Silicon reduces sodium uptake in rice (Oryza sativa L.) in saline conditions and this is accounted for by a reduction in the transpirational bypass flow

Rice is relatively sensitive to salinity and is classified as a silicon accumulator. There have been reports that silicon can reduce sodium uptake in crop grasses in saline conditions, but the mechanism by which silicon might alleviate salinity damage is unclear. We report on the effects of silicon on growth, gas exchange and sodium uptake in rice genotypes differing in salt tolerance. In non-saline media there were no effects of supplementary silicate upon shoot fresh or dry weight or upon root dry weight, indicating that the standard culture solution was not formally deficient with respect to silicon. Plants grown with supplementary silicate had slightly, but significantly, shorter leaves than plants grown in a standard culture solution. Salinity reduced growth and photosynthetic gas exchange. Silicate supplementation partly overcame the reduction in growth and net photosynthesis caused by salt. This amelioration was correlated with a reduction in sodium uptake. Silicate supplementation increased the stomatal conductance of salt-treated plants, showing that silicate was not acting to reduce sodium uptake via a reduction in the transpiration rate. Silicate reduced both sodium transport and the transport of the apoplastic tracer trisodium-8-hydroxy-1,3,6-pyrenetrisulphonic acid (PTS). This implies that the mode of action of silicate was by partial blockage of the transpirational bypass flow, the pathway by which a large proportion of the uptake of sodium in rice occurs. Mechanisms by which silicate might reduce the transpirational bypass flow directly are discussed.     

A new screening technique for salinity resistance in rice (Oryza sativa L.) seedlings using bypass flow

A lack of screening techniques delays progress in research on salinity resistance in rice. In this study, we report our test of the hypothesis that an apoplastic pathway (the so-called bypass flow) causes a difference in salt resistance between rice genotypes and can be used in screening for salinity resistance. Fourteen-day-old seedlings of low- and high-Na(+) -transporting recombinant inbred lines (10 of each) of rice IR55178 were treated with 50 mm NaCl and 0.2 mm trisodium-8-hydroxy-1,3,6-pyrenetrisulphonic acid (PTS), a bypass flow tracer, for short (4 d) and long (90 d) periods of time. The results showed that the average shoot Na(+) concentration and bypass flow for high-Na(+) -transporting lines were 1.4 and 2.4 times higher than those of low-Na(+) -transporting lines, respectively. There was a positive linear correlation between the percentage of bypass flow and Na(+) concentrations in the shoots, suggesting that the difference in Na(+) transport in rice is a consequence of different degrees of bypass flow. Moreover, a high correlation was found between bypass flow and seedling survival after prolonged salt stress: the lower the magnitude of bypass flow, the greater the seedling survival. We conclude that bypass flow could be used as a new screening technique for salt resistance in rice.     

The role of lateral roots in bypass flow in rice (Oryza sativa L.)

Although an apoplastic pathway (the so‐called bypass flow) is implicated in the uptake of Na⁺ by rice growing in saline conditions, the point of entry of this flow into roots remains to be elucidated. We investigated the role of lateral roots in bypass flow using the tracer trisodium‐8‐hydroxy‐1,3,6‐pyrenetrisulphonic acid (PTS) and the rice cv. IR36. PTS was identified in the vascular tissue of lateral roots using both epifluorescence microscopy and confocal laser scanning microscopy. Cryo‐scanning electron microscopy and epifluorescence microscopy of sections stained with berberine‐aniline blue revealed that the exodermis is absent in the lateral roots. We conclude that PTS can move freely through the cortical layers of lateral roots, enter the stele and be transported to the shoot via the transpiration stream.     

Net sodium fluxes change significantly at anatomically distinct root zones of rice (Oryza sativa L.) seedlings

Casparian bands of endodermis and exodermis play crucial roles in blocking apoplastic movement of ions and water into the stele of roots through the cortex. These apoplastic barriers differ considerably in structure and function along the developing root. The present study assessed net Na⁺ fluxes in anatomically distinct root zones of rice seedlings and analyzed parts of individual roots showing different Na⁺ uptake. The results indicated that anatomically distinct root zones contributed differently to the overall uptake of Na⁺. The average Na⁺ uptake in root zones in which Casparian bands of the endo- and exo-dermis were interrupted by initiating lateral root primordia (root zone III) was significantly greater than that at the root apex, where Casparian bands were not yet formed (root zone I), or in the region where endo- and exo-dermis with Casparian bands were well developed (root zone II). The measurement of net Na⁺ fluxes using a non-invasive scanning ion-selective electrode technique (SIET) demonstrated that net Na⁺ flux varied significantly in different positions along developing rice roots, and a net Na⁺ influx was obvious at the base of young lateral root primordia. Since sodium fluxes changed significantly along developing roots of rice seedlings, we suggest that the significantly distinct net Na⁺ flux profile may be attributed to different apoplastic permeability due to lateral root primordia development for non-selective apoplastic bypass of ions along the apoplast.     

