Leaf expansion in sunflower as influenced by salinity and short-term changes in carbon fixation

Abstract With a view to defining factors regulating the growth responses of sunflower to salinity, plants were grown in solution culture (0, 50 or 100 mol m⁻³ NaCl) and under natural light, and the areas of every leaf measured once or twice daily from 22 until 38 d after germination. During this period, carbon availability for growth was manipulated by changing light levels and by the use of a photosynthesis inhibitor, DCMU. Salinity reduced relative leaf expansion rates per plant (RLER) by an average of 0.04 (50 mol m⁻³) and 0.08 (100 mol m⁻³) m² m⁻² d⁻¹ compared with control plants of equivalent leaf area: the effects were found in expanding leaves regardless of age or size. Control plants expanded faster during the day than the night, but plants grown in salt had an almost constant RLER throughout the 24 h, indicating that salt influences the rate of utilization of assimilates independently of their production. DCMU reduced RLER considerably in both control and salt-treated plants and reduced the advantage of control plants during the day. Conditions of low light also reduced the differences in RLER between control and salt-treated plants. When salt was removed from the root medium of non-DCMU plants, the expansion rates equalled that of the controls within 24 h and remained at the same levels for the following 3 d measurement period: this recovery applied to leaves of all ages. Salt-grown plants with no photosynthesis (DCMU treatments) also increased their expansion rates upon removal of salt from the root medium, thus providing further evidence that growth was not limited by carbohydrate status, i.e. that salt influences growth primarily via its effects on the rate of utilization of stored assimilates.     

Combined effects of long-term salinity and soil drying on growth, water relations, nutrient status and proline accumulation of Sesuvium portulacastrum

The interaction between soil drying and salinity was studied in the perennial halophyte, Sesuvium portulacastrum. Rooted cuttings were individually cultivated for three months in silty-sandy soil under two irrigation modes: 100 and 25% of field capacity (FC). The amount of the evapotranspirated water was replaced by a nutrient solution containing either 0 or 100 mM NaCl. Whole-plant growth, leaf water content, leaf water potential (Psi(w)), and Na+, K+, and proline concentrations in the tissues were measured. When individually applied, both drought and salinity significantly restricted whole-plant growth, with a more marked effect of the former stress. However, the effects of the two stresses were not additive on whole-plant biomass or on leaf expansion. Root growth was more sensitive to salt than to soil drying, the latter being even magnified by the adverse impact of salinity. Leaf water content was significantly reduced following exposure to water-deficit stress, but was less affected in salt-treated plants. When simultaneously submitted to water-deficit stress and salinity, plants displayed higher values of water and potassium use efficiencies, leaf proline and Na+ concentrations, associated with lower leaf water potential (-1.87 MPa), suggesting the ability of S. portulacastrum to use Na+ and proline for osmotic adjustment.     

Signalling mechanisms involved in the response of two varieties of Humulus lupulus L. to soil drying: I. changes in xylem sap pH and the concentrations of abscisic acid and anions

Background and aims

Soil drying leads to the generation of chemical signals in plants that regulate water use via control of the stomatal aperture. The aim of our work was to identify the presence and identity of potential chemical signals, their dynamics, and their relationship with transpiration rate during soil drying in hop (Humulus lupulus (L.)) plants.

Methods

We used pressure chamber technique for measurement of shoot water potential and collection of shoot xylem sap. We analyzed concentrations of abscisic acid (ABA), nitrate, phosphate, sulphate and malate in sap and also the rate of whole plant transpiration.

Results

Transpiration rate decreased prior to changes in shoot water potential. The concentration of ABA in xylem sap continuously increased from early to later stages of water stress, whereas in leaves it increased only at later stages. Shoot sap pH increased simultaneously with the decrease of transpiration rate. Xylem sap alkalization was in some cases accompanied by a decrease in nitrate concentration and an increase in malate concentration. Concentration of sulphate increased in xylem sap during drying and sulphate in combination with a higher ABA concentration enhanced stomatal closure.

