A kinetic study of phosphorus absorption by excised maize roots from flowing solutions influenced by 2,4 dinitrophenol and viscosity of solution

The effect of 2,4 dinitrophenol and increased viscosity of the absorption solution on the absorption of phosphorus by excised roots of maize plants was investigated. The concentration of the solution was 0.1 mM KH₂PO₄, the activity of³²P was 52 µCi l^-1. The temperature of the absorption solution was 26 °C, pH 5.5, aeration prior to the experiment. There was 11 of solution for every 1 g of roots. Two basic variants were used for comparison: with non-flowing solution and with solution flow (circulation) of 0.162 cm s^-1, respectively. In all cases, 2,4 dinitrophenol reduced the rate of phosphorus absorption by the roots regardless of the mechanism of phosphorus supply to the roots (diffusion, mass flow). If it is proved that 2,4 dinitrophenol inhibits the active uptake of phosphorus, then the uptake of phosphorus by the roots increased under the influence of mass flow will be active,i.e., connected with energy metabolism. Raising the viscosity of the absorption solution 3.3 times over that of water by means of potato starch substantially reduced the absorption of the phosphorus transported to the absorption zone by diffusion and practically did not affect the rate of absorption, or the amount of anions transported to the absorption area by mass flow.     

Phosphorus absorption by intact maize plants from flowing solutions influenced by 2.4-dinitrophenol and viscosity of solution

The investigation of phosphorus absorption by intact plants during a short period has above all confirmed the validity of the results obtained in the foregoing study of the kinetics of absorption by excised roots. Further, the results show the unquestionably important role of mass flow in transporting ions to plant roots, mainly at lower and medium concentrations, that is, from about 0.1 to 10 mM. Under conditions of growth close to the optimum, the supply by means of mass flow can be sufficient even at lower concentrations of phosphorus, such as 1.47 mM KH₂PO₄, or the absorption of phosphorus by plants can be higher than in the case of ions being transported to roots by diffusion. With a higher absorption the phosphorus distribution somewhat changes as well, relatively more of it being accumulated in the roots. 2.4-DNP applied to the nutrient solution at a concentration of 10^-5 M reduces the phosphorus absorption.     

Effect of adsorbed cations on phosphorus uptake by excised roots

Pretreatment of excised roots of Hordeum vulgare, Zea mays, and Glycine max with various salt solutions affected their subsequent rate of phosphorus absorption from 2 × 10⁻⁵m KH₂PO₄. The rate of absorption was greatest for roots pretreated with trivalent cations, intermediate with divalent cations and lowest with monovalent cations. It appeared that the pretreatment involved a rapid exchange reaction at the root surface which was reversible. A 1 min pretreatment was effective for more than 20 min. The acceleration of phosphorus uptake by roots produced by polyvalent cations may be due partly or entirely to a greater reduction in the electrical potential at the root surface or within the pores of the negatively charged cell wall by polyvalent cations than by monovalent cations.     

Increasing phosphorus concentration in the extraradical hyphae of <Emphasis Type="Italic">Rhizophagus irregularis</Emphasis> DAOM 197198 leads to a concomitant increase in metal minerals

Plants associated with arbuscular mycorrhizal fungi (AMF) acquire phosphorus via roots and extraradical hyphae. How soil P level affects P accumulation within hyphae and how P in hyphae influences the accumulation of metal minerals remains little explored. A bi-compartmented in vitro cultivation system separating a root compartment (RC), containing a Ri T-DNA transformed carrot root associated to the AMF Rhizophagus irregularis DAOM 197198, from a hyphal compartment (HC), containing only the extraradical hyphae, was used. The HC contained a liquid growth medium (i.e., the modified Strullu-Romand medium containing P in the form of KH₂PO₄) without (0 μM) or adjusted to 35, 100, and 700 μM of KH₂PO₄. The accumulation of P and metal minerals (Ca, Mg, K, Na, Fe, Cu, Mn) within extraradical hyphae and AMF-colonized roots, and the expression of the phosphate transporter gene GintPT were assessed. The expression of GintPT in the extraradical hyphae did not differ in absence of KH₂PO₄ or in presence of 35 and 100 μM KH₂PO₄ in the HC but was markedly reduced in presence of 700 μM KH₂PO₄. Hyphal P concentration was significantly lowest in absence of KH₂PO₄, intermediate at 35 and 100 μM KH₂PO₄ and significantly highest in presence of 700 μM KH₂PO₄ in the HC. The concentrations of K, Mg, and Na were positively associated with the concentration of P in the extraradical hyphae developing in the HC. Similarly, P concentration in extraradical hyphae in the HC was related to P concentration in the growth medium and influenced the concentration of K, Mg, and Na. The accumulation of the metal mineral K, Mg, and Na in the extraradical hyphae developing in the HC was possibly related to their function in neutralizing the negative charges of PolyP accumulated in the hyphae.     