Phosphorylation upon cold stress in rice (Oryza sativa L.) seedlings

The response of plants to cold stress is not well understood at the biochemical level, although it has been studied extensively at the ecological level. To investigate whether protein phosphorylation may play an important role in cold stress, we exposed rice seedlings to low temperatures, prepared protein extracts from the leaves and incubated these in the presence of [γ-³²P]ATP. The proteins were then separated by two-dimensional polyacrylamide gel electrophoresis. While several proteins were found to be phosphorylated upon cold stress one protein, pp35, which has an isoelectric point of 8.0, was more phosphorylated than the others. The pp35 protein was found to be phosphorylated when rice seedlings were incubated for 6 h at 5°C before the leaf protein extract was prepared and radioactive labeling was performed. The pp35 was, however, significantly more phosphorylated in cold-tolerant rice varieties. Antibodies were raised against purified pp35 in adult rabbits. Using this pp35 antibody, which can recognize the RuBisCO large-chain subunit (LSU), and from amino acid sequencing of pp35, we were able to identify and confirm the pp35 protein as the fragment of RuBisCO LSU (EC 4.1.1.39). Phosphorylation of the RuBisCO LSU may be important in cold tolerance. Received: 7 July 1998 / Accepted: 19 December 1998     

Exogenous sodium nitroprusside and glutathione alleviate copper toxicity by reducing copper uptake and oxidative damage in rice (Oryza sativa L.) seedlings

Nitric oxide (NO) and glutathione (GSH) regulate a variety of physiological processes and stress responses; however, their involvement in mitigating Cu toxicity in plants has not been extensively studied. This study investigated the interactive effect of exogenous sodium nitroprusside (SNP) and GSH on Cu homeostasis and Cu-induced oxidative damage in rice seedlings. Hydroponically grown 12-day-old seedlings were subjected to 100 μM CuSO₄ alone and in combination with 200 μM SNP (an NO donor) and 200 μM GSH. Cu exposure for 48 h resulted in toxicity symptoms such as stunted growth, chlorosis, and rolling in leaves. Cu toxicity was also manifested by a sharp increase in lipoxygenase (LOX) activity, lipid peroxidation (MDA), hydrogen peroxide (H₂O₂), proline (Pro) content, and rapid reductions in biomass, chlorophyll (Chl), and relative water content (RWC). Cu-caused oxidative stress was evident by overaccumulation of reactive oxygen species (ROS; superoxide (O₂ ^?–) and H₂O₂). Ascorbate (AsA) content decreased while GSH and phytochelatin (PC) content increased significantly in Cu-stressed seedlings. Exogenous SNP, GSH, or SNP?+?GSH decreased toxicity symptoms and diminished a Cu-induced increase in LOX activity, O₂ ^?–, H₂O₂, MDA, and Pro content. They also counteracted a Cu-induced increase in superoxide dismutase (SOD), ascorbate peroxidase (APX), glutathione reductase (GR), monodehydroascorbate reductase (MDHAR), and glyoxalase I and glyoxalase II activities, which paralleled changes in ROS and MDA levels. These seedlings also showed a significant increase in catalase (CAT), glutathione peroxidase (GPX), dehydroascorbate reductase (DHAR), glutathione S-transferase (GST) activities, and AsA and PC content compared with the seedlings stressed with Cu alone. Cu analysis revealed that SNP and GSH restricted the accumulation of Cu in the roots and leaves of Cu-stressed seedlings. Our results suggest that Cu exposure provoked an oxidative burden while reduced Cu uptake and modulating the antioxidant defense and glyoxalase systems by adding SNP and GSH play an important role in alleviating Cu toxicity. Furthermore, the protective action of GSH and SNP?+?GSH was more efficient than SNP alone.     

