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Rice is one of the most important crops in the world and is considered a strategic crop for food security. Furthermore, the excessive use of chemical fertilizers to obtain high yields causes environmental problems. A sustainable alternative includes taking advantage of beneficial bacteria that promote plant growth. Here, we investigate the effect of five bacterial biofertilizers from halophytes on growth, and we investigate photosynthetic efficiency in rice plants grown under saline conditions (0 and 85 mmol L -1 NaCl) and future climate change scenarios, including increased CO 2 concentrations and temperature (400/700 ppm and 25/+4 °C, respectively). Biofertilizers 1-4 increased growth by 9-64% in plants grown with and without salt in both CO 2 - temperature combinations, although there was no significant positive effect on the net photosynthetic rate of rice plants. In general, biofertilizer 1 was the most effective at 400 ppm CO 2 and at 700 ppm CO 2 +4 °C in the absence of salt. Inocula 1-5 also stimulated plant length at high CO 2 levels without salt. Finally, the positive effect of biofertilization was attenuated in the plants grown under the interaction between salt and high CO 2 . This highlights the significance of studying biofertilization under stress interaction to establish the real potential of biofertilizers in the context of climate change conditions.

The Earth is undergoing alterations at a high speed, which causes problems such as environmental pollution and difficulty in food production. This is where halophytes are interesting, due to their high potential in different fields, such as remediation of the environment and agriculture. For this reason, it is necessary to deepen the knowledge of the development of halophytes and how plant growth-promoting bacteria (PGP) can play a fundamental role in this process. Therefore, in this work were tested the effects of five PGP bacteria on its rhizosphere and other endophytic bacteria at different concentrations of NaCl on seed germination, plant growth (0 and 171 mM) and cutting growth (0 mM) of Arthrocaulon macrostachyum . The growth promotion in this strict halophyte is highlighted due to the presence of PGP bacteria and the fact that no salt is needed. Thus, without salt, the bacterial strains Kocuria polaris Hv16, Pseudarthrobacter psychrotolerans C58, and Rahnella aceris RTE9 enhanced the biomass production by more than 60% in both stems and roots. Furthermore, germination was encouraged by more than 30% in the presence of both R. aceris RTE9 and K. polaris Hv16 at 171 mM NaCl; the latter also had a biocontrol effect on the fungi that grew on the seeds. Additionally, for the first time in cuttings of this perennial species, the root biomass was improved thanks to the consortium of K. polaris Hv16 and P. psychrotolerans C58. Finally, this study demonstrates the potential of PGPs for optimising the development of halophytes, either for environmental or agronomic purposes.

Soil contamination with petroleum-derived substances such as diesel fuel has become a major environmental threat. Phytoremediation is one of the most studied ecofriendly low-cost solutions nowadays and halophytes species has been proved to have potential as bio-tools for this purpose. The extent to which salinity influences diesel tolerance in halophytes requires investigation. A greenhouse experiment was designed to assess the effect of NaCl supply (0 and 85 mM NaCl) on the growth and photosynthetic physiology of Juncus acutus plants exposed to 0, 1 and 2.5% diesel fuel. Relative growth rate, water content and chlorophyll a derived parameters were measured in plants exposed to the different NaCl and diesel fuel combinations. Our results indicated that NaCl supplementation worsened the effects of diesel toxicity on growth, as diesel fuel at 2.5% reduced relative growth rate by 25% in the absence of NaCl but 80% in plants treated with NaCl. Nevertheless, this species grown at 0 mM NaCl showed a high tolerance to diesel fuel soil presence in RGR but also in chlorophyll fluorescence parameters that did not significantly decrease at 1% diesel fuel concentration in absence of NaCl. Therefore, this study remarked on the importance of knowing the tolerance threshold to abiotic factors in order to determine the bioremediation capacity of a species for a specific soil or area. In addition, it showed that NaCl presence even in halophytes does not always have a positive effect on plant physiology and it depends on the pollutant nature.