Kinetics of phosphorus absorption by mycorrhizal and nonmycorrhizal tomato roots

Kinetics of P absorption were investigated in mycorrhizal (Glomus fasciculatus) and nonmycorrhizal tomato (Lycopersicon esculentum) roots to determine why increased ion absorption by mycorrhizae occurs. Initial rates of absorption of ³²P were measured at 1 to 100 micromolar KH₂PO₄ (pH 4.6). Absorption rates of mycorrhizae were about twice those of control roots. Augustinsson-Hofstee analysis yielded two linear phases; V_max and K_m were calculated for each phase. In the low phase (1 to 20 micromolar), V_max values for the mycorrhizal and nonmycorrhizal roots were each 0.10 micromoles P per gram fresh weight per hour while K_m values were 1.6 and 3.9 micromolar KH₂PO₄, respectively. For the high phase (30 to 100 micromolar), V_max values for mycorrhizal and nonmycorrhizal roots were 0.32 and 0.25 micromoles P per gram fresh weight per hour and K_m values were 35 and 42 micromolar, respectively. These results indicate that at the lower phase concentrations, similar to those expected in most soil solutions, a major factor contributing to the increased uptake was an apparent greater affinity of the absorbing sites for H₂PO₄⁻ (lower K_m).     

Solubility of phosphorus added to four Manitoba soils with different calcium and magnesium contents

Summary The solubility of phosphorus was found to approximate that of dicalcium phosphate dihydrate and/or dimagnesium phosphate trihydrate when KH₂-PO₄, H₃PO₄ and K₂HPO₄ were added to four Manitoba soils. Eighty to one hundred, seventy to ninety and sixty to eighty per cent of the phosphorus added remained in solution when H₃PO₄, KH₂PO₄ and K₂HPO₄ were added, respectively. The solubility of the added phosphorus was high in all samples and relatively soluble compounds, dicalcium phosphate dihydrate and dimagnesium phosphate trihydrate, were most likely formed in the samples indicating that phosphorus added to these soils would be readily available to plants. Associate Professor and Professor respectively.     

Effect of seed pretreatments on phosphorus uptake by alfalfa

Summary A pot experiment was made to study the effect of two seed pretreatments on the phosphorus uptake by alfalfa in two different soils. The results showed that the dry weight of both the tops and the roots as well as the total P-uptake of alfalfa in both soils increased when the seeds were previously soaked in 0.5 mol/l KH₂PO₄ or 150 ppm indoleacetic acid for 6 hours when compared with the control. The increase was much higher with the seeds soaked in the former than in the later solution. Both soil and fertilizer phosphorus taken up by alfalfa increased by seed soaking in the above mentioned solutions and the increase was most pronounced in the fraction that derived from the soil.Professor of Soils, Professor of Plant Nutrition and a Lecturer in the Faculty of Agriculture, Ain Shams University, Cairo, respectively.     

Influence of ectomycorrhizal fungi on the response of Sitka spruce and Japanese larch to forms of phosphorus

