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1.
The effects of copper (CuCl2) on active and passive Rb+(86Rb+) influx in roots of winter wheat grown in water culture for 1 week were studied. External copper concentrations in the range of 10–500 μ M in the uptake nutrient solution reduced active Rb+ influx by 20–70%, while passive influx was unaffected (ca 10% of the Rb+ influx in the Cu-free solution). At external Rb+ concentrations of up to 1 m M , Cu exposure (50 μ M decreased Vmax to less than half and increased Km to twice the value of the control. Short Cu exposure reduced the K+ concentration in roots of low K+ status. Pretreatment for 5 min in 50 μ M CuCl2 prior to uptake experiments reduced Rb+ influx by 26%. After 60 min pretreatment with Cu, the corresponding reduction was 63%. Cu in the cultivation solution impeded growth, especially of the roots. The Cu concentration in the roots increased linearly with external Cu concentration (0–100 μ M ) while Cu concentration in the shoots was relatively unchanged. The K+ concentration in both roots and shoots decreased significantly with increased Cu in the cultivation solutions. Possible effects of Cu on membranes and ion transport mechanisms are discussed.  相似文献   

2.
Influx of Rb+(86Rb+) and Ca2+ (45Ca2+) in roots of intact winter wheat (Triticum aestivum L. cv. Weibulls Starke II) was determined at intervals before, during and after exposure to cold acclimation conditions (2°C and 8 h light period). The plants were grown in nutrient medium of two ionic strengths. During the initial two weeks of growth at 16°C and 16 h light period, Rb+ influx into roots decreased with increasing age, probably as a consequence of a decreasing proportion of metabolically active roots. The presence of 10?4M 2,4-dinitrophenol (DNP) reduced Rb+ influx to a low and constant level, indicating that metabolic influx was the dominant process. In contrast, Ca2+ influx in plants grown in full strength nutrient solution was higher in the presence than in the absence of DNP. This effect may have been due to an active extrusion mechanism mediating re-export of absorbed Ca2+(45Ca2+) during the uptake experiment. With the metabolic uncoupler inhibiting such extrusion the Ca2+(45Ca2+) influx mesured would increase. During cold treatment, Rb+ influx remained at a low level, and was further decreased when DNP was present in the uptake solution. This effect may have been due to inhibition of residual active influx of Rb+ at 2°C by the uncoupler and/or to a decrease in membrane permeability. In contrast to Rb+, Ca2+ influx increased during cold treatment, which could again be explained as inhibition of re-export. The presence of DNP reduced Ca2+ influx at 2°C, indicating decreased membrane permeability by DNP at low temperature. After transfer of plants from cold acclimation conditions to 16°C, Rb+ and Ca2+ influx increased in plants grown at both ionic strengths. Influx levels were independent of the length of the cold acclimation period (1, 6 and 8 weeks), but the patterns were different for the two ions. After each of the cold acclimation periods, Rb+ influx increased during the first week and decreased or remained at the same level during the second week, while Ca2+ influx always decreased during the second week of post-cold treatment.  相似文献   

3.
The effects of cadmium and lead on the internal concentrations of Ca2+ and K+, as well as on the uptake and translocation of K(86Rb+) were studied in winter wheat (Triticum aestivum L. a. MV-8) grown hydroponically at 2 levels of K+ (100 uM and 10 mM). Cd2+ and Pb2+ were applied in the nutrient solution in the range of 0.3 to 1000 u.M. Growth was more severely inhibited by Cd2+ and in the high-K+ plants as compared to Pbz+ and low-K+ plants. Ions of both heavy metals accumulated in the roots and shoots, but the K+ status influenced their levels. Ca2+ accumulation was increased by low concentrations of Cd2+ mainly in low-K+ shoots, whereas it was less influenced by Pb2+. The distribution of Cd2+ and Ca2+ in the plant and in the growth media indicated high selectivity for Cd2+ in the root uptake, while Ca2+ was preferred in the radial and/or xylem transport. Cd2+ strongly inhibited net K+ accumulation in high-K+ plants but caused stimulation at low K+ supply. In contrast, the metabolis-dependent influx of K+(86Rb+) was inhibited in low-K+ plants, while the passive influx in high-K+ plants was stimulated. Translocation of K+ from the roots to the shoots was inhibited by Cd2+ but less influenced in Pb2+-treated plants. It is concluded that the effects of heavy metals depend upon the K+-status of the plants.  相似文献   

