ (Quercus) grown in a nursery mix substrate within large 57-L containers using an X-ray computed tomography (CT) approach through time. Individual root identification was performed manually on two-dimensional slices of CT scans. Our data show high variation in species total root number through time with Carpinus exhibiting the largest root population throughout the study period. However, species exhibited differences in root distribution patterns as exemplified by the shallow and horizontally more uniform rooting pattern of Acer in comparison with the highly plastic root distribution in space through time in Gleditsia. Root frequencies within 1-mm root diameter class distributions shifted by species with the most drastic differences found between high frequencies of relatively small diameter roots in Acer vs. pronounced shifts in dominate root diameter size class as found in Gleditsia and lesser so in Carpinus during a growing season. Our findings demonstrate differences in whole tree root systems space occupation non-destructively through time and highlight a disparity in how species fill a container volume during growth.

Aims Plant tolerance to herbivory has often been linked to plant growth rate, with faster growing plants that present high tissue turnover rates expected to be more tolerant than slower-growing plants. We tested whether this relationship also holds for rootstock growth rate and tolerance to apple replant disease (ARD).

Substrate moisture sensors offer an affordable monitoring system for containerized tree production. However, root system distribution can vary greatly among species within ornamental container production systems, resulting in variation within substrate readings among sensors within a container. The aim of this study was to examine the relationship of substrate moisture sensor readings in six ornamental trees to their root distribution patterns within a container. Following root anatomical analysis, tree root systems were dissected by root order as a means to separate fine (uptake) roots and coarse (transport) roots. Substrate moisture variability was measured through the deployment of 12 substrate moisture sensors per container. Of the tree species studied, we found the following two patterns of root distribution: a shallow, “conical-shaped,” root system, with the broadest portion of the root system in the shallow soil layer, and a more evenly distributed “cylindrical-shaped” root system. Root system distribution type influenced substrate moisture reading variability. Conical root systems had lower substrate moisture variability and high fine root variability, whereas the opposite was true for cylindrical root systems—most likely due to the larger, coarse woody mass of roots. We were unable to find any correlations between fine root morphological features including root diameter, length, or surface area and substrate moisture variability. However, higher specific root length was associated with higher substrate moisture variability. Classifying a tree’s root system by its growth and distribution within a container can account for variation in substrate moisture readings and help inform future decisions on sensor placement within containerized systems.

Root development may exert control on plant-pathogen interactions with soil-borne pathogens by shaping the spatial and temporal availability of susceptible tissues and in turn the impact of pathogen colonization on root function. To evaluate the relationship between root development and resistance to apple replant disease (ARD) pathogens, pathogen abundance was compared across root branching orders in a bioassay with two rootstock genotypes, M.26 (highly susceptible) and CG.210 (less susceptible). Root growth, anatomical development and secondary metabolite production were evaluated as tissue resistance mechanisms. ARD pathogens primarily colonized first and second order roots, which corresponded with cortical tissue senescence and loss in second and third order roots. Defense compounds were differentially allocated across root branching orders, while defense induction or stress response was only detected in first order and pioneer roots. Our results suggest disease development is based largely on fine-root tip attrition. In accordance, the less susceptible rootstock supported lower ARD pathogen abundance and altered defense compound production in first order and pioneer roots and maintained higher rates of root growth in both the ARD soil and pasteurized control compared to the more susceptible. Thus, this rootstock's ability to maintain shoot growth in replant soil may be attributable to relative replant pathogen resistance in distal root branches as well as tolerance of infection based on rates of root growth.

PREMISE OF THE STUDY Plant phenology influences resource utilization, carbon fluxes, and interspecific interactions. Although controls on aboveground phenology have been studied to some degree, controls on root phenology are exceptionally poorly understood. METHODS We used minirhizotrons to examine the timing of grape root production over 5 yr in Fredonia, New York, USA, in a humid continental climate; and over 3 yr in Oakville, California, USA, in a Mediterranean climate. We used data from previous experiments to examine the relationship of root phenology with aboveground phenology. We compared interannual variability in root and shoot growth and determined the influence of abiotic factors on the timing of root initiation, peak root standing crop, peak root growth rate, and cessation of root growth. KEY RESULTS Root phenology was not tightly coupled with aboveground phenological periods. Both sites typically had one yearly root flush and high interannual variability in root growth. Root phenology was more variable in California than in New York. In this and other published studies, interannual variation in root phenology was greater than variation in aboveground phenology. The three phenological phases of root growth-root initiation, peak root growth, and root cessation-were related to different suites of abiotic factors. CONCLUSIONS Root phenology is highly variable among years. Analysis of potential controlling factors over several years suggest that belowground phenological phases should be analyzed separately from each other. If aboveground grape phenology responds differently than belowground phenology to changes in air temperature, global warming may further uncouple the timing of aboveground and belowground growth.