Ectomycorrhizal fungi (Paxillus involutus, Suillus grevillei and two unidentified basidiomycetes from excised Sitka spruce mycorrhizas) were isolated from stands of Sitka spruce either in monoculture or in a mixture with Japanese larch in an Irish conifer plantation. The growth of these fungi and their mycorrhizal formation in Sitka spruce and Japanese larch were examined after incubation in modified Melin-Norkrans medium containing either KH₂PO₄, Ca₃(PO₄)₂ or Fe phytate as the phosphorus (P) source. P. involutus and S. grevillei utilized all three P sources. The unidentified basidiomycetes had limited ability to utilize Fe phytate. Basidiomycete 1 showed poor growth on KH₂PO₄ whereas growth of basidiomycete 2 was low on Ca₃(PO₄)₂. Pure culture synthesis studies confirmed that P. involutus and the two basidiomycetes formed mycorrhizas with both tree species but S. grevillei was mycorrhizal only on Japanese larch. P. involutus formed more mycorrhizas in both conifers than the other fungi. Following inoculation with each of the four fungi, shoot and root dry mass of both Sitka spruce and Japanese larch seedlings was enhanced compared with uninoculated/nonmycorrhizal controls. On Fe phytate, Paxillus-inoculated Sitka spruce seedlings had the lowest primary root length and on KH₂PO₄, Suillus-inoculated Japanese larch had the greatest number of short roots. The only differences when Sitka spruce seedlings were grown in either monoculture or in a mixture with Japanese larch mycorrhizal with S. grevillei were primary root length and number of short roots after growth on media containing Fe phytate.     

Differential expression of genes involved in alternative glycolytic pathways,phosphorus scavenging and recycling in response to aluminum and phosphorus interactions in <Emphasis Type="Italic">Citrus</Emphasis> roots

The objective was to determine the possible links between the expression levels of genes involved in alternative glycolytic pathways, phosphorus (P) scavenging and recycling and Citrus tolerance to aluminum (Al) and/or P-deficiency. ‘Xuegan’ (Citrus sinensis) and ‘Sour pummelo’ (Citrus grandis) seedlings were irrigated for 18 weeks with nutrient solution containing 0 and 1.2 mM AlCl₃·6H₂O × 0, 50 and 200 μM KH₂PO₄. C. sinensis displayed more tolerant to Al and P-deficiency than C. grandis. Under Al stress, C. sinensis accumulated more Al in roots and less Al in shoots than C. grandis. P concentration was higher in C. sinensis shoots and roots than in C. grandis ones. C. sinensis roots secreted more malate and citrate than C. grandis ones when exposed to Al. Al-induced-secretion of malate and citrate by excised roots from Al-treated seedlings decreased with increasing P supply. Al-induced-secretion of malate and citrate from roots and Al precipitation by P in roots might be responsible for Al-tolerance of C. sinensis. qRT-PCR analysis showed that Al-activated malate transporter (ALMT1), ATP-dependent phosphofructokinase (ATP-PFK), pyrophosphate-dependent phosphofructokinase (PPi-PFK), tonoplast adenosine-triphosphatase subunit A (V-ATPase A), tonoplast pyrophosphatase (V-PPiase), pyruvate kinase (PK), acid phosphatase (APase), phosphoenolpyruvate carboxylase (PEPC), malic enzyme (ME) and malate dehydrogenase (MDH) genes might contribute to the tolerance of Citrus to Al and/or P-deficiency, but any single gene could not explain the differences between the two species. Citrus tolerance to Al and/or P-deficiency might be caused by the coordinated regulation of gene expression involved in alternative glycolytic pathways, P scavenging and recycling.     

Direct assessment of mineral phosphorus availability to tropical crops using 32P labelled compounds

The availability to plants of phosphorus (P) derived from sparingly soluble iron and aluminium phosphates was directly assessed with ³²P labelled compounds in two glasshouse trials.In the first experiment, the comparative availability of all mineral phosphate (P) compounds to maize increased with time (14 to 42 days post emergence) and plant total P uptake, but P source did not affect the growth or total plant uptake of P. The comparative availability of the amorphous AlPO₄ (Al-P), crystalline AlPO₄ (variscite), amorphous FePO₄ (Fe-P), and crystalline FePO₄ (strengite) compared to KH₂PO₄ (=100) was 53.1, 3.4, 38.9, and 1.9%, respectively. In the second experiment, the availability of Fe-P, strengite, and KH₂PO₄ to several topical crop species was examined. There was no difference between maize, sorghum, mungbean, cowpea or soybean in their ability to utilise Fe-P or KH₂PO₄, although maize utilised strengite more than the other species. The major difference between these species in their ability to acquire P appears to be a difference in ability to locate soluble soil P rather than differences in their ability to access different pools of soil P.The advantages of using neutron irradiation to directly measure P absorption from mineral P compounds over traditional methodologies is discussed.     