4.
Ice crystal formation temperature was determined in the region of the crown in one group of 7-day-old intact unhardened high-salt plants of winter wheat (Triticum aestivum L. cv. Weibulls Starke II) with TA (Thermal Analysis) and DTA (Differential Thermal Analysis) methods. After exposure of another group of plants, grown for the first 7 days in the same way as the first group, to various sub-zero temperatures (-1 to 5°C), influx in roots of Rb+(86Rb+) and Ca2+(45Ca2+) and contents of K+ and Ca2+ were determined at intervals during 7 days of recovery. Ice crystal formation in the crown tissue was probably extracellular and took place at about -4°C. There was a large loss of K+ from the roots after treatment at sub-zero temperatures. This loss increased as the temperature of the sub-zero treatment decreased. During recovery, roots of plants exposed to -1, -2 and -3°C gradually reabsorbed K+. Reabsorption of K+ in roots of plants exposed to -4°C was greatly impaired. Rb+ influx decreased and Ca2+ influx increased after sub-zero temperature treatments of the plants. Active Rb+ influx mechanisms and active extrusion of Ca2+ were impaired or irreversibly damaged by the exposure. While Rb+ influx mechanisms were apparently repaired during recovery in plants exposed to temperatures down to -3°C, Ca2+ extrusion mechanisms were not. The temperature for ice crystal formation in the region of the crown tissue coincides with the temperature at which the plants lost the ability to reabsorb K+ and to repair Rb+ influx mechanisms during the recovery period. Plants were lethally damaged at temperatures below ?4°C.  相似文献   

5.
Betula papyrifera Marsh, seedlings adapted very poorly to flooding for up to 60 days. Responses to flooding included increased ethylene production; stomatal closure; leaf senescence; drastic inhibition of shoot growth, cambial growth, and root growth; decay of roots, and death of many seedlings. Flooding inhibited growth of leaves that formed prior to flooding, inhibited formation of new leaves, and induced abscission of old leaves. As a result of extensive leaf abscission, fewer leaves were present after flooding than before flooding was initiated. The drastic reduction in leaf area was associated with greatly decreased growth of the lower stem and roots. No evidence was found of adaptive morphological changes to flooding. The data indicate that intolerance of B. papyrifera seedlings to flooding is an important barrier to regeneration of the species on sites subject to periodic inundation.  相似文献   

6.
When 1 m M spermidine or spermine was included in an absorption solution which contained 20 m M Na+ and 1 m M Rb+, Na+ influx into excised maize roots ( Zea mays L. cv. Golden Cross Bantam) was reduced. Rb+ influx was reduced in the presence of spermidine and uneffected in the presence of spermine when compared with control solutions. When 1 m M Ca2+ replaced the polyamines, Na+ influx was strongly reduced and Rb+ influx was promoted. Rb+ influx from 1 m M Rb+ solutions which did not contain Na+ was also promoted by 1 m M Ca2+, but was inhibited by 1 m M spermidine. This Ca2+ promotion of Rb+ influx could be reversed by 10 times greater concentration of spermidine in the absorption solution. H+ efflux from excised roots was inhibited by spermidine when compared with Ca2+ or control solutions, however, the plasma membrane ATPase was not inhibited by spermidine. It is concluded that external Ca2+ plays two separate roles in membrane function, only one of which can be substituted for by polyamines. The first role, maintenance of membrane integrity, can be substituted for by spermidine or spermine. The second function, maintenance of the Rb+ transport mechanism, is Ca2+ specific and cannot be substituted for by spermidine or spermine. The results of this study are discussed in terms of electrostatic interactions between the plasma membrane and the Ca2+ or polyamines.  相似文献   