Conclusions

Several early chemical signals appear in sap of hop plants during soil drying and their impact on transpiration may vary according to the stage of soil drying.     

Zinc Extraction potential of two common crop plants,Nicotiana tabacum and Zea mays

White spruce [Picea glauca (Moench) Voss] seedlings were inoculated with Hebeloma crustuliniforme and treated with 25 mM NaCl to examine the effects of salinized soil and mycorrhizae on root hydraulic conductance and growth. Mycorrhizal seedlings had significantly greater shoot and root dry weights, number of lateral branches and chlorophyll content than non-mycorrhizal seedlings. Salt treatment reduced seedling growth in both non-mycorrhizal and mycorrhizal seedlings. However, needles of salt-treated mycorrhizal seedlings had several-fold higher needle chlorophyll content than that in non-mycorrhizal seedlings treated with salt. Mycorrhizae increased N and P concentrations in seedlings. Na levels in shoots and roots of salt-treated mycorrhizal seedlings were significantly lower and root hydraulic conductance was several-fold higher than in non-mycorrhizal seedlings. A reduction of about 50% in root hydraulic conductance of mycorrhizal seedlings was observed after removal of the fungal hyphal sheath. Transpiration and root respiration rates were reduced by salt treatments in both groups of seedlings compared with the controls, however, both transpiration and respiration rates of salt-treated mycorrhizal seedlings were as high as those in the non-mycorrhizal seedlings that had not been subjected to salt treatment. The reduction of shoot Na uptake while increasing N and P absorption and maintaining high transpiration rates and root hydraulic conductance may be important resistance mechanisms in ectomycorrhizal plants growing in salinized soil.     

Effects of short-term salinity on leaf gas exchange of the fig (Ficus carica L.)

The influence of short-term salinity (day 1–day 2: 50 mol m^–3 NaCl, day 3–day 7: 100 mol m^–3 NaCl in the nutrient solution) on leaf gas exchange characteristics were studied in two fig clones (Ficus carica L.), whose root mass had been varied in relation to the leaf area. The stomatal conductance was diminished by NaCl in the first week of treatment. NaCl slightly reduced the calculated intercellular partial pressure of CO₂. The net photosynthetic rate of plants with many roots was stimulated by NaCl on some days of the first week of treatment, whereas the net assimilation rate of the plants with few roots remained unaltered or decreased by NaCl. Only the assimilation of the salt-treated plants of one clone for some days during the first week of treatment seemed to be influenced by stomatal conductance. Nonstomatal factors were primarily responsible for the changes in CO₂ uptake in response to salt and/or root treatment. The water use efficiency increased during several days of the first week of NaCl treatment. Decreased stomatal conductance, increased water use efficiency and stimualtion of the net CO₂ assimilation rate appear to enhance salt tolerance during the first few days of salinity. ei]H Lambers     

Water relations of pine seedlings in relation to root and shoot growth

The effects of water stress on growth and water relations of loblolly and white pine seedlings were studied during series of drying cycles. As mean soil water potential decreased, growth of roots, needles, and buds decreased. Growth of roots during successive severe drying cycles was not uniform, however. A study of needle and root extension showed that of the total growth of roots for 3 7-day drying cycles, only 6% occurred during the third cycle, while needle extension was uniform for the 3 cycles. The difference in response of needles and roots to drying cycles may be attributed primarily to the effect of water stress on the growing region. When subjected to a severe stress, roots matured toward the tip and became dormant, resulting in less growth during subsequent drying cycles. The intercalary growing region of needles, however, was not altered seriously enough by the stress to cause a difference in amount of growth during each drying cycle.

