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1.

Background and aims

Soil compaction strongly affects water uptake by roots. The aim of the work was to examine soil—plant interactions with focus on the impact of distribution of compacted soil layers on growth and water uptake by wheat roots.

Methods

The growth-chamber experiment was conducted on wheat growth in soil with compacted soil layers. The system for maintaining constant soil water potential and measurement of daily water uptake from variously compacted soil layers was used.

Results

Layered soil compaction differentiated vertical root distribution to higher extent for root length than root mass. The propagation rate of a water extraction front was the highest through layers of moderately compacted soil. The root water uptake rate was on average 67 % higher from moderately than heavily compacted soil layers. Correlations between water uptake and the length of thick roots were increasing with increasing level of soil compaction.

Conclusions

The study shows that root amount, water uptake, propagation of water extraction and shoot growth strongly depend on the existence of compacted layers within soil profile. The negative effects of heavily compacted subsoil layer on water uptake were partly compensated by increased uptake from looser top soil layers and significant contribution of thicker roots in water uptake.  相似文献   

2.

Background

It is known that the soil near roots, the so-called rhizosphere, has physical and chemical properties different from those of the bulk soil. Rhizosphere properties are the result of several processes: root and soil shrinking/swelling during drying/wetting cycles, soil compaction by root growth, mucilage exuded by root caps, interaction of mucilage with soil particles, mucilage shrinking/swelling and mucilage biodegradation. These processes may lead to variable rhizosphere properties, i.e. the presence of air-filled gaps between soil and roots; water repellence in the rhizosphere caused by drying of mucilage around the soil particles; or water accumulation in the rhizosphere due to the high water-holding capacity of mucilage. The resulting properties are not constant in time but they change as a function of soil condition, root growth rate and mucilage age.

Scope

We consider such a variability as an expression of rhizosphere plasticity, which may be a strategy for plants to control which part of the root system will have a facilitated access to water and which roots will be disconnected from the soil, for instance by air-filled gaps or by rhizosphere hydrophobicity. To describe such a dualism, we suggest classifying rhizosphere into two categories: class A refers to a rhizosphere covered with hydrated mucilage that optimally connects roots to soil and facilitates water uptake from dry soils. Class B refers to the case of air-filled gaps and/or hydrophobic rhizosphere, which isolate roots from the soil and may limit water uptake from the soil as well water loss to the soil. The main function of roots covered by class B will be long-distance transport of water.

Outlook

This concept has implications for soil and plant water relations at the plant scale. Root water uptake in dry conditions is expected to shift to regions covered with rhizosphere class A. On the other hand, hydraulic lift may be limited in regions covered with rhizosphere class B. New experimental methods need to be developed and applied to different plant species and soil types, in order to understand whether such dualism in rhizosphere properties is an important mechanism for efficient utilization of scarce resources and drought tolerance.  相似文献   

3.

Background and aims

Wetland plants have been widely used in constructed wetlands for the clean-up of metal-contaminated waters. This study investigated the relationship between rate of radial oxygen loss (ROL), root porosity, Zn uptake and tolerance, Fe plaque formation in wetland plants.

Methods

A hydroponic experiment and a pot trial with Zn-contaminated soil were conducted to apply different Zn level treatments to various emergent wetland plants.

Results

Significant differences were found between plants in their root porosities, rates of ROL, Zn uptake and Zn tolerance indices in the hydroponic experiment, and concentrations of Fe and Mn on roots and in the rhizosphere in the pot trial. There were significant positive correlations between root porosities, ROL rates, Zn tolerance, Zn, Fe and Mn concentrations on roots and in the rhizosphere. Wetland plants with higher root porosities and ROL tended to have more Fe plaque, higher Zn concentrations on roots and in their rhizospheres, and were more tolerant of Zn toxicity.

Conclusions

Our results suggest that ROL and root porosity play very important roles in Fe plaque formation, Zn uptake and tolerance, and are useful criteria for selecting wetland plants for the phytoremediation of Zn-contaminated waters and soils/sediments.  相似文献   

4.

