Neutron imaging reveals internal plant water dynamics

Background and aims

Knowledge of plant water fluxes is critical for assessing mechanistic processes linked to biogeochemical cycles, yet resolving root water transport dynamics has been a particularly daunting task. Our objectives were to demonstrate the ability to non-invasively monitor individual root functionality and water fluxes within Zea mays L. (maize) and Panicum virgatum L. (switchgrass) seedlings using neutron imaging.

Methods

Seedlings were propagated for 1–3 weeks in aluminum chambers containing sand. Pulses of water or deuterium oxide were then tracked through the root systems by collecting consecutive radiographs during exposure to a cold-neutron source. Water flux was manipulated by cycling on a growth lamp to alter foliar demand for water.

Results

Neutron radiography readily illuminated root structure, root growth, and relative plant and soil water content. After irrigation there was rapid root water uptake from the newly wetted soil, followed by hydraulic redistribution of water through the root system to roots terminating in dry soil. Water flux within individual roots responded differentially to foliar illumination based on supply and demand of water within the root system.

Conclusions

Sub-millimeter scale image resolution revealed timing and magnitudes of root water uptake, redistribution within the roots, and root-shoot hydraulic linkages—relationships not well characterized by other techniques.     

Uptake of water from a Kandosol subsoil. II. Control of water uptake by roots

Aim

To test for the presence of an impediment to water flow at the soil-root interface.

Methods

Wheat plants were grown in repacked and undisturbed field soil. Their transpiration rate, E, was varied in several steps from low to high and then back to low again, while the hydrostatic pressure in the leaf xylem, ψ _xylem, was measured non-destructively and continuously. These measurements were compared to a mathematical model that calculated ψ _xylem by assuming that the hydraulic resistance across the plant was constant and that the radial flow of water to unit length of a typical plant root generated gradients in pressure in the soil water.

Results

For the repacked soil, the radial flow model could not match the experiment during the falling phase of E, unless it was assumed that either an additional, constant, interfacial resistance between the soil and the roots had developed when E was large and ψ _xylem was rapidly falling, or that the resistance within the plant had changed. For the undisturbed field soil, the radial flow model did not agree with the experiment. Plausible agreement was achieved when plant water uptake was accounted for using a distributed sink model in HYDRUS-1D, with E integrated across the rootzone. This approach was based on the measured large variation in the vertical distribution of roots.

Conclusions

There was no strong evidence of large drawdowns of soil water in the rhizosphere, even when ψ _xylem was falling rapidly when E was large and the soil was moderately dry. Thus, there seems to have been an additional impediment to water flow from soil to plant, either within the plant, or at the interface between the two.     

Uptake of water from a Kandosol subsoil: I. Determination of soil water diffusivity

Aims

To determine soil water diffusivity, D(θ), on undisturbed field soil at medium to low water content (suction range from 10 to 150 m of water), for the purpose of modeling the uptake of water by plant roots.

Methods

The method is based on the analysis of one-step outflow induced by a turbulent stream of dry air over the exposed end of a soil core, with the other end of the core enclosed. The outflow is measured through time as the change in the weight of the core as it sits on a recording balance. D(θ) is calculated by deconvoluting the measured outflow function.

Results

Over the suction range of 10 to 150 m of water, D(θ) calculated on the undisturbed soil ranged from 20?×?10^?9 to 10?×?10^?9 [m²?s^?1], substantially higher than other published estimates over this range in suction.

Conclusions

These unusually large values cast doubt on the view that flow of water to roots limits uptake of water from the targeted subsoil.     

Photosynthetic responses to soil water stress in summer in two Japanese urban landscape tree species (<Emphasis Type="Italic">Ginkgo biloba</Emphasis> and <Emphasis Type="Italic">Prunus yedoensis</Emphasis>): effects of pruning mulch and irrigation management

Key message

Stomatal regulation involves beneficial effects of pruning mulch and irrigation on leaf photosynthesis in Prunus yedoensis and Ginkgo biloba under moderate drought. G. biloba showed conservative water use under drought.