7.
Purified plasmalemma vesicles were isolated in the presence of 250 m M sucrose from roots of 14-day-old seedlings of winter wheat ( Triticum aestivum L. Martonvásári-8) by phase partitioning of salt-washed microsomal fractions in a Dextran-polyethylene glycol two-phase system, and both Mg2+- and Ca2+-ATPase activities were detected. Orthovanadate-sensitive Mg2+-ATPase activity associated with the inside of right side-out plasmalemma (PM) vesicles (latency 98%) was inhibited 76% by 0.3 m M Ca2+, Ca2+-dependent ATPase activity located partly on the inside and partly on the outside of plasmalemma vesicles (latency 47%) was not affected by Mg2+.
Mg2+-ATPase activity was inhibited by 68% and inhibition of Mg2+ activation by 0.3 m M Ca2+ partly disappeared in the presence of 10 p M tentoxin, a fungal phytotoxin. Mg2+-ATPase activity remained inhibited up to 10 n M tentoxin while at 1 μ M tentoxin Mg2+ activation was as high as without tentoxin. K+-stimulation and vanadate inhibition was increased and decreased, respectively, by 100 p M -10 n M tentoxin. Ca2+-dependent ATPase activity was continuously increased by 1 p M -10 n M tentoxin, but at 1 μ M tentoxin the stimulation disappeared. The effects of p M tentoxin on plasma-lemma Mg2+-ATPase are discussed in relation to its influence on K+ transport in wheat seedlings.  相似文献   

8.
Seedlings of spring wheat ( Triticum aestivum L. cv. Svenno) were cultivated at 20°C in continuous light or darkness with the roots in nutrient solutions for six days. The plants were starved for K+ during different periods of time to produce plants with various K+ status. In one cultivation light-grown plants were pretreated in darkness, and vice versa, before the uptake experiment. In all experiments, roots were put in a complete nutrient medium containing 2.0 m M K+ radiolabelled with 86Rb. The uptake time was varied (5, 60 or 120 min).
The K+ concentration in the roots, [K+]root, increased during the course of the uptake experiments, especially in light and at initially low [K+]root, At the same time K+ (86Rb) influx in the roots decreased. The simoidal relationship obtained between K+ (86Rb) influx and [K+]root was affected by these changes, and Hill plots gave various Hill coefficients, nH, depending on the duration of the uptake experiments. nH from three apparently straight line segments of the same plot, in different [K+]root - intervals, indicated a falling degree of interaction between the binding sites as [K+]root increased. For the dark-grown plants negative cooperativity could not be demonstrated.  相似文献   

9.
Root development was studied in winter wheat ( Triticum aestivum L. cv Starke II) grown at 5,10, 15 and 20°C in nutrient solutions with phosphate concentrations of 10, 100 or 1000 μM . The plants were grown for 38 days (5 and 10°C), 19 days (15°C) or 14 days (20°C). At the end of the cultivation period the phosphate influx in the roots was determined with 32P-phosphate. Root development (lateral and seminal roof length and number) was monitored throughout the cultivation period on the same individuals by repeated (approximately every second day) photocopying of the roots for measurements with digitizer and appropriate software. The 5°C treatment yielded no laterals, and the seminals were only slightly affected by the different phosphate treatments. The 10 μM phosphate treatment gave high root:shoot dry weight ratio, high average lateral root length and high specific root length [m root (g root fresh weight)-1]. The 1000 μM phosphate treatment yielded the highest number of laterals per m seminal root, and usually also the highest absolute numbers. Phosphate influx decreased with increased P status of the roots. It is argued that phosphate influx is dependent on factors such as P status, root geometry and relative root extension rate.  相似文献   

10.
Effects of interrupted K+ supply on different parameters of growth and mineral cation nutrition were evaluated for spring wheat (Triticum aestivum L. cv. Svenno). K+ (2.0 mM) was supplied to the plants during different periods in an otherwise complete nutrient solution. Shoot growth was reduced before root growth after interruption in K+ supply. Root structure was greatly affected by the length of the period in K+ -free nutrient solution. Root length was minimal, and root branching was maximal within a narrow range of K+ status of the roots. This range corresponded to cultivation for the last 1 to 3 days, of 11 in total, in K+ -free nutrient solution, or to continuous cultivation in solution containing 0.5 to 2 mM K+. In comparison, both higher and lower internal/external K+ concentrations had inhibitory effects on root branching. However, the differing root morphology probably had no significant influence on the magnitude of Ca2+, Mg2+ and Na+ uptake. Uptake of Ca2+ and especially Mg2+ significantly increased after K+ interruption, while Na+ uptake was constant in the roots and slowly increased in the shoots. The two divalent cations could replace K+ in the cells and maintain electroneutrality down to a certain minimal range of K+ concentrations. This range was significantly higher in the shoot [110 to 140 μmol (g fresh weight)?1] than in the root [20 to 30 μmol (g fresh weight)?1]. It is suggested that the critical K+ values are a measure of the minimal amount of K+ that must be present for physiological activity in the cells. At the critical levels, K+ (86Rb) influx and Ca2+ and Mg2+ concentrations were maximal. Below the critical K+ values, growth was reduced, and Ca2+ and Mg2+ could no longer substitute for K+ for electrostatic balance. In a short-term experiment, the ability of Ca2+ to compete with K+ in maintaining electroneutrality in the cells was studied in wheat seedlings with different K+ status. The results indicate that K+, which was taken up actively and fastest at the external K+ concentration used (2.0 mM), partly determines the size of Ca2+ influx.  相似文献   