Background and aims

Biochar has been proposed as a tool to enhance phytostabilisation of contaminated soils but little data are available to illustrate the direct effect on roots in contaminated soils. This work aimed to investigate specific root traits and to assess the effect of biochar amendment on contaminant availability.

Methods

Amendment with two different types of biochar, pine woodchip and olive tree pruning, was assessed in a rhizobox experiment with maize planted in a soil contaminated with significant levels of copper and arsenic.

Results

Amendment was found to significantly improve root traits compared to the control soil, particularly root mass density and root length density. Copper uptake to plants and ammonium sulphate extractable copper was significantly less in the biochar amended soils. Arsenic uptake and extractability varied with type of biochar used but was not considered to be the limiting factor affecting root and shoot development.

Conclusions

Root establishment in contaminated soils can be enhanced by biochar amendment but choice of biochar is key to maximising soil improvement and controlling contaminant availability.  相似文献   

5.

Aims

The research aimed at studying the effect of flooding with sulfate-rich water on the activity, abundance and diversity of sulfate-reducing micro-organisms present in the root zone of an oxygen-releasing plant growing on two riparian grassland soils with contrasting amounts of iron.

Methods

A series of microcosms was used to investigate the effects. Plants were grown under controlled conditions in microcosms containing a rhizosphere and bulk soil compartment for a period of 12 weeks in the presence of sulfate-rich flood water. Molybdate-treated systems served as non-sulfate-reducing controls.

Results

At harvest, activity and numbers of sulfate-reducing micro-organisms were higher in the absence of molybdate, but a rhizosphere effect and an impact of the presence of high levels of iron were not observed on activity and numbers. Both soils had in common a diverse community of sulfate-reducing micro-organisms covering all major cultured bacterial taxa. The appearance of members of the Desulfovibrionaceae exclusively in the rhizosphere of G. maxima was the only unambiguous indication of a plant effect.

Conclusion

The presence of sulfate-rich flood water stimulated the activity and growth of a part of the sulfate-reducing community leading to a change in community composition. The proximity of aerenchymatous plant roots and the abundance of iron in the soil had a negligible effect on the sulfate-reducing community.  相似文献   

6.

Background and aims

Vineyards harbour a variety of weeds, which are usually controlled since they compete with grapevines for water and nutrients. However, weed plants may host groups of fungi and bacteria exerting important functions.

Methods

We grew three different common vineyard weeds (Taraxacum officinalis, Trifolium repens and Poa trivialis) in four different soils to investigate the effects of weeds and soil type on bacterial and fungal communities colonising bulk soil, rhizosphere and root compartments. Measurements were made using the cultivation-independent technique Automated Ribosomal Intergenic Spacer Analysis (ARISA).

Results

Weeds have a substantial effect on roots but less impact on the rhizosphere and bulk soil, while soil type affects all three compartments, in particular the bulk soil community. The fungal, but not the bacterial, bulk soil community structure was affected by the plants at the late experimental stage. Root communities contained a smaller number of Operational Taxonomic Units (OTUs) and different bacterial and fungal structures compared with rhizosphere and bulk soil communities.

Conclusions

Weed effect is localised to the rhizosphere and does not extend to bulk soil in the case of bacteria, although the structure of fungal communities in the bulk soil may be influenced by some weed plants.  相似文献   

7.
Yilin Li  Xingxiang Wang 《Plant and Soil》2013,365(1-2):115-126

Aims

To evaluate the external and internal morphological differences of roots that might influence rice root radial oxygen loss (ROL) and the corresponding rhizosphere nitrification activity, growth characteristics and nitrogen nutrition of rice.

Methods

The root ROL and rhizosphere oxygen profile were determined using a miniaturised Clark-type oxygen microelectrode system, and the rhizosphere nitrification activity was studied with a short-term nitrification activity assay.