Transpiration of loblolly pine was lower in the second drying cycle than in the first. Needle water potential after rewatering was as high as that of control plants watered daily; root resistance was apparently not important in restricting transpiration during a second drying cycle. Needle diffusion resistance of loblolly pine, measured with a low-resistance diffusion porometer, was slightly higher during the second drying cycle than during the first. In addition, many primary needles were killed during the first period of stress. These factors contributed to the reduction of transpiration during the second drying cycle. Diffusion resistance of Coleus increased and transpiration ceased during the first drying cycle while water potential remained relatively high. After rewatering, both leaf resistance and transpiration returned to the control level, presumably because the stress during the first period of drying was not severe. The diffusion resistances observed for well-watered plants were 30 to 50 sec·cm⁻¹ for loblolly pine, 3 to 5 sec·cm⁻¹ for Coleus, and 4 to 6 sec·cm⁻¹ for tomato. These values agree closely with those reported by other workers.

     

Growth and Osmotic Adjustment of Cultured Suspension Cells from Alternanthera philoxeroides (Mart.) Griseb After an Abrupt Increase in Salinity

We have developed a cell suspension culture from alligator weed(Alternanthera philoxeroides [Mart.] Griseb), a C₃ member ofthe Amaranthaceae. Intact plants of alligator weed can growat 400 mol m^–3 NaCl. Growth of alligator weed suspensionswas compared to growth of tobacco (Nicotiana tabacum L. cv.Wisconsin 38) suspensions after subculture to 200 mol m^–3NaCl. Fresh weight and cell density of salt-treated alligatorweed suspensions more than doubled by 7 d after subculture,but the fresh weight of salt-treated tobacco suspensions didnot double during the 21 d experiment. Correspondingly, cellviability dropped from about 90% to 77% in alligator weed andto 41% in tobacco, at 1 d after subculture to 200 mol m^–3NaCl. The symplastic volume of alligator weed cells declined36% by 2 h after subculture to 200 mol m^–3 NaCl, but cellcontents became iso-osmotic with the media at this point. Between2 h and 6 h there was a further decrease in osmotic potential,an increase in turgor potential and a partial recovery of symplasticvolume. Turgor potential was similar to that in control cellsby 24 h, indicating significant osmotic adjustment. Turgor potentialsremained similar in both treatments from 24 h through 21 d butthe average symplastic volume of salt-treated cells was 11 %less than in control cells. Therefore, alligator weed suspensioncells exhibit a rapid recovery of water balance and cell growthafter an abrupt and substantial increase in salinity. Key words: Cell culture, growth, osmotic adjustment, salinity, turgor potential     

The Relationship Between Plant Water Potential and Transpiration Rate in Young Cabbage Plants Growing in Wet Soil

Water potentials in the plant stem were monitored together withtranspiration rates in young cabbage plants growing in potsof soil and subjected to cyclic transpirational demands. Whenthe soil was very wet plant water potential was approximatelyproportional to transpiration rate. At lower soil water contentswater potentials remained almost constant despite considerablechanges in transpiration rate. When the transpiration rate washigh, water potentials in plants were lower in wet soil thanin soil having a slightly lower moisture content. It is arguedthat the results can best be explained in terms of root conductivityincreasing as the plant becomes more water stressed when thegradient between soil and plant water potentials steepens. Key words: Cabbage, Stem water potential, Transpiration rate     

KCl和NaCl处理对盐生植物碱蓬幼苗生长和水分代谢的影响

研究了用不同浓度KCl和等浓度NaCl处理的盐生植物碱蓬幼苗水分代谢的变化.结果表明,NaCl处理显著促进碱蓬生长,根吸水增加,含水量高于对照.而相同浓度KCl处理却严重抑制碱蓬生长,含水量显著下降,400 mmol/L的 KCl处理后6 d,地上部分发生萎蔫,尔后死亡.KCl处理伤害碱蓬的原因之一是抑制根系吸水,而对蒸腾速率无显著影响,即水分吸收速率与蒸腾速率比值下降,植株缺水引起伤害.     

