. 2023 Aug 3;12(15):2861. doi: 10.3390/plants12152861

Table 2.

Reported effect of phosphorus on plant tolerance to drought, salinity, temperature, high CO₂ and heavy metal stresses.

Abiotic Stresses	Plant	P application	Alleviation Mechanisms	References
Drought	Echinacea purpurea	Split plot experiment was performed with drought stress in term of available water depletion (25, 50 and 75%) to retain the filed capacity. P fertilizer was used as control without P fertilizer, 100% plant required P from triple super-phosphate, sole application of mycorrhizal arbuscular fungi (AMF) and Pseudomonas fluorescens bacteria (PFB) and AMF or PFB combination with 50% of the plant’s P requirement.	Application of P increased root biomass and yield by 57% and 47%, respectively, under drought conditions. In 25% of AWD, the highest root cynarin (0.583 mg/g dry matter) was observed in the joint application of phosphorus + AMF. Together with the AMF, P improved the drought tolerance traits.	[136]
Water deficit	Rapeseed	Plants were grown under different irrigation levels (70-, 100-, 130- and 160-mm evaporation from class A pan, respectively), which were treated with five fertilizers (control, chemical fertilizer including N and P (about 300 and 150 kg ha⁻¹, respectively, based on soil analysis)).	P fertilizers especially in combination with other fertilizers decreased proline content and leaf temperature, with an increase in antioxidants and enzymatic activities including chlorophyll content, leaf water content, membrane stability index and stomatal conductance with improved yield.	[137]
Drought	Alnus cremastogyne	P fertilizer was applied in the form of NaH₂PO₄ (25.5% P) to each of the pots in each 30-day interval 3 times in the entire crop duration.	Application of P improved the relative water content, photosynthesis rate, increase in antioxidative enzymes including SOD, CAT, POD, osmolytes accumulation, soluble proteins and decrease in lipid peroxidation levels.	[138]
Drought	Eucalyptus grandis	P fertilizer was applied in the form of NaH₂PO₄ (25.5% P) to each of the pots in each 30-day interval 3 times in the entire crop duration.	P application alleviated the drought stress effect by improving the leaf relative water content, net photosynthesis, quantum efficiency of photosystem II and amelioration in some other physiological traits related to drought stress tolerance.	[139]
Drought	Phoebe zhennan	P fertilizer was applied in the form of NaH₂PO₄ (25.5% P) to each of the pots in each 30-day interval.	P alleviated the drought effect by increasing the relative water content, net photosynthesis rate, higher quantum efficiency, higher rooting systems, enhanced root biomass, decrease in MDA and upregulations of photosynthetic pigments, osmolytes and nitrogenous compounds.	[140]
Drought	Pisum sativum	P was applied in the form of KH₂PO₄ at two different rates: 15 (P15) and 60 mg P kg⁻¹ (P60), mixed evenly throughout the soil.	Optimal P application enhanced the water use efficiency, soluble sugars and relative water content. Application of P increased the salt tolerance index with higher root length, nodule number, stomatal conductance and uptake of nitrogen.	[14]
Drought	Soybean	The P levels (added as KH₂PO₄) were 0, 15 and 30 mg P kg⁻¹ soil.	Addition of P enhanced the concentration and accumulation of nitrogen in shoots and seeds. P application mitigated the drought effects on plant by increasing the protein concentration.	[141]
Salinity	Wheat	P was used in the form of Ortho P fertilizer, phosphoric acid-based fertilizers with K and N containing 52% and 62% of P2O5, respectively, with 100% orthoP for each one.	Sufficient P application alleviated the salinity stress by promoting growth and reducing salt toxicity. Fertilized plants have higher shoot and root dry weights under salinity stress.	[142]
Salinity	Quinoa	Filed experiment was set up with different EC levels of irrigation water (5, 12 and 17 dS·m⁻¹). P was added in the form of P2O5 at the rate of 0, 60 and 70 kg of P₂O₅ ha⁻¹.	P application under saline conditions minimized the effect of salinity and improved the yield with higher water and nutrient uptake. The results suggested that P application minimizes the adverse effects of high soil salinity and can be adopted as a coping strategy under saline conditions.	[143]
Salinity	Sugar beet	Plants were grown under salinity water with different EC values of 0.7, 4, 8 and 12 dS·m⁻¹). P was applied in the form of P2O5 with a rate of 100, 120 and 140 kg P₂O₅·ha⁻¹.	P improved the rate of yield and sugar content of sugar beets under the tested salinity levels.	[144]
Salinity	Aeluropus littoralis	Plants were grown at moderate salinity (100 mM NaCl). P fertilizers were applied at different doses: low, moderate and high (5 mM, 60 mM and 180 mM KH₂PO₄).	P fertilization improved the salinity tolerant characteristics including increase in leaf hair and trichome densities, total polyphenol content and total antioxidant capacity in plants cultivated.	[145]