Identification of salt-stress responsive genes in rice (Oryza sativa L.) by cDNA array

To identify salt stress-responsive genes, we constructed a cDNA library with the salt-tolerant rice cultivar, Lansheng. About 15000 plasmids were extracted and dotted on filters with Biomeck 2000 HDRT system or by hand. Thirty genes were identified to display altered expression levels responding to 150 mmol/L NaCl. Among them eighteen genes were up-regulated and the remainders down-regulated. Twenty-seven genes have their homologous genes in Gen-Bank Databases. The expression of twelve genes was studied by Northern analysis. Based on the functions, these genes can be classified into five categories, including photosynthesis-related gene, transport-related gene, metabolism-related gene, stress- or resistance-related gene and the others with various functions. The results showed that salt stress influenced many aspects of rice growth. Some of these genes may play important roles in plant salt tolerance.     

Saturated humidity accelerates lateral root development in rice (Oryza sativa L.) seedlings by increasing phloem-based auxin transport

Auxin transport plays a significant role modifying plant growth and development in response to environmental signals such as light and gravity. However, the effect of humidity on auxin transport is rarely documented. It is shown here that the transport of labelled indole-3-acetic acid (IAA) from the shoot to the root is accelerated in rice (Oryza sativa L. ssp. indica cv. IR8) seedlings grown under saturated humidity (SH-seedlings) compared with plants grown under normal humidity (NH-seedlings). The development of lateral roots in SH-seedlings was greatly enhanced compared with NH-seedlings. Removal of the shoot from SH-seedlings reduced the density of lateral roots, and the application of IAA to the cut stem restored the lateral root density, while the decapitation of NH-seedlings did not alter lateral root development. Phloem-based auxin transport appeared responsible for enhanced lateral root formation in SH-seedlings since (i) the rate of IAA transport from the shoot to the root tip was greater than 3.5 cm h-1 and (ii) naphthylphthalamic acid (NPA)-induced reduction of polar auxin transport in the shoot did not influence the number of lateral roots in SH-seedlings. It is proposed that high humidity conditions accelerate the phloem-based transport of IAA from the leaf to the root, resulting in an increase in the number of lateral roots.     

Effect of silicate on the growth and arsenate uptake by rice (Oryza sativa L.) seedlings in solution culture

A solution culture experiment was conducted to investigate the effect of silicate on the yield and arsenate uptake by rice. Rice seedlings (Oryza sativaL. cv. Weiyou 77) were cultured in modified Hoagland nutrient solution containing three arsenate levels (0, 0.5 and 1.0 mg L ^–1 As) and four silicate levels (0, 14, 28 and 56 mg L ^–1 Si). Addition of Si significantly increased shoot dry weight (P=0.001) but had little effect on root dry weight (P=0.43). Addition of As had no significant effect on shoot dry weight (P=0.43) but significantly increased root dry weight (P=0.01). Silicon concentrations in shoots and roots increased proportionally to increasing amounts of externally supplied Si (P < 0.001). The presence of As in the nutrient solution had little effect on shoot Si concentration (P=0.16) but significantly decreased root Si concentration (P=0.005). Increasing external Si concentration significantly decreased shoot and root As concentrations and total As uptake by rice seedlings (P <0.001). In addition, Si significantly decreased shoot P concentration and shoot P uptake (P <0.001). The data clearly demonstrate a beneficial effect of Si on the growth of rice seedlings. Addition of Si to the growth medium also inhibited the uptake of arsenate and phosphate by the rice seedlings.     

Influence of external zinc and phosphorus supply on Cd uptake by rice (Oryza sativa L.) seedlings with root surface iron plaque