Land salinization, resulting from the ongoing climate change phenomena, is having an increasing impact on coastal ecosystems like salt marshes. Although halophyte species can live and thrive in high salinities, they experience differences in their salt tolerance range, being this a determining factor in the plant distribution and frequency throughout marshes. Furthermore, intraspecific variation to NaCl response is observed in high-ranging halophyte species at a population level. The present study aims to determine if the environmental history, namely heavy metal pre-conditioning, can have a meaningful influence on salinity tolerance mechanisms of Spartina patens , a highly disperse grass invader in the Mediterranean marshes. For this purpose, individuals from pristine and heavy metal contaminated marsh populations were exposed to a high-ranging salinity gradient, and their intraspecific biophysical and biochemical feedbacks were analyzed. When comparing the tolerance mechanisms of both populations, S. patens from the contaminated marsh appeared to be more resilient and tolerant to salt stress, this was particularly present at the high salinities. Consequently, as the salinity increases in the environment, the heavy metal contaminated marsh may experience a more resilient and better adapted S. patens community. Therefore, the heavy metal pre-conditioning of salt mash populations appears to be able to create intraspecific physiological variations at the population level that can have a great influence on marsh plant distribution outcome.

Salicornia ramosissima is a C 3 halophyte that grows naturally in South Western Spain salt marshes, under soil salinity and heavy metal pollution (mostly Cu, Zn, As, and Pb) caused by both natural and anthropogenic pressure. However, very few works have reported the phytoremediation potential of S. ramosissima . In this work, we studied a microbe-assisted phytoremediation strategy under greenhouse conditions. We inoculated plant growth promoting (PGP) and heavy metal resistant bacteria in pots with S. ramosissima and natural non-polluted and polluted sediments collected from Spanish estuaries. Then, we analyzed plant ecophysiological and metal phytoaccumulation response. Our data suggested that inoculation in polluted sediments improved S. ramosissima plant growth in terms of relative growth rate (RGR) (32%) and number of new branches (61%). S. ramosissima photosynthetic fitness was affected by heavy metal presence in soil, but bacteria inoculation improved the photochemical apparatus integrity and functionality, as reflected by increments in net photosynthetic rate (21%), functionality of PSII (F m and F v /F m ) and electron transport rate, according to OJIP derived parameters. Beneficial effect of bacteria in polluted sediments was also observed by augmentation of intrinsic water use efficiency (28%) and slightly water content (2%) in inoculated S. ramosissima . Finally, our results demonstrated that S. ramosissima was able to accumulate great concentrations of heavy metals, mostly at root level, up to 200 mg Kg -1 arsenic, 0.50 mg Kg -1 cadmium, 400 mg Kg -1 copper, 25 mg Kg -1 nickel, 300 mg Kg -1 lead, and 300 mg Kg -1 zinc. Bioaugmentation incremented S. ramosissima heavy metal phytoremediation potential due to plant biomass increment, which enabled a greater accumulation capacity. Thus, our results suggest the potential use of heavy metal resistant PGPB to ameliorate the capacity of S. ramosissima as candidate for phytoremediation of salty polluted ecosystems., (Copyright © 2020 Mesa-Marín, Pérez-Romero, Redondo-Gómez, Pajuelo, Rodríguez-Llorente and Mateos-Naranjo.)

Plant growth promoting bacteria' (PGPB) beneficial role on plant tolerance to salinity stress has previously been well recognized. However, bacteria-triggered plant physiological mechanisms involved in this response require investigation, especially in plants with innate salt tolerance. A glasshouse experiment was designed to investigate the effect of the PGPB Vibrio spartinae on Halimione portulacoides growth, physiological performance and ion homeostasis in plants exposed to 0, 171, 510 and 1020 mM NaCl for 100 days. Bacterial inoculation alleviated ~28% of the deleterious impact of salinity excess on the relative growth rate (RGR) in plants grown at 510 mM and led to 30% and 44% enhancements in those exposed to 0 and 171 mM NaCl, respectively. This effect was linked to a reduction in Na tissue concentrations which improved plant ion homeostasis at elevated NaCl concentration, and to the overall protective effects on various steps in the photosynthetic pathway between 0 and 510 mM NaCl. Thus, inoculated plants were able to maintain higher net photosynthesis (A N ) than their non-inoculated counterparts. Hence, A N differences under saline conditions were ascribed to inoculation amelioration NaCl-induced CO 2 diffusion limitations, as reflected in the greater g s and C i values recorded at 171 and 510 mM NaCl, together with an enhancement of photochemical apparatus functionality (in terms of energy absorption, transformation and transport), as indicated by a higher electron transport rate (ETR) and energy fluxes derived from Kautsky curves, compared with their non-inoculated counterparts., (Copyright © 2020 Elsevier Masson SAS. All rights reserved.)