11.
Microsomal fractions from wheat tissues exhibit a higher level of ATP hydrolytic activity in the presence of Ca2+ than Mg2+. Here we characterise the Ca2+-dependent activity from roots of Triticum aestivum lev. Troy) and investigate its possible function. Ca2+-dependent ATP hydrolysis in the microsomal fraction occurs over a wide pH range with two slight optima at pH 5.5 and 7.5. At these pHs the activity co-migrates with the major peak of nitrate-inhibited Mg2+. Cl-ATPase on continuous sucrose gradients indicating that it is associated with the vacuolar membrane. Ca2+-dependent ATP hydrolysis can be distinguished from an inhibitory effect of Ca2+ on the plasma membrane K+, Mg2+-ATPase following microsomal membrane separation using aqueous polymer two phase partitioning. The Ca2+-dependent activity is stimulated by free Ca2+ with a Km of 8.1 μM in the absence of Mg2+ ([CaATP] = 0.8 mM). Vacuoiar membrane vacuolar preparations contain a higher Ca2+-dependent than Mg2+-dependent ATP hydrolysis, although the two activities are not directly additive. The nucleotide specificity of the divalent ion-dependent activities in vacuolar membrane-enriched fractions was low. hydrolysis of CTP and UTP being greater than ATP hydrolysis with both Ca2+ and Mg2+ The Ca2+-dependent activity did discriminate against dinucleotides, and mononucleotides. and failed to hydrolyse phosphatase substrates. Despite low nucleotide specificity the Mg2+-dependent activity functioned as a bafilomycin sensitive H+-pump in vacuolar membrane vesicles. Ca2+-dependent ATP hydrolysis was not inhibited by the V-, P-, or F-type ATPase inhibitors bafilomycin. vanadate and azide, respectively. nor by the phosphatase inhibitor molybdate, but was inhibited 20% at pH 7.5 by K+. Possible functions of Ca2+-dependent hydrolysis as a H+-pump or a Ca2+-pump was investigated using vacuolar membrane vesicles. No H+ or Ca2+ translocating activity was observed under conditions when the Ca2+-dependent ATP hydrolysis was active.  相似文献   

12.
Uptake of Rb+ from a complete nutrient solution with 2.0 mM Rb+ was studied in roots of spring wheat seedlings ( Triticum aestivum L. cv. Svenno) with different K+ levels. The relationship between Rb+ uptake and concentration of K+ in the roots indicated a negative feedback mechanism operating through allosteric control. The Rb+ uptake process in root cells was divided into two steps: (1) binding of the ion in the free space, and (ii) transmembrane transport into the cytoplasm. Metabolic and non-metabolic components of uptake were separated by addition of the metabolic inhibitor 2,4-dinitrophenol (DNP) to the nutrient solution. It is suggested that metabolic Rb+ uptake requires energy in two uptake steps (for binding to the carrier entity in the free space and for transmembrane transport) or in one step only (for transmembrane transport), dependent on the K+ status of the roots. The change from metabolic to non-metabolic binding in the free space is accomplished by changing the conformational state of the carrier (slow/fast transitions). There may be a hysteretic effect on metabolic Rb+ uptake through a slow transition between carrier states. This is superimposed on the negative cooperativity, strengthening further cooperativity at intermediate K+ levels in the roots. Non-metabolic Rb+ uptake probably consists of two components, a carrier-mediated (facilitated diffusion) and a parallel diffusive component.  相似文献   