Abscisic Acid is not the only stomatal inhibitor in the transpiration stream of wheat plants

Xylem sap was collected from the transpiration stream of wheat (Triticum aestivum L.) plants and assayed for the presence of an inhibitor of transpiration using leaves detached from well-watered plants. Transpiration of detached leaves was reduced by nearly 60% by sap collected from plants in drying soil, and to a lesser extent (about 25%) by sap from plants in well-watered soil. As the soil dried the abscisic acid (ABA) concentration in the sap increased by about 50 times to 5 × 10⁻⁸ molar. However, the ABA in the sap did not cause its inhibitory activity. Synthetic ABA of one hundred times this concentration was needed to reduce transpiration rates of detached leaves to the same extent. Furthermore, inhibitory activity of the sap was retained after its passage through an immunoaffinity column to remove ABA. Xylem sap was also collected by applying pressure to the roots of plants whose leaf water status was kept high as the soil dried. Sap collected from these plants reduced transpiration to a lesser extent than sap from nonpressurised plants. This suggests that the inhibitory activity was triggered partly by leaf water deficit and partly by root water deficit.     

Non-hydraulic signals from maize roots in drying soil: inhibition of leaf elongation but not stomatal conductance

Conditions of soil drying and plant growth that lead to non-hydraulic inhibition of leaf elongation and stomatal conductance in maize (Zea mays L.) were investigated using plants grown with their root systems divided between two containers. The soil in one container was allowed to dry while the other container was kept well-watered. Soil drying resulted in a maximum 35% inhibition of leaf elongation rate which occurred during the light hours, with no measurable decline in leaf water potential (_w). Leaf area was 15% less than in control plants after 18 d of soil drying. The inhibition of elongation was observed only when the soil _w declined to below that of the leaves and, thus, the drying soil no longer contributed to transpiration. However, midday root _w in the dry container (-0.29 MPa) remained much higher than that of the surrounding soil (-1.0 MPa) after 15 d of drying, indicating that the roots in drying soil were rehydrated in the dark.To prove that the inhibition of leaf elongation was not caused by undetectable changes in leaf water status as a result of loss of half the watergathering capacity, one-half of the root system of control plants was excised. This treatment had no effect on leaf elongation or stomatal conductance. The inhibition of leaf elongation was also not explained by reductions in nutrient supply.Soil drying had no effect on stomatal conductance despite variations in the rate or extent of soild drying, light, humidity or nutrition. The results indicate that non-hydraulic inhibition of leaf elongation may act to conserve water as the soil dries before the occurrence of shoot water deficits.Symbol _w water potential Contribution from the Missouri Agricultural Experiment Station, Journal Series No. 10881     

Substitution of Na+ by K+ in Tissues and Root Vacuoles of Barley (Hordeum vulgare L. cv. Aramir)

The aim of this paper was to investigate whether Na⁺ accumulated in the vacuoles of barley roots can be replaced by K⁺, or else remain irreversibly trapped.Barley seedlings (Hordeum vulgare L. cv. Aramir) were grown for 2 days on nutrient solutions and then salinized over 24 h by a linear increase in NaCl concentration (50 mol · m⁻³). NaCl was replaced by an equimolar concentration of KCl and 1 day later the KCl concentration was gradually decreased over 23 hours. After this the plants were grown for 2 further days on basic nutrient solution. Plant material was harvested at each experimental step in order to measure plant growth and K⁺ and Na⁺ concentrations in vacuoles and tissues.The results from quantitated, energy dispersive X-ray analyses on frozen, hydrated vacuoles of the rhizodermls and 3 subsequent cortical cell rows, point to a proportional replacement of K⁺ by Na⁺ after NaCl stress. A replacement of Na⁺ by K⁺ was clearly found in the rhizodermal vacuoles after KCl treatment. Flame emission spectrometric data from corresponding root segments support these results.     