Many halophytic physiological traits related to the tolerance of plants to salinity excess have been extensively studied, with a focus on biomass and/or gas exchange parameters. To gain a more complete understanding of whether salinity excess affects the physiological performance of halophytes, an experiment was performed using the halophyte Atriplex halimus L. as a model. A. halimus plants were subjected to two salinity treatments (171 and 513 mM NaCl) over 60 days in a controlled environment. After this period, dry biomass, specific stem conductivity, water potential at turgor loss point, osmotic potential, gas exchange parameters, and the fluorescence of chlorophyll a derived parameters were assessed in order to obtain knowledge about the differences in vulnerability that these parameters can show when subjected to salinity stress. Our results showed a decrease in belowground and aboveground biomass. The decrement in biomass seen at 513 mM NaCl was related to photosynthetic limitations and specific stem conductivity. Turgor loss point did not vary significantly with the increment of salinity. Therefore, the parameter that showed less vulnerability to saline stress was the turgor loss point, with only a 5% decrease, and the more vulnerable trait was the stem conductivity, with a reduction of nearly 50%.

Climate change would increase frequency and intensity of extreme events as heat and cold waves. There is a lack of studies that consider the co-occurrence of these waves with other abiotic factors relevant on a climate change scenario as salinity. Therefore, it could be interesting to improve our knowledge about the effects that this co-occurrence could have in different species due to the species specific effect of the photosynthesis tolerance to extreme temperatures. A controlled condition experiment was performed using the salt marsh species Sarcocornia perrnis with eight experimental blocks combining temperature ranges (40-28/22-15/13-5 °C) and salinity concentration on the growth solution (171/1050 mM NaCl). After 3 days of treatment, gas exchange, chlorophyll a fluorescence, pigment profile and water state measurement were applied. Photosynthetic machinery function of this perennial species decreased on for both high and low temperature range. Nevertheless, at 13-5 °C the effect of the salinity was mainly due to diffusion limitations more than to damage on the photosystems. At 40-28 °C, in presence of optimal salinity S. fruticosa was not altered overall. However, high temperatures in combination with high salinity reduced the photosynthetic capacity mainly by reducing the efficiency of the electron transport chain., Competing Interests: Declaration of competing interest The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2019 Elsevier Masson SAS. All rights reserved.)

Polycyclic aromatic hydrocarbons (PAH) have become a threat for the conservation of wetlands worldwide. The halophyte Spartina densiflora has shown to be potentially useful for soil phenanthrene phytoremediation, but no studies on bacteria-assisted hydrocarbon phytoremediation have been carried out with this halophyte. In this work, three phenanthrene-degrading endophytic bacteria were isolated from S. densiflora tissues and used for plant inoculation. Bacterial bioaugmentation treatments slightly improved S. densiflora growth, photosynthetic and fluorescence parameters. But endophyte-inoculated S. densiflora showed lower soil phenanthrene dissipation rates than non-inoculated S. densiflora (30% below) or even bulk soil (23% less). Our work demonstrates that endophytic inoculation on S. densiflora under greenhouse conditions with the selected PAH-degrading strains did not significantly increase inherent phenanthrene soil dissipation capacity of the halophyte. It would therefore be advisable to provide effective follow-up of bacterial colonization, survival and metabolic activity during phenanthrene soil phytoremediation., (Copyright © 2019 Elsevier Inc. All rights reserved.)

