Osmotic Pressure Research Articles

On the laminar solutions and stability of accelerating and decelerating channel flows

We study the effect of acceleration and deceleration on the stability of channel flows. To do so, we derive an exact solution for laminar profiles of channel flows with an arbitrary, time-varying wall motion and pressure gradient. This solution then allows us to investigate the stability of any unsteady channel flow. In particular, we restrict our investigation to the non-normal growth of perturbations about time-varying base flows with exponentially decaying acceleration and deceleration, with comparisons to growth about a constant base flow (i.e. the time-invariant simple shear or parabolic profile). We apply this acceleration and deceleration through the velocity of the walls and through the flow rate. For accelerating base flows, perturbations never grow larger than perturbations about a constant base flow, while decelerating flows show massive amplification of perturbations – at a Reynolds number of $500$ , properly timed perturbations about the decelerating base flow grow $ {O}(10^5)$ times larger than perturbations grow about a constant base flow. This amplification increases as we raise the rate of deceleration and the Reynolds number. We find that this amplification arises due to a transition from spanwise perturbations leading to the largest amplification to streamwise perturbations leading to the largest amplification that only occurs in the decelerating base flow. By evolving the optimal perturbations through the linearized equations of motion, we reveal that the decelerating base flow achieves this massive amplification through the Orr mechanism, or the down-gradient Reynolds stress mechanism, which accelerating and constant base flows cannot maintain.

Hypertonic water reabsorption with a parallel-current system via the glandular and saccular renal tubules of Ruditapes philippinarum.

We investigated the renal function of the brackish water clam, Ruditapes philippinarum, employing magnetic resonance imaging (MRI). The R. philippinarum kidney consists of two renal tubules, a glandular (GT) and a saccular (ST) tubule. After exposure to seawater containing manganese ion (Mn2+) at 20°C, the intensity of the T1-weighted MRI and longitudinal relaxation rates (1/T1=R1) of the kidney were increased. In the ST, haemolymph containing Mn2+ entered directly from the auricle, and the Mn2+ concentration ([Mn2+]) increased in the initial part of the ST. Thereafter, [Mn2+] was almost constant until the posterior end of the kidney. The GT received haemolymph from the pedal sinus via the visceral sinus. The GT runs parallel inside the ST, and [Mn2+] increased progressively until it merged with the ST. In a range of seawater with [Mn2+] from 1 to 30 µmoll-1, the [Mn2+] increased 12-fold in the posterior part of the ST, compared with the ambient [Mn2+]. Based on these results, the epithelium of the initial part of the ST reabsorbs water from luminal fluid, building up a higher osmotic pressure. Using this osmotic gradient, hypertonic water is reabsorbed via the epithelium of the GT to the ST, and then transferred to the haemolymph via the epithelium of the ST. Excess water is excreted as urine. This model was supported by the increases in [Mn2+] in the ST when the clams were exposed to seawater containing Mn2+ at salinity from 26.0 to 36.0‰, showing that the parallel-current system works in hypotonic seawater.

Wall pressure control of a 3D cavity with lateral apertures and wall proximity

Wall pressure control of a 3D cavity with lateral apertures and wall proximity

Salicylic acid priming before cadmium exposure increases wheat growth but does not uniformly reverse cadmium effects on membrane glycerolipids.

Cadmium (Cd) is an abiotic stressor negatively affecting plant growth and reducing crop productivity. The effects of Cd (25 μM) and of pre-soaking seeds with salicylic acid (SA) (500 μM) on morphological, physiological, and glycerolipid changes in two cultivars of wheat (Triticum aestivum L. 'Tosunbey' and 'Cumhuriyet') were explored. Parameters measured were length, fresh and dry biomass, Cd concentration, osmotic potential (ψ), lipid peroxidation, and polar lipid species in roots and leaves, as well as leaf chlorophyll a, carotenoids, and fv/fm. Fresh biomass of roots and leaves and leaf length were strongly depressed by Cd treatment compared to the control, but significantly increased with SA + Cd compared to Cd alone. Cd reduced leaf levels of chlorophyll a, carotenoids, and fv/fm, compared to controls. Treatment with SA + Cd increased pigment levels and fv/fm compared to Cd alone. Cd treatment led to a decrease in DW of total membrane lipids in leaves and depressed levels of monogalactosyldiacylglycerol and phosphatidic acid in leaves and roots of both cultivars. The effects of SA priming and SA + Cd treatment on lipid content and composition were cultivar-specific, suggesting that lipid metabolism may not be a primary target underlying SA remediation of the damaging effects of Cd on wheat growth and development.

