Hydraulic lift: consequences of water efflux from the roots of plants

Hydraulic lift is the passive movement of water from roots into soil layers with lower water potential, while other parts of the root system in moister soil layers, usually at depth, are absorbing water. Here, we review the brief history of laboratory and field evidence supporting this phenomenon and discuss some of the consequences of this below-ground behavior for the ecology of plants. Hydraulic lift has been shown in a relatively small number of species (27 species of herbs, grasses, shrubs, and trees), but there is no fundamental reason why it should not be more common as long as active root systems are spanning a gradient in soil water potential (Ψ_s) and that the resistance to water loss from roots is low. While the majority of documented cases of hydraulic lift in the field are for semiarid and arid land species inhabiting desert and steppe environments, recent studies indicate that hydraulic lift is not restricted to these species or regions. Large quantities of water, amounting to an appreciable fraction of daily transpiration, are lifted at night. This temporary partial rehydration of upper soil layers provides a source of water, along with soil moisture deeper in the profile, for transpiration the following day and, under conditions of high atmospheric demand, can substantially facilitate water movement through the soil-plant-atmosphere system. Release of water into the upper soil layers has been shown to afford the opportunity for neighboring plants to utilize this source of water. Also, because soils tend to dry from the surface downward and nutrients are usually most plentiful in the upper soil layers, lifted water may provide moisture that facilitates favorable biogeochemical conditions for enhancing mineral nutrient availability, microbial processes, and the acquisition of nutrients by roots. Hydraulic lift may also prolong or enhance fine-root activity by keeping them hydrated. Such indirect benefits of hydraulic lift may have been the primary selective force in the evolution of this process. Alternatively, hydraulic lift may simply be the consequence of roots not possessing true rectifying properties (i.e., roots are leaky to water). Finally, the direction of water movement may also be downward or horizontal if the prevailing Ψ_s gradient so dictates, i.e., inverse, or lateral, hydraulic lift. Such downward movement through the root system may allow growth of roots in otherwise dry soil at depth, permitting the establishment of many phreatophytic species. Received: 2 June 1997 / Accepted: 24 September 1997     

Does hydraulic lift exist in shallow-rooted species? A quantitative examination with a half-shrub Gutierrezia sarothrae

Hydraulic lift occurs in some deep-rooted shrub and herbaceous species. In this process, water taken up by deep roots from the moist subsoil is delivered to the drier topsoil where it is later reabsorbed by shallow roots. However, little is known about the existence of hydraulic lift in shallow-rooted xeric species. The objectives of this study were 1) to ascertain whether hydraulic lift exists in Gutierrezia sarothrae (broom snakeweed), a widespread North American desert species with a shallow root system, grown in pot and field conditions and 2) if it does, how much water can be transferred from the subsoil to the 30 cm topsoil during the night. Snakeweed seedlings were transplanted in buried pots allowing the deeper roots to grow into the subsoil 30 cm below the surface. Soil water content inside and outside of the pot was measured seasonally and diurnally with time domain reflectometry technique (TDR). An increase in water content was detected in the pot after the plant was covered for 3 h by an opaque plastic bag during the day, suggesting hydraulic lift from deeper depths and exudation of water into the drier topsoil. Root exudation was also observed on native range sites dominated by snakeweed. Water efflux in the pot was 271 g per plant per night. which was equivalent to 15.3% of the extrapolated, porometer-derived whole-plant daily transpiration. Hydraulic lift observed in Gutierrezia improved water uptake during the day when evaporative demand is high and less water is available in the topsoil. We concluded that hydraulic lift might help snakeweed to alleviate the effect of water stress.     