A 45-days long climatic chamber experiment was design to evaluate the effect of 400 and 700 ppm atmospheric CO 2 treatments with and without soil water logging in combination with 171 and 510 mM NaCl in the halophyte Salicornia ramosissima. In order to ascertain the possible synergetic impact of these factors associate to climatic change in this plant species physiological and growth responses. Our results indicated that elevated atmospheric CO 2 concentration improved plant physiological performance under suboptimal root-flooding and saline conditions plants. Thus, this positive impact was mainly ascribed to an enhancement of energy transport efficiency, as indicated the greater P G , N and Sm values, and the maintaining of carbon assimilation capacity due to the higher net photosynthetic rate (A N ) and water use efficiency ( i WUE). This could contribute to reduce the risk of oxidative stress owing to the accumulation of reactive oxygen species (ROS). Moreover, plants grown at 700 ppm had a greater capacity to cope with flooding and salinity synergistic impact by a greater efficiency in the modulation in enzyme antioxidant machinery and by the accumulation of osmoprotective compounds and saturated fatty acids in its tissues. These responses indicate that atmospheric CO 2 enrichment would contribute to preserve the development of Salicornia ramosissima against the ongoing process of increment of soil stressful conditions linked with climatic change., (Copyright © 2018 Elsevier Masson SAS. All rights reserved.)

Increasing extreme temperature climatic events could exert an important effect on plant photosynthetic performance, which could be modulated by the co-occurrence with other environmental factors, such as salinity, in estuarine ecosystems. Therefore, a mesocosm experiment was designed to assess the impact of temperature events for three days (13/5 °C, 25/13 °C and 40/28 °C) in combination with two NaCl concentrations (171 and 1050 mM NaCl) on the physiological performance of Salicornia ramosissima. Extreme temperature events had a negative impact on S. ramosissima photosynthetic efficiency, this effect being more marked with cold wave at both salinities, compared with heat wave, even in presence of NaCl excess. This differential thermotolerance in the photosynthetic apparatus was ascribed to the greater integrity and functioning of its photosynthetic pathway at high temperature, as indicated by constant g s, V c,max values at optimal salinity and the higher values of those parameters and g m recorded in combination with NaCl excess. Moreover, S. ramosissima was able to upregulate the energy sink capacity of its photochemical apparatus at elevated temperature and salinity by a greater energy excess dissipation capacity. This could have contributed to reducing the risk of oxidative stress, along with the recorded higher capacity for antioxidant enzyme activity modulation under these conditions.

The research on the plant population metal intra-specific tolerance variability is of paramount importance for the design of phytoremediation restoration. The aim of this study was to asses if any variability exists in the copper stress response during seed germination and seedling development in Juncus acutus depending on provenance habitat. Our results showed that J. acutus were able to germinate until Cu concentration of 23 mM Cu, but at 15 and 23 mM Cu, the final percentage of germination were 100 and 68% for seeds derived from polluted area and were 86 and 40% for those collected in non-polluted one, respectively. Moreover, the germination kinetic was more impaired by Cu concentration in those no historically exposed to metal excess. Provenance effect was also reflected in seedlings survival and development; thus at 9 mM Cu higher survival percentage, total height and dry mass were recorded in seedlings derived from no polluted area compared with their historically exposed counterparts. Therefore, we can conclude that the variability of Cu tolerance in J. acutus should be considered for the design of restoration projects, since it allows use of provenances with greater potential as a source of propagules highly adapted to metal excess.

The potential importance of Juncus acutus for remediation of Zn-contaminated lands has been recognized, because of its Zn tolerance and capacity to accumulate Zn. Since it is also a halophyte, the extent to which salinity influences its Zn tolerance requires investigation. A factorial greenhouse experiment was designed to assess the effect of NaCl supply (0 and 85 mM NaCl) on the growth, photosynthetic physiology and tissue ions concentrations of plants exposed to 0, 30 and 100 mM Zn. Our results indicated that NaCl supplementation alleviated the effects of Zn toxicity on growth, as Zn at 100 mM reduced relative growth rate (RGR) by 60% in the absence of NaCl but by only 34% in plants treated also with NaCl. This effect was linked to a reduction in Zn tissue concentrations, as well as to overall protective effects on various stages in the photosynthetic pathway. Thus, at 85 mM NaCl plants were able to maintain higher net photosynthesis (A N ) than in the absence of added NaCl, although there were no differences in stomatal conductance (g s ). This contributed to preserving the trade-off between CO 2 acquisition and water loss, as indicated by higher intrinsic water use efficiency ( i WUE). Hence, A N differences were ascribed to limitation in the RuBisCO carboxylation, manifested as higher intercellular CO 2 concentration (C i ), together with dysfunction of PSII photochemistry (in term of light harvest and energy excess dissipation), as indicated by higher chronic photoinhibition percentages and variations in the photosynthetic pigment profiles in presence of Zn under non-saline conditions., (Copyright © 2018 Elsevier Inc. All rights reserved.)