The confined stresslet for suspensions in a spherical cavity. Part 1. Traceless elements

The confined Stokesian dynamics (CSD) algorithm recently reported equilibrium properties but was missing hydrodynamic functions required for suspension stress and non-equilibrium properties. In this first of a two-part series, we expand the CSD algorithm to model the traceless part of the stress tensor. To obtain quantities needed to solve the integral expressions for the stress, we developed a general method to solve Stokes’ equations in bispherical coordinates. We calculate the traceless stress tensor for arbitrary particle-to-enclosure size ratio. We next compute rheology of a confined suspension by implementing the stresslet hydrodynamic coefficients into CSD, yielding the deviatoric part of the many-body hydrodynamic stresslet. We employed energy methods to relate this stresslet to the high-frequency viscosity of the confined suspension, finding an increase in viscous dissipation with crowding and confinement well beyond the unconfined value. We show that confinement effects on viscosity are dominated by near-field interactions between the particles that reside very near the cavity wall (rather than particle–wall interactions). Surprisingly, this near-field effect is stronger than the viscosity of an unconfined suspension, showing that entropic exclusion driven by the wall sets up many lubrication interactions that then generate strong viscous dissipation. The limiting case of a particle near a flat wall reveals a correction to prior literature. The theory presented in this work can be expanded to study the Brownian contribution to the viscosity of confined suspensions in and away from equilibrium. In part 2, we report the osmotic pressure, via the trace of the stress tensor.

Generalized Effective Stress of Dual-Porosity Geomaterials Considering Thermochemical Effects

The accurate calculating of effective stress is crucial to predicting soil deformation and ensuring structural safety under complex mechanical, thermal, and chemical loads. In this study, the thermodynamic pressures of the solid phase, and liquid phases in macropores and micropores of dual-porosity geomaterials are determined on the basis of thermodynamics and conservation equations with considering for the impact of temperature and solute concentration on soil-water interactions. In particular, thermal pressurization related to temperature, and generalized osmotic pressure related to solute concentration were incorporated into the liquid-phase thermodynamic pressure equations. Thereafter, an equation for the generalized effective stress that could be applied specifically to dual-porosity geotechnical materials with consideration for environmental loads was derived from the definition of the stress tensor. The deformation behavior of clay under mechanical, thermal, and chemical coupling loading conditions was predicted using the newly proposed generalized effective stress. Predicted results revealed a stronger correlation between the generalized effective stress and void ratio, thereby confirming the accuracy and effectiveness of the proposed approach. The derived equation establishes a solid theoretical foundation for designing impermeable buffer barriers in geotechnical engineering and geo-environmental engineering safety assessments.

Bacterial-host adhesion dominated by collagen subtypes remodelled by osmotic pressure.

Environmental osmolarity plays a crucial role in regulating the functions and behaviors of both host cells and pathogens. However, it remains unclear whether and how environmental osmotic stimuli modulate bacterial‒host interfacial adhesion. Using single-cell force spectroscopy, we revealed that the interfacial adhesion force depended nonlinearly on the osmotic prestimulation of host cells but not bacteria. Quantitatively, the adhesion force increased dramatically from 25.98 nN under isotonic conditions to 112.45 or 93.10 nN after the host cells were treated with the hypotonic or hypertonic solution. There was a strong correlation between the adhesion force and the number of host cells harboring adherent/internalized bacteria. We further revealed that enhanced overexpression levels of collagen XV and II were responsible for the increases in interfacial adhesion under hypotonic and hypertonic conditions, respectively. This work provides new opportunities for developing host-directed antibacterial strategies related to interfacial adhesion from a mechanobiological perspective.

