The arbuscular mycorrhizal symbiosis regulates aquaporins activity and improves root cell water permeability in maize plants subjected to water stress

Studies have suggested that increased root hydraulic conductivity in mycorrhizal roots could be the result of increased cell‐to‐cell water flux via aquaporins. This study aimed to elucidate if the key effect of the regulation of maize aquaporins by the arbuscular mycorrhizal (AM) symbiosis is the enhancement of root cell water transport capacity. Thus, water permeability coefficient (P_f) and cell hydraulic conductivity (L_pc) were measured in root protoplast and intact cortex cells of AM and non‐AM plants subjected or not to water stress. Results showed that cells from droughted‐AM roots maintained P_f and L_pc values of nonstressed plants, whereas in non‐AM roots, these values declined drastically as a consequence of water deficit. Interestingly, the phosphorylation status of PIP2 aquaporins increased in AM plants subjected to water deficit, and P_f values higher than 12 μm s^?1 were found only in protoplasts from AM roots, revealing the higher water permeability of AM root cells. In parallel, the AM symbiosis increased stomatal conductance, net photosynthesis, and related parameters, showing a higher photosynthetic capacity in these plants. This study demonstrates a better performance of AM root cells in water transport under water deficit, which is connected to the shoot physiological performance in terms of photosynthetic capacity.     

施磷和接种AM真菌对玉米耐盐性的影响

在盆栽条件下研究了不同施磷水平（２５,５０,１００,１５０ｍｇ／ｋｇ）,不同盐水平（ＮａＣｌ０,１．２ｇ／ｋｇ）和不同接种ＡＭ真菌处理（接种和不接种）对玉米生长的影响。结果表明,施磷量为５０ｍｇ／ｋｇ时基本满足玉米生长的需要,１．２ｇ／ｋｇ ＮａＣｌ胁迫显著抑制了玉米的生长;施磷明显促进玉米在盐胁迫条件下的生长,施磷水平和接种菌根真菌的交互作用对玉米耐盐性具有显著影响;盐胁迫条件下,接种ＡＭ真菌在     

Differences in the effects of three arbuscular mycorrhizal fungal strains on P and Pb accumulation by maize plants

The effect of arbuscular mycorrhizal (AM) fungi on the accumulation and transport of lead was studied in a pot experiment on maize plants grown in anthropogenically-polluted substrate. The plants remained uninoculated or were inoculated with different Glomus intraradices isolates, either indigenous to the polluted substrate used or reference from non-polluted soil. A considerably lower tolerance to the conditions of polluted substrate was observed for the reference isolate that showed significantly lower frequency of root colonisation as well as arbuscule and vesicule abundance. Plants inoculated with the reference isolate also had significantly lower shoot P concentrations than plants inoculated with the isolate from polluted substrate. Nevertheless, inoculation with either indigenous or reference G. intraradices isolate resulted in higher shoot and root biomass and inoculated plants showed lower Pb concentrations in their shoots than uninoculated plants, regardless of differences in root colonisation. Root biomass of maize plants was divided according to AM-induced colouration into brightly yellow segments intensively colonised by AM fungus and non-colonised or only slightly colonised whitish ones. Intensively colonised segments of the isolate from polluted substrate contained significantly higher concentrations of phosphorus and lead than non-colonised ones, which suggest significant participation of fungal structures in element accumulation. Responsible Editor: Peter Christie.     

Changes in antioxidant activities and phenol content in tomato plants subjected to partial root drying and regulated deficit irrigation

Abstract

Partial rootzone drying (PRD) and regulated deficit irrigation (RDI) are water saving irrigation systems that have been developed to increase water use efficiency (WUE) without significant yield reduction. To examine whether tomato responded differently to RDI and PRD, we compared the changes in antioxidative defenses in tomato plants using a split-root system. Tomato plants were grown for 21 days under controlled conditions with their roots separated equally between two soil compartments. Three irrigation treatments were imposed: Control, receiving an amount of water equivalent to 100% of plant transpiration; PRD in which one compartment was watered with 50% of the amount of water supplied to the controls, allowing one-half of the root system to be exposed to dry soil, and switching irrigation between sides weekly; RDI in which 50% of the amount of water given to the controls was supplied, half to each side of the root system. Relative water content (RWC), midday leaf Ψ and chlorophyll content decreased largely in RDI-treated plants, whereas the PRD plants exhibited relatively higher Ψ and RWC values. An enhanced level of lipid peroxidation in both roots and leaves indicated that PRD and RDI caused oxidative stress in tomato plants. In leaves, superoxide dismutase (SOD), soluble peroxidase (POX) and polyphenol oxidase (PPO) activities showed an increase in the early phase of water deficit, and then decreased in the remaining phase of the drying cycle. However, the increase was more pronounced under RDI. Catalase (CAT) activity declined continuously from the onset of PRD and RDI treatments to below the control level, and the reduction was less under PRD than RDI. POX cell-wall associated activities exceeded the control level by 450% and 230%, respectively, under RDI and PRD. At the root level, while CAT activity also decreased under both PRD and RDI, the activities of SOD, POX and PPO significantly increased and their activities showed an alternating increase/decrease paralleling the alternating irrigation in PRD-treated roots. As a result of the difference in POX and PPO activities between the two water treatments applied, PRD-treated plants accumulated more soluble and cell-wall bound phenolic compounds.     