Abstract

Leaf photosynthesis is highly sensitive to soil water stress via stomatal and/or biochemical responses, which markedly suppress the growth of landscape trees. Effective irrigation management to maintain leaf photosynthesis and information on species-specific photosynthetic responses to soil water stress are essential for the sustainable management of landscape trees in Japan, in which summer drought often occurs. In order to investigate effective irrigation management, we used plants with moderate soil water stress as controls, and examined the effects of daily irrigation and pruning mulch on leaf photosynthesis in container-grown Ginkgo biloba and Prunus yedoensis, which are the first and second main tall roadside trees in Japan. Stomatal conductance was significantly increased by pruning mulch and daily irrigation, with similar increases in leaf photosynthesis being observed in P. yedoensis and G. biloba. In order to obtain information on species-specific photosynthetic responses to soil water stress, we compared the responses of leaf photosynthesis and leaf water status to reductions in soil water content (SWC) between the two species. G. biloba maintained a constant leaf water potential, leaf water content, maximum carboxylation rate, and electron transport rate with reductions in SWC, whereas reductions were observed in P. yedoensis. We concluded that pruning mulch and irrigation effectively offset the negative impact of moderate water stress on leaf photosynthesis in summer in P. yedoensis and G. biloba via stomatal regulation, and also that G. biloba maintained its photosynthetic biochemistry and leaf water status better than P. yedoensis under severe water stress.

     

Longleaf and loblolly pine seedlings respond differently to soil compaction,water content,and fertilization

Aims

Longleaf pine (Pinus palustris Mill.) is being restored across the U.S. South for a multitude of ecological and economic reasons, but our understanding of longleaf pine’s response to soil physical conditions is poor. On the contrary, our understanding of loblolly pine (Pinus taeda L.) root and shoot growth response to soil conditions is well established.

Methods

We performed a comparative greenhouse study which modeled root length density, total seedling biomass, and the ratio of aboveground:belowground mass as functions of volumetric water content, bulk density and soil fertility (fertilized or not).

Results

Root length density was about 35 % greater in longleaf pine seedlings compared to loblolly pine seedlings, and was reasonably well modeled (R ²?=?0.54) for longleaf pine by bulk density (linear), volumetric water content (quadratic), soil fertility, and the interactions of bulk density, volumetric water content, species, and soil fertility. The aboveground:belowground mass ratio (ABR) increased at both extremes of water content.

Conclusions

This research indicates that young longleaf pine seedling root systems respond more negatively to extremes of soil physical conditions than loblolly pine, and compacted or dry loamy soils should be ameliorated in addition to normal competition control, especially on soils degraded by past management.     

Impact of soil texture and water availability on the hydraulic control of plant and grape-berry development

Aims

All components of the soil-plant-atmosphere (s-p-a) continuum are known to control berry quality in grapevine (Vitis vinifera L.) via ecophysiological interactions between water uptake by roots and water loss by leaves. The scope of the present work was to explore how the main hydraulic components of grapevine influence fruit quality through changes in liquid- and gas-phase hydraulic conductance.

Methods

To reach our objectives, determinations of shoot growth, berry size and sugar content, leaf gas exchange, predawn leaf water potential (as a proxy of soil water potential), midday stem water potential and leaf water potential were performed in conjunction with anatomical measurements of shoot xylem. All measurements were conducted in two different cultivars (Cabernet franc and Merlot) and on three different soil types (clayey, gravelly, and sandy).

Results

Shoot xylem morphometric characteristics and whole-plant hydraulic conductance were influenced by cultivar and soil type. Differences in leaf gas exchange parameters and water potentials were determined by soil type significantly more than by cultivar. Between the two extremes (gravelly soil imposing drought conditions and sandy soil with easily accessible water) the clayey soil expressed an intermediate plant water consumption and highest sugar accumulation in berry.

