Effect of Salt Stress on Different Parts of Plants Review

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Peer-reviewed

Enquiry Article

Effects of salt stress on the photosynthetic physiology and mineral ion assimilation and distribution in white willow (Salix alba 50.)

• Xin Ran,
• Xiao Wang,
• Xiaokuan Gao,
• Haiyong Liang,
• Bingxiang Liu,
• Xiaoxi Huang

x

• Published: November 18, 2021
• https://doi.org/10.1371/journal.pone.0260086

Figures

Abstract

Objective

The purpose of this study was to explore the adaptive mechanism underlying the photosynthetic characteristics and the ion absorption and distribution of white willow (Salix alba L.) in a common salt stress environment in cut seedlings. The results lay a foundation for further understanding the distribution of sodium chloride and its effect on the photosynthetic organization.

Method

A table salt stress environment was false in a hydroponics organization with dissimilar NaCl concentrations in one-year-sometime Salix alba 50.branches every bit the examination materials. Their growth, ion absorption, transport and distribution in the roots and leaves, and the changes in the photosynthetic fluorescence parameters were studied after 20 days under hydroponics.

Results

The results show that The formation and elongation of roots are promoted in the presence of 171mM NaCl, but root growth is comprehensively inhibited under increasing salt stress. Under salt stress, Na⁺ accumulates significantly in the roots and leaves, and the Na⁺ content and the Na⁺/Thou⁺ and Na⁺/Ca²⁺ root ratios are significantly greater than those in the leaves. When the NaCl concentration is ≤ 342mM, Salix alba can maintain relatively stable K⁺ and Caⁱⁱ⁺ contents in its leaves by improving the selective absorption and accumulation of K⁺ and Ca^two+ and adjusting the transport capacity of mineral ions to aboveground parts, while One thousand⁺ and Ca²⁺ levels are clearly decreased under high table salt stress. With increasing salt concentrations, the cyberspace photosynthetic charge per unit (P_{due north}), transpiration rate (Eastward) and stomatal conductance (k_south) of leaves decrease gradually overall, and the intercellular CO₂ concentration (C_i) first decreases and so increases. When the NaCl concentration is < 342mM, the decrease in foliage P_n is primarily restricted by the stomata. When the NaCl concentration is > 342mM, the decrease in the P_n is largely inhibited by non-stomatal factors. Due to the common salt stress environs, the OJIP bend (Rapid chlorophyll fluorescence) of Salix alba turns into an OKJIP curve. When the NaCl concentration is > 171mM, the fluorescence values of points I and P subtract significantly, which is accompanied by a clear inflection point (1000). The quantum yield and energy distribution ratio of the PSⅡ reaction center change significantly (φPo, Ψo and φEo show an overall downward trend while φDo is promoted). The functioning index and driving force (PI_ABS, PI_CSm and DF_CSm) decrease significantly when the NaCl concentration is > 171mM, indicating that table salt stress causes a fractional inactivation of the PSII reaction center, and the functions of the donor side and the recipient side are damaged.

Conclusion

The in a higher place results bespeak that Salix alba can respond to salt stress by intercepting Na⁺ in the roots, improving the selective assimilation of M⁺ and Ca²⁺ and the transport chapters to the above ground parts of the plant, and increasing φDo, thus shows an ability to self-regulate and arrange.

Citation: Ran X, Wang X, Gao Ten, Liang H, Liu B, Huang X (2021) Furnishings of salt stress on the photosynthetic physiology and mineral ion absorption and distribution in white willow (Salix alba L.). PLoS Ane 16(11): e0260086. https://doi.org/10.1371/journal.pone.0260086

Editor: Mayank Gururani, United Arab Emirates University, UNITED ARAB EMIRATES

Received: July 16, 2021; Accepted: November 3, 2021; Published: November 18, 2021

Copyright: © 2021 Ran et al. This is an open admission article distributed nether the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

Information Availability: All relevant data are within the paper and its S1 Appendix files.

Funding: This written report was jointly supported past grants from the Natural Science foundation of Hebei Province(17226320D-iv). The funders had no function in study pattern, data drove and assay, decision to publish, or preparation of the manuscript.

Competing interests: The authors accept alleged that no competing interests be.