Experiential learning can be an important part of an undergraduate curriculum in the sciences. A new course, The Nature of Plants, was developed to provide students across a broad range of majors with an in-depth study of plant science both basic and applied. The course was enriched by using a local natural area as an informal learning environment. We examined whether experiential learning improved homework scores among students who participated in a field trip by asking if simply attending the field trip increased the homework score or if participation in the tree climbing exercise had any additional benefit. Our results show participating in a field trip experience when coupled with a homework assignment increased student homework scores. Moreover, the tree climbing portion of the field trip increased homework scores particularly for students not in a science major. This research supports experiential learning and the value of field trips within science courses focused on a comprehensive exploration of plants.

Fungal endophytes have significant effects on host performance including but not limited to changes in reproductive output, as well as, biotic and abiotic stress tolerance. The ultimate mechanisms for resistance to herbivory have been documented in a number of fungal-plant symbiota and involve the production of alkaloids by the fungus. Alkaloids have thus been defined as a currency responsible for increased host resistance to herbivory. We provide support for the hypothesis that another currency may be responsible for increased host tolerance to abiotic stress resulting from endophyte colonization; namely antioxidants. We report comparatively higher activity of antioxidants in endophyte colonized hosts resulting from abiotic stress and propose antioxidants are another currency via which mutualistic interactions between fungal endophytes and their hosts can occur. We recognize fungal endophyte interactions are diverse and complex and include antagonisms, commensalisms, and mutualisms.

Summary 1. Root herbivores and pathogens interfere with basic below-ground plant function, and can thereby affect plant fitness and spatial and temporal patterns in natural plant communities. However, there has been little development of concepts and theories on below-ground plant defence, a deficit that is in contrast to the abundance of theorizing for above-ground plant parts. 2. A review of the past 10 years of research on below-ground plant–herbivore interactions has revealed that, similar to above-ground tissues, root defences can be expressed constitutively or induced upon herbivore attack, and can be classified into direct and indirect traits, tolerance, and escape. Indeed, it has been shown that roots tolerate herbivory by outgrowing or re-growing lost tissues, or resist it by producing secondary metabolites that are toxic to herbivores or attract natural enemies of herbivores. 3. We propose that, similar to above-ground plant–herbivore theories, the partition of abiotic and biotic factors over ecological succession can serve as the basis for predicting investment in defence strategies below-ground. 4. Investigation of herbivore pressure and root responses along primary and secondary successional gradients suggests that: (i) roots are often fast growing, thinner and softer in early compared to later succession. (ii) Insect and nematode herbivore pressure increases until mid-succession and later decreases. (iii) Mycorrhizal abundance increases with succession, and the composition of fungal species changes through succession, often shifting from arbuscular mycorrhizae to ecto-mycorrhizae. 5. Based on these findings, and on classical (above-ground) plant defence theory, we suggest the following set of testable hypotheses for below-ground plant defence: (i) During succession, early plants invest most of their resources in growth and less in defences (associated with a general lack of herbivores and pathogens, and with limited availability of resources in the system), therefore relying more on re-growth (tolerance) strategies. (ii) During mid-succession, a buildup of herbivore pressure facilitates replacement by plant species that exhibit greater direct and indirect defence strategies. (iii) Constitutive and inducible levels of defences may trade-off, and early successional plants should rely more on induction of defences after herbivore attack, whereas late successional plants will increasingly rely on constitutively produced levels of physical and chemical defence. (iv) Successional changes in microbial associations have consequences for root defence by improving plant nutrition and defence expression as well as directly competing for root space; however, toxic or impenetrable root defences may also limit association with root symbionts, and so may constrain the expression of root defence.