13.
Plasma membrane preparations of high purity were obtained from roots of dark-grown wheat (Triticum aestivum L. cv. Drabant) by aqueous polymer two-phase partitioning. These preparations mainly contained sealed, right-side-out vesicles (ca 90% exposing the original outside out). By subjecting the preparations to 4 freeze/thaw cycles the proportion of sealed, inside-out (cytoplasmic side out) vesicles increased to ca 30%. Inside-out and right-side-out plasma membrane vesicles were then separated by partitioning the freeze/thawed plasma membranes in another aqueous polymer two-phase system. In this way, highly purified, sealed, inside-out (>60% inside-out) vesicles were isolated and subsequently used for characterization of the Ca2+ transport system in the wheat plasma membrane. The capacity for 45Ca2+ accumulation, nonlatent ATPase activity and proton pumping (the latter two markers for inside-out plasma membrane vesicles) were all enriched in the inside-out vesicle fraction as compared to the right-side-out fraction. This confirms that the ATP-binding site of the 45Ca2+ transport system, similar to the H+-ATPase, is located on the inner cytoplasmic surface of the plant plasma membrane. The 45Ca2+ uptake was MgATP-dependent with an apparent Km for ATP of 0.1 mM and a high affinity for Ca2+ [Km(Ca2+/EGTA) = 3 μM]. The pH optimum was at 7.4–7.8. ATP was the preferred nucleotide substrate with ITP and GTP giving activities of 30–40% of the 45Ca2+ uptake seen with ATP. The 45Ca2+ uptake was stimulated by monovalent cations; K? and Na+ being equally efficient. Vanadate inhibited the 45Ca2+ accumulation with half-maximal inhibitions at 72, 57 and 2 μM for basal, total (with KCI) and net K+-stimulated uptake, respectively. The system was also highly sensitive to erythrosin B with half-maximal inhibition at 25 nM and total inhibition at 1μM. Our results demonstrate the presence of a primary Ca2+ transport ATPase in the plasma membrane of wheat roots. The enzyme is likely to be involved in mediating active efflux (ATP-binding sites on the cytoplasmic side) to the plant cell exterior to maintain resting levels of cytoplasmic free Ca2+ within the cell.  相似文献   

14.
The regulatory role of abscisic acid (ABA) and kinetin on influx of K+(86RB+) IN tools of 7day old intact winter wheat which plant (Fritieun aestivum I ass starke 1 and 11) Was studied the inhibitory effect of 40,80 μM ABA in the uptake solution on K+(86RB+)influx was transiently stipulated pretreatment of the plants with ABA kinetin content enacted inhibitors effect caused by ABA. At low water potential in the uptake solution (05MPa)K+(86RB+) influx was slights higher in the presence of ABA than in is absence High humidity 123kpa ca 100% relative humidity (RID)around the shoots counteracted the inhibitory effect on k+(86RB+) influx caused by A,B,A IN the uptake solution the present data contain the hypothesis that when plants are subjected to conditions such as low water potential and low temperature. ABA stimulates K influx to facilitate water uptake.  相似文献   

15.
The red light (R)-induced swelling of mesophyll protoplasts, isolated from dark-grown wheat ( Triticum aestivum L. cv. Arminda) leaves, was inhibited by guanosine-5'-0-(2-thiodiphosphate) (GDP-β-S). In darkness or after control irradiation with far-red light (FR), guanosine-5'-O-(3-thiotriphosphate) (GTP-γ-S) induced swelling to the same extent as after R. Both GDP-β-S and GTP-γ-S were introduced into the cytoplasm by means of electroporation. The possibility of R-induced activation of the phosphatidylinositol pathway of transmembrane signalling was investigated. Neomycin, Li+ and l-(5-isoquinolinesulfonyl)-2-methylpiperazine (H7) inhibited the R-induced swelling. Phorbol 12-myristate 13-acetate (PMA) induced swelling after control irradiation with FR. Neomycin and Li+ also inhibited GTP-γ-S-induced swelling. These results suggest that a GTP-binding protein is involved in the phytochrome-regulated swelling response. Addition of N6, 2'-0-dibutyryladenosine 3':5'-cyclic monophosphate (DB-cAMP) induced swelling to the same extent as R-irradiation. The calmodulin antagonist N-(6-aminohexyl)5-chloro-l-naphthalenesulfonamide (W7) induced swelling after FR, while R-induced swelling was not affected. The less active analogue N-(6-aminohexyl)-l-naphthalenesulfonamide (W5) induced no swelling after FR. It is speculated that the protoplast volume is correlated with the cytoplasmic concentration of free Ca2+.  相似文献   