Influence of the phosphorus and nitrogen source on nisin production in Lactococcus lactis subsp. lactis batch fermentations using a complex medium

The influence of different phosphorus and nitrogen sources on Lactococcus lactis subsp. lactis NIZO 22186 growth and nisin production was studied in batch fermentations using a complex medium. KH₂PO₄ was found to be the best phosphorus source for nisin production. Increasing initial phosphate concentrations from 0 to 5% KH₂PO₄ exerted a double effect, creating favourable pH conditions and particularly stimulating the nisin production levels, which were highest at 5% KH₂PO₄. Up to now, no such high initial phosphate concentrations have been reported for the production of other antibiotics or bacteriocins. Nisin, a lanthionine-containing peptide antibiotic with bacteriocin properties, clearly behaved as a primary metabolite, since its formation was linked with active growth and was not suppressed by phosphate concentrations up to 5%. A complex medium supplemented with cotton seed meal as nitrogen source also gave very high nisin yields. Correspondence to: L. De Vuyst     

The Effect of Root Temperatures on Water and Sulphate Absorption in Intact Sunflower Plants

The effect of 15, 25, and 35°C root temperature on waterabsorption, transpiration, and sulphate uptake by the rootsand transport to the shoots of intact sunflower plants has beenstudied using 0.5, 5.0, and 50.0 mM sulphate concentrationsat two rates of transpiration induced (1) by light and low relativehumidity and (2) by darkness and high relative humidity. Root temperatures and sulphate concentrations did not significantlyaffect the water absorption and transpiration and both theseprocesses were approximately similar at the different treatments.There was a nearly twofold increase in water absorption andtranspiration in the light and low relative humidity as comparedto the dark and high relative humidity irrespective of the roottemperatures and sulphate concentrations. The A.F.S. uptake in the roots was found to be independent ofthe root temperatures, sulphate concentrations, and transpirationrates, and amounted to 15 to 21 per cent based on the root weight.Sulphate accumulation in the roots was not significantly influencedby the root temperatures at 0.5 and 5.0 mM sulphate concentrations,but nearly doubled with temperature at 50.0 mM sulphate concentrationof the external solution. The slow nature of accumulation ofsulphate, the high sulphate status of the experimental plants,and the short duration of the experiments are considered aslikely reasons for the absence of a clear effect of temperatureson accumulation of sulphate at the two lower concentrationsof the external solution. Effects of high concentration on permeabilityand metabolism of the cells are suggested as the reasons forthe decreased accumulation with an increase in temperature at50.0 mM sulphate concentration. Accumulation of sulphate inthe roots was not significantly influenced by the transpirationrates. Unlike root accumulation, sulphate transport to the shoots increasedwith increasing transpiration. However, a major part of thesulphate transport (70 to 75 per cent at 0.5 and 5.0 mM sulphateconcentrations and 80 to 85 per cent at 50.0 mM sulphate concentration)appeared to have occurred at the low transpiration. The similarityof this transport to the accumulation of sulphate in the rootsindicates that it was due to an active transport process sensitiveto root temperatures and sulphate concentrations. A low concentrationof sulphate in the xylem and an increased permeability of theroot cells to ion movement induced by an increased suction inthe xylem are considered as reasons for a small increase inthe sulphate transport at high transpiration rate. The evidencefor the existence of a barrier—probably endodermis—preventingthe passive diffusion of sulphate and sensitivity of the TranspirationStream Concentration to root temperatures and sulphate concentrationsfavour that the increased transport with increased transpirationwas due to an active process.     

Alleviation of phosphorus deficiency stress by moderate salinity in the halophyte <Emphasis Type="Italic">Hordeum maritimum</Emphasis> L.

Hordeum maritimum (Poacea) is a facultative halophyte potentially useful for forage production in saline zones. Here, we assessed whether moderate NaCl-salinity can modify the plant response to phosphorus (P) shortage. Plants were cultivated for 55 days under low or sufficient P supply (5 or 60 μmol plant⁻¹ week⁻¹ KH₂PO₄, respectively), with or without 100 mM NaCl. When individually applied, salinity and P deficiency significantly restricted whole-plant growth, with a more marked effect of the latter stress. Plants subjected to P deficiency showed a significant increase in root growth (as length and dry weight) and root/shoot DW ratio. Enhanced root growth and elongation presumably correspond to the well-known root adaptive response to mineral deficiency. However, leaf relative water content, leaf P concentration, and leaf gas exchange parameters were significantly restricted. The interactive effects of salinity and P deficiency were not added one to another neither on whole plant biomass nor on plant nutrient uptake. Indeed, 100 mM NaCl-addition to P-deficient plants significantly restored the plant growth and improved CO₂ assimilation rate, root growth, K⁺/Na⁺ ratio and leaf proline and soluble sugar concentrations. It also significantly enhanced leaf total antioxidant capacity and leaf anthocyanin concentration. This was associated with significantly lower leaf osmotic potential, leaf Na⁺ and malondialdehyde (MDA) concentration. Taken together, these results suggest that mild salinity may mitigate the adverse effects of phosphorus deficiency on H. maritimum by notably improving the plant photosynthetic activity, the osmotic adjustment capacity, the selective absorption of K⁺ over Na⁺ and antioxidant defence.     