Conclusions

Hydraulic and non hydraulic limitations to vine/berry interactions supported the conclusion that water availability in the soil overrides differences due to cultivar in determining the productive potential of the vineyard. Non hydraulic stomatal control was expected to be an important component on plants grown on the clayey soil, which experienced a moderate water stress. Possible links between hydraulic traits and berry development and quality are discussed.     

Mycorrhization and phosphorus nutrition affect water relations and CAM induction by drought in seedlings of Clusia minor

Background and Aims

Crassulacean acid metabolism (CAM) is currently viewed as an adaptation to water deficit. In plants of Clusia minor, which grow mostly on acidic, P-deficient soils, CAM is induced by water deficit. The symbiosis between plants and mycorrhizal fungi alleviates the symptoms of P deficiency and may influence drought resistance. Therefore, the effect of P supply, modified by three different experimental treatments, on the induction of CAM by drought in C. minor was investigated to test the hypothesis that P deficiency will produce greater CAM activity and, in addition, that treatment will modify drought tolerance.

Methods

Seedlings were grown in forest soil sterilized and inoculated with Scutellospora fulgida (SF treatment), sterilized and supplemented with P (Ph treatment) or non-sterilized and containing native mycorrhizae (Nat treatment). Leaf turgor potential (ψ_T) was determined psychrometrically, and CAM activity as nocturnal acid accumulation (ΔH⁺) by titration of dawn and dusk leaf sap.

Key Results

Plant mass and P content were higher in SF and Ph than in Nat seedlings. After 21 d of water deficit, ψ_T increased in SF, decreased in Ph and remained unchanged in Nat, and, after 7 and 14 d of water deficit, ΔH⁺ in Nat was three times higher than at the beginning of drought, whereas in SF and Ph ΔH⁺ was lower than on day 0.

Conclusions

P deficiency in Nat seedlings was ameliorated by inoculation or P addition. The SF and Nat seedlings showed greater tolerance of drought than Ph. P deficiency promoted the induction of CAM by drought in Nat seedlings, whereas P fertilization and mycorrhization did not. Nocturnal acid accumulation was highly and negatively correlated with plant P and N contents, indicating that P and N deficiencies are promoters of CAM in droughted plants of C. minor.Key words: Clusia minor, crassulacean acid metabolism, CAM, mycorrhiza, drought, phosphorus deficiency, nitrogen–water relations     

Modelling the impact of heterogeneous rootzone water distribution on the regulation of transpiration by hormone transport and/or hydraulic pressures

Aims

A simulation model to demonstrate that soil water potential can regulate transpiration, by influencing leaf water potential and/or inducing root production of chemical signals that are transported to the leaves.

Methods

Signalling impacts on the relationship between soil water potential and transpiration were simulated by coupling a 3D model for water flow in soil, into and through roots (Javaux et al. 2008) with a model for xylem transport of chemicals (produced as a function of local root water potential). Stomatal conductance was regulated by simulated leaf water potential (H) and/or foliar chemical signal concentrations (C; H?+?C). Split-root experiments were simulated by varying transpiration demands and irrigation placement.

Results

While regulation of stomatal conductance by chemical transport was unstable and oscillatory, simulated transpiration over time and root water uptake from the two soil compartments were similar for both H and H?+?C regulation. Increased stomatal sensitivity more strongly decreased transpiration, and decreased threshold root water potential (below which a chemical signal is produced) delayed transpiration reduction.

Conclusions

Although simulations with H?+?C regulation qualitatively reproduced transpiration of plants exposed to partial rootzone drying (PRD), long-term effects seemed negligible. Moreover, most transpiration responses to PRD could be explained by hydraulic signalling alone.     

Precipitation variability does not affect soil respiration and nitrogen dynamics in the understorey of a Mediterranean oak woodland

Background and aims

Future climate scenarios for the Mediterranean imply increasing precipitation variability. This study presents a large-scale water manipulation experiment simulating changes in precipitation variability, aiming at a better understanding of the effects of rainfall patterns on soil C and N cycling and understorey productivity in a Mediterranean oak woodland.