An experiment was conducted to evaluate the combined effect of temperature (26 and 30 °C) and Cr toxicity (0, 100 and 1000 μM Cr) on growth, photosynthesis, water content, Cr and nutrients uptake and translocation. The role of antioxidative enzyme towards stresses tolerance was also investigated. Results showed that the maximum relative growth rate and leaf area per plant of Atriplex halimus L. were recorded at 100 μM Cr and 26 °C. However, presence of Cr reduced net photosynthetic and stomatal conductance rates. Overall, temperature rise enhanced the toxic effect of Cr by reducing growth and photosynthesis and inducing antioxidant enzymes activities. Furthermore, temperature rise increased nutrient uptake, as well as nutrient translocation to aboveground tissues; while it diminished Cr translocation. Finally, roots were the main sink for Cr accumulation in A. halimus. At 1000 μM Cr, root Cr concentrations reached 7.2 and 9.1 mg g -1 at 26 and 30 °C, respectively; while shoot Cr concentrations were 0.45 and 0.44 mg g -1 (26 and 30 °C, respectively). The high Cr-accumulation in roots suggests that A. halimus presents a great potential for phytoremediation, especially phytostabilisation of Cr contaminated soils., (Copyright © 2018 Elsevier Masson SAS. All rights reserved.)

A mesocosm experiment was designed to assess the effect of atmospheric CO 2 increment on the salinity tolerance of the C 3 halophyte Salicornia ramosissima. Thus, the combined effect of 400 ppm and 700 ppm CO 2 at 0, 171 and 510 mM NaCl on plants growth, gas exchange, chlorophyll fluorescence parameters, pigments profiles, antioxidative enzyme activities and water relations was studied. Our results highlighted a positive effect of atmospheric CO 2 increment on plant physiological performance under suboptimal salinity concentration (510 mM NaCl). Thus, we recorded higher net photosynthetic rate (A N ) values under saline conditions and 700 ppm CO 2 , being this effect mainly mediated by a reduction of mesophyll (g m ) and biochemical limitation imposed to salt excess. In addition, rising atmospheric CO 2 led to a better plant water balance, linked with a reduction of stomatal conductante (g s ) and an overall increment of osmotic potential (Ѱ o ) with NaCl concentration increment. In spite of these positive effects, there were no significant biomass variations between any treatments. Being this fact ascribed by the investment of the higher energy fixed for salinity stress defence mechanisms, which allowed plants to maintain more active the photochemical machinery even at high salinities, reducing the risk of ROS production, as indicated an improvement of the electron flux and a rise of the energy dissipation. Finally, the positive effect of the CO 2 was also supported by the modulation of pigments profiles (mainly zeaxhantin and violaxhantin) concentrations and anti-oxidative stress enzymes, such as superoxide dismutase (SOD) and ascorbate peroxidase (APx)., (Copyright © 2018 Elsevier Masson SAS. All rights reserved.)

A glasshouse experiment was designed to investigate the effect of the co-occurrence of 400 and 700ppm CO 2 at 0, 15 and 45mM Cu on the Cu-tolerance of C 4 cordgrass species Spartina densiflora, by measuring growth, gas exchange, efficiency of PSII, pigments profiles, antioxidative enzyme activities and nutritional balance. Our results revealed that the rising atmospheric CO 2 mitigated growth reduction imposed by Cu in plants grown at 45mM Cu, leading to leaf Cu concentration bellow than 270mgKg -1 Cu, caused by an evident dilution effect. On the other hand, non-CO 2 enrichment plants showed leaf Cu concentration values up to 737.5mgKg -1 Cu. Furthermore, improved growth was associated with higher net photosynthetic rate (A N ). The beneficial effect of rising CO 2 on photosynthetic apparatus seems to be associated with a reduction of stomatal limitation imposed by Cu excess, which allowed these plants to maintain greater i WUE values. Also, plants grown at 45mM Cu and 700ppm CO 2 , showed higher ETR values and lower energy dissipation, which could be linked with an induction of Rubisco carboxylation and supported by the recorded amelioration of N imbalance. Furthermore, higher ETR values under CO 2 enrichment could lead to an additional consumption of reducing equivalents. Idea that was reflected in the lower values of ETR max /A N ratio, malondialdehyde (MDA) and ascorbate peroxidase (APx), guaiacol peroxidase (GPx) and superoxide dismutase (SOD) activities under Cu excess, which could indicate a lower production of ROS species under elevated CO 2 concentration, due to a better use of absorbed energy., (Copyright © 2017 Elsevier GmbH. All rights reserved.)