Active transport of ions across sunflower roots

Summary The electrical potential difference across exuding roots of Helianthus annuus in two strengths of complete culture solution was measured. The determination of the concentration of the major nutrient ions in the outside solution and the xylem sap enabled the Nernst potential for each ion to be calculated. A comparison of the measured and calculated potentials indicated that the anions NO₃, SO₄, H₂PO₄ and HPO₄ were actively transported into the sap against the electrochemical potential gradient. The cations Ca and Mg, on the other hand, appeared to move passively into the sap. The behaviour of potassium depended on its concentration in the medium. With a relatively low external concentration (0.75 mM) it appeared to be actively tansported into the sap, whilst at higher outside concentrations (7.5 mM) it was apparently moving passively into the xylem down the electrochemical potential gradient. The possibility of potassium being pumped out of the sap with relatively high external concentrations is discussed.     

Growth and Phosphate Transport in Barley and Tomato Plants During the Development of, and Recovery from, Phosphate-stress

Barley and tomato plants were cultured in nutrient solutionsincluding 0.15 mol m^–3 H²PO^–₄. The phosphate supplywas discontinued and the subsequent effects on growth, internalphosphorus concentrations, phosphate absorption and translocationwere measured at frequent intervals. Growth rates were at firstunchanged and the internal phosphorus concentration decreased.During this phase the rate of phosphate transport by the rootssometimes increased significantly. Growth slowed more in shootsthan in roots during a second phase of stress development andvisual symptoms of deficiency appeared in tomato but not inbarley. During this phase, enhancement of phosphate uptake capacityreached a maximum in both species. The subsequent decline inuptake capacity was associated with visible symptoms of deficiencydeveloping in barley and intensifying in tomato. When stressedplants were returned to a solution containing 0.15 mol m^–3H₂PO^–₄ rapid absorption continued for several days afterthe internal phosphorus concentration had returned to the levelof the controls. Phosphate toxicity may have been the causeof leaf lesions and necrosis during the ‘recovery’phase. Stomatal conductance in tomato was decreased at an early stageof stress development. Foliar-applied phosphate was absorbedmore rapidly by P-stressed barley leaves than by their controlsand much larger amounts were translocated from the leaves tothe roots.     

Effect of byproduct,nitrogen fertilizer,and zeolite on phosphate rock dissolution and extractable phosphorus in acid soil

The relative plant availability of selenate versus selenite depends on the concentrations of competing ions, specifically sulfate and phosphate, respectively. In solution culture, the concentration of phosphate is typically 100- to 1000-fold greater than in soil solution, an artifact that could lead to underestimation of the phytoavailability of selenite. A nutrient solution study was conducted to compare the availability of selenite and selenate to perennial ryegrass (Lolium perenne L. cv. Evening Shade) and strawberry clover (Trifolium fragiferrum L. cv. O'Conner) at basal concentrations of SO₄ (0.5 mM) and PO₄ (2 μM) similar to those found in soil solution. Concentrations up to 5 mM SO₄ and 200 μM PO₄ allowed quantitative comparison of the efficacy of the competing ions. In both species, selenite was more phytotoxic than selenate, especially for shoot growth. Selenate was less toxic, and tended to preferentially inhibit root growth. Translocation percentages were much higher with selenate (≥84%) than with selenite (≤47%). A 10-fold increase in sulfate decreased uptake from selenate by >90% in both species. In ryegrass, 10-fold increases in phosphate caused 30% to 50% decreases in Se accumulation from selenite, but in clover such decreases only occurred in the roots. Sulfate-selenate antagonisms were thus stronger than phosphate-selenite antagonisms. Nonetheless, conventional nutrient solutions with millimolar phosphate will significantly underestimate Se availability from selenite, and moderate levels of sulfate salinity can inhibit selenate uptake sufficiently to reverse the apparent relative availability of the two forms of Se. This revised version was published online in June 2006 with corrections to the Cover Date.     