Methods

We used rain-out shelters to achieve (1) a normal dry period (7 days), and (2) a dry period increased three-fold (21 days), without altering total annual precipitation inputs.

Results

The temporal patterns of soil respiration (R _s) and soil inorganic N were not affected by treatment. However, water infiltration and N leaching increased with large infrequent watering events. R _s and soil NH₄ ⁺-N correlated with soil temperature, with soil NO₃ ^?-N being influenced by leaching.

Conclusions

The lack of significant treatment effects on either R _s or soil inorganic N can be explained by (1) minor differences in plant productivity between the treatments, suggesting equal plant N demand, and (2) the absence of moisture dependence of R _s and soil NH₄ ⁺-N. Increased N leaching with large infrequent precipitation events may have longer-term consequences for ecosystem functioning. Our results contribute to an improved understanding of possible climate change effects on key ecosystem processes in Mediterranean ecosystems.     

Shoot hydraulic characteristics, plant water status and stomatal response in olive trees under different soil water conditions

Aims

To evaluate the impact of the amount and distribution of soil water on xylem anatomy and xylem hydraulics of current-year shoots, plant water status and stomatal conductance of mature ‘Manzanilla’ olive trees.

Methods

Measurements of water potential, stomatal conductance, hydraulic conductivity, vulnerability to embolism, vessel diameter distribution and vessel density were made in trees under full irrigation with non-limiting soil water conditions, localized irrigation, and rain-fed conditions.

Results

All trees showed lower stomatal conductance values in the afternoon than in the morning. The irrigated trees showed water potential values around ?1.4 and ?1.6 MPa whereas the rain-fed trees reached lower values. All trees showed similar specific hydraulic conductivity (K _s) and loss of conductivity values during the morning. In the afternoon, K _s of rain-fed trees tended to be lower than of irrigated trees. No differences in vulnerability to embolism, vessel-diameter distribution and vessel density were observed between treatments.

Conclusions

A tight control of stomatal conductance was observed in olive which allowed irrigated trees to avoid critical water potential values and keep them in a safe range to avoid embolism. The applied water treatments did not influence the xylem anatomy and vulnerability to embolism of current-year shoots of mature olive trees.     

Impact of the plant community composition on labile soil organic carbon, soil microbial activity and community structure in semi-natural grassland ecosystems of different productivity

Aims

The main objective was to describe the effects of plant litter on SOC and on soil microbial activity and structure in extensively managed grasslands in Central Germany that vary in biomass production and plant community composition.

Methods

The decomposition of shoot and root litter was studied in an incubation experiment. Labile C and N were isolated by hot water extraction (C_HWE, N_HWE), while functional groups of microbes were identified by PLFA analysis and microbial activity was measured using a set of soil exo-enzymes.

Results

The plant community composition, particulary legume species affected SOC dynamics and below-ground microbial processes, especially via roots. This was reflected in about 20% lower decomposition of root litter in low productivity grassland soil. The C_HWE soil pool was found to be a key driver of the below-ground food web, controlling soil microbial processes.

Conclusions

Below-ground responses appear to be related to the presence of legume species, which affected the microbial communities, as well as the ratio between fungal and bacterial biomass and patterns of soil enzyme activity. Low productivity fungal-dominated grasslands with slow C turnover rates may play an important role in SOC accumulation. The approach used here is of particular importance, since associated biological and biochemical processes are fundamental to ecosystem functioning.     

Enhanced nodulation and symbiotic effectiveness of Medicago truncatula when co-inoculated with Pseudomonas fluorescens WSM3457 and Ensifer (Sinorhizobium) medicae WSM419

Aims

Low numbers of rhizobia in soil or inoculants delay nodulation and decrease symbiotic legume productivity. This study investigated the effect of co-inoculation with a helper bacterium, Pseudomonas fluorescens WSM3457 on the Medicago truncatula - Ensifer (Sinorhizobium) medicae WSM419 symbiosis challenged by a low inoculum dose.