The responses of stomata and leaf gas exchange to vapour pressure deficits and soil water content

The responses of leaf conductance, leaf water potential and rates of transpiration and net photosynthesis at different vapour pressure deficits ranging from 10 to 30 Pa kPa^-1 were followed in the sclerophyllous woody shrub Nerium oleander L. as the extractable soil water content decreased. When the vapour pressure deficit around a plant was kept constant at 25 Pa kPa^-1 as the soil water content decreased, the leaf conductance and transpiration rate showed a marked closing response to leaf water potential at-1.1 to-1.2 MPa, whereas when the vapour pressure deficit around the plant was kept constant at 10 Pa kPa^-1, leaf conductance decreased almost linearly from-0.4 to-1.1 MPa. Increasing the vapour pressure deficit from 10 to 30 Pa kPa^-1 in 5 Pa kPa^-1 steps, decreased leaf conductance at all exchangeable soil water contents. Changing the leaf water potential in a single leaf by exposing the remainder of the plant to a high rate of transpiration decreased the water potential of that leaf, but did not influence leaf conductance when the soil water content was high. As the soil water content was decreased, leaf conductances and photosynthetic rates were higher at equal levels of water potential when the decrease in potential was caused by short-term increases in transpiration than when the potential was decreased by soil drying.As the soil dried and the stomata closed, the rate of photosynthesis decreased with a decrease in the internal carbon dioxide partial pressure, but neither the net photosynthetic rate nor the internal CO₂ partial pressure were affected by low water potentials resulting from short-term increases in the rate of transpiration. Leaf conductance, transpiration rate and net photosynthetic rate showed no unique relationship to leaf water potential, but in all experiments the leaf gas exchange decreased when about one half of the extractable soil water had been utilized. We conclude that soil water status rather than leaf water status controls leaf gas exchange in N. oleander.     

Salt-tolerance in plants. I. Ions, compatible organic solutes and the stability of plant ribosomes

Abstract Polysomes and ribosomes recovered from a number of plant species were tested for stability when incubated at 25°C in salt solutions in the absence of ATP and initiation factors. Stability was assessed by sucrose density gradient analysis. The stability was inversely proportional to salt concentrations above 125 mol m⁻³ KCl. Polysomes were less stable in the presence of Na⁺ than K⁺ salts, and were much less stable in Cl⁻ than in acetate salts. Polysomes from Triticum aestivum. Hordeum vulgare, Capsicum annuum, Helianthus annuus. Pisum sativum, Atriplex nummularia, Beta vulgaris, Cladophora sp., Enteromorpha sp. and Corallina cuvieri were similarly sensitive to KCl. Polysomes from Ulva lactuca were more sensitive than the other species. Cytoplasmic and plastid polysomes from T. aestivum were similarly unstable in 500 mol m⁻³ KCl. Unprogrammed ribosomal subunit couples from T. aestivum, B. vulgaris and U. lactuca showed Mg²⁺-dependent conformational instability and dissociation in KCl. Slight differences in ribosomal stability were observed between species, but these were unrelated to the salt tolerances of the plants. The ‘compatible’ organic solutes, glycinebetaine and proline, failed to reduce ion-induced instability. Ribosome yield and polysome profiles were similar in leaves of B. vulgaris containing significantly different levels of both Na⁺ and Cl⁻ after growth in media containing 50 or 200 mol m⁻³ NaCl. The results are consistent with the hypothesis that plants maintain a cytoplasmic solute environment that is compatible with ribosomal stability.     

Effects of PAR intensity and NaCl concentration on growth of <Emphasis Type="Italic">Salicornia europaea</Emphasis> plants as relevant to artificial ecological systems