Rice seedlings were grown in hydroponic culture to determine the effects of external Zn and P supply on plant uptake of Cd in the presence or absence of iron plaque on the root surfaces. Iron plaque was induced by supplying 50 mg l⁻¹ Fe²⁺ in the nutrient solution for 2 day. Then 43-day-old seedlings were exposed to 10 μmol l⁻¹ Cd together with 10 μmol l⁻¹ Zn or without Zn (Zn–Cd experiment), or to 10 μmol l⁻¹ Cd with 1.0 mmol l⁻¹ P or without P (P–Cd experiment) for another 2 day. The seedlings were then harvested and the concentrations of Fe, Zn, P and Cd in dithionite–citrate–bicarbonate (DCB) extracts and in roots and shoots were determined. The dry weights of roots and shoots of seedlings treated with 50 mg l⁻¹ Fe were significantly lower than when no Fe was supplied. Adsorption of Cd, Zn and P on the iron plaque increased when Fe was supplied but Cd concentrations in DCB extracts were unaffected by external Zn or P supply levels. Cd concentrations in shoots and roots were lower when Fe was supplied. Zn additions decreased Cd concentrations in roots but increased Cd concentrations in shoots, whereas P additions significantly increased shoot and root Cd concentrations and this effect diminished when Fe was supplied. The percentage of Cd in DCB extracts was significantly lower than in roots or shoots, accounting for up to 1.8–3.8% of the plant total Cd, while root and shoot Cd were within the ranges 57–76% and 21–40% respectively in the two experiments. Thus, the main barrier to Cd uptake seemed to be the root tissue and the contribution of iron plaque on root surfaces to plant Cd uptake was minor. The changes in plant Cd uptake were not due to Zn or P additions altering Cd adsorption on iron plaque, but more likely because Zn or P interfered with Cd uptake by the roots and translocation to the shoots.     

Silicon-mediated amelioration of zinc toxicity in rice (Oryza sativa L.) seedlings

Background and aims

Silicon (Si) was suggested to enhance plant resistance to toxic elements, and its beneficial role was mainly based on external and internal plant mechanisms. This work aimed at investigating the internal effect of Si on zinc (Zn) detoxification to rice (Oryza sativa L., cv. Tian You 116) seedlings.

Methods

In a hydroponic experiment, we examined the uptake, xylem loading and localization of Zn in rice seedlings under the condition of 200?μM Zn contamination with the additional silicate supply at three levels ( 0, 0.5 and 1.8?mM).

Results

The silicate addition significantly increased the seedling biomass, and decreased Zn concentration in both root and shoot of seedlings and in xylem sap flow. Zinpyr-1 fluorescence test and Energy-dispersive X-ray spectroscopy analysis showed the concentration of biologically active Zn²⁺ decreased, and Zn and Si co-localized in the cell wall of metabolically less active tissues, especially in sclerenchyma of root. The fractionation analysis further supported silicate supply increased about 10% the cell wall bound fraction of Zn.

Conclusions

This study suggests the Si-assisted Zn tolerance of rice is mainly due to the reduction of uptake and translocation of excess Zn, and a stronger binding of Zn in the cell wall of less bioactive tissues might also contribute to some degree.     

A new precipitation technique provides evidence for the permeability of Casparian bands to ions in young roots of corn (Zea mays L.) and rice (Oryza sativa L.)

Using an insoluble inorganic salt precipitation technique, the permeability of cell walls and especially of endodermal Casparian bands (CBs) for ions was tested in young roots of corn (Zea mays) and rice (Oryza sativa). The test was based on suction of either 100 µm CuSO₄ or 200 µm K₄[Fe(CN)₆] into the root from its medium using a pump (excised roots) or transpirational stream (intact seedlings), and subsequent perfusion of xylem of those root segments with the opposite salt component, which resulted in precipitation of insoluble brown crystals of copper ferrocyanide. Under suction, Cu²⁺ could cross the endodermis apoplastically in both plant species (although at low rates) developing brown salt precipitates in cell walls of early metaxylem and in the region between CBs and functioning metaxylem vessels. Hence, at least Cu²⁺ did cross the endodermis dragged along with the water. The results suggested that CBs were not perfect barriers to apoplastic ion fluxes. In contrast, ferrocyanide ions failed to cross the mature endodermis of both corn and rice at detectable amounts. The concentration limit of apoplastic copper was 0.8 µm at a perfusion with 200 µm K₄[Fe(CN)₆]. Asymmetric development of precipitates suggested that the cation, Cu²⁺, moved faster than the anion, [Fe(CN)₆]^4–, through cell walls including CBs. Using Chara cell wall preparations (‘ghosts’) as a model system, it was observed that, different from Cu²⁺, ferrocyanide ions remained inside wall-tubes suggesting a substantially lower permeability of the latter which agreed with the finding of an asymmetric development of precipitates. In both corn and rice roots, there was a significant apoplastic flux of ions in regions where laterals penetrated the endodermis. Overall, the results show that the permeability of CBs to ions is not zero. CBs do not represent a perfect barrier for ions, as is usually thought. The permeability of CBs may vary depending on growth conditions which are known to affect the intensity of formation of bands.     