Cadmium (Cd) is a non-essential element for plants, and its excess impairs plant performance. Physiological impacts of Cd excess are well known in non-tolerant plants, however this information is scarce for Cd-tolerant plants. A glasshouse experiment was designed to investigate the effect of five different Cd levels (0, 0.05, 0.20, 0.65 and 1.35 mM Cd) on the growth, photosynthetic apparatus (PSII chemistry), gas exchange characteristics, photosynthetic pigments profiles, water relations and nutritional balance of the Cd-accumulator Salicornia ramosissima. Ours results confirmed the accumulation capacity of S. ramosissima, as indicated the bioaccumulation factor (BC) greater than 1.0 for all Cd levels. Furthermore, after 21 days of treatment S. ramosissima growth was not highly affected by Cd. Total photosynthetic limitation increased from 38% at 0.05 mM Cd to 70% at 1.35 mM Cd. CO 2 diffusion restriction imposed the main contribution to total photosynthetic limitation. Mesophyll conductance reduction was of major importance (with between 69 and 86%), followed by stomatal conductance (with between 9 and 20%). Maximum carboxylation rate (V c,max ), remained stable until 0.2 mM Cd, and chlorophyll fluorescence parameters (F v /F m , qP) and pigments concentrations were not significantly decreased by increased Cd supply. Finally, S. ramosissima water relations (intrinsic water use efficiency and relative water content) and nutritional level did not highly vary between Cd treatments. Thus, our finding suggested that Cd tolerance S. ramosissima is in certain degree supported by the tolerance of its carbon assimilation enzyme (RuBisCO) and with the high functionality and integrity of the PSII reaction center under Cd excess., (Copyright © 2016 Elsevier Masson SAS. All rights reserved.)

The xero-halophyte Atriplex halimus L., recently described as Cd-hyperaccumulator, was examined to determine Cd toxicity threshold and the physiological mechanisms involved in Cd tolerance. An experiment was conducted to investigate the effect of cadmium from 0 to 1350 μM on chlorophyll fluorescence parameters, gas exchange, photosynthetic pigment concentrations and antioxidative enzyme activities of A. halimus. Cadmium, calcium, iron, manganese, magnesium, potassium, phosphorous, sodium and zinc concentrations were also analyzed. Plants of A. halimus were not able to survive at 1350 μM Cd and the upper tolerance limit was recorded at 650 μM Cd; although chlorosis was observed from 200 μM Cd. Cadmium accumulation increased with increase in Cd supply, reaching maxima of 0.77 and 4.65 mg g(-1) dry weight in shoots and roots, respectively, at 650 μM Cd. Dry mass, shoot length, specific leaf area, relative growth rate, net photosynthetic rate, stomatal conductance, pigments contents and chlorophyll fluorescence were significantly reduced by increasing Cd concentration. However, the activities of superoxide dismutase (SOD; EC1.15.1.1), catalase (CAT; EC1.11.1.6) and guaiacol peroxidase (GPx; EC1.11.1.7) were significantly induced by Cd. Exposures to Cd caused also a significant decrease in P contents in roots, Mg and Mn contents in shoots and Fe and K contents in roots and shoots and had no effect on Ca, Na and Zn contents. The tolerance of A. halimus to Cd stress might be related with its capacity to avoid the translocation of great amounts of Cd in its aboveground tissues and higher activities of enzymatic antioxidants in the leaf., (Copyright © 2016 Elsevier Masson SAS. All rights reserved.)