1 Introduction

As one of abiotic stresses, table salt stress tin can significantly affect plant growth and yield. Today, i.125 billion hectares of farmland around the globe are threatened by table salt stress, which is an of import issue for agriculture [1]. China also has 367 billion hectares afflicted past common salt stress, accounting for ane,230 hectares of farmland soil [two]. Amid the areas of business organisation, the coastal surface area, one of the primary types of saline-alkali land, has frequent water-salt interactions and secondary salinization as it is close to the sea [3]. Considering the ecological environment of the salinization expanse is fragile and natural conditions are limited past many factors, it is highly pregnant to develop and apply saline-alkali country scientifically and rationally while under pressure from a rapid population increase and a sharp decline in land resources; the goal is to advance towards the sustainable and salubrious development of China'southward forestry and ecological environment [iv].

Choosing and cultivating first-class salt-tolerant tree species through biotechnology is currently one of the nigh economical, effective, ecological and environment-friendly biological measures to solve the soil salinization problem [5]. Salix babylonica is a deciduous tree or shrub belonging to Salix in the family Salicaceae. It has strong ecological adaptability and tin can grow well nether saline-brine, drought and barren soil conditions [6]. Previous studies on the salt tolerance of Salix plants were generally focused on the physiological responses of seedlings to salt stress [7–9], but there are few studies on the adaptability of plants to common salt stress specific to seedlings. High table salt stress will cause plant h2o loss, ion imbalance and nutrient chemical element deficiency through osmotic stress and ion poisoning [10], which volition affect the normal growth and morphology of plants. A serial of physiological growth changes in plants under salt stress are the comprehensive embodiment of their salt tolerance ability, among which the growth status of plant roots, the ion accumulation in dissimilar organs and the alter in photosynthetic fluorescence parameters are important factors affecting the salt tolerance ability of plants [11–13]. These indicators not only tin represent the extent of the effects of stress factors on plants, but they can also reflect the growth of plants under salt stress, the selective absorption and ship of ions, and the photosynthesis power. Willow has the characteristics of antipyretic, analgesic, anti-inflammatory, anti-rheumatism, astringent, drought resistance [14] and anti-corrosion [15], among which the bark of White Willow contains salicin [xvi, 17] with antibacterial, bactericidal, antioxidant, antipyretic, analgesic and other functions, and is a good natural food condiment and food resource of wellness intendance products [18]. Its roots can too absorb harmful elements, reduce the affect of harmful elements on the surrounding soil [19, 20], and play a role in purifying polluted water [21]. Salix alba has strong adaptability [22], so it has swell potential for employ and promotion in the ecological management of coastal saline-alkali soil. Therefore, this experiment involved Salix alba branches as the object and used hydroponics to simulate the seedling raising process of cuttings on coastal saline-alkali country to study the growth of their roots, the changing ion contents in the roots and leaves, and the irresolute photosynthetic fluorescence parameters under different salt concentrations. Exploring the common salt tolerance of Salix alba can not only provide theoretical reference for the study of its salt stress adaptation mechanism, only besides provide suggestions for the utilization of salix salix salix in coastal saline-alkali areas.

2 Materials and methods

2.one Exam materials and test blueprint

The test materials were collected from the germplasm resource nursery of Golden Beach Forest Farm in Huai'an County, Hebei Province. The branches of the Salix alba were basically the same in terms of growth, and those that were robust and costless of diseases and insect pests were selected after the leaves barbarous in December. The middle ii-thirds of the selected branches were cutting into 20 cm-long cuttings. The uppermost bud was 0.5–1 cm from the pinnacle of the cuttings. The upper cut was a apartment cutting, and the lower cut was an oblique cutting. The experiment was performed in the Artificial Climate Room of Hebei Agricultural University, Baoding Urban center, Hebei Province on Dec 23, 2019. The temperature of the climate room was set to 28°C/25°C (calorie-free/nighttime); the LED common cold light source maintained the light intensity at 1000 μmol·yard^-2·s^-1; the photoperiod was 14 h/10 h (light/dark); and the humidity was 60%.

