Accounting for sap flow from different parts of the root system improves the prediction of xylem ABA concentration in plants grown with heterogeneous soil moisture

When soil moisture is heterogeneous, sap flow from, and ABA status of, different parts of the root system impact on leaf xylem ABA concentration ([X-ABA]leaf). The robustness of a model for predicting [X-ABA]leaf was assessed. 'Two root-one shoot' grafted sunflower (Helianthus annuus L.) plants received either deficit irrigation (DI, each root system received the same irrigation volumes) or partial rootzone drying (PRD, only one root system was watered and the other dried the soil). Irrespective of whether relative sap flow was assessed using sap flow sensors in vivo or by pressurization of de-topped roots, each root system contributed similarly to total sap flow during DI, while sap flow from roots in drying soil declined linearly with soil water potential (Psisoil) during PRD. Although Psisoil of the irrigated pot determined the threshold Psisoil at which sap flow from roots in drying soil decreased, the slope of this decrease was independent of the wet pot Psisoil. Irrespective of whether sap was collected from the wet or dry root system of PRD plants, or a DI plant, root xylem ABA concentration increased as Psisoil declined. The model, which weighted ABA contributions of each root system according to the sap flow from each, almost perfectly explained [X-ABA] immediately above the graft union. That the model overestimated measured [X-ABA]leaf may result from changes in [X-ABA] along the transport pathway or an artefact of collecting xylem sap from detached leaves. The implications of declining sap flow through partially dry roots during PRD for the control of stomatal behaviour and irrigation scheduling are discussed.     

胡杨根系水力提升作用的证据及其生态学意义

 生长在塔里木河流域的荒漠河岸林植被虽长期忍受着高温和干旱的威胁, 然而它们却能够一直延续并保存至今。除了植物深根系吸水作用外, 另一个更主要的原因可能就是荒漠河岸林植被存在水力提升的效应。该文采用HRM热比率法茎流仪对3株胡杨(Populus euphratica)主根和侧根的液流速率分别进行了为期4 d的连续监测; 利用自动气象站对微气象因子：风速、空气相对湿度、叶面温度和地表温度进行连续监测; 同时采用了烘干法对不同深度土层在不同时刻的土壤含水率进行了取样分析。试验结果表明：胡杨主根液流在白天和夜间均表现为正值, 相反的, 胡杨侧根液流速率则出现了明显的夜间负向流动。胡杨根系0~120 cm土层土壤水分含量具有下湿上干的变化趋势; 胡杨侧根在夜间发生负向流动后, 土壤含水率显著升高, 尤其在60~120 cm土层中, 4:00土壤含水率上升幅度达到4:00时刻土壤含水率的22%~26%。影响胡杨侧根液流速率的主要气象因子主要是叶面水汽压亏缺。     

Nitrogen fixation and metabolism by groundwater-dependent perennial plants in a hyperarid desert

The Central Asian Taklamakan desert is characterized by a hyperarid climate with less than 50 mm annual precipitation but a permanent shallow groundwater table. The perched groundwater (2–16 m) could present a reliable and constant source of nitrogen throughout the growing season and help overcome temporal nitrogen limitations that are common in arid environments. We investigated the importance of groundwater and nitrogen fixation in the nitrogen metabolism of desert plants by assessing the possible forms and availability of soil N and atmospheric N and the seasonal variation in concentration as well as isotopic composition of plant N. Water availability was experimentally modified in the desert foreland through simulated flooding to estimate the contribution of surface water and temporally increased soil moisture for nutrient uptake and plant–water relations. The natural vegetation of the Taklamakan desert is dominated by plants with high foliar nitrogen concentrations (2–3% DM) and leaf nitrate reductase activity (NRA) (0.2–1 mol NO₂^– g^–1 FW h^–1). There is little evidence that nitrogen is a limiting resource as all perennial plants exhibited fast rates of growth. The extremely dry soil conditions preclude all but minor contributions of soil N to total plant N so that groundwater is suggested as the dominant source of N with concentrations of 100 M NO₃^–. Flood irrigation had little beneficial effect on nitrogen metabolism and growth, further confirming the dependence on groundwater. Nitrogen fixation was determined by the ¹⁵N natural abundance method and was a significant component of the N-requirement of the legume Alhagi, the average contribution of biologically fixed nitrogen in Alhagi was 54.8%. But nitrogen fixing plants had little ecological advantage owing to the more or less constant supply of N available from groundwater. From our data we conclude that the perennial species investigated have adapted to the environmental conditions through development of root systems that access groundwater to satisfy demands for both water and nutrients. This is an ecologically favourable strategy since only groundwater is a predictable and stable resource.     