Methods

In a glasshouse experiment the effect of co-inoculation with WSM3457 on the kinetics of nodule initiation and development was assessed 5, 7, 10, 14, 17, 21, and 42 days after inoculation of M. truncatula cv. Caliph with 10³ cells/plant of E. medicae WSM419.

Results

Co-inoculated plants had enhanced rate of nodule initiation and development, greater numbers of larger crown nodules, and by day 42 accumulated more N than plants inoculated with E. medicae WSM419 alone. Nodule development was altered by co-inoculation. Approximately 25% of nodule initials on co-inoculated plants formed in closely associated pairs, young nodules were larger with multiple meristems and developed into cluster-like multi-lobed nodules compared to those on WSM419 inoculated plants. Molecular typing showed WSM3457 occupied a significant proportion of root nodules on co-inoculated plants.

Conclusion

Co-inoculation with P. fluorescens WSM3457 enhanced symbiotic effectiveness of M. truncatula when inoculated with a low inoculum dose of E. medicae WSM419.     

Effects of soil acidity and water stress on corn and soybean performance under a no-till system

Background and Aims

Field studies have demonstrated that aluminum (Al) toxicity is low in no-till systems during cropping seasons that have adequate and well-distributed rainfall. This study evaluated the performance of corn (Zea mays L.) and soybean (Glycine max L. Merrill) on an acid loamy soil under a long-term no-till system, in response to surface liming and as affected by genotypic tolerance to Al and water stress.

Methods

A field trial examined the effect of surface application of lime (0, 4, 8, and 12 Mg ha^?1) on no-till corn and soybean nutrition and yield. Trials were also carried out in undisturbed soil columns taken from the unlimed and limed plots. Two hybrids/cultivars of corn and soybean, one sensitive and the other moderately sensitive to Al were grown at two soil moisture levels with and without water stress (50 % and 80 % water filled pore space).

Results

Alleviating soil acidity by liming improved nutrition and increased grain yields of corn and soybean. The benefits of liming on root length density, nutrient uptake and shoot biomass production of corn and soybean were more pronounced in Al-sensitive genotypes under water stress.

Conclusions

The results suggest that plants exposed to drought stress under no-till systems are more affected by Al toxicity.     

Root dynamics of Carex stricta-dominated tussock meadows

Background and aims

The roots of tussock-forming plants contribute to the formation of microtopographic features in many ecosystems, but the dynamics of such roots are poorly understood. We examined the spatial heterogeneity of tussock fine root dynamics to investigate allocation patterns and the role of root productivity in the persistence of tussock structures.

Methods

We compared the spatial variability of fine root (<1 mm, 1–2 mm) density, biomass, % live, allocation, turnover rate (using bomb 14C), and productivity of four Carex stricta Lam.-dominated tussock meadows in the upper Midwest, USA (3 reference, 1 restored site).

Results

Relative to underlying microsites, tussocks were warm, dry, and high in root density, productivity, % live biomass, and turnover. Root productivity averaged 649 g?m^?2 yr^?1 (±208) in reference sites, comprised 57 % (±10) of total net production, and was concentrated in tussocks (70 %?±?4). Root turnover rate averaged 0.63 yr^?1 (±0.08), but tussocks had ~50 % faster root turnover than the underlying soil, and <1 mm roots turned over ~40 % faster than 1–2 mm roots.

Conclusions

Our detailed analysis of the spatial heterogeneity of tussock root dynamics suggests that high allocation and elevated turnover of tussock roots facilitates organic matter accumulation and tussock persistence over time.     