The test material was placed in a 55 cm×38 cm×15 cm (length ×width ×height) plastic box for hydroponic culture (Fig one). The experiment consisted of 5 treatments, and each handling was repeated iii times. We ready the concentration of Nacl and the limerick of civilisation medium by referring to existing studies [23–25]. A one/2 dilution of Hoagland's complete food solution was used every bit the base of operations to set up hydroponic solutions with NaCl concentrations of 171mmol、342mmol、513mmol、684mmol, and 1/2 Hoagland's complete nutrient solution (PH = seven.2) was used as a control (CK). i/2 Hoagland'southward complete nutrient solution includes: Ca(NO₃)₂·4H_iiO 472.5mg、K₂Then_iv 303.5mg、NH₄H₂PO₄ 57.5mg、MgSO₄ 246.5mg、NaFeC_xH₁₂North₂O₈·3H₂O 30mg、FeSO₄ 15mg、H_threeBO₃ 2.86mg、Na₂B₄O₇·10H₂O 4.5mg、MnSO₄ 2.13mg, CuSO₄ 0.05mg、ZnSO₄ 0.22mg、H₈Mon₂O₄ 0.02mg. In that location were 25 cuttings in each treatment, and they were soaked directly in the solution; the height of the solution was approximately more than than half of the height of the cuttings. The nutrient solution was inverse every 5 days during the growth process. Before the nutrient solution was changed, the cuttings were removed and the roots were rinsed with water to wash abroad the last residue salt and prevent excessive table salt accumulation. The contents of Na⁺, K⁺ and Ca²⁺ ions and the photosynthetic parameters and chlorophyll fluorescence kinetic curve parameters in the roots and leaves were determined following 20 days of treatment.

2.2 Measured items and methods

2.2.i Measurement of root growth parameters.

During the growth of Salix alba, the number of root sprouting days and the rooting charge per unit of all the cuttings were counted, and the rooting index was calculated according to the number of root sprouting days (rooting alphabetize = ∑Gt/Dt, where Dt, the day of the rooting test; Gt, the number of rooting branches on the day, and the rooting index is the number of rooting branches on the mean solar day/sum of days). After 20 days of salt stress handling, 5 uniformly growing cuttings were selected to measure out the average root number and boilerplate root length.

2.2.2 Determination of ion contents in the roots and leaves, adding of the ion selective assimilation and transport ratio.

The measurement method for tracking the ion content was slightly modified relative to the method by Yang Sheng et al. [26] and Yu Bingjun et al. [27]. The sample was first broiled at 105°C for 30 min and so stale at 70–80°C to a constant weight. Afterwards the sample was ground and passed through a sieve (the aperture was 0.425 mm), the fixed mass was weighed. Thirty mL of deionized water was added to the sample, which was then shaken well and placed in a boiling water bath for 2 h. After cooling, the sample was filtered and diluted to l mL. The Na⁺, K⁺ and Ca²⁺ contents were determined by atomic absorption method (Atomic absorption spectrometer: ZEEnit700-700P; analytikjena computer in Germany). The methods of Zheng Qingsong et al. [28] and Yang Xiaoying et al. [29] were used to calculate the selective absorption and transport coefficients of ions X (K⁺ and Ca²⁺) by the roots and leaves co-ordinate to the post-obit formula. Ion assimilation coefficient SA _{x, Na} = root ([X]/[Na⁺])/medium ([X]/[Na⁺]); ion transport coefficient ST 10, _Na = leaf ([10]/[Na⁺])/Root ([X]/[Na⁺]). In the formula, the K⁺ content in the medium (civilisation broth) was 272 mg, and the Caⁱⁱ⁺ content was 230 mg.

ii.ii.3 Determination of photosynthetic parameters in the leaves.

Following 20 days of table salt stress treatment, the photosynthetic gas exchange parameters of the Salix alba leaves were measured. Five uniformly growing cuttings were selected for each handling group in the test. After the cuttings were left under normal illumination in the climate room for 3 hours, we selected the 3rd to 5th leaves from the tiptop to bottom with the same position, size, and light-receiving direction and with fully expanded functional leaves. Using a Li-6800 portable photosynthesis meter (LI-COR, USA), the P_n, E, g_s and C_i can be determined. The measurement conditions were as follows: the PAR was thousand μmol·m^-2 ·s^-one, the CO₂ concentration in the stock-still system was 400 μmol·mol^-ane, and the relative humidity was threescore%.

two.two.4 Rapid decision of the chlorophyll fluorescence induction kinetic curve.

After twenty days of salt stress treatment, 5 Salix alba cuttings with average growth were selected from each treatment for measurement. Before the measurement, the leaves were dark-adapted for 15 minutes, and so the rapid chlorophyll fluorescence induction kinetic bend and related parameters were measured using a Pocket PEA institute efficiency analyzer (Pocket PEA, Hansatech, United kingdom of great britain and northern ireland). The resulting O-K-J-I-P curve was used for rapid chlorophyll fluorescence induction curve information assay (JIP-examination) and adding [30, 31].

2.iii Data processing

One-way ANOVA and the LSD method were used to exam the significance of the differences (α = 0. 05).

3 Results

three.1 Effects of common salt stress on Salix albicans root growth

The examination results show that although plants under salt stress can reach a 100% rooting charge per unit between treatments, the average root number, average root length and rooting index are quite unlike amongst the treatments, and the overall tendency is basically the aforementioned. The trend is that low-common salt stress stimulates root germination and elongation, high-salt stress inhibits root growth, and the intensity of the inhibition is positively correlated with the salt concentration.