The role of root systems in drought tolerance is a subject of very limited information compared with above-ground responses. Adjustments to the ability of roots to supply water relative to shoot transpiration demand is proposed as a major means for woody perennial plants to tolerate drought, and is often expressed as changes in the ratios of leaf to root area (A(L):A(R)). Seasonal root proliferation in a directed manner could increase the water supply function of roots independent of total root area (A(R)) and represents a mechanism whereby water supply to demand could be increased. To address this issue, seasonal root proliferation, stomatal conductance (g(s)) and whole root system hydraulic conductance (k(r)) were investigated for a drought-tolerant grape root system (Vitis berlandieri×V. rupestris cv. 1103P) and a non-drought-tolerant root system (Vitis riparia×V. rupestris cv. 101-14Mgt), upon which had been grafted the same drought-sensitive clone of Vitis vinifera cv. Merlot. Leaf water potentials (ψ(L)) for Merlot grafted onto the 1103P root system (-0.91±0.02 MPa) were +0.15 MPa higher than Merlot on 101-14Mgt (-1.06±0.03 MPa) during spring, but dropped by approximately -0.4 MPa from spring to autumn, and were significantly lower by -0.15 MPa (-1.43±0.02 MPa) than for Merlot on 101-14Mgt (at -1.28±0.02 MPa). Surprisingly, g(s) of Merlot on the drought-tolerant root system (1103P) was less down-regulated and canopies maintained evaporative fluxes ranging from 35-20 mmol vine(-1) s(-1) during the diurnal peak from spring to autumn, respectively, three times greater than those measured for Merlot on the drought-sensitive rootstock 101-14Mgt. The drought-tolerant root system grew more roots at depth during the warm summer dry period, and the whole root system conductance (k(r)) increased from 0.004 to 0.009 kg MPa(-1) s(-1) during that same time period. The changes in k(r) could not be explained by xylem anatomy or conductivity changes of individual root segments. Thus, the manner in which drought tolerance was conveyed to the drought-sensitive clone appeared to arise from deep root proliferation during the hottest and driest part of the season, rather than through changes in xylem structure, xylem density or stomatal regulation. This information can be useful to growers on a site-specific basis in selecting rootstocks for grape clonal material (scions) grafted to them.

Understanding factors controlling root dynamics and functioning can lead to more efficient and profitable vineyard management. However, our current understanding of root dynamics and their regulation by plant and environmental factors is limited, particularly under field conditions. This paper presents current understanding of grape root dynamics, highlighting studies using minirhizotron cameras, which directly assess root dynamics, and experiments on roots of known age, which link root phenology and function. Data summarised here show timing of grape root production varies widely among different regions, as well as among rootstocks and canopy management systems in the same region. Timing of production can be responsive to differences in soil moisture. Lifespan of grape roots, however, appears less affected by soil moisture because of nocturnal hydraulic redistribution. Root function, such as capacity for P and N uptake, declines rapidly with root age. Differences in timing and spatial distribution of root production can effect above-ground growth and vineyard water-use efficiency. Improving our understanding of when roots grow and are functionally active in agricultural systems can lead to improved water and fertiliser applications, and more precise vineyard management. Because both environmental and biological factors affect root dynamics, simple predictions of timing of root production or standing populations with shoot development are unlikely to be achieved. However, with multi-year data on root dynamics, and environmental and biological factors, regionally specific models of root populations and their functioning may be possible to develop.

Aims: The objectives of this study were to evaluate (1) the fertilizer potential of bone char, (2) the effects of wood biochar on plant-available phosphorus (P), and (3) the role of root-mycorrhizae-biochar interactions in plant P acquisition from a P-fixing soil. Methods: Incubation and pot experiments were conducted with a P-fixing soil and maize with or without root hairs and arbuscular mycorrhizae (AM) inoculation. Olsen-, resin-P and plant P accumulation were used to estimate P availability from bone char, co-pyrolyzed bone char-wood biochar, and separate bone char and wood biochar additions produced at 60, 350 and 750 °C, and Triple Superphosphate (TSP). Results: Maize inoculated with AM showed similar P accumulation when fertilized with either 750 °C bone char or TSP. Pyrolyzing bone did not increase extractable P in soil in comparison to unpyrolyzed bone, apart from a 67 % increase in resin-extractable P after additions of bone char pyrolyzed at 350 °C. Despite greater Olsen-P extractability, co-pyrolysis of bone with wood reduced maize P uptake. Wood biochars reduced resin-P from bone char by 14–26 %, whereas oven-dried wood increased resin-P by 23 %. Conclusions: Bone char is an effective P fertilizer, especially if root-AM interactions are simultaneously considered. Biochar influences plant access to soil P and requires careful management to improve P availability.