Numerical investigation of noise reduction of a pump-jet propulsor using porous metal

A novel strategy for noise reduction in pump-jet propulsor (PJP) is proposed through the design of porous metal trailing edges on stator blades. The interference between stator wake vortices and rotor blades is one of the primary mechanisms for noise generation in PJP. The goal is to use a porous medium to diminish the intensity of turbulent vortices behind the stator blades, control the interaction between these shedding vortices and the rotor blades downstream of the stator, and ultimately suppress the rotor noise generation source. Large eddy simulation and acoustic analogy analysis of the entire propulsion system based on the Ffowcs-Williams and Hawkings equation are performed. It is found that with the use of porous stator trailing edges, the wake flow pattern of the stator is notably altered, the strong impact of the turbulent vortices on the rotor blades is weakened, and the wall pressure fluctuations on the rotor blades and duct are significantly reduced. As a result, the far-field radiation noise and tonal noise are both effectively reduced, with the maximum reduction in total and tonal sound pressure levels reaching 3.8 dB and 12.53 dB, respectively, corresponding to percentage reductions of 6.0% and 21.69%. These findings demonstrate that porous media have great potential for practical engineering applications in PJP noise control.

Viscosity regulates cell spreading and cell-extracellular matrix interactions.

Fluid viscosity and osmolarity are among some of the underappreciated mechanical stimuli that cells can detect. Abnormal changes of multiple fluidic factors such as viscosity and osmolarity have been linked with diseases such as cystic fibrosis, cancer, and coronary heart disease. Changes in viscosity have been recently suggested as a regulator of cell locomotion. These novel studies focus on cell migration and spreading on glass substrates and through microchannels, and it remains a question whether viscosity impacts the cellular remodeling of extracellular matrices (ECMs). Here, we demonstrate that elevated viscosity induces cellular remodeling of collagen substrates and enhances cell spreading on ECM-mimetic substrates. Our results expand on recent work showing that viscosity induces increased cellular forces and demonstrates that viscosity can drive local ECM densification. Our data further show that microtubules, Ras-related C3 botulinum toxin substrate 1 (Rac1), actin-related protein 2/3 (Arp2/3) complex, Rho-associated protein kinase 1 (ROCK), and myosin are important regulators of viscosity-induced ECM remodeling. In the context of viscosity-induced cell spreading, cells cultured on glass and collagen substrates exhibit markedly different responses to pharmacological treatments, indicating that microtubules, Rac1, and Arp2/3 play distinct roles in regulating cellular spreading depending on the substrate. In addition, our results demonstrate that high osmotic pressures override viscosity-induced cell spreading by suppressing membrane ruffling. Our results demonstrate viscosity as a regulator of ECM remodeling and cell spreading in a fibrillar microenvironment. We also reveal a complex interplay between viscosity and osmolarity. We anticipate that our research can pave the way for future investigations into the crucial roles played by viscosity in both physiological and pathological conditions.

Computational Fluid Dynamics Modeling of Pressure-Retarded Osmosis: Towards a Virtual Lab for Osmotic-Driven Process Simulations

Pressure-Retarded Osmosis (PRO) is an osmotically driven membrane-based process that has recently garnered significant attention from researchers due to its potential for clean energy harvesting from salinity gradients. The complex interactions between mixed-mode channel flows and osmotic fluxes in real PRO membrane modules necessitate high-fidelity modeling approaches. In this work, an efficient CFD framework is developed for the 3D simulation of osmotically driven membrane processes. This approach is based on a two-way coupling between a CFD solver, which captures external concentration polarization (ECP) effects, and an analytical representation of internal concentration polarization (ICP). Consequently, the osmotic water flux and reverse salt flux (RSF) can be accurately determined, accounting for all CP effects without any limitations on the geometrical complexity of the membrane chamber or its flow mode/regime. The proposed model is validated against experimental data, showing good agreement across various PRO case studies. Additionally, the model’s flexibility to simulate other types of osmotically driven processes such as forward osmosis (FO) is examined. Thus, the contributions of ECP and ICP effects in local osmotic pressure drop along the membrane chamber are comprehensively compared for FO and PRO modes. Finally, the capability of the CFD model to simulate a lab-scale PRO module is demonstrated across a range of Reynolds numbers from low-speed laminar up to turbulent flow regimes.