Figs 2 and 3 show that when the NaCl concentration was 171mM, the average root number and average root length were significantly increased compared with those of the command. This upshot may exist a stimulating effect of low-salt stress on root growth and then announced again as the stress intensifies, with a gradual downward trend. When the NaCl concentration was 513mM, the root number and length were significantly lower than those of their corresponding controls by 48.7% and 39.nine%, and the root growth was significantly inhibited at that fourth dimension. Compared with the command, the rooting index did non change significantly when the NaCl concentration was 171mM, but with the increase in stress, the number of days for root formation was delayed, and the rooting alphabetize decreased significantly. When the NaCl concentrations were 342mM, 513mM and 684mM, the rooting indexes were significantly lower than that of the control (ix.vi%, eighteen.one% and 27.7%).

3.2 Effects of table salt stress on ion content, absorption and transport in the roots and leaves of Salix alba

The ion content measurements (Fig 4) showed that under different concentrations of NaCl, the Na⁺ contents in the roots and leaves of Salix alba were significantly higher than that in the command group, and the range of Na⁺ change was positively correlated with the stress concentration. The comparing of Na⁺ contents in the roots and leaves shows that the Na⁺ content of the roots is much college than that in the leaves. Under 684mM NaCl stress, the Na⁺ content in the roots could accomplish twice that in the leaves. With increasing stress concentration, the Thou⁺ content in the leaves starting time increased and so decreased, reaching a peak at a concentration of 171mM NaCl, which was a significant increase of fourteen.0% compared to the control group. Nonetheless, after the NaCl concentration was greater than 342mM, the concentration was significantly lower than that of the control. As the stress concentration increased, the K⁺ contents in the roots of each treatment group showed a gradual decrease, which were all significantly lower than that of the command. The Ca^two+ content in the leaves of Salix alba increased get-go and then decreased with increasing salt concentration. At 342mM NaCl, compared with the command group, the concentration significantly increased past 13.6% and so showed a meaning down tendency. The Caⁱⁱ⁺ content in the roots decreased continuously with increasing stress, and when the NaCl concentration was 684mM, the Ca²⁺ content dropped to 35.6% of the control.

Figs 5 and 6 show that both the Na⁺/Thousand⁺ and Na⁺/Ca²⁺ in the roots and leaves increased significantly with increasing NaCl stress concentration. This finding shows that as the stress intensifies, the relative absorption of Na⁺ by Salix alba increases profoundly, but the assimilation of Yard⁺ and Ca^two+ decreases. The Na⁺/K⁺ and Na⁺/Ca²⁺ contents of all the treatments gradually decreased from root to foliage, and the rising Na⁺/1000⁺ and Na⁺/Ca²⁺ in the roots were significantly (F = 1263.766, df = 4, Sig.<0.001; F = 10485.256, df = iv, Sig.<0.001) college than those in leaves (F = 1235.223, df = 4, Sig.<0.001; F = 2335.783, df = 4, Sig.<0.001), suggesting that Salix alba could reduce the salt stress damage to immature tissues by regulating ion ship.

As shown in Fig 7, with increasing NaCl stress, the SA _k, _Na, ST _k, _Na, SA _Ca, _Na, and ST _Ca, _Na all showed a tendency of first increasing and so decreasing. When the NaCl concentration was less than or equal to 342mM, the selective absorption capacity of the roots for Thousand⁺ and Ca^two+ and the selective ship chapters of the leaves for K⁺ () and Caⁱⁱ⁺ were enhanced and reached a significant level (F = 998.922, df = 4, Sig.<0.001;F = 1018.689, df = 4, Sig.<0.001; F = 168.047, df = 4, Sig.<0.001; F = 29.925, df = 4, Sig.<0.001). The selective absorption capacity of roots for K⁺ is greater than that of Caⁱⁱ⁺, but the selective transport capacity of the leaves to Ca²⁺ is greater than that of K⁺. These results indicated that Salix alba could adjust the upward transport capacity of K⁺ and Caⁱⁱ⁺ via the selective absorption and accumulation of mineral ions to compensate for the alter in concentration under salt stress, to preclude the impacts of nutrient deficiency and ion toxicity on the shoot growth.