Hydraulic lift through transpiration suppression in shrubs from two arid ecosystems: patterns and control mechanisms

Hydraulic lift (HL) is the passive movement of water through the roots from deep wet to dry shallow soil layers when stomata are closed. HL has been shown in different ecosystems and species, and it depends on plant physiology and soil properties. In this study we explored HL patterns in several arid land shrubs, and developed a simple model to simulate the temporal evolution and magnitude of HL during a soil drying cycle under relatively stable climatic conditions. This model was then used to evaluate the influence of soil texture on the quantity of water lifted by shrubs in different soil types. We conducted transpiration suppression experiments during spring 2005 in Chile and spring 2008 in Spain on five shrub species that performed HL, Flourensia thurifera, Senna cumingii and Pleocarphus revolutus (Chile), Retama sphaerocarpa and Artemisia barrelieri (Spain). Shrubs were covered with a black, opaque plastic fabric for a period of 48–72 h, and soil water potential was recorded at different depths under the shrubs. While the shrubs remained covered, water potential continuously increased in shallow soil layers until the cover was removed. The model output indicated that the amount of water lifted by shrubs is heavily dependent on soil texture, as shrubs growing in loamy soils redistributed up to 3.6 times more water than shrubs growing on sandy soils. This could be an important consideration for species growing in soils with different textures, as their ability to perform HL would be context dependent.     

Soil salinity determines the relative abundance of C3/C4 species in Argentinean grasslands

Aim The C₄ and crassulacean acid metabolism (CAM) pathways are adaptations to compensate for high rates of photorespiration and water and carbon deficiency. This is the first attempt to compare the relative abundance of C₃ vs. C₄ + CAM species in temperate and subtropical grasslands across a latitudinal gradient in central Argentina. We predict that under the same rainfall regime, C₄ + CAM plants will have larger soil coverage in highly saline soils than in neighbouring non‐saline ones. Location Data were taken from three phytogeographical provinces in the Santa Fe province of Argentina: Chaquenian, Pampean and Espinal. Methods The salinity of the soil was estimated through aqueous solution conductivity. The proportions of species belonging to C₃/C₄ + CAM photosynthetic pathways were compared among halophyte and non‐halophyte communities with a χ² homogeneity test. The sum of cover percentages corresponding to the C₃ and C₄ + CAM photosynthetic pathways were calculated and compared using analysis of variance (ANOVA). Results The soil conductivity values were higher in the halophyte than in the non‐halophyte communities for the same phytogeographical area. The C₄ + CAM plants had much higher soil coverage values in halophyte than in non‐halophyte communities in the Pampean and Espinal phytogeographical provinces. The differences were not statistically significant in the Chaquenian province. Main conclusions Soil drought provoked by soil salinity results in a much higher soil cover by C₄ + CAM plants in regions with positive to neutral water balance (i.e. Pampean and Espinal). This differential abundance pattern in C₄ + CAM functional group is not observed in areas where a pronounced water deficit exists per se (Chaquenian region), and therefore C₄ + CAM plants predominate in all environments regardless of soil salinity. Our results suggest that one of the main environmental forces driving the upsurge of C₄ species in Argentinean grasslands might have been the strong local soil salinity gradient.     