Results

The rice biomass, nitrogen accumulation and nitrogen use efficiency (NUE) of ZH (high yield) were significantly higher than those of HS (low yield). The root biomass, number, diameter and porosity of ZH were also much greater than those of HS. The inner and surface oxygen concentrations of the root of ZH were significantly higher than those of HS. The order of paddy soil oxygen penetration depth was ZH?>?HS?>?CK, and the order of the oxygen concentrations detected in the water layer and rhizosphere soil was the same. The rhizosphere nitrification activity and nitrate concentration of ZH were significantly higher than those of HS.

Conclusions

More porous and thicker roots improved the individual root ROL, and more adventitious root numbers enhanced the entire plant ROL and correspondingly improved the rhizosphere nitrification activity, which might influence the growth and nitrogen nutrition of rice.  相似文献   

8.

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.  相似文献   

9.
Chia-Cheng Fan 《Plant and Soil》2012,355(1-2):103-119

Aims

This paper presents a displacement-based model for predicting the relationship between the increase in shear resistance and shear displacement for soils permeated with an entire plant root system.

Methods

The root force in the root system is estimated based on the shear deformation developed in the soil. This displacement-based model takes a number of factors into account, including the distribution of the shear deformation in the soil, the root orientation, the mobilized root forces, and the root properties.

Results

The proposed model reasonably captures the relationship between the increase in the shear resistance (ΔS) and the shear displacement, as shown by a comparison of the predicted results with data from in situ shear tests.

Conclusions

Major findings are the following: (1) the ΔS value increases considerably with increasing b coefficients, which are used to describe the deformed shape of the shear zone, and Young’s moduli of roots at the early stage of shearing; (2) the ΔS value increases significantly with the τ value at large shear deformations; (3) short roots play an important role in the contribution of root systems to the shear resistance of the soil. However, the success of the model relies on the appropriate estimate of the deformation characteristics on the shear zone and the soil-root bond strength.  相似文献   

10.

Aims and background

Despite increasing knowledge of the role of allelochemicals in the productivity decline of replanted Chinese fir plantations, relatively little is known about the levels and sources of allelochemicals in relation to autoinhibition.

Methods

Allelopathic potential of litter, root exudates, and soils in successive rotations of Chinese fir plantations were detected. An allelochemical cyclic dipeptide (6-hydroxy-1,3-dimethyl-8-nonadecyl-[1,4]-diazocane-2,5-dione) from litter, root exudates, and soils in successive rotations was quantified.

Results

Extracts of leaf litter, fine root, and root exudates significantly inhibited the growth of Chinese fir germinants, and inhibition increased with successive rotations. Similar results were observed in the rhizosphere soil, basal soil, and bulk soil. The largest observed inhibition occurred in the rhizosphere soil. Furthermore, cyclic dipeptide was found in litter, root exudates, and soils, and the concentrations increased with successive rotations. The rhizosphere soil had the highest cyclic dipeptide level, followed by basal soil, while bulk soil contained the lowest concentration. There was a significant positive relationship between the inhibition of radicle growth of Chinese fir germinants and the concentration of cyclic dipeptide. Annual release of cyclic dipeptide through root exudation was 2.08–9.78 mol ha?1 annum, but the annual release of cyclic dipeptide through leaf litter decomposition was lowered to 0.32–1.41 mol ha?1 annum.

Conclusions

Cyclic dipeptide which caused autoinhibition of Chinese fir may be released into the soil through litter decomposition and root exudation. Root exudates provided more contributions to soil cyclic dipeptide levels than litter in Chinese fir plantations.  相似文献   

11.

Background

In this issue, Estrada-Medina and coworkers described the diversity of materials in the rhizosphere of the Yucatán karst, México, and quantified the distribution of roots across karst features.