three.3 Effects of salt stress on photosynthetic parameters in Salix alba Leaves

Figs 8 and 9 testify that the photosynthetic parameters of Salix alba leaves were afflicted to different degrees under different table salt concentrations. When the NaCl concentration was 171mM, the P_n of the leaves increased, but there was no significant deviation from the control. Later, every bit the salt stress intensified, the photosynthetic carbon assimilation power of Salix alba leaves was significantly inhibited (F = 95.66, df = 4, Sig.<0.001); when the NaCl concentration was greater than 171mM, both the Due east (F = 100.091, df = iv, Sig.<0.001)and g_south (F = 69.346, df = 4, Sig.<0.001) were significantly lower than the command and became stronger; but at a low salt concentration (171mM NaCl), in that location is no meaning difference from the command. With the increased salt concentration, the leaf C_i showed a tendency of outset decreasing and then increasing, reaching the lowest when the table salt concentration was 342mM, which was significantly lower than the control by ten.4%, and then information technology gradually increased. The C_i of leaves (F = xx.50, df = 4, Sig.<0.001) under 513mM and 684mM NaCl treatments were not significantly different from that of the control, but they were significantly college than the lowest value by 12.4% and fourteen.six%, respectively.

3.4 Effect of salt stress on the rapid chlorophyll fluorescence induction kinetic bend (OJIP) of Salix alba leaves

The OJIP curve tin can provide a keen deal of photochemical information nigh PSⅡ and accurately reflect the state of the plant photosynthetic apparatus and the electron redox state of the PSⅡ donor side, acceptor side and PSⅡ reaction heart in the photoreaction [32], thus representing the effects of external stress on the plant photosynthesis ability and even the caste of damage to the photosynthetic organs. Fig 10 shows that with the increasing NaCl concentration, the OJIP bend of Salix alba leaves changes to unlike degrees. Compared with the control grouping, under the 171mM NaCl treatment, the fluorescence value of JIP does not modify significantly; when the NaCl concentration reaches 342mM and college, the fluorescence values of I and P drop significantly and in that location is an obvious inflection point K (approximately 300 μs), and the OJIP curve changes to the O-G-J-I-P bend. The K-phase fluorescence value nether high salt handling is higher than that nether low salt handling, and the maximum fluorescence can exist reached faster, which indicates that the higher the salt handling concentration is, the greater the damage to the leaves of Salix alba.

3.5 Effects of table salt stress on quantum yield and free energy distribution ratio

Fig eleven shows that under different common salt stresses, the energy absorbed, transformed, used for electron transfer, and dissipated by thermal radiation in the leaves of Salix alba changes. Compared with the control grouping, with the increasing NaCl concentration, the maximum photochemical efficiency (φPo) of Salix alba leaves after dark accommodation gradually decreased. Under the 342mM NaCl treatment, the φPo was significantly lower than that of the control. At that fourth dimension, salt stress triggered photoinhibition, and the photosynthetic capacity of the leaves was reduced.

The excitons captured past the reaction center transfer electrons to the electron transport chain, and the ratio of excitons that exceed Q_A's other electron acceptors to promote Q_A reduction excitons (Ψo) and the low-cal energy captivated by the reaction center are used for electron transfer. The breakthrough yields (φEo) all increased offset and so decreased with the increasing common salt stress. At 171mM NaCl, although the Ψo and φEo increased, they were not significantly different from the control. After, as the stress intensified, both the Ψo and φEo were significantly lower than those of the control. When the NaCl concentration was 342mM, the Ψo and φEo were significantly lower than the 11.ane% and 11.9% of the command grouping, respectively. Compared with the command group, salt stress increased the quantum ratio (φDo) of Salix alba leaves for oestrus dissipation. When the NaCl concentration was 513mM, φDo was significantly higher than that of the command.

3.6 Influence of salt stress on the operation index and driving forcefulness

The operation index and driving force can accurately reflect the changes in the country of the photosynthetic appliance of plants nether stress. PI_ABS refers to the performance index based on the absorption of light free energy, PI_CSm refers to the functioning index based on the unit of measurement expanse, and DF_CSm refers to the driving force based on the unit expanse of the material. Figs 12 and 13 show that as the NaCl stress concentration increases, PI_ABS, PI_CSm and DF_CSm all show a gradual refuse. PI_ABS showed no meaning difference from the control when the NaCl concentration was 171mM, and and then with the increased salt concentrations, the departure became more significant (F = 61.074, df = 4, Sig.<0.001), indicating that the Salix alba leaves experienced photoinhibition, the PSⅡ was damaged, and the measurement at the 684mM NaCl concentration was significantly lower than that of the control, by lx.two%. When the NaCl concentrations were 342mM, 513mM and 684mM, the PI_CSm values were significantly lower (F = 202.821, df = iv, Sig.<0.001) than that of the control past xx.1%, 43.9% and 66.iv%; when the NaCl concentrations were 513mM and 684mM, the DF_CSm values were significantly lower (F = xl.755, df = 4, Sig.<0.001) than the control by 6.3% and xi.2%. Common salt stress seriously affects the absorption of light energy by plants and leads to a pass up in the bones driving force.