Hydraulic lift in drought-tolerant and -susceptible maize hybrids

Hydraulic lift was investigated in a greenhouse study involving two drought-tolerant maize (Zea mays L.) hybrids (TAES176 and P3223) and a drought-susceptible hybrid (P3225) during the flowering stage. Root systems were grown in two soil compartments – a drier upper soil and a wetter deep soil. The plants were shaded for 3 h during the daytime. Soil volumetric water content (Ø_v) in the upper pots was measured with time domain reflectometry (TDR) before and after shading. An increase in Ø_v in the upper pot was detected with TDR in the drought-tolerant hybrids following 3 h of shading, but not in the drought-susceptible hybrid. Furthermore, water exuded from roots in the top soil layers was greater in the more drought-tolerant TAES176 than in P3223 (489 vs. 288 g per pot in 3 h, P<0.005). The sizable amount of water from hydraulic lift allowed TAES176 to reach a peak transpiration rate 27–42% higher than the drought-susceptible hybrid P3225 on the days when the evaporative demand was high. To our knowledge, this is the first experiment that reveals a significant surge of transpiration due to hydraulic lift following midday shading. Hydraulic lift also prevented soil moisture depletion in the upper pots with TAES176, but not with P3223 or P3225. Root characteristics may be responsible for differences in hydraulic lift of the three maize hybrids. There were 2.3–3.3-fold more primary roots in the deep moist soil in P3223 and TAES176 than in P3225 that may enable these hybrids to absorb and transport water at faster rates. Therefore, more water can be exuded into the upper drier soil when transpiration is suppressed by shading. Larger primary roots (20–28% larger diameter) and a higher root volume in the upper soil in TAES176 and P3223 than in P3225 may contribute to higher root hydraulic conductance and greater water efflux from the roots. The negligible hydraulic lift in P3225 may also relate to higher night-time transpiration of the hybrid. This report has documented, for the first time, the existence of genetic variations in hydraulic lift among maize hybrids and links between hydraulic lift and drought tolerance within maize plants. It appears that one of drought tolerance mechanisms in maize may lie in the extent of hydraulic lift.     

Hydraulic lift: a potentially important ecosystem process

Hydraulic lift is the process by which some deep-rooted plants take in water from lower soil layers and exude that water into upper, drier soil layers. Hydraulic lift is beneficial to the plant transporting the water, and may be an important water source for neighboring plants. Recent evidence shows that hydraulically lifted water can promote greater plant growth, and could have important implications for net primary productivity, as well as ecosystem nutrient cycling and water balance.     

Fluctuations in nitrate reductase activity,and nitrate and organic nitrogen concentrations of succulent plants under different nitrogen and water regimes

The CAM (Crassulacean acid metabolism) succulent species Kalanchoe daigremontiana, K. tubiflora and Crassula argentea, and the succulent C₃ species Peperomia obtusifolia, were cultivated in pure culture in open-air conditions under two different regimes of nitrogen and water supply. At specified intervals during the course of vegetative growth, biomass, nitrate reductase activity (NRA), nitrate concentration, and organic nitrogen concentration of whole plants were measured. After 100 days of cultivation the leaf conductance of Crassula and Peperomia was measured at intervals for the duration of a day. Behaviour of all four species was strongly influenced by the cultivation regime. This was apparent in terms of productivity and variable flucturations in NRA, nitrate concentration, and organic nitrogen concentration during the vegetative period. Increase in biomass was mostly connected with a decrease in all other investigated parameters, especially under conditions of water and/or nitrogen deficiency. The typical reaction of the CAM species Crassula to limited netrogen but adequate soil water was to reduce leaf conductance during light, whereas the C₃ plant Peperomia increased conductance in comparison with plants having a nitrogen suppy. The NRA of all plant species was reduced by both soil nitrate deficiency and drought. The succulent plant species, which are specially adapted to drought, neither took up nor used nitrate when water was limited. This was particularly the case for the CAM species, but less so for the C₃ Peperomia, which showed very high concentrations of nitrate and organic nitrogen, but low NRA and biomass gain. A formula was derived to express the nitrogen use efficiency (NUE) of the species, i.e. the ability of a plant to use nitrogen over a specific period of growth. NUE was shown to increase with age for the crassulacean species but to decrease for the C₃ Peperomia. Furthermore, NUE varied with the different nutrient levels in a species-specific manner, with high values for NUE not necessarily coupled to high productivity, and with NUE of the C₃ species generally higher than that of CAM species.     

Le vie fotosintetiche C3, C4 e CAM nel contesto ecologico

Abstract

Ecological aspects of C₃, C₄ and CAM photosynthetic pathways. - Three different photosynthetic CO₂ fixation pathways are known to occur in higher plants. However all three pathways ultimately depend on the Calvin-Benson cycle for carbon reduction. The oxygenase activity of RuBP carboxilase is responsible for photorespiratory CO₂ release. Both C₄ and CAM pathways behave as a CO₂ concentrating mechanism which prevent photorespiration. The CO₂-concentrating mechanism in C₄ plants is based on intracellular symplastic transport of C₄ dicarboxylic acids from mesophyll-cells to the adjacent bundle-sheath cells. On the contrary in CAM plants the CO₂-concentrating mechanism is based on the intracellular transport of malic acid into and out of the vacuole.