Prediction of splash noise in a rectangular tank under longitudinal periodic excitation

Sloshing phenomena within fuel tanks have emerged as a significant contributor to noise in hybrid and high-end vehicles, particularly as other sources of noise, such as those from the engine and transmission system, minimized. The manifestation of noise during sloshing results from the dynamic interplay between the fluid within the tank and both its surrounding structures and the fluid itself. “Splash noise” is an acoustic phenomenon which arises due to pressure fluctuations induced by fluid-fluid interactions during sloshing. Thus, the generation of splash noise encompasses a multi-physics process involving fluid flow and acoustic radiation. Accurate prediction of splash noise is crucial for implementing measures to mitigate it during the design phase. The current paper proposes a hybrid approach for predicting splash noise within a rectangular tank. The first step in the methodology is to predict the flow field within the tank, from which acoustic sources are computed. These sources are then utilized to calculate the sound propagation based on the acoustic analogy. To emulate a regime dominated by fluid-fluid interactions, periodic longitudinal excitations are imposed on the rectangular tank, both with and without baffles. Parameters such as fluid free-surface profile, tank wall pressures, and radiated sound pressure levels are systematically calculated and subsequently validated against experimental results available in the existing literature.

Instability of isolator shocks to fuel flow rate modulations in a strut-stabilised scramjet combustor

Abstract Reynolds-Averaged Navier–Stokes (RANS) simulations, both steady and unsteady, are used to investigate supersonic, chemically reacting, flow fields inside a strut-stabilised supersonic combustion ramjet (scramjet) engine operating under different fuel flow rates. Fully supersonic, fully subsonic and mixed modes of operations inside the combustor, obtained at different fuel flow rates, are studied numerically through shock wave visualisations and top-wall static-pressure probing. The effect of changing fuel flow rates, imposed both suddenly and gradually, on the behaviour of shock waves and wall pressure profiles are studied in detail. For certain modes of combustion characterised by the presence of oblique shocks at the strut, shockwaves in the combustor respond predictably to an increase or decrease in fuel flow rate attaining the steady state flow fields as predicted by RANS simulations for those fuel flow rates. For certain other modes of combustion, characterised by the presence of shockwaves in the isolator and the absence of oblique shocks at the leading edge of the strut, shockwaves in the flow field appear unstable to fuel flow rate modulations. For such cases, any change in fuel flow rates, sudden or gradual, increase or decrease, causes the isolator shocks to immediately move upstream and eventually out of the isolator. A plausible physics-based explanation of the observed phenomena is presented.

A simple and cost-effective real-time PCR method using diluted and heat-treated whole blood lysate

Although DNA extraction is a crucial step before polymerase chain reaction (PCR), attempts to perform PCR without DNA isolation have been ongoing since the 1990s. While partial success has been achieved with direct conventional PCR, the DNA isolation step has remained indispensable for real-time PCR. Here, we developed a method that does not require complete DNA isolation by lysing EDTA-treated whole blood samples through the application of osmotic pressure and heat, followed by centrifugation to get a clear lysate. Blood samples were mixed with distilled water, incubated at 95 °C for 20 min before centrifugation at 14,000 rpm for 5 min. The resulting lysates were used as templates for real-time PCR. Real-time PCRs were performed under identical conditions with lysates and DNA samples using 9 different primer sets. Target genes were successfully amplified using 1:10 and 1:5 diluted blood lysates both at 60 °C and 61 °C. The PCR efficiency for ACTB and PIK3CA differed by 20% and 14%, respectively, between the DNA samples and blood lysates. We successfully amplified all selected genes using “GG-RT PCR”, greater temperature, greater speed method, without the need for additional enzymes or buffers. GG-RT PCR presents a cost effective option for applications such as SNP analysis and deletion detection if appropriate primer designs are made.

Astrocyte aquaporin mediates a tonic water efflux maintaining brain homeostasis.

Brain water homeostasis not only provides a physical protection, but also determines the diffusion of chemical molecules key for information processing and metabolic stability. As a major type of glia in brain parenchyma, astrocytes are the dominant cell type expressing aquaporin water channel. How astrocyte aquaporin contributes to brain water homeostasis in basal physiology remains to be understood. We report that astrocyte aquaporin 4 (AQP4) mediates a tonic water efflux in basal conditions. Acute inhibition of astrocyte AQP4 leads to intracellular water accumulation as optically resolved by fluorescence-translated imaging in acute brain slices, and in vivo by fiber photometry in mobile mice. We then show that aquaporin-mediated constant water efflux maintains astrocyte volume and osmotic equilibrium, astrocyte and neuron Ca2+ signaling, and extracellular space remodeling during optogenetically induced cortical spreading depression. Using diffusion-weighted magnetic resonance imaging (DW-MRI), we observed that in vivo inhibition of AQP4 water efflux heterogeneously disturbs brain water homeostasis in a region-dependent manner. Our data suggest that astrocyte aquaporin, though bidirectional in nature, mediates a tonic water outflow to sustain cellular and environmental equilibrium in brain parenchyma.