Effects of competition on induction of crassulacean acid metabolism in a facultative CAM plant

Abiotic drivers of environmental stress have been found to induce CAM expression (nocturnal carboxylation) in facultative CAM species such as Mesembryanthemum crystallinum. The role played by biotic factors such as competition with non-CAM species in affecting CAM expression, however, remains largely understudied. This research investigated the effects of salt and water conditions on the competition between M. crystallinum and the C₃ grass Bromus mollis with which it is found to coexist in California’s coastal grasslands. We also investigated the extent to which CAM expression in M. crystallinum was affected by the intensity of the competition with B. mollis. We found that M. crystallinum had a competitive advantage over B. mollis in drought and saline conditions, while B. mollis exerted strong competitive effects on M. crystallinum in access to light and soil nutrients in high water conditions. This strong competitive effect even outweighed the favorable effects of salt or water additions in increasing the biomass and productivity of M. crystallinum in mixture. Regardless of salt conditions, M. crystallinum did not switch to CAM photosynthesis in response to this strong competitive effect from B. mollis. Disturbance (i.e., grass cutting) reduced the competitive pressure by B. mollis and allowed for CAM expression in M. crystallinum when it was grown mixed with B. mollis. We suggest that moderate competition with other functional groups can enhance CAM expression in M. crystallinum, thereby affecting its plasticity and ability to cope with biological stress.     

Does geographic origin dictate ecological strategies in Acacia senegal (L.) Willd.? Evidence from carbon and nitrogen stable isotopes

Background and aims

Acacia senegal, a leguminous dryland tree, is economically and ecologically important to sub-Saharan Africa. Water-use efficiency (WUE) and biological nitrogen fixation (BNF) are fundamental to plant productivity and survival. We quantify provenance differences in WUE, BNF, photosynthesis, biomass and gum arabic production from A. senegal assessing genetic improvement potential.

Methods

Using stable isotope ratios, we determined WUE (δ¹³C) and BNF (δ¹⁵N) from provenances of mature A. senegal in field-trials (Senegal), sampling leaves at the beginning (wet) and end (dry) of the rainy season. Seedling provenance trials (UK) determined photosynthesis, and biomass and δ¹³C in relation to water table. Environmental data were characterised for all provenances at their sites of origin.

Results

Provenances differed in both δ¹³C and δ¹⁵N. Gum yield declined with increasing WUE. Virtually no BNF was detected during the dry season and seedlings and mature trees may have different WUE strategies. Wind speed and soil characteristics at provenance origin were correlated with isotope composition and gum production.

Conclusion

Provenance differences suggest that selection for desirable traits, e.g., increased gum production, may be possible. As ecological strategies relate to native locality, the environmental conditions at plantation site and provenance origin are important in assessing selection criteria.     

Alpine plant functional group responses to fertiliser addition depend on abiotic regime and community composition

Background and aims

We ask how productivity responses of alpine plant communities to increased nutrient availability can be predicted from abiotic regime and initial functional type composition.

Methods

We compared four Caucasian alpine plant communities (lichen heath, Festuca varia grassland, Geranium-Hedysarum meadow, snow bed community) forming a toposequence and contrasting in productivity and dominance structure for biomass responses to experimental fertilization (N, P, NP, Ca) and irrigation for 4–5?years.

Results

The dominant plants in more productive communities monopolized added N and P, at the expense of their neighbors. In three out of four communities, N and P fertilizations gave greater aboveground biomass increase than N or P fertilization alone, indicating overall co-limitation of N and P, with N being most limiting. Relative biomass increase in NP treatment was negatively related to biomass in control plots across the four communities. Grasses often responded more vigorously to P, but sedges to N alone. Finally, we present one of the rare examples of a forb showing a strong N or NP response.

Conclusion

Our findings will help improve our ability to predict community composition and biomass dynamics in cool ecosystems subject to changing nutrient availability as induced by climate or land-use changes.     

Water use and water-use efficiency of coppice and seedling Eucalyptus globulus Labill.: a comparison of stand-scale water balance components

Aims

Growers of Eucalyptus globulus Labill. plantations can establish second and later rotations from coppice or by replanting with seedlings. At most locations where E. globulus is grown commercially, water availability is a major driver for productivity. Thus growers must consider which reestablishment technique will maximize productivity whilst sustaining site resources for subsequent rotations. In this study we aimed to compare the stand-scale water balance components of young coppice and seedling E. globulus.