The C₄ photosynthetic pathway as compared to the C₃ pathway permits higher rates of CO₂ fixation in high light and high temperature environments at low costs in terms of water loss, given the stability of the photosynthetic apparatus under such conditions.

CAM is interpreted as an adaptation to arid environments because it enables carbon assimilation to take place at very low water costs during the night when the evaporative demand is low. Nevertheless many aquatic species of Isoetes and some relatives are CAM, suggesting the adaptive role of CAM to environments which become depleted in CO₂.

The photosynthetic carbon fixation pathway certainly contributes to the ecological success of plants in different environments. However the distribution of plants may also reflect their biological history. On the other hand plants with different photosynthetic pathways coexist in many communities and tend to share resources in time. In any case some generalizations are possible: C₄ plants enjoy an ecological advantage in hot, moist, high light regions while the majority of species in desert environments are C₃; CAM plants are more frequent in semiarid regions with seasonal rainfall, coastal fog deserts, and in epiphytic habitats in tropical rain forests.     

沙丘多枝柽柳灌丛根层土壤含水量变化特征与根系水力提升证据

分析干旱区深根型荒漠植物的根层土壤水分是揭示荒漠植物与土壤水分关系机理的重要方面。在黑河中游一片风沙侵蚀区域的多枝柽柳(Tamarix ramosissima)人工林地中, 对表层0.3 m到3 m深的土壤不同深度的含水量进行了连续的动态观测。结果显示, 多枝柽柳根系层土壤含水量可以分为明显不同的3层: 浅层(0.2-1.7 m深)相对湿润层、中间(1.7-2.7 m深)相对干层和深层(2.7 m以下)有效含水层。在多枝柽柳生长盛期, 浅层相对湿润层土壤含水量呈现明显的昼夜变化特征, 同时, 在晚上植物根系与浅层土壤之间存在正水势梯度, 这说明存在根系水力提升现象。水力提升是干旱气候下根层浅层土壤含水量保持相对湿润的主要原因, 并因此维系浅层根系的发育, 也为多枝柽柳具备的防风固沙功能提供了可能的解释。据初步估算, 多枝柽柳根系水力提升占每天耗水量的5%-8%, 耗水的主要水分来源仍然是充足的土壤深层有效含水层。     

Do hydraulic redistribution and nocturnal transpiration facilitate nutrient acquisition in Aspalathus linearis?

The significance of soil water redistribution by roots and nocturnal transpiration for nutrient acquisition were assessed for deep-rooted 3-year-old leguminous Aspalathus linearis shrubs of the Cape Floristic Region (South Africa). We hypothesised that hydraulic redistribution and nocturnal transpiration facilitate nutrient acquisition by releasing moisture in shallow soil to enable acquisition of shallow-soil nutrients during the summer drought periods and by driving water fluxes from deep to shallow soil powering mass-flow nutrient acquisition, respectively. A. linearis was supplied with sub-surface (1-m-deep) irrigation rates of 0, 2 or 4 L day^?1 plant^?1. Some plants were unfertilized, whilst others were surface- or deep-fertilized (1 m depth) with Na¹⁵NO₃ and CaP/FePO₄. We also supplied deuterium oxide (²H₂O) at 1 m depth at dusk and measured its predawn redistribution to shallow soil and plant stems. Hydraulic redistribution of deep water was substantial across all treatments, accounting for 34–72 % of surface-soil predawn moisture. Fourteen days after fertilization, the surface-fertilized plants exhibited increased hydraulic redistribution and increased ¹⁵N and P acquisition with higher rates of deep-irrigation. Deep-fertilization also increased hydraulic redistribution to surface soils, although these plants additionally accumulated ²H₂O in their stem tissue overnight, probably due to nocturnal transpiration. Plants engaged in nocturnal transpiration also increased ¹⁵N and P acquisition from deep fertilizer sources. Thus, both nocturnal transpiration and hydraulic redistribution increased acquisition of shallow soil N and P, possibly through a combination of increased nutrient availability and mobility.     