Scope

This commentary explores the implications of their work for below-ground competition and the dynamics of plant-available water on seasonal to inter-annual timescales. Though details differ, seasonal dynamics of water use were consistent with a two-layer model, characterized by water uptake from shallow soil and rock layers during the wet season and deeper soil pockets and rock layers during the dry season. Soil pockets were more densely rooted than rock and experienced large fluctuations in soil moisture, suggesting intense below-ground competition. Total water storage capacity in the rhizosphere was far greater than actual storage in the year of the study. This raises the question whether some storage components in the karst rhizosphere fluctuate on time scales exceeding 1 year.

Conclusions

Despite the significant global extent of karst and their larger than proportional contribution to global biodiversity, vegetation models have ignored their unique rhizosphere structure. Differences in water storage could affect the responses of karst ecosystems and communities to climate change.  相似文献   

12.

Aims

Aluminum-tolerant wheat plants often produce more root exudates such as malate and phosphate than aluminum-sensitive ones under aluminum (Al) stress, which provides environmental differences for microorganism growth in their rhizosphere soils. This study investigated whether soil bacterial community composition and abundance can be affected by wheat plants with different Al tolerance.

Methods

Two wheat varieties, Atlas 66 (Al-tolerant) and Scout 66 (Al-sensitive), were grown for 60 days in acidic soils amended with or without CaCO3. Plant growth, soil pH, exchangeable Al content, bacterial community composition and abundance were investigated.

Results

Atlas 66 showed better growth and lower rhizosphere soil pH than Scout 66 irrespective of CaCO3 amendment or not, while there was no significant difference in the exchangeable Al content of rhizosphere soil between the two wheat lines. The dominant bacterial community composition and abundance in rhizosphere soils did not differ between Atlas 66 and Scout 66, although the bacterial abundance in rhizosphere soil of both wheat lines was significantly higher than that in bulk soil. Sphingobacteriales, Clostridiales, Burkholderiales and Acidobacteriales were the dominant bacteria phylotypes.

Conclusions

The difference in wheat Al tolerance does not induce the changes in the dominant bacterial community composition or abundance in the rhizosphere soils.  相似文献   

13.

Aims

We investigated how rhizosphere factors (total rhizosphere, roots, arbuscular mycorrhizal fungal hyphae [AMF], and soil solution) and water availability affect interactions between neighboring Medicago sativa plants.

Methods

A three-compartment mesocosm was used to test the effects of rhizosphere factors on plant–plant interactions. A relative interaction index (RII) was calculated to indicate whether effects of neighbor plant on target plant were positive or negative (facilitative or competitive). Isotope tracers were used to test whether AMF hyphae mediated competition for nitrogen (N) between target and neighbor plants.

Results

The effects of rhizosphere factors on the interactions between neighboring M. sativa plants depended on water availability. The effects of total rhizosphere shifted RII from negative to positive as water availability increased. Interaction with the roots and rhizosphere soil solution of neighbor plants shifted RII from negative to positive as water availability increased but the opposite was true for AMF hyphae. AMF hyphae helped neighbor plants compete for 15N when water was available but not when water was limiting.

Conclusions

The effect of total rhizosphere on plant–plant interaction of M. sativa shifted from competitive to facilitative as water availability increased. Competition was reduced by neighboring soil solution and roots but was increased by AMF hyphae.  相似文献   

14.

Aims

This study analyzed the extent to which root exudates diffuse from the root surface towards the soil depending on topsoil and subsoil properties and the effect of arbuscular mycorrhizal fungal hyphae on root-derived C distribution in the rhizosphere.

Methods

Alfalfa was grown in three-compartment pots. Nylon gauze prevented either roots alone or roots and arbuscular mycorrhizal fungal hyphae from penetrating into the rhizosphere compartments. 14CO2 pulse labeling enabled the measurement of 14C-labeled exudates in dissolved (DOC) and total organic carbon (TOC) in the rhizosphere, distributed either by diffusion alone or by diffusion, root hair and hyphal transport.