Salinity-induced enhancement of L-myo-inositol 1-phosphate synthase in rice (Oryza sativa L.)

The salt-tolerant varieties of rice (Oryza sativa L.) exhibit enhanced activity of the chloroplast form of L-myo-inositol 1-phosphate synthase (EC 5.5.4.1) under NaCl treatment either during the seedling stage or in fully grown plants during field growth. The salt-induced enhancement was noticeable only in chloroplasts from light-grown plants. The effects of these treatments on the cytosolic inositol synthase activity were less pronounced. While the effect of salt on the activity of the two forms was marginal in the salt-sensitive varieties during seedling growth, salinity affected the chloroplast inositol synthase activity adversely in these varieties during growth of the plants under field conditions. The salt-enhanced activities of inositol synthase(s) in the highly salt-tolerant varieties studied were found to be comparable to that observed in Porteresia coarctata, a halophytic wild rice species. The implications of these findings, which suggest a role of the inositol pathway in osmoregulation, are discussed.     

The involvement of the transpirational bypass flow in sodium uptake by high- and low-sodium-transporting lines of rice developed through intravarietal selection

We report the characterization of high- and low-sodium-transporting lines developed by intravarietal selection within a cultivar, IR36, of rice (Oryza sativa L.). The purpose was to investigate the mechanistic basis of sodium uptake in material in which differences in salt uptake could be isolated from the many other morphological and physiological characteristics that affect the phenotypic expression of salt tolerance. The lines differed in mean sodium transport by a factor of 2. They differed in vigour and water use efficiency, which are characters that modify the effects of salt transport, by only 12% or 13%. The lines did not differ significantly in other physiological traits that are components of salt resistance: compartmentalization at the leaf and cellular levels. There was a strong correlation between the transport of sodium and a tracer for apoplastic pathways (trisodium, 3-hydroxy-5,8,10-pyrene trisulphonic acid, PTS) in both lines. The regression coefficient for sodium transport on PTS transport was the same in both lines. The individual variation in PTS transport was similar to that in sodium transport, and the variation in the transport of both was very much greater than the variation in any other character studied. The high-sodium-transporting line took up proportionately more PTS than the low-sodium-transporting line. It is concluded that the transpirational bypass flow is of major importance in sodium uptake by rice and that selection for differences in sodium transport has been brought about by selection for heritable differences in the bypass flow.     

Studies on sodium bypass flow in lateral rootless mutants lrt1 and lrt2, and crown rootless mutant crl1 of rice (Oryza sativa L.)

An apoplastic pathway, the so‐called bypass flow, is important for Na⁺ uptake in rice (Oryza sativa L.) under saline conditions; however, the precise site of entry is not yet known. We report the results of our test of the hypothesis that bypass flow of Na⁺ in rice occurs at the site where lateral roots emerge from the main roots. We investigated Na⁺ uptake and bypass flow in lateral rootless mutants (lrt1, lrt2), a crown rootless mutant (crl1), their wild types (Oochikara, Nipponbare and Taichung 65, respectively) and in seedlings of rice cv. IR36. The results showed that shoot Na⁺ concentration in lrt1, lrt2 and crl1 was lower (by 20–23%) than that of their wild types. In contrast, the bypass flow quantified using trisodium‐8‐hydroxy‐1,3,6‐pyrenetrisulphonic acid (PTS) was significantly increased in the mutants, from an average of 1.1% in the wild types to 3.2% in the mutants. Similarly, bypass flow in shoots of IR36 where the number of lateral and crown roots had been reduced through physical and hormonal manipulations was dramatically increased (from 5.6 to 12.5%) as compared to the controls. The results suggest that the path of bypass flow in rice is not at the sites of lateral root emergence.     