Hydraulic lift and its influence on the water content of the rhizosphere: an example from sugar maple,Acer saccharum

Hydraulic lift, the transport of water from deep in the soil through plant root systems into the drier upper soil layers, has been demonstrated in several woody plant species. Here the volume of water involved in hydraulic lift by a mature sugar maple tree is estimated. Twenty-four intact soil cores were collected from the vicinity of a sugar maple tree at the same positions at which thermocouple psychrometers had been placed. Desorption measurements were made on the soil cores and the data were fitted to the Campbell relation for soil matric potential versus soil water content . The psychrometer data were filtered to obtain the diurnal component contributed by hydraulic lift. The diurnal component in was combined with the Campbell relation for each soil core to obtain the increase in soil water content due to hydraulic lift. The additional water contents were numerically integrated to obtain a volume of 102±54 1 of water which was hydraulically lifted each night. The volume of hydraulically lifted water (HLW) is sufficiently great that in ecosystems where hydraulic lift occurs it should be included in models for calculating the water balance. However, a previous analysis of the stable hydrogen isotope composition (D) of water in understory plants around trees conducting hydraulic lift implies a much greater volume of HLW than that calculated from the analysis performed above. To reconcile these differences, it is hypothesized that some understory plants preferentially extract HLW due to its higher matric potential and that the proportion of this water source within the xylem sap of at least some understory plants that use HLW was so great that the roots of these plants must therefore be in close proximity to the tree roots from which the HLW comes. The results of this study have implications for studies of plant competition where positive associations may exist as well as for ion uptake, nutrient cycling and the design of agroforestry systems.     

干旱区水力提升的生态作用

水力提升是某些植物通过深层根系从较湿的深层土壤中吸取水分再通过浅层根系在较干的浅层土壤中释出的过程。水力提升所释出水量及其释出过程具有积极的生态学意义,它不但为伴生植物提供新的水源、改善其邻近植物的蒸腾作用、缓解水分亏缺、提高水分利用效率．而且促进植物的养分吸收、促进上层干土中分解、硝化和矿化等生化过程．尤其是发生在克隆植物植株间的水分共享这种特殊的水力提升,还影响群落的形成。然而,对于一般的水力提升而言,水力提升所释出水量到底有多少。其生态学作用有多大．有待深入研究。结合研究实践认为．要研究水力提升具有多大的生态学作用．必须准确地区分水;勺提升的水量和毛管水的上升水量。还建议了研究水力提升新的研究方法。     

Photosynthetic Pathways and Life Form Types for Native Plant Species from Hulunbeier Rangelands,Inner Mongolia,North China

Photosynthetic pathway Types (C₃, C₄, and CAM) and life forms of native species from Hulunbeier rangelands, north China were studied. Of the total 258 species, 216 species in 132 genera and 42 families had C₃ photosynthetic pathway, including dominant herbs, e.g. Stipa baicalensis Roshev. and Leymus chinensis (Trin.) Tzvel., Filifolium sibiricum Kitam. and Arudinella hirta (Thunb.) Koidz. 38 species in 28 genera and 10 families were found with C₄ photosynthesis, and 4 species in 2 genera and 1 family had CAM photosynthetic pathway. The occurrence of C₄ species was common in Gramineae and Chenopodiaceae, and the two families were leading ones within C₄ plants. More than 52 % of the total 258 species were in H form, 21 % in Th form, 19 % in G form; the other life form Types, e.g. Ch, M, N, and HH, formed less than 3 %. 68 % of C₄ species were in Th form and 24 % in H form, indicating that these Types were the dominant life forms for C₄ species in the rangeland region. The occurrence of C₄ species was closely related with plant habitats, disturbed lands had the highest C₄ abundance (55 % of the total C₄ species), followed by grasslands and sandy soil, and forests had the lowest C₄ abundance (8 %). Hence the occurrence of C₄ species could be efficient indicator for rangeland dynamics in Hulunbeier rangelands.     