Results

Root exudation and microbial decomposition of exudates was higher in the rhizosphere with topsoil compared to subsoil properties. Exudates extended over 28 mm (DOC) and 20 mm (TOC). Different soil properties and mycorrhization, likely caused by the low arbuscular mycorrhizal colonization of roots (13?±?4 % (topsoil properties) and 18?±?5 % (subsoil properties)), had no effect.

Conclusions

Higher microbial decomposition compensated for higher root exudation into the rhizosphere with topsoil properties, which resulted in equal exudate extent when compared to the rhizosphere with subsoil properties. Higher 14C activity used for labeling compared with previous studies enabled the detection of low exudate concentrations at longer distances from the root surface.  相似文献   

15.

Background and aims

Australian herbaceous native species have evolved in phosphorus (P) impoverished soils. Our objective was to explore shoot and root adaptations of two of these species with potential to be developed as pasture plants, at low, moderate and high P supply after 4 and 7?weeks of growth.

Methods

A glasshouse experiment examined the effect of 5, 20 and 80?mg?P?kg?1 air-dry soil on growth, rhizosphere carboxylate content, and mineral nutrition of two Australian native perennials, Kennedia nigricans (Fabaceae) and Ptilotus polystachyus (Amaranthaceae), and the exotic Medicago sativa (Fabaceae).

Key results

Leaf P concentrations at P80 were 6, 14 and 52?mg?P?g?1 leaf dry weight for M. sativa, K. nigricans and P. polystachyus, respectively. As soil P concentration increased, rhizosphere carboxylate content decreased for M. sativa, increased and then decreased for K. nigricans and was unchanged for P. polystachyus. For all species, the contribution of malic acid declined at the second harvest. For all species and P treatments, the amount of rhizosphere carboxylates per unit root length decreased as root length of a plant increased. Plant P content was determined more by P uptake rate per unit root length and time than by root length. Uptake of Mo for all species, and uptake of K, Mg and Mn for P. polystachyus, increased with soil P concentration. Uptake of Fe and S was higher when the content of carboxylates in the rhizosphere was higher.

Conclusion

Root physiological adaptations (i.e. rhizosphere carboxylate content and P-uptake rate) are more important than morphological adaptations (i.e. root length and diameter) to enhance the uptake of P and cations.  相似文献   

16.

Aims

Zinc (Zn) and phosphorus (P) deficiency often occurs at the same time and limits crop production in many soils. It has been suggested that citrate root exudation is a response of plants to both deficiencies. We used white lupin (Lupinus albus L.) as a model plant to clarify if citrate exuded by roots could increase the bioavailability of Zn and P in calcareous soils.

Methods

White lupin was grown in nutrient solution and in two calcareous soils in a rhizobox. Rhizosphere soil solution was sampled to determine citrate, metals and P. Based on the measured citrate concentrations, a soil extraction experiment with citrate as extractant was done.

Results

Absence of Zn triggered neither cluster root formation nor citrate exudation of white lupin grown in nutrient solution, whereas low P supply did. The maximum citrate concentration (~1.5?mM) found in the cluster rhizosphere soil solution of one soil mobilized P, but not Zn. In the other soil the highest citrate concentration (~0.5?mM) mobilized both elements.

Conclusions

White lupin does not respond to low Zn bioavailability by increasing citrate exudation. Such a response was observed at low P supply only. Whether Zn and P can be mobilized by citrate is soil-dependent and the possible controlling mechanisms are discussed.  相似文献   

17.

Aims

The rhizosphere is a dynamic system strongly influenced by root activity. Roots modify the pH of their surrounding soil causing the soil pH to vary as a function of distance from root surface, location along root axes, and root maturity. Non-invasive imaging techniques provide the possibility to capture pH patterns around the roots as they develop.

Methods

We developed a novel fluorescence imaging set up and applied to the root system of two lupin (Lupinus albus L., Lupinus angustifolius L.) and one soft-rush (Juncus effusus L.) species. We grew plants in glass containers filled with soil and equipped with fluorescence sensor foils on the container side walls. We gained highly-resolved data on the spatial distribution of H+ around the roots by taking time-lapse images of the samples over the course of several days.