4 Conclusions and word

4.ane Influence of common salt stress on the root growth status of Salix alba

As the primary organ responsible for plant material exchange, the root system and its growth status are closely related to the growth and development of the aboveground plant parts, whether the root system tin can function normally, and the plant'south h2o and nutrient utilization efficiency [33]. Under salt stress, the root system is the first to experience the arduousness stress signal, and it is also the most directly affected role [34]. Its ring-stripe inhibition is primarily manifested in the low levels of the root length, area and other parameters, and the root arrangement grows slowly. A high-salt surround will cause plants to feel osmotic stress and ion toxicity, which will lead to changes in membrane permeability, which volition in plough affect the absorption of water and nutrient elements past the roots, causing the plants to lose a large amount of water; the ions near the roots will be unbalanced, the physiological functions of the roots will eventually be lowered, and even the construction volition be destroyed. Some of the aboveground leaves wilt, and photosynthetic product cannot be performed unremarkably, which causes plant growth and metabolic disorders until the loss of physiological functions.

The modify in root growth and the time of the root sprouting period can straight reflect the degree of harm to plants by salt stress and represent the strength of plant salt tolerance [35]. This report showed that the 171mM NaCl concentration significantly promoted the increment in the boilerplate number of roots and the elongation of the average root length of Salix alba cuttings, and information technology tin can promote the rooting of the root system in advance, to a certain extent, which is consequent with Wang Shufeng et al. [36] and Ci Dun. The inquiry results of Wei et al. [35] were basically the same. This growth response may be due to the decrease in water potential outside the roots nether table salt stress, which stimulates the growth of the roots instead of moderate osmotic stress to ensure the normal absorption of water and nutrients to encounter the physiological and metabolic needs of the aboveground parts.

Some plants do have the miracle that low salt promotes the increase of some indicators, such equally: promoting the germination of sorghum seeds [37], the roots of the seedlings of wolfberry [38] and rice [39], and the growth indicators of corn [40, 41]. Both ChorophyⅡin chrysanthemum [42] and proline content of cherry seedlings [43] are increased, while the net photosynthetic charge per unit of wild chrysanthemum [44] and hazel trees increased [45]. The reason for the depression table salt concentration may exist that the common salt stress has a dual effect of stimulus and inhibition on plants. The strong and weak human relationship between stimulus and inhibition triggers changes in various plant indicators, resulting in the aforementioned low salt. It can promote growth, and information technology will be inhibited later loftier salt. This finding shows that Salix alba has some ability to arrange and adapt to salt stress, and this adaptability is of great significance to the survival and continuation of the plant itself under arduousness. Still, every bit the common salt stress intensifies, the power of plants to coordinate their own growth is destroyed, root germination and elongation are significantly inhibited and become more intense, the root functions are destroyed, and the plants cannot maintain their normal growth and development.

iv.2 Effects of salt stress on ion content, absorption and transport in Salix alba

Ions play an important role in the normal growth of plants, but salt stress can destroy the dynamic balance of ions in plants [46], hinder the absorption of nutrients, and cause establish metabolism disorders. The alter in the distribution of ions reflects the degree of damage to plant cells past the external adverse environment. Additionally, plants can maintain balanced nutrition past improving the absorption and ship of ions, which also represents the level of plant resistance to stress. When measuring the ion contents of plant roots and leaves, it is helpful to reveal the salt tolerance or salt damage machinery of plants.

In this study, when the salt concentration was depression, the growth of Salix alba was basically normal, the symptoms of salt damage were not significant, and the harm was obvious under severe stress. Na⁺ accumulates significantly in the roots and leaves of Salix alba under table salt stress, but the Na⁺ content in different organs is significantly different, and it is primarily concentrated in the roots. This consequence shows that the willow root system has a compensation mechanism that tin can reduce the transportation of salt to aboveground parts by enriching Na⁺ in the root, thereby effectively reducing or delaying the occurrence of common salt damage in the aboveground parts. This conclusion is consistent with the report past Hao Han et al. [47]. When the salt stress is too high, this residue is broken, and growth is blocked.