Photosynthetic Pathways and Life Forms in Different Grassland Types from North China

Photosynthetic pathways (C₄, C₃, and CAM species) and plant life forms of three grassland types in North China were compared. Of the total 201 species, 144 species in 78 genera and 34 families had C₃ photosynthetic pathway, 56 species in 35 genera and 11 families had C₄ photosynthetic pathway, and 1 species had CAM photosynthetic pathway. The number of C₄ species in Songnen meadow was 70–80 % greater than that in Xilinguole steppe and Hunshandak desert grassland, but that for C₃ species did not differ significantly among the three grassland types. The number of therophytes in the Songnen meadow was relatively greater than that of the other two grassland types, but that of hemicryptophytes was lower. Thus the distribution of C₄ species and plant life form is probably related to precipitation.     

Soil water partitioning contributes to species coexistence in tallgrass prairie

The majority of tallgrass prairie root biomass is located in the upper soil layers (0–25 cm), but species differences exist in reliance on soil water at varying depths. These differences have led to the hypothesis that resource partitioning belowground facilitates species co‐existence in this mesic grassland. To determine if plant water relations can be linked to soil water partitioning as a potential mechanism allowing C₃ species to persist among the more dominant C₄ grasses, we measured differences in the source of water‐use using the isotopic signature of xylem water, volumetric soil water content at 4 depths, and leaf water potentials. Data were collected for seven species representing C₄ grasses, C₃ forbs and C₃ shrubs over three growing seasons at the Konza Prairie (Kansas, USA) to encompass a range of natural climatic conditions. C₄ grasses relied on shallow soil water (5 cm) across the growing season and had midday leaf water potentials that were highly correlated with shallow soil water regardless of soil water availability at other portions of the soil profile (20, 40 and 90 cm). In contrast, C₃ species only used shallow soil water when plentiful at this depth; these species increased their dependence on soil water from greater depths as the upper soil layers dried. Structural equation models describing plant water relations were very similar for the three C₄ species, whereas a unique set of models and drivers were identified for each of the C₃ species. These results support soil water partitioning as a mechanism for species coexistence, as C₄ species in this grassland have relatively consistent dependence on water in shallow soil layers, whereas C₃ species show niche differentiation in water use strategies to avoid competition with C₄ grasses for water in shallow soil layers when this resource is limiting and leaf water stress is high.     