Respiration of the external mycelium in the arbuscular mycorrhizal symbiosis shows strong dependence on recent photosynthates and acclimation to temperature

* Although arbuscular mycorrhizal (AM) fungi are a major pathway in the global carbon cycle, their basic biology and, in particular, their respiratory response to temperature remain obscure. * A pulse label of the stable isotope (13)C was applied to Plantago lanceolata, either uninoculated or inoculated with the AM fungus Glomus mosseae. The extra-radical mycelium (ERM) of the fungus was allowed to grow into a separate hyphal compartment excluding roots. We determined the carbon costs of the ERM and tested for a direct temperature effect on its respiration by measuring total carbon and the (13)C:(12)C ratio of respired CO(2). With a second pulse we tested for acclimation of ERM respiration after 2 wk of soil warming. * Root colonization remained unchanged between the two pulses but warming the hyphal compartment increased ERM length. delta(13)C signals peaked within the first 10 h and were higher in mycorrhizal treatments. The concentration of CO(2) in the gas samples fluctuated diurnally and was highest in the mycorrhizal treatments but was unaffected by temperature. Heating increased ERM respiration only after the first pulse and reduced specific ERM respiration rates after the second pulse; however, both pulses strongly depended on radiation flux. * The results indicate a fast ERM acclimation to temperature, and that light is the key factor controlling carbon allocation to the fungus.     

Citrus root responses to localized drying soil: A new approach to studying mycorrhizal effects on the roots of mature trees

Because fine roots tend to be concentrated at the soil surface, exposure to dry surface soil can have a large influence on patterns of root growth, death and respiration. We studied the effects of arbuscular mycorrhizas (AM) formation on specific root length (SRL), respiration and mortality of fine roots of bearing red grapefruit (Citrus paradisi Macf.) trees on Volkamer lemon (C. volkameriana Tan. & Pasq.) rootstock exposed to drying soil. For each tree, the fine roots were removed from two woody lateral roots, the roots were surface sterilized and then each woody root was placed in a separate pair of vertically divided and independently irrigated soil compartments. The two split-pot systems were filled with sterilized soil and one was inoculated with arbuscular mycorrhizal fungi (Glomus etunicatum/G. intraradices). New fine lateral roots that emerged from the woody laterals were permitted to grow inside the pots over a 10-month period. Irrigation was then removed from the top compartment for a 15-week period. At the end of the study, roots inoculated with AM fungi exhibited about 20% incidence of AM formation, whereas the uninoculated roots were completely void of AM fungi. Arbuscular mycorrhizal roots exhibited lower SRL, lower root/soil respiration and about 10% lower fine root mortality than nonmycorrhizal roots after 15 weeks of exposure to dry surface soil. This study demonstrates the feasibility of examining mycorrhizal effects on the fine roots of adult trees in the field using simple inexpensive methods.     

Effects of different arbuscular mycorrhizal fungal backgrounds and soils on olive plants growth and water relation properties under well‐watered and drought conditions

The adaptation capacity of olive trees to different environments is well recognized. However, the presence of microorganisms in the soil is also a key factor in the response of these trees to drought. The objective of the present study was to elucidate the effects of different arbuscular mycorrhizal (AM) fungi coming from diverse soils on olive plant growth and water relations. Olive plants were inoculated with native AM fungal populations from two contrasting environments, that is, semi‐arid – Freila (FL) and humid – Grazalema (GZ) regions, and subjected to drought stress. Results showed that plants grew better on GZ soil inoculated with GZ fungi, indicating a preference of AM fungi for their corresponding soil. Furthermore, under these conditions, the highest AM fungal diversity was found. However, the highest root hydraulic conductivity (Lp_r) value was achieved by plants inoculated with GZ fungi and growing in FL soil under drought conditions. So, this AM inoculum also functioned in soils from different origins. Nine novel aquaporin genes were also cloned from olive roots. Diverse correlation and association values were found among different aquaporin expressions and abundances and Lp_r, indicating how the interaction of different aquaporins may render diverse Lp_r values.     