As an important inorganic solute, M⁺ is essential for reducing the cell osmotic potential and maintaining the water residual. By and large, plants accept an antagonistic effect on the absorption of Na⁺ and Chiliad⁺ [48], and the competition between the two usually leads to a subtract in the K⁺ content. The loss of K⁺ will cause changes in the physical structure of the stomata, frustrating photosynthesis [49]. In addition, G⁺ participates in the metabolism of various enzymes in plants [50]. As common salt stress increases, an excessive loss of K⁺ will lead to Grand⁺ dependent enzymes in Salix alba The enzyme activity decreases, which affects the metabolic reactions in which information technology participates. Therefore, if plants are to grow in a salty environs, the selective assimilation of K⁺ by the root organization and the transportation of K⁺ to the ground are particularly important. This written report showed that the K⁺ content in the roots of Salix alba significantly decreased with increasing stress, but the K⁺ in the leaves could be maintained at a high level at a 342mM NaCl concentration and below and even increased significantly when the NaCl concentration was 171mM, according to Zhou Qi et al. [51] A study on Carpinus chinensis also confirmed this result. At this time, the value and increase of Na⁺/K⁺ in the roots of the Salix alba were greater than that of the leaves, and the SA _k, _Na and ST _k, _Na all increased significantly. Studies have shown that under salt stress, the Na⁺/K⁺ value can stand for the caste of table salt damage to the plant, and the lower Na⁺/G⁺ value of the leaves can aid the found better maintain its growth and photosynthetic function [52], and the SA _g, _Na and ST _k, _Na indicates that the plants can better tolerate salt stress [53]. This result shows that at that time, Salix alba could maintain a relatively stable leaf K⁺ content and the normal progress of photosynthesis by restricting the transportation of Na⁺ from the root to the leaves, increasing the selective absorption of K⁺ through the plant roots and the power to transport K⁺ to the ground. The accumulation of Na⁺ causes impairment to plants, which may be an important mechanism by which Salix alba copes with common salt stress. Afterward, with the increment in table salt stress, the Grand⁺ in the roots and leaves clearly flowed out. A loftier concentration of Na⁺ will supersede the Ca²⁺ leap to the membrane system, which volition harm the integrity of the membrane structure and membrane function, thereby destroying the ion balance in the plant body and causing a large amount of organic solute extravasation [54]. The establishment of Ca²⁺ homeostasis in the cytoplasm is a cardinal condition for table salt adaptation [55]. This experiment showed that every bit the salt stress intensified, the Ca²⁺ content in the Salix alba roots continued to decrease, but it could accrue in the leaves when the NaCl concentration was ≤342mM. The results of Jia Yin et al. [56] were similar; the Na⁺/Ca²⁺ value of white Salix roots was higher than that of the leaves, and the Sa _Ca, _Na and ST _ca, _Na were all significantly increased. This result may exist due to the big influx of Na⁺ into the root arrangement under salt stress, activating Ca²⁺ indicate transduction, triggering the sodium elimination system to reduce the damage of Na⁺, and enhancing the selective assimilation of Ca²⁺ in leaves, thereby enhancing the selective transport of Ca^two+ from root to shoot to maintain the low cell osmotic potential and the stability of the cell membrane. In addition, studies have shown that the increase in intracellular Ca²⁺ contents under salt stress can inhibit the outflow of K⁺, thereby alleviating the impairment of salt stress to plants [57]. Therefore, the upward transportation of Ca²⁺ in the roots of Salix alba may be an important mechanism for information technology to maintain the balance of K⁺ and Na⁺ in the aerial function, establish ion homeostasis in the aeriform office, and accommodate to salt stress. Even so, due to the limited power of the roots of Salix alba to blot Ca²⁺, nether high salt stress, the absorption of the roots will not be able to offset the loss of food elements caused by ion poisoning.

iv.3 Effects of table salt stress on photosynthetic parameters of Salix alba

Photosynthesis is a central metabolic process that provides material energy for plants. Loftier salt stress volition comprehensively affect the photosynthesis of plants through osmotic stress, ion toxicity, and feedback inhibition acquired past the aggregating of photosynthetic products [58]. These furnishings volition cause the devastation of the membrane structure and the imbalance of ions in tissue cells, affecting the assimilation of light energy past plants and the procedure of carbon absorption [59]. This change inhibits the formation of leaf primordia and reduces the photosynthetic area and carbon assimilation of individual plants, resulting in physiological metabolic disorders and the accumulation of toxic substances. In fact, the free energy supply related to photosynthesis, carbohydrate metabolism, and the TCA cycle are all inhibited by salt stress [threescore].