Salinity	Maize	P was introduced to the nutrient medium in the form of KH₂PO₄, with concentrations of 5 μM considered as low P and 200 μM as high P.	Higher P application prevented leaf chlorosis under salinity stress with improved	[146]
Salinity	Woody species	Plants were grown in a factorial 2 × 2 × 2 (presence or absence of salt, AMF, or P_i), totaling eight treatments: control, P_i, AMF, AMF ∗ P_i, salt, salt ∗ P_i, salt ∗ AMF and salt ∗ AMF ∗ P_i, with eight replicates each and one plant per pot, totaling 64 experimental units.	P fertilization increased biomass and photosynthetic pigments under salinity conditions. Metabolites were also positively impacted due to fertilization.	[147]
Salinity	Phaseolus vulgaris	Plants were grown under different salinity conditions (1.56, 4.78 and 8.83 dS·m⁻¹) and P fertilizer at the rate of 0, 30, 60 and 90 kg ha⁻¹.	P application significantly increased the total chlorophyll content, total soluble sugars, carotenoids, total free amino acid and proline with higher accumulation of K⁺, Ca²⁺, Mg²⁺ and higher tolerance and yield.	[148]
Salinity	Okra	Plants were grown under two different salinity conditions (4 and 8 dS·m⁻¹) and combined with different rates of P fertilizers (0, 30, 60 and 90 kg ha⁻¹ from triple super phosphate).	Application of P increased the green and dry pod yield under salinity stress.	[149]
Salinity	Cicer arietinum L.	Plants were grown under two salinity conditions (0 and 150 mM NaCl) and treated with three P fertilizers (0, 90, 200 kg h⁻¹ of P₂O₅ in the form of super triple phosphate).	P application improved plant growth. P fertilization increased the leghemoglobin (92%), reduced proline content (−69%) and protected membranes against peroxidation compared to saline conditions. Also, yield was increased due to the P fertilizers.	[150]
Low temperatures	Wheat	Pot experiment was conducted with tolerant and sensitive cultivar and was treated under chilling (T1 at 4 °C) and freezing treatment (T2 at −4 °C) as well as ambient temperature (CK at 11 °C) during the anther differentiation treated with P fertilizers.	Application of P alleviated low-temperature stress by increasing the stomatal conductance, dry matter accumulation, transportation of assimilates, grain number per spike, 1000 grain weights, yield per plant.	[151]
Low and high temperatures	Soybean	Plants were grown at different temperatures (22, 26, 30 and 34 °C corresponds to moderately low, optimum, moderately high and high temperature) with combination of 0.5 mM and 0.08 mM P nutrition.	Sufficient P fertilization improved the temperature stress tolerance by increasing the plant efficiency in utilization and biomass partitioning to pods.	[152]
High-temperature stress	Rice	Plants were grown in controlled growth conditions with high day and night temperatures (35 °C ± 2 and 32 °C ± 2, respectively). P fertilizers were added singly or combined with biochar.	P fertilization ameliorated the adverse impact of high temperatures with higher grain formation and grain quality.	[153]
Heavy metals	-	Five different types of treatments were added: control (C), heavy metal pollution (H), heavy metal pollution + nitrogen (HN), heavy metal pollution + phosphorus (HP), heavy metal pollution + nitrogen and phosphorus (HNP)	P addition of HP and HNP treatments restored plant species richness and increased plant diversity under heavy metal pollution. P addition had a better performance in restoring the species composition and relative dominance of plant communities.	[15]
Heavy metals	Maize	Maize plants were grown in soil collected from paddy field located near recycle area and fertilizers were applied as two amendments (calcium–magnesium phosphate fertilizer, PF; commercial organic fertilizer, OF) tested in single and in combination.	Application of P increased the maize shoot and root biomass and enzymatic activities such as urease and catalase. Additionally, microbial community structure was improved.	[154]
Heavy metals	Lespedeza bicolor; Lespedeza cuneata	Two species, Lespedeza bicolor and L. cuneata, were grown for 30 d with alternate Al and P treatments in a hydroponics system.	P application improved root growth and heavy metal tolerance by lowering the Al uptake and accumulation. Enhancement of Al resistance by P in the resistant species might be associated with its more efficient P accumulation and translocation to shoots and greater Al exclusion from root tips after P application.	[155]
Waterlogging	Brachlaria grass	Tolerant and sensitive cultivars were grown under control and waterlogged conditions with two different fertilizer levels (low and high).	Higher availability of P under waterlogged soil imparted the tolerance.	[156]
Drought and elevated CO₂	Field pea (Pisum sativum)	Plants were grown in P-deficient vertisol, supplied with two doses of P (15 mg and 60 mg Kg⁻¹) and treated with ambient and elevated CO₂ (380–400 ppm and 550–580 ppm).	P improved water use efficiency under elevated CO₂ levels.	[14]