Maize growth and ion absorption in Richter's solution at different flow rates

By comparing maize plants cultivated in standing nutrient solution with those from solutions flowing at different flow rates it has been established that absorption of nitrogen, potassium and especially of phosphorus was increased owing to the flow. There was likewise a relative rise in the distribution of nutrients to the overground parts of the plants. The content expressed per unit dry matter was increased only in the case of phosphorus; with nitrogen and potassium it was slightly lower than in the standing solution. Increasing amounts of iron were required under the conditions of flowing nutrient solutions to prevent chlorosis of the plants. The production of dry matter,NAR andRGR was also increased because of the flow. The flow considerably changed the habitus of the primary roots of the maize plants. The roots were longer, thinner and on the whole they contained relatively less dry matter (RWR). The lengthening of the roots is explained as a response to stimulation by the solution flow—the rheotropism.     

Development and application of a nutrient-diffusing bioassay for large rivers

1. Laboratory and field experiments were performed to develop and then apply a nutrient-diffusing substratum (NDS) design suitable for use in large, fast-flowing rivers. 2. Initial laboratory experiments quantified diffusion of PO₄ and NO₃ from new and previously used clay pots, which were soaked in deionized distilled water. Mean release rates initially exceeded 2.4 and 725 μmol l^–1 day^–1 P and 0.22 and 18 μmol l^–1 day^–1 N from new and used pots, respectively, but declined rapidly with increasing time spent in deionized distilled water and were below detectable levels after about 18 and 29 days, respectively. 3. A phosphorus (P) dose–response experiment in a P-limited reach of the Athabasca River, Alberta, Canada showed that epilithic biomass and macroinvertebrate density on NDS increased with increasing concentrations of KH₂PO₄ up to about 0.5 m . Beyond this threshold, biomasses and densities were unaffected by initial KH₂PO₄ concentration. Coefficients of variation of epilithic biomass estimates declined with increasing KH₂PO₄ whereas invertebrate density appeared to be unaffected by KH₂PO₄ levels. 4. Release rates of both P and N from NDS filled with 0.5 m KH₂PO₄ or 0.5 m NaNO₃ declined at a log-negative rate from about 5000 μmol N-NO₃ l^–1 day^–1 and 3500 μmol P-PO₄ l^–1 day^–1 on day 2, to 200 μmol l^–1 day^–1 for both N and P on day 32. 5. After development, we used the diffusing substrata to identify spatial patterns in nutrient limitation at seven sites along a 120 km reach in the Athabasca River, that receives two known point-source nutrient inputs. NDS consisting of N, P, N + P and unenriched controls were attached to the river bottom for 22–23 days and then retrieved and sampled for epilithic chlorophyll a. Physicochemical parameters and epilithic biomasses on upper stone surfaces were also quantified when NDS were deployed and retrieved from each site. 6. Sites located immediately downstream of the two point source inputs had higher water column concentrations of PO₄ and epilithic biomasses than the site immediately upstream; epilithic biomass was positively related to PO₄ in the late autumn (r²⁼ 0.58) but not in early autumn. Sites located immediately below nutrient inputs were not nutrient-limited, whereas upstream reference sites were P-limited.     

Effects of water stress on phosphorus transport to the xylem

Summary Tomato plants were treated for one hour in nutrient solutions at-10.4 atm. Roots were excised, transferred to solutions at-0.4 atm and put into a pressure chamber to induce rates of water flow similar to those in transpiring plants.For roots continuously at-0.4 atm, the xylem sap had much higher phosphorus concentrations than the external solution, which contained 6 p.p.m. phosphorus.Roots previously treated at-10.4 atm had much lower concentrations in the sylem sap than in the external solution and the amount of phosphorus transported and the water flow were linearly related. This phosphorus transport was due to passive movement as shown by measuring transport of both ³²P and ¹⁴C mannitol. Thus transport to the xylem mediated by active processes was abolished even though uptake by the roots remained substantial. These results obtained after plasmolysis support the view that radial transport to the xylem includes uptake into and movement through the symplast.