Effects of variable levels of photosynthetically active radiation (PAR) and NaCl concentrations, typical of closed ecological life support systems, on growth of Salicornia europaea L. plants, CO₂ exchange, mineral composition, and the content of malondialdehyde (MDA) and photosynthetic pigments were investigated. The plants were grown for 25 days at different salinities of nutrient Knop solution (171, 342, and 513 mM NaCl) under two PAR levels (690 and 1150 μmol/(m² s)). At PAR of 690 μmol/(m² s), the plant productivity did not show significant changes at increasing salinities; at 1150 μmol/(m² s), the maximal productivity was observed at NaCl concentrations of 171 and 342 mM. The increase in NaCl concentration from 171 to 513 mM in the nutrient solution led to a substantial increase in the relative Na content in aboveground organs at PAR level of 1150 μmol/(m² s). The MDA content in aboveground organs by the end of the growth period was independent of PAR intensity. The content of photosynthetic pigments in the assimilatory tissue decreased with the increase in salinity from 342 to 513 mM NaCl at PAR level of 1150 μmol/(m² s) but not at the lower irradiance. The combination of 1150 μmol/(m² s) PAR intensity with the salinity as high as 342 mM NaCl was found to be the most effective for optimal productivity of S. europaea plants.     

Hardening of root cell walls: a growth inhibitory response to salinity stress

Primary roots of intact maize plants (Zea mays L.) grown for several days in nutrient solutions containing 100 mol m⁻³ NaCl and additional calcium, had relatively inhibited rates of elongation. Possible physical restraints underlying this salt induced inhibition were investigated. The inhibition did not involve reductions in osmotic potential gradients and turgor in the tip tissues responsible for root elongation growth. The apparent yield threshold pressure, which is related to capacity of cell walls to undergo loosening by stress relaxation, was estimated psychrometrically in excised root tips. Salinity increased yield threshold values. Comparative root extensibility values were obtained for intact plants by determining the initial (1 min) increase in root elongation rate induced by an 0.1 MPa osmotic jump. Comparative extensibility was significantly reduced in the salinized root tips. Salinity did not reduce capacities for water efflux and associated elastic contraction in root tip tissues of intact plants exposed to hypertonic mannitol. We conclude that cell wall hardening in the elongating root tips is an important component of root growth inhibition induced by long-term salinization.     

The influence of arbuscular mycorrhizal colonization on soil–root hydraulic conductance in Agrostis stolonifera L. under two water regimes

The hypothesis that mycorrhizal colonization improves the soil–root conductance in plants was experimentally tested in a growth chamber using pot cultures of Agrostis stolonifera L. colonized by Glomus intraradices. Plants were grown in 50-l pots filled with autoclaved sand/silt soil (1:1), with and without the mycorrhizal fungus. Within the mycorrhizal treatment, half of the pots remained well watered, while the other half was subjected to a progressive water deficit. Soil water potential (estimated as plant water potential measured at the end of the dark period), xylem water potential measured at the tiller base, transpiration rate, and soil water content were monitored throughout the experiment. Soil–root hydraulic conductance was estimated as the ratio between the instantaneous transpiration rate and the soil and xylem water potential difference. To obtain cultures with similar nutritional status, the P in the modified Hoagland’s nutrient solution was withheld from the inoculated pots and applied only once a month. Even though there were no differences on growth or nutrient status for the mycorrhizal treatments, water transport was enhanced by the inoculum presence. Transpiration rate was maintained at lower xylem water potential values in the presence of mycorrhizae. The analysis of the relationship between soil–root hydraulic resistance and soil water content showed that mycorrhizal colonization increased soil–root hydraulic conductance as the soil dried. For these growing conditions, this effect was ascribed to the range of 6–10%.     

Influence of soil drying on root development,water relations and leaf growth of Ceratonia siliqua L.

Summary Seedlings of Ceratonia siliqua L., an evergreen sclerophyll species native to the Mediterranean region, were grown in 30-cm deep tubes of John Innes II potting compost in a growth cabinet maintained at 15° C during a 12-h day where PAR was 400 mol m^–2 s^–1. After a period of acclimatisation to the conditions in the cabinet during which plants were watered every day, water was withheld from the soil in some tubes for 24 days. These conditions may be regarded as a simulation of the natural situation. Estimates of leaf and root water potential and solute potential, leaf growth and root development were made at intervals during the soil drying cycle on both watered and unwatered plants. Water potential and solute potential measurements were made both on young expanding and on fully expanded leaves. During the experimental period, root growth of C. siliqua was not much affected by soil drying, and roots in both the watered and the unwatered columns penetrated to the bottom of the soil tubes by the end of the drying treatment. Expanded leaves showed significant limitation in stomatal conductance as soil drying progressed. Leaf water potential of fully expanded leaves of unwatered plants declined substantially. In contrast, water potential of young expanding leaves on unwatered plants declined to only a limited extent and turgor was sustained. As the soil dried, stomatal conductance of young leaves was always higher than that of mature leaves; also, placticity and elasticity of young leaves slowly decreased whereas mature leaves became stiff. Changing leaf cell wall properties may determine different patterns of water use as the leaves age. A mechanism of continuous diffusion of water through the soil towards the tip and pumping towards the young leaves is proposed.     