16.
17.
The development of SO42- influx in roots and sulfur transport to shoots was followed in 35S-tracer experiments for sulfur-deficient spring wheat (Triticum aestivum L. cv. Svenno) seedlings pretreated for various time periods (0–24 h) in nutrient solutions with SO42-. Effects of the metabolic inhibitor 2,4-dinitrophenol (DNP) and the protein synthesis inhibitor cycloheximide (CH) on SO42- influx were also evaluated. The SO42- influx appears feedback-regulated by the internal sulfur level of the roots. Regulation may be achieved solely by a rapidly changed SO42- carrier activity through an allosteric effect by the intracellular SO42- concentration of the roots, followed first by induction of carrier synthesis and then by repression of carrier synthesis after transfer of the roots from SO42--deficient nutrient solutions to solutions with SO42-. A Hill plot of the partly sigmoidal relationship between SO42- influx and intracellular sulfur concentration in the roots gave a Hill coefficient of -4.2, indicating negative cooperativity between a minimum number of four interacting allosteric binding sites for sulfur on each carrier entity. DNP-experiments showed that SO42- influx was mainly metabolic, especially after short pretreatment in SO42- at an external SO42- concentration of 0.1 mM. Pretreatment with CH rapidly prevented new SO42- carriers from being formed. Long CH pretreatment (24 h) and different SO42- pretreatments reduced SO42- influx below the non-metabolic level obtained by uptake experiments with DNP, indicating the existence of SO42- carriers mediating passive SO42- transport across the plasmalemma of the root cells. SO42- influx was further decreased for the CH pretreated (24 h) plants by the presence of both CH and DNP in the experimental nutrient solution. This probably indicates the diffusive part of the non-metabolic SO42- influx in the present experiments. Finally, it is suggested that there is a feedback signal between root and shoot, regulating sulfur transport upwards.  相似文献   

18.
19.
The influx of Rb+ into the roots of two barley varieties (Hordeum vulgare L. cv. Salve and cv. Ingrid) from a K+-free 86Rb-labelled nutrient solution with 2.0 mM Rb+, was checked at intervals from day 6 to day 18. The control plants were continuously grown in complete nutrient solution containing 5.0 mM K+, while two other groups of plants were grown in K+-free nutrient solution starting on day 6 and between day 6 and day 9, respectively. The pattern of Rb+ influx was similar for both varieties, although their efficiencies in absorbing Rb+ were different. The relationship between Rb+ influx and K+ concentration of the root could be interpreted in terms of negative feedback through allosteric control of uptake across the plasmalemma of the root cells. Hill plots were bimodal, but in the opposite direction. The Hill coefficients, reflecting the minimum number of interacting allosteric binding sites for K+ (Rb+), were low (≤–3.0). It is discussed whether the threshold value, that is the breaking point in the Hill plot, is indicative of a changed efficiency of transporting units for K+ (Rb+) transport to the xylem. Moreover, feedback regulation might be involved in transport of K+ between root and shoot. The variation in K+ concentrations in the roots and shoots of control plants were cyclic but in phase opposition despite an exponential growth. The average K+ concentration varied only slightly with age.  相似文献   

20.
Six cultivars of spring barley ( Hordeum vulgare L. cvs Salve, Nümberg II, Bomi, Risø 1508, Mona and Sv 73 608) were grown in water culture for three weeks with various combinations of mineral supply and differential roots/shoot temperatures during the growth period. Most important for growth and accumulation of N, K+, Ca2+ and Mg2+ was the mineral supply, followed by the root temperature and the choice of cultivar. Treatments with low mineral supply or low root temperature induced a uniform reduction in growth and accumulation of the ions studied. The effects of low mineral supply and low root temperature on growth and N accumulation was additive, which indicates that these factors exert their influence independently of each other.
Roots grown at 10°C were smaller and Rb+(86Rb) influx was higher than in roots grown at 20°C. It is suggested that the control of Rb+(86Rb) influx is affected by the root temperature and the age of the plants. The higher 86Rb+ (86Rb) influx into the low temperature roots could not compensate for the smaller root size. However, the lower total mineral accumulation made up for the needs of the smaller plants and cannot explain the reduction in growth.  相似文献   

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