Dynamics of water and ion transport driven by corn canopy in the Yellow River basin

Water and ion balance in a corn field in the semi-arid region of the upper Yellow River basin (Inner Mongolia, China) was analyzed with special reference to transpiration stream and selective nutrient uptake driven by the crop canopy. During the crop development stage (June 7 to July 17, 2005), crop transpiration and soil evaporation were evaluated separately on a daily basis, and concentrations of NO ₃ ^? , PO ₄ ^3? , K⁺, Na⁺, Ca²⁺, Mg²⁺ and Cl^? ions in the Yellow River water, irrigation water, ground water, soil of the root zone and xylem sap of the crop were analyzed.The crop transpiration accounted for 83.4% of the evapotranspiration during the crop development stage. All ions except for Na⁺ were highly concentrated in the xylem sap due to the active and selective uptake of nutrients by roots. In particular, extremely high concentrations of the major essential nutrients were found in the nighttime stem exudate, while these concentrations in the river water, the irrigation water, the ground water and the root-zone soil were lower. On the other hand, Na⁺, which is not the essential element for crop growth, was scarcely absorbed by roots and was not highly concentrated in the xylem sap. Consequently, Na⁺ remained in the ground water and the root-zone soil at higher concentrations. These results indicate that during the growing season, crop transpiration but not soil evaporation induces the most significant driving force for mass flow (capillary rise) transporting the ground water toward the rhizosphere, where the dynamics of ion balance largely depends on the active and selective nutrient uptake by roots.     

Abscisic acid signalling when soil moisture is heterogeneous: decreased photoperiod sap flow from drying roots limits abscisic acid export to the shoots

To investigate the contribution of different parts of the root system to total sap flow and leaf xylem abscisic acid (ABA) concentration ([X-ABA]_leaf), individual sunflower ( Helianthus annuus L.) shoots were grafted onto the root systems of two plants grown in separate pots and sap flow through each hypocotyl measured below the graft union. During deficit irrigation (DI), both pots received the same irrigation volumes, while during partial root zone drying (PRD) one pot ('wet') was watered and another ('dry') was not. During PRD, once soil water content ( θ ) decreased below a threshold, the fraction of sap flow from drying roots declined. As θ declined, root xylem ABA concentration increased in both irrigation treatments, and [X-ABA]_leaf increased in DI plants, but [X-ABA]_leaf of PRD plants actually decreased within a certain θ range. A simple model that weighted ABA contributions of wet and dry root systems to [X-ABA]_leaf according to the sap flow from each, better predicted [X-ABA]_leaf of PRD plants than either [X-ABA]_dry, [X-ABA]_wet or their mean. Model simulations revealed that [X-ABA]_leaf during PRD exceeded that of DI with moderate soil drying, but continued soil drying (such that sap flow from roots in drying soil ceased) resulted in the opposite effect.     

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.     

Xylem sap composition: A tool for investigating mineral uptake and cycling in adult spruce

Xylem sap composition of spruce is influenced by several factors, such as the sampled organ, the sampling period, the availability of soil nutrients, and the soil water potential. Based on literature data and ongoing investigations carried out with adult trees, we present an overview on the main factors influencing xylem sap concentrations of Norway spruce. Direct measurements of nutrient fluxes in the xylem sap are then used to suggest a general scheme of mineral element cycling within adult trees. In Norway spruce (Picea abies Karst.), nutrient concentration in the xylem sap was higher in twigs and fine roots compared to the bottom of the trunk, the highest concentrations beeing observed in spring during the shoot elongation. Xylem sap concentrations were higher in spruce growing at nutrient rich sites than at poor sites. The combination of twig and trunk xylem sap analysis, together with xylem flow measurements in the trunk during the course of a vegetation period allowed the quantification of mineral fluxes via xylem sap flow in the trunk and twigs. These results were compared to gross mineral uptake measurements at the same site. Ca flux in the trunk xylem sap was lower than the gross uptake of Ca. Mg flux in trunk sap was approximately equivalent to Mg gross uptake whereas P and K fluxes in trunk sap were much higher than the gross uptake. Fluxes of Ca, Mg, K and P in the twig sap were much higher than that in trunk sap. Data suggest that internal cycling is responsible for a large part of the nutrient fluxes in the xylem sap of the crown. Xylem sap composition thus appears to be a tool which can complement other sources of information on mineral uptake and cycling in adult spruce     