Performance of two Picea abies (L.) Karst. stands at different stages of decline

Summary The water relations of Picea abies in a healthy stand with green trees only and a declining stand with trees showing different stages of needle yellowing were investigated in northern Bavaria. The present study is based on observations of trees differing in their nutritional status but apparently green on both sites in order to identify changes in the response pattern which might be caused by atmospheric concentrations of air pollutants and could lead to the phenomenon of decline. Transpiration was measured as water flow through the hydroactive xylem using an equilibrium mass-flow measurement system. Total tree transpiration was monitored diurnally, from July 1985 until October 1985 at both sites. The relationship between transpiration and meteorological measurements indicated that transpiration was a linear function of the vapor pressure deficit. No differences in transpiration of green trees were observed between the two sites. Canopy transpiration was 57%–68% of total throughfall and 41%–54% of total rainfall. Due to this positive water balance, soil water potential at 10 and 20 cm depths remained close to-0.02 MPa (max.-0.09 MPa) for most of the summer. Soil water potential was correlated with the difference between the weekly precipitation and transpiration. No differences in the water relations of apparently healthy trees in the two P. abies stands were observed. It is concluded that differences between green trees at the two sites in terms of nutrient relations or growth rate cannot be explained by changes in whole-tree transpiration or soil water status.     

Alleviation of salt stress by arbuscular-mycorrhizal Glomus species in Lactuca sativa plants

Improved salt tolerance of mycorrhizal plants is commonly attributed to their better mineral nutrition, particularly phosphorus. However, the effect of arbuscular-mycorrhizal (AM) fungi on salt tolerance may not be limited to this mechanism. We investigated the possibility that non-nutritional effects of AM fungi, based on proline accumulation or increased photosynthesis and related parameters, can influence the tolerance of lettuce (Lactuca sativa L.) to salinity. Three levels of salt (3, 4 and 5 g NaCl kg^-1 dry soil) were applied and plants were maintained under these conditions for 7 weeks. The salt-treated AM plants produced greater root and shoot dry weights than unfertilized or P-fertilized non-AM controls. With increasing salinity, both shoot and root dry weights were reduced, but this decrease was greater in uninoculated plants. In particular, shoot dry weight was not reduced in G. fasciculatum-colonized plants as a consequence of salt, whereas in uninoculated plants it was reduced by about 35% at the highest salt level. Proline accumulation was considerably lower for P-amended non-AM and for AM plants except for G. mosseae-colonized plants than was the case for unamended plants. Transpiration, carbon dioxide exchange rate (CER), stomatal conductance and water use efficiency (WUE) were higher in mycorrhizal plants. At 5 g NaCl kg^-1, both photosynthesis and WUE increased by more than 100% in mycorrhizal treatment relative to uninoculated plants. The contents of phosphorus of P-fertilized non-AM plants was similar to or higher than those of G. mosseae- and G. fasciculatum-colonized plants. Plants colonized by G. deserticola had the highest P-content regardless of salt level. Hence, the effect of G. mosseae and G. fasciculatum on salt tolerance in this experiment could not be attributed to a difference in the P content. The mechanisms by which these two fungi alleviated salt stress appeared to be based on physiological processes (increased CER, transpiration, stomatal conductance and WUE) rather than on nutrient uptake (N or P).