水稻砷污染及其对砷的吸收和代谢机制

水稻是当今世界大部分地区(尤其是东南亚)的主要的粮食作物之一,同时也是砷(As)进入食物链的主要途径之一。日益严重的水稻田As污染,不但影响了稻米的产量和品质,而且通过食物链威胁着人体健康。如何减少水稻地上部(尤其是米粒)As的含量和降低其毒性,及提高水稻As耐性是亟需解决的世界食品安全问题。深入了解水稻对As的吸收、积累和代谢的生理及分子生物学机制是解决水稻As污染的关键途径。综述国内外研究,对今后深入研究提出建议。     

Metabolic responses of etiolated rice (Oryza sativa L.) seedlings to streptomycin

This paper reports experiments performed to investigate the influence of various concentrations of streptomycin sulphate on a few parameters of importance in the metabolism of rice (Oryza sativa L.) seedlings. It was shown that respiration rate was accelerated by streptomycin. The specific activities of catalase and peroxidase decreased whereas IAA oxidase increased with increasing streptomycin concentrations. Increased activities of all these enzymes were apparent on a dry weight basis suggesting increased succulence caused by streptomycin treatment. There was a considerable rise in the water soluble protein content following streptomycin application. It may be suggested that growth inhibition by streptomycin results from reduction in the auxin level owing to enhanced auxin destruction.     

Zinc ameliorates copper-induced oxidative stress in developing rice (Oryza sativa L.) seedlings

Rice (Oryza sativa L.) seedlings were treated with different concentrations of copper (Cu) either in presence or absence of zinc (Zn), and different events were investigated to evaluate the ameliorative effect of Zn on Cu stress. In presence of high Cu concentration, growth of both root and shoots were considerably reduced. Decline in elongation and fresh mass was observed in root and shoot. Zn alone did not show any considerable difference as compared to control, but when supplemented along with high concentration Cu, it prompted the growth of both root and shoot. After 7 days, root growth was 9.36 and 9.59 cm, respectively, at 200 and 500 μM of Cu alone as compared to 10.59 and 12.26 cm at similar Cu concentrations, respectively, in presence of Zn. Cu accumulation was considerably high after 7 days of treatment. In absence of Zn, significant accumulation of Cu was observed. Zn supplementation ameliorated the toxic impact of Cu and minimized its accumulation. Cu treatment for 1 and 7 days resulted in a dose-dependent increase in hydrogen peroxide (H₂O₂). When Cu was added in presence of Zn, the H₂O₂ production in root and shoot was reduced significantly. The increase in H₂O₂ production under Cu stress was accompanied by augmentation of lipid peroxidation. In absence of Zn, Cu alone enhanced the malondialdehyde (MDA) production in both root and shoot after 1 and 7 days of treatment. The MDA content drastically reduced in root and shoot as when Zn was added during Cu treatment. The activities of antioxidant enzymes like superoxide dismutase (SOD), catalase (CAT), and guaiacol peroxidase (GPX) were elevated under Cu stress both in root and shoot. Addition of Zn further stimulated the activities of these enzymes. Both ascorbate (AsA) and glutathione (GSH) contents were high under Cu stress either in presence or absence of Zn. The results suggests that Zn supplementation improves plant survival capacity under high Cu stress by modulating oxidative stress through stimulation of antioxidant mechanisms and restricts the accumulation of toxic concentrations of Cu.     

Stagnant deoxygenated growth enhances root suberization and lignifications, but differentially affects water and NaCl permeabilities in rice (Oryza sativa L.) roots

It has been shown that rice roots grown in a stagnant medium develop a tight barrier to radial oxygen loss (ROL), whereas aerated roots do not. This study investigated whether the induction of a barrier to ROL affects water and solute permeabilities. Growth in stagnant medium markedly reduced the root growth rate relative to aerated conditions. Histochemical studies revealed an early deposition of Casparian bands (CBs) and suberin lamellae (SL) in both the endodermis (EN) and exodermis, and accelerated lignification of stagnant roots. The absolute amounts of suberin, lignin and esterified aromatics (coumaric and ferulic acid) in these barriers were significantly higher in stagnant roots. However, correlative permeability studies revealed that early deposition of barriers in stagnant roots failed to reduce hydraulic conductivity (Lp(r) ) below those of aerated roots. In contrast to Lp(r) , the NaCl permeability (P(sr) ) of stagnant roots was markedly lower than that of aerated roots, as indicated by an increased reflection coefficient (σ(sr) ). In stagnant roots, P(sr) decreased by 60%, while σ(sr) increased by 55%. The stagnant medium differentially affected the Lp(r) and P(sr) of roots, which can be explained in terms of the physical properties of the molecules used and the size of the pores in the apoplast.