Nutrient limitation of plant growth on the floodplain of the Narran River, Australia: growth experiments and a pilot soil survey

Water resource development has altered the hydrological regime on the Lower Balonne River in Queensland, Australia. Concerns have been raised about possible impacts to floodplain plant communities, which support a pastoral industry and a range of native fauna. Water and nutrients commonly limit plant growth in south central Queensland, where the climate is semi-arid and the soils are infertile. Floodplain plant productivity is boosted by inundation with water, but the role of flooding in nutrient provision is not known. Growth experiments and a pilot soil survey were conducted to help determine if soil nutrient deficiencies exist and if regular flooding is required to maintain floodplain soil fertility. Soils were sampled from areas representing three flood frequency classes: high, moderate, and low. Chemical extractions were performed as a surrogate for `bioavailable' nutrients. Soil nitrogen (N) but not phosphorus (P) limited the growth of seedlings of wheat (Triticum aestivum L. Gardia) based on responses to nutrient additions: plants supplied with N had greater shoot length and total biomass than plants without N. Clear evidence of an effect of flood frequency on fertility was not revealed. Neither soil P, soil N, nor plant growth varied significantly with flood frequency. However, this analysis had low statistical power. There were trends for greater biomass of seedlings grown on moderately flooded soils and thinner roots for seedlings grown on frequently flooded soils, but neither of these growth responses was clearly linked to nutrient limitation. Nevertheless, the possibility that flooding provides a nutrient subsidy to plants cannot be ruled out because of a number of factors, including the statistical power of this analysis and the possibility that short-term nutrient subsidies occur with floods.     

Reverse flow of sap in tree roots and downward siphoning of water by Grevillea robusta

1. Constant-power heat-balance sap flow gauges were used to compare sap flow in vertical and lateral roots of Grevillea robusta trees growing without access to ground water at a semiarid site in Kenya.
2. Reversal of sap flow occurred when root systems crossed gradients in soil water potential. Measurement of changes in the direction of flow was possible because of the symmetrical construction of the sap flow gauges; gradients in temperature across the gauges, and thus computed rates of sap flow, changed sign when reverse flow occurred.
3. Reverse flow in roots descending vertically from the base of the tree occurred, while uptake by lateral roots continued, when the top of the soil profile was wetter than the subsoil. The transfer of water downwards by root systems, from high to low soil water potential, was termed 'downward siphoning'; this is the reverse of hydraulic lift.
4. Downward siphoning was induced by the first rain at the end of the dry season and by irrigation of the soil surface during a dry period.
5. Downward siphoning may be an important component of the soil water balance where there are large gradients in water potential across root systems, from a wet soil surface downwards. By transferring water beyond the reach of shallow-rooted neighbours, downward siphoning may enhance the competitiveness of deep-rooted perennials.     

Nutrient status and mycorrhizal enhancement of water transport in soybean

Mycorrhizal soybean (Glycine max L. Merr. var. Harosoy-63) plants have lower resistances to water transport than nonmy-corrhizal plants after 4.5 weeks of growth. Although resistances of whole plants differ by 40%, there were no differences in the resistances of stems plus leaves, indicating that the major effect of the mycorrhizae was to reduce the resistance of the roots. Since the fungitoxicant, p-chloronitrobenzene, had no effect on resistances to water transport, reduced resistances were probably not caused by a direct modification of the transport pathway by the fungus. Differences in resistance between mycorrhizal and nonmycorrhizal soybean were essentialy eliminated by the application of nutrients to the soil. Thus, lowered resistances of mycorrhizal roots growing in soil with low levels of nutrients probably resulted from the enhanced nutrient status of the plant brought about by the fungus. Mycorrhizal infection increased growth at both low and high nutrient levels.