CAM photosynthesis in submerged aquatic plants

Crassulacean acid metabolism (CAM) is a CO₂-concentrating mechanism selected in response to aridity in terrestrial habitats, and, in aquatic environments, to ambient limitations of carbon. Evidence is reviewed for its presence in five genera of aquatic vascular plants, includingIsoëtes, Sagittaria, Vallisneria, Crassula, andLittorella. Initially, aquatic CAM was considered by some to be an oxymoron, but some aquatic species have been studied in sufficient detail to say definitively that they possess CAM photosynthesis. CO₂-concentrating mechanisms in photosynthetic organs require a barrier to leakage; e.g., terrestrial C₄ plants have suberized bundle sheath cells and terrestrial CAM plants high stomatal resistance. In aquatic CAM plants the primary barrier to CO₂ leakage is the extremely high difrusional resistance of water. This, coupled with the sink provided by extensive intercellular gas space, generates daytime CO₂(pi) comparable to terrestrial CAM plants. CAM contributes to the carbon budget by both net carbon gain and carbon recycling, and the magnitude of each is environmentally influenced. Aquatic CAM plants inhabit sites where photosynthesis is potentially limited by carbon. Many occupy moderately fertile shallow temporary pools that experience extreme diel fluctuations in carbon availability. CAM plants are able to take advantage of elevated nighttime CO2 levels in these habitats. This gives them a competitive advantage over non-CAM species that are carbon starved during the day and an advantage over species that expend energy in membrane transport of bicarbonate. Some aquatic CAM plants are distributed in highly infertile lakes, where extreme carbon limitation and light are important selective factors. Compilation of reports on diel changes in titratable acidity and malate show 69 out of 180 species have significant overnight accumulation, although evidence is presented discounting CAM in some. It is concluded that similar proportions of the aquatic and terrestrial floras have evolved CAM photosynthesis. AquaticIsoëtes (Lycophyta) represent the oldest lineage of CAM plants and cladistic analysis supports an origin for CAM in seasonal wetlands, from which it has radiated into oligotrophic lakes and into terrestrial habitats. Temperate Zone terrestrial species share many characteristics with amphibious ancestors, which in their temporary terrestrial stage, produce functional stomata and switch from CAM to C₃. Many lacustrineIsoëtes have retained the phenotypic plasticity of amphibious species and can adapt to an aerial environment by development of stomata and switching to C₃. However, in some neotropical alpine species, adaptations to the lacustrine environment are genetically fixed and these constitutive species fail to produce stomata or loose CAM when artificially maintained in an aerial environment. It is hypothesized that neotropical lacustrine species may be more ancient in origin and have given rise to terrestrial species, which have retained most of the characteristics of their aquatic ancestry, including astomatous leaves, CAM and sediment-based carbon nutrition.     

On the assessment of root and soil respiration for soils of different textures: interactions with soil moisture contents and soil CO2 concentrations

Estimates of root and soil respiration are becoming increasingly important in agricultural and ecological research, but there is little understanding how soil texture and water content may affect these estimates. We examined the effects of soil texture on (i) estimated rates of root and soil respiration and (ii) soil CO₂ concentrations, during cycles of soil wetting and drying in the citrus rootstock, Volkamer lemon (Citrus volkameriana Tan. and Pasq.). Plants were grown in soil columns filled with three different soil mixtures varying in their sand, silt and clay content. Root and soil respiration rates, soil water content, plant water uptake and soil CO₂ concentrations were measured and dynamic relationships among these variables were developed for each soil texture treatment. We found that although the different soil textures differed in their plant-soil water relations characteristics, plant growth was only slightly affected. Root and soil respiration rates were similar under most soil moisture conditions for soils varying widely in percentages of sand, silt and clay. Only following irrigation did CO₂ efflux from the soil surface vary among soils. That is, efflux of CO₂ from the soil surface was much more restricted after watering (therefore rendering any respiration measurements inaccurate) in finer textured soils than in sandy soils because of reduced porosity in the finer textured soils. Accordingly, CO₂ reached and maintained the highest concentrations in finer textured soils (> 40 mmol CO₂ mol⁻¹). This study revealed that changes in soil moisture can affect interpretations of root and soil measurements based on CO₂ efflux, particularly in fine textured soils. The implications of the present findings for field soil CO₂ flux measurements are discussed. This revised version was published online in June 2006 with corrections to the Cover Date.     

Does hydraulic lift or nighttime transpiration facilitate nitrogen acquisition?