Results

We showed how the soil pH in the vicinity of roots developed over time to different values from that of the original bulk soil. The soil pH in the immediate vicinity of the root surface varied greatly along the root length, with the most acidic point being at 0.56–3.36 mm behind the root tip. Indications were also found for temporal soil pH changes due to root maturity.

Conclusion

In conclusion, this study shows that this novel optical fluorescence imaging set up is a powerful tool for studying pH developments around roots in situ.  相似文献   

18.

Aims and background

The ability to suppress soil nitrification through the release of nitrification inhibitors from plant roots is termed ‘biological nitrification inhibition’ (BNI). Earlier, we reported that sorghum roots release higher BNI-activity when grown with NH 4 + , but not with NO 3 - as N source. Also for BNI release, rhizosphere pH of <5.0 is needed; beyond this, a negative effect on BNI release was observed with nearly 80% loss of BNI activity at pH >7.0. This study is aimed at understanding the inter-functional relationships associated with NH 4 + uptake, rhizosphere-pH and plasma membrane H+-ATPase (PM H+-ATPase) activity in regulating the release of BNIs (biological nitrification inhibitors) from sorghum roots.

Methods

Sorghum was grown hydroponically and root exudates were collected from intact plants using a pH-stat system to separate the secondary acidification effects by NH 4 + uptake on BNIs release. A recombinant luminescent Nitrosomonas europaea bioassay was used to determine BNI-activity. Root plasma membrane was isolated using a two-phase partitioning system. Hydrolytic H+-ATPase activity was determined. Split-root system setup was deployed to understand the localized responses to NH 4 + , H+-ATPase-stimulator (fusicoccin) or H+-ATPase-inhibitor (vanadates) on BNI release by sorghum.

Results

Presence of NH 4 + in the rhizosphere stimulated the expression of H+-ATPase activity and enhanced the release of BNIs from sorghum roots. Fusicoccin, which stimulates H+-ATPase activity, also stimulated BNIs release in the absence of NH 4 + ; vanadate, which suppresses H+-ATPase activity, also suppressed the release of BNIs. NH 4 + levels (in rhizosphere) positively influenced BNIs release and root H+-ATPase activity in the concentration range of 0-1.0 mM, indicating a close relationship between BNI release and root H+-ATPase activity with a possible involvement of carrier-mediated transport for the release of BNIs in sorghum.

Conclusion

Our results suggest that NH 4 + uptake, PM H+-ATPase activity, and rhizosphere acidification are functionally inter-connected with BNI release in sorghum. Such knowledge is critical to gain insights into why BNI function is more effective in light-textured, mildly acidic soils compared to other soil types.  相似文献   

19.

Background and aims

Trees allocate a high proportion of assimilated carbon belowground, but the partitioning of that C among ecosystem components is poorly understood thereby limiting our ability to predict responses of forest C dynamics to global change drivers.

Methods

We labeled sugar maple saplings in natural forest with a pulse of photosynthetic 13C in late summer and traced the pulse over the following 3 years. We quantified the fate of belowground carbon by measuring 13C enrichment of roots, rhizosphere soil, soil respiration, soil aggregates and microbial biomass.

Results

The pulse of 13C contributed strongly to root and rhizosphere respiration for over a year, and respiration comprised about 75 % of total belowground C allocation (TBCA) in the first year. We estimate that rhizosphere carbon flux (RCF) during the dormant season comprises at least 6 % of TBCA. After 3 years, 3.8 % of the C allocated belowground was recovered in soil organic matter, mostly in water-stable aggregates.

Conclusions

A pulse of carbon allocated belowground in temperate forest supplies root respiration, root growth and RCF throughout the following year and a small proportion becomes stabilized in soil aggregates.  相似文献   

20.

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.  相似文献   

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