Impact of defoliation intensities on plant biomass,nutrient uptake and arbuscular mycorrhizal symbiosis in Lotus tenuis growing in a saline‐sodic soil

The impact of different defoliation intensities on the ability of Lotus tenuis plants to regrowth, mobilise nutrients and to associate with native AM fungi and Rhizobium in a saline‐sodic soil was investigated. After 70 days, plants were subjected to 0, 25, 50, 75 and 100% defoliation and shoot regrowth was assessed at the end of subsequent 35 days. Compared to non‐defoliated plants, low or moderate defoliation up to 75% did not affect shoot regrowth. However, 100% treatment affected shoot regrowth and the clipped plants were not able to compensate the growth attained by non‐defoliated plants. Root growth was more affected by defoliation than shoot growth. P and N concentrations in shoots and roots increased with increasing defoliation while Na⁺ concentration in shoots of non‐defoliated and moderately defoliated plants was similar. Non‐defoliated and moderately defoliated plants prevented increases of Na⁺ concentration in shoots through both reducing Na⁺ uptake and Na⁺ transport to shoots by accumulating Na⁺ in roots. At high defoliation, the salinity tolerance mechanism is altered and Na⁺ concentration in shoots was higher than in roots. Reduction in the photosynthetic capacity induced by defoliation neither changed the root length colonised by AM fungi nor arbuscular colonisation but decreased the vesicular colonisation. Spore density did not change, but hyphal density and Rhizobium nodules increased with defoliation. The strategy of the AM symbiont consists in investing most of the C resources to preferentially retain arbuscular colonisation as well as inoculum density in the soil.     

Opening the black box: outcomes of interactions between arbuscular mycorrhizal (AM) and non‐host genotypes of Medicago depend on fungal identity,interplay between P uptake pathways and external P supply

We investigated the physiology that underlies the influence of arbuscular mycorrhizal (AM) colonization on outcomes of interactions between plants. We grew Medicago truncatula A17 and its AM‐defective mutant dmi1 in intragenotypic (two plants per pot of the same genotype, x2) or intergenotypic (one plant of each genotype, 1 + 1) combinations, inoculated or not with Rhizophagus irregularis (formerly Glomus intraradices) or Gigaspora margarita. We measured plant growth, colonization, contributions of AM and direct P uptake pathways using ³²P, and expression of plant P_i transporter genes at two levels of P supply. A17 (x2) responded positively to inoculation only at low P. The response was enhanced with 1 + 1 even at high P where colonization in A17 was reduced. With R. irregularis P uptake by the AM pathway was unaffected by P supply, whereas with G. margarita, the AM pathway was lower at high P, and direct uptake higher. Gene expression varied and was unrelated to P uptake through the two pathways. There was no evidence of plant control of P uptake via R. irregularis at high P but there was via G. margarita. Importantly, growth responses of plant genotypes grown alone did not predict outcomes of intergenotypic interactions.     

Different effects of phospholipase Dζ2 and non‐specific phospholipase C4 on lipid remodeling and root hair growth in Arabidopsis response to phosphate deficiency

Phosphate (Pi) deficiency in soils is a major limiting factor for plant growth. In response to Pi deprivation, one prominent metabolic adaptation in plants is the decrease in membrane phospholipids that consume approximately one‐third cellular Pi. The level of two phospholipid‐hydrolyzing enzymes, phospholipase Dζ2 (PLDζ2) and non‐specific phospholipase C4 (NPC4), is highly induced in Pi‐deprived Arabidopsis. To determine the role of PLDζ2 and NPC4 in plant growth under Pi limitation, Arabidopsis plants deficient in both PLDζ2 and NPC4 (npc4pldζ2) were generated and characterized. Lipid remodeling in leaves and roots was analyzed at three different durations of Pi deficiency. NPC4 affected lipid changes mainly in roots at an early stage of Pi deprivation, whereas PLDζ2 exhibited a more overt effect on lipid remodeling in leaves at a later stage of Pi deprivation. Pi deficiency‐induced galactolipid increase and phospholipid decrease were impeded in pldζ2 and npc4pldζ2 plants. In addition, seedlings of npc4pldζ2 had the same root hair density as pldζ2 but shorter root hair length than pldζ2 in response to Pi deficiency. The loss of NPC4 decreased root hair length but had no effect on root hair density. These data suggest that PLDζ2 and NPC4 mediate the Pi deprivation‐induced lipid remodeling in a tissue‐ and time‐specific manner. PLDζ2 and NPC4 have distinctively different roles in root hair growth and development in response to Pi deprivation; PLDζ2 negatively modulates root hair density and length, whereas NPC4 promotes root hair elongation.