Water movement from roots to soil at night in the process of hydraulic lift (redistribution) rehydrates the rhizosphere and has been proposed to improve plant nutrient acquisition. Another process that has now been found in many plant species is nighttime transpiration and this could also affect nutrient relations by influencing supply of mobile nutrients to roots at night. The effects of these soil/root water relations interactions have not been adequately tested. We chose ten Sarcobatus vermiculatus (Hook.) Torrey shrubs with different magnitudes of hydraulic lift (i.e. diel range in soil water potential) to test the hypothesis that the magnitude of lift would be positively related to the amount of nitrogen (N) uptake over a period of days. A ¹⁵N tracer was injected in the 20–30 cm soil layer at locations with hydraulic lift to determine plant ¹⁵N acquisition by shallow roots conducting hydraulic lift. Half of the plants were also placed in large humidified tents (i.e. “bagged”), which suppressed nighttime transpiration, and thus were expected to have greater magnitudes of hydraulic lift, although they did not. All plants took up the ¹⁵N tracer, but contrary to our hypothesis the magnitude of hydraulic lift had no significant effect on the amount of ¹⁵N acquired over a 9-day period following labeling. However, plants that were bagged tended to have lower ¹⁵N acquisition (P = 0.07). These data indicate that decreased nighttime transpirational water loss or some other effect of bagging may decrease nutrient acquisition by these nutrient-limited phreatophytic shrubs and more generally suggests a possible nutritional benefit of nighttime transpiration by plants. This suggestion needs more thorough testing to elucidate an important potential link between plant water and nutrient relations.     

Estimating heat tolerance among plant species by two chlorophyll fluorescence parameters

The heat tolerance of 8 temperate- and 1 subtropical-origin C₃ species as well as 17 tropical-origin ones, including C₃, C₄, and CAM species, was estimated using both F₀-T curve and the ratio of chlorophyll fluorescence parameters, prior to and after high temperature treatment. When leaves were heated at the rate of ca. 1 °C min⁻¹ in darkness, the critical temperature (T_c) varied extensively among species. The T_c's of all 8 temperate-origin species ranged between 40–46 °C in winter (mean temperature 16–19 °C), and between 32–48 °C in summer (mean temperature ca. 30 °C). Those for 1 subtropical- and 12 tropical-origin C₃ species ranged between 25–44 °C and 35–48 °C, and for 1 CAM and 4 C₄ species were 41–47 and 45–46 °C, respectively. Acclimating three C₃ herbaceous plants at high temperature (33/28 °C, day/night) for 10 d in winter caused their T_c's rising to nearly the values measured in summer. When leaves were exposed to 45 °C for 20 min and then kept at room temperature in darkness for 1 h, a significant correlation between RF_v/m (the ratio of F_v/F_m before and after 45 °C treatment) and T_c was observed for all tested temperate-origin C₃ species as well as tropical-origin CAM and C₄ species. However, F₀ and F_v/F_m of the tropical-origin C₃ species were less sensitive to 45 °C treatment, regardless of a large variation of T_c; thus no significant correlation was found between their RF_v/m and T_c. Thus T_c might not be a suitable index of heat tolerance for plants with wide range of environmental adaptation. Nevertheless, T_c's of tropical origin C₃ species, varying and showing high plasticity to seasonal changes and temperature treatment, appeared suitable for the estimation of the degree of temperature acclimation in the same species.     

Evidence of hydraulic lift in a young beech and oak mixed forest using 18O soil water labelling

Hydraulic lift (HL) by tree roots in a young, broad-leaved, mixed temperate European forest was investigated during the 2008 growing season by injecting ¹⁸O-enriched soil water at a depth of 75–90 cm under drought conditions experimentally imposed in a rain-exclusion system. Based on sap flow, leaf water potential, 2-D root distribution measurements, soil isotope profiles, and xylem water isotope composition, water acquisition and use by two tree species, beech (Fagus sylvatica) and oak (Quercus petraea) was compared. We showed that, unlike oak, beech experienced a marked decrease in sap flow and predawn leaf water potential with increasing soil drought. This behaviour was logical considering the shallower root system in beech than in oak. Six days after ¹⁸O-labelling, we observed isotopic enrichment in the shallower soil layers. Since the intermediate soil layers did not display any enrichment, our results clearly pointed to hydraulic lift by tree roots. The superficial enrichment that was observed in the vicinity of oak trunks and the increase in the isotopic signature of xylem sap in the oak trees but not in the beech trees confirmed the predominant role of oak in the hydraulic lift at our site. Even though facilitation for water acquisition among species was not observed here, our results suggest a potential positive contribution of species like oak toward maintaining species diversity in mixed forest ecosystems submitted to severe drought events.