Astrocyte aquaporin mediates a tonic water efflux maintaining brain homeostasis

Brain water homeostasis not only provides a physical protection, but also determines the diffusion of chemical molecules key for information processing and metabolic stability. As a major type of glia in brain parenchyma, astrocytes are the dominant cell type expressing aquaporin water channel. How astrocyte aquaporin contributes to brain water homeostasis in basal physiology remains to be understood. We report that astrocyte aquaporin 4 (AQP4) mediates a tonic water efflux in basal conditions. Acute inhibition of astrocyte AQP4 leads to intracellular water accumulation as optically resolved by fluorescence-translated imaging in acute brain slices, and in vivo by fiber photometry in mobile mice. We then show that aquaporin-mediated constant water efflux maintains astrocyte volume and osmotic equilibrium, astrocyte and neuron Ca2+ signaling, and extracellular space remodeling during optogenetically induced cortical spreading depression. Using diffusion-weighted magnetic resonance imaging (DW-MRI), we observed that in vivo inhibition of AQP4 water efflux heterogeneously disturbs brain water homeostasis in a region-dependent manner. Our data suggest that astrocyte aquaporin, though bidirectional in nature, mediates a tonic water outflow to sustain cellular and environmental equilibrium in brain parenchyma.

New insights into a sensitive life stage: hydraulics of tree seedlings in their first growing season.

The first year in a tree's life is characterized by distinct morphological changes, requiring constant adjustments of the hydraulic system. Despite their importance for the natural regeneration of forests and future vegetation composition, little has been known about the hydraulics of tree seedlings. At different times across the first growing season, we analysed xylem area-specific (Kshoot_Axyl) and leaf area-specific (Kshoot_L) shoot hydraulic conductance, as well as embolism resistance of three temperate conifer trees, two angiosperm trees and one angiosperm shrub, and related findings to cell osmotic parameters and xylem anatomical traits. Over the first 10 wk after germination, Kshoot_Axyl and Kshoot_L sharply decreased, then remained stable until the end of the growing season. Embolism resistance was remarkably low in the youngest stages but, coupled with an increase in cell wall reinforcement, significantly increased towards autumn. Contemporaneously, water potential at turgor loss and osmotic potential at saturation decreased. Independent of lineage, species and growth form, the transition from primary to secondary xylem resulted in a less efficient but increasingly more embolism-resistant hydraulic system, enabling stable water supply under increasing risk for low water potentials.

Effect of environmental factors on seed germination and seedling emergence of weedy rice (Oryza sativa f. spontanea) in China

Abstract Weedy rice (Oryza sativa f. spontanea Auct. ex Backer) is a troublesome annual weed from the Gramineae family and infests rice (Oryza sativa L.) fields globally, with a notable presence in China. However, limited information is available regarding the effects of diverse environmental factors on its germination and emergence. A better understanding of the seed biology and ecology of weedy rice is crucial for developing effective weed management strategies. Experiments were conducted to evaluate the effects of temperature, light, soil burial depth, wheat (Triticum aestivum L.) crop residue amount, salt stress, osmotic stress, and radiant heat on the germination and seedling emergence of weedy rice. Weedy rice exhibited robust germination (>98%) when exposed to varying day/night temperatures (20/15 to 35/30 C) and remained unaffected by light conditions. Seedling emergence was not influenced within the top 5-cm soil layer, where 100% of the seedlings emerged. However, emergence decreased as the soil burial depth increased, eventually resulting in no emergence from a burial depth of 11 cm. The soil burial depth required for 50% of the maximum emergence was 8.3 cm. Seedling emergence ranged from 97% to 100% across different amounts of the wheat straw residue cover (0 to 10,000 kg ha−1). The sodium chloride concentration and osmotic potential required for 50% were 230.8 mM and −0.5 MPa, respectively. No germination was observed when weedy rice seeds were exposed to >110 C pretreatment (radiant heat for 5 min), indicating that residue burning could reduce infestation of weedy rice. The insights gained from this study contribute valuable knowledge to enhance the integrated management of weedy rice in China.