Herbivory tolerance has been linked to plant growth rate where plants with fast growth rates are hypothesized to be more tolerant of herbivory than slower-growing plants. Evidence supporting this theory has been taken primarily from observations of aboveground organs but rarely from roots. Grapevines differing in overall rates of new root production, were studied in Napa Valley, California over two growing seasons in an established vineyard infested with the sucking insect, grape phylloxera (Daktulosphaira vitifoliae Fitch). The experimental vineyard allowed for the comparison of two root systems that differed in rates of new root tip production (a 'fast grower', Vitis berlandieri x Vitis rupestris cv. 1103P, and a slower-growing stock, Vitis riparia x Vitis rupestris cv. 101-14 Mgt). Each root system was grafted with a genetically identical shoot system (Vitis vinifera cv. Merlot). Using minirhizotrons, we did not observe any evidence of spatial or temporal avoidance of insect populations by root growth. Insect infestations were abundant throughout the soil profile, and seasonal peaks in phylloxera populations generally closely followed peaks in new root production. Our data supported the hypothesis that insect infestation was proportional to the number of growing tips, as indicated by similar per cent infestation in spite of a threefold difference in root tip production. In addition, infested roots of the fast-growing rootstock exhibited somewhat shorter median lifespans (60 d) than the slower-growing rootstock (85 d). Lifespans of uninfested roots were similar for the two rootstocks (200 d). As a consequence of greater root mortality of younger roots, infested root populations in the fast-growing rootstock had an older age structure. While there does not seem to be a trade-off between potential growth rate and relative rate of root infestation in these cultivars, our study indicates that a fast-growing root system may more readily shed infested roots that are presumably less effective in water and nutrient uptake. Thus, differences in root tip production may be linked to differences in the way plants cope with roots that are infested by sucking insects.

Research in the field of plant biology has recently demonstrated that inter- and intra-specific interactions belowground can dramatically alter root growth. Our aim was to answer questions related to the effect of inter- vs. intra-specific interactions on the growth and utilization of undisturbed space by fine roots within three dimensions (3D) using micro X-ray computed tomography. To achieve this, Populus tremuloides (quaking aspen) and Picea mariana (black spruce) seedlings were planted into containers as either solitary individuals, or inter-/intra-specific pairs, allowed to grow for two months, and 3D metrics developed in order to quantify their use of belowground space. In both aspen and spruce, inter-specific root interactions produced a shift in the vertical distribution of the root system volume, and deepened the average position of root tips when compared to intra-specifically growing seedlings. Inter-specific interactions also increased the minimum distance between root tips belonging to the same root system. There was no effect of belowground interactions on the radial distribution of roots, or the directionality of lateral root growth for either species. In conclusion, we found that significant differences were observed more often when comparing controls (solitary individuals) and paired seedlings (inter- or intra-specific), than when comparing inter- and intra-specifically growing seedlings. This would indicate that competition between neighboring seedlings was more responsible for shifting fine root growth in both species than was neighbor identity. However, significant inter- vs. intra-specific differences were observed, which further emphasizes the importance of biological interactions in competition studies.

Growers plan many of their horticultural activities around certain shoot phenological stages such as bloom and veraison. Timing of root growth in relation to these stages of the shoot is of interest in fertilization scheduling and understanding carbon allocation demands of the root system. With the recent use of minirhizotron root observation tubes, a much greater understanding of patterns of root growth has been made possible. In this paper some published work and present data from ongoing studies on timing of root production are reviewed. Studies discussed include ongoing work in grape vineyards in Fredonia, New York, USA (Vitis lambruscana), and Oakville, California, USA (V. vinifera), and in apple orchards (Malus × domestica) in Summerland, British Columbia, Canada. Root production mainly occurs in the summer, but the specific timing varies widely between bloom and harvest from year to year. Usually there is little root growth in the few weeks prior to harvest and when the vines are dormant. Cultural practices may affect root production, usually from a combination of changes in the soil environment (moisture, temperature) and changes in photosynthetic capacity.

Eight red maple (Acer rubrum L.) provenances, four each from wet and dry sites, were grown under the same conditions and their physiological responses to soil water availability investigated. Under well-watered conditions, seedlings of wet-site provenances grew faster and had consistently higher net photosynthesis, leaf conductance, maximum carboxylation rate, maximum rate of coupled photosynthetic electron transport, apparent quantum use efficiency, light-saturated photosynthesis and dark respiration than seedlings of dry-site provenances. Under conditions of low soil water availability, only dry-site provenances responded with decreased osmotic potential at full hydration and at the turgor loss point; however, provenances from wet sites showed a smaller reduction in absolute growth rate, a greater reduction in gas exchange and a greater increase in abscisic acid concentrations than dry-site provenances.