Methods

A second rotation E. globulus coppice and seedling trial was monitored for two successive seasonal cycles. Coppice and seedling plots were instrumented with sap flow- and meteorological-sensors so that stand-scale water balance components could be estimated on a daily time step.

Results

Stand-scale transpiration rate (E) and rate of interception (E _I) were larger in coppice compared to seedlings, but the rate of soil evaporation (E _S) was lower. At approximately 2?years of age each coppice stump was reduced to a single dominant stem, a standard management practice for E. globulus growers, which reduced stem biomass by approximately 70% and caused E to fall to a value approximating that in seedlings. The cumulative transpiration of coppice was 425?mm greater than seedlings up to 34?months of age. Without the coppice reduction (down to one stem/stump), we estimate that the difference would have been much greater. The water-use efficiency of stem production (WUE_stem) was greater in young coppice compared to seedlings but this benefit is likely to be offset by the loss of biomass (and thus transpired water) during coppice stem reduction.

Conclusion

Under the conditions of this study, which included reducing coppice to a single stem, reestablishment with seedling E. globulus resulted in a higher water-use efficiency of stem biomass production compared to coppice of a similar age.     

Root plasticity maintains growth of temperate grassland species under pulsed water supply

Background and aims

The frequency of rain is predicted to change in high latitude areas with more precipitation in heavy, intense events interspersed by longer dry periods. These changes will modify soil drying cycles with unknown consequences for plant performance of temperate species.

Methods

We studied plant growth and root traits of juveniles of four grasses and four dicots growing in a greenhouse, when supplying the same total amount of water given either regular every other day or pulsed once a week.

Results

Pulsed water supply replenished soil moisture immediately after watering, but caused substantial drought stress at the end of the watering cycle, whereas regular watering caused more moderate but consistent drought. Grasses had lower water use efficiency in the pulsed watering compared to regular watering, whereas dicots showed no difference. Both grasses and dicots developed thinner roots, thus higher specific root length, and greater root length in the pulsed watering. Growth of dicots was slightly increased under pulsed watering.

Conclusions

Temperate species coped with pulsed water supply by eliciting two responses: i) persistent shoot growth, most likely by maximizing growth at peaks of soil moisture, thus compensating for slower growth during drought periods; ii) plasticity of root traits related to increased resource uptake. Both responses likely account for subtle improvement of growth under changed water supply conditions.     

Changes in stem water content influence sap flux density measurements with thermal dissipation probes

Key message

Stem WC may decline during the day. Zero-flow dT _m increases when WC decreases. Use of nighttime dT _m in the calculation of sap flux density during the day might introduce errors.

Abstract

There is increasing evidence of diel variation in water content of stems of living trees as a result of changes in internal water reserves. The interplay between dynamic water storage and sap flow is of current interest, but the accuracy of measurement of both variables has come into question. Fluctuations in stem water content may induce inaccuracy in thermal-based measurements of sap flux density because wood thermal properties are dependent on water content. The most widely used thermal method for measuring sap flux density is the thermal dissipation probe (TDP) with continuous heating, which measures the influence of moving sap on the temperature difference between a heated needle and a reference needle vertically separated in the flow stream. The objective of our study was to investigate how diel fluctuations in water content could influence TDP measurements of sap flux density. We analysed the influence of water content on the zero-flow maximum temperature difference, dT _m, which is used as the reference for calculating sap flux density, and present results of a dehydration experiment on cut branch segments of American sycamore (Platanus occidentalis L.). We demonstrate both theoretically and experimentally that dT _m increases when stem water content declines. Because dT _m is measured at night when water content is high, this phenomenon could result in underestimations of sap flux density during the day when water content is lower. We conclude that diel dynamics in water content should be considered when TDP is used to measure sap flow.