Compatibility of dry incubator on in vitro production of bovine embryos

Embryo culture is crucial to achieve successful outcomes in in vitro production-embryo transfer for cattle. This study explored the innovative use of dry incubators for bovine embryo culture, building on their advantages in human medicine, such as reduced contamination risk, stable temperature control, and lower gas consumption. In this study, we examined changes in osmotic pressure, the in vitro developmental potential of IVP embryos including the cleavage rate, blastocyst development rate, blastocyst diameter, and blastocyst cell number, morphokinetics, and the transcriptional profile of the blastocysts between humidified and dry incubators. Our research demonstrates the feasibility of this approach, showing that although the osmotic pressure gradually increases over the culture period (on day 8: 271.7 vs. 299.0, respectively; P < 0.09), it did not negatively affect the blastocyst formation rate (62.4% vs. 69.8%) and the morphological quality of blastocysts (diameter: 237.4 vs. 242.8, total cell number: 189.2 vs. 242.8). Embryos cultured in dry incubators exhibited morphokinetics comparable to those cultured in conventional humidified incubators. Furthermore, RNA-seq revealed that while a few genes showed changes, the transcriptomic profiles of blastocysts cultured in dry incubators were largely similar to those of blastocysts cultured in humidified incubators. These findings highlight the considerable potential of dry incubators for the in vitro production of bovine embryos.

Hidden Water's Influence on Rhodopsin Activation.

Structural biology relies on several powerful techniques, but these tend to be limited in their ability to characterize protein fluctuations and mobility. Over-reliance on structural approaches can lead to omission of critical information regarding biological function. Currently there is a need for complementary biophysical methods to visualize these mobile aspects of protein function. Here we review hydrostatic and osmotic pressure-based techniques to address this shortcoming for the paradigm of rhodopsin. Hydrostatic and osmotic pressure data contribute important examples which are interpreted in terms of an energy landscape for hydration-mediated protein dynamics. We find that perturbations of rhodopsin conformational equilibria by force-based methods are not unrelated phenomena; rather they probe various hydration states involving functional proton reactions. Hydrostatic pressure acts on small numbers of strongly interacting structural or solvent-shell water molecules with relatively high energies, while osmotic pressure acts on large numbers of weakly interacting bulk-like water molecules with low energies. Local solvent fluctuations due to the hydration shell and collective water interactions affect hydrogen-bonded networks and domain motions that are explained by a hierarchical energy landscape model for protein dynamics.

Melatonin application enhances salt stress-induced decreases in minerals, betalains, and phenolic acids in beet (Beta vulgaris L.) cultivars.

Melatonin is a potentially active signaling molecule and plays a crucial role in regulating the growth and development of plants under stress conditions, alleviating oxidative damage, enhancing antioxidant defence mechanisms and regulating ion homeostasis. This study examined the effects of exogenous melatonin application on leaf biomass, ion concentrations, betalains, phenolic acid and endogenous melatonin contents comparing red beet (Beta vulgaris L. 'Ruby Queen' and 'Scarlet Supreme') and white beet ('Rodeo' and 'Ansa') cultivars under increasing salinity levels of 50, 150, and 250 mM NaCl. Exogenous melatonin increased salinity-induced reductions in fresh and dry weights and osmotic potential in leaves. Na+ concentrations rose significantly with increasing salinity, but cultivar-specific decreases were observed in K+ and Ca2+ concentrations. Additionally, melatonin application improved betalain, betanin and neobetanin contents induced by salt stress. Furthermore, melatonin application caused salt stress and cultivar-specific changes in phenolic acid contents e.g., ferulic acid, sinapic acid, or m-coumaric acid, in soluble free, ester- and glycoside-conjugated and cell wall-bound forms. In addition, antioxidant enzyme activities and compound contents increased significantly in the beets and were subsequently lowered in a cultivar-specific manner by salt stress + melatonin treatment. The current findings indicate that exogenous melatonin improved plant stress tolerance suppressing reactive oxygen species levels, increasing the antioxidant enzyme activities and compound contents and reducing the levels of Na+, maintaining an ionic homeostasis in the selected red and white sugar beet cultivars. It appears that melatonin application may help improve cultivar-specific salt tolerance by enhancing ion homeostasis and betalain and phenolic acid production levels in beets.