Because stomata are directly connected to the external environment, their coordinated response under stress determines whether the photosynthetic chapters of the plant is normal [61]. In this experiment, the P_{due north}, E, and g_s did not change significantly when the NaCl concentration was 171mM. Equally the table salt concentration further increased, each index decreased significantly, which is basically consistent with the results of previous studies [62, 63]. When the NaCl concentration was less than 342mM, the C_i of the Salix alba leaves decreased with decreasing g_southward. Thus, the diffusion resistance of CO₂ in the leaves increases, and the carbon sequestration power weakens. The stoma cistron is the dominant factor restricting the decline in Salix alba leaf photosynthesis. Later on, equally the degree of salt stress farther intensified, the C_i increased with the decreasing g_s, and the photosynthetic system action of the mesophyll cells decreased, resulting in a decrease in the assimilation capacity, which is a typical non-stomatal limiting factor. Previous studies have shown that nether adverse stress, stomatal restriction and non-stomatal restriction and the interaction of the two will reduce the photosynthetic rate of plants; under mild stress, stomatal restriction is ascendant; and under severe stress, stomatal restriction leads to non-stomatal restriction [64, 65]. Our experiment likewise supports this view.

four.4 Effects of salt stress on chlorophyll fluorescence kinetics of Salix alba

The OJIP curve contains a great deal of data nearly the original photochemical reaction of the PSII reaction heart [66]. When environmental conditions alter, chlorophyll fluorescence tin can directly or indirectly affect the photosystem operation of plants [67]. The changes in the PSII can reflect the touch of changes in the stress environment on the photosynthetic chapters of plants and the adaptation mechanism of photosynthetic mechanism to environmental changes. Loftier common salt stress can inhibit or destroy parts of the functions of PSⅡ, hinder the original photochemical reaction and electron transfer process of PSⅡ, and reduce the photosynthetic capacity of Salix alba leaves. This consequence may exist the outcome of the aggregating of Na⁺. The typical fast fluorescence kinetics curve generally has O, J, I, and P phases during the ascension phase of fluorescence [68]. This written report shows that when the concentration of NaCl is ≥ 342mM, the OJIP curve of Salix alba will exist deformed to OKJIP, the fluorescence values of points I and P will decrease significantly, and obvious inflection point K will announced. The occurrence of the G signal is caused by harm to the PSII donor side oxygen release circuitous (OEC) due to the inhibition of the water lysis arrangement and the receptor-side part before Q_A, and the relatively variable fluorescence of the K point can represent the degree of OEC damage [69, 70]. In addition, the loftier table salt treatment greatly shortened the time required to reach the P indicate (the maximum fluorescence value). This result indicates that the higher the degree of common salt stress, the greater the damage to the stability of the PSⅡ reaction center and the OEC on the PSⅡ donor side of Salix alba leaves, the weaker the ability to provide electrons downstream and the stronger the reduction of the PSⅡ acceptor side is hindered.

The φPo, Ψo, φEo, φDo reflect the free energy distribution ratio of plants. In this study, when the NaCl concentration was 171mM, at that place was no pregnant difference among the indicators. As the stress intensified, the φPo, Ψo and φEo decreased significantly while the φDo increased significantly, which is different from the results of Huang Qinqin et al. [71]. This finding shows that Salix alba adjusted the energy distribution ratio of the PSII reaction center under unlike degrees of stress. This adjustment occurs to increment the quantum ratio used for heat dissipation and reduce the proportion of energy in photochemical reactions, which is an adaptive regulation mechanism of Salix alba under salt stress. The decrease in the φPo, Ψo and φEo indicates that the photosynthetic mechanism is clearly damaged, the ability to reduce the Q_B and P_Q on the PSII receptor side is diminished, and the electron transfer process is inhibited. Plants are prone to occur or beal photoinhibition in adverse environments [69]. In this report, when the concentration of NaCl was greater than 171mM, the PI_ABS, PI_CSm and DF_CSm all showed a pregnant downwards trend. This trend shows that Salix alba leaves exhibit photoinhibition, the PSII reaction center is reversibly inactivated or irreversibly degraded, the conversion efficiency of light energy is reduced, and the role of the photosynthetic appliance is impaired, which restricts the normal progress of photosynthesis.

In this study, 171mM NaCl stress had no significant effect on the growth status of the Salix alba root system, ion distribution or photosynthetic fluorescence characteristics and even increased these parameters to a certain extent. As the table salt treatment concentration gradually increased, the boilerplate root number and length, and rooting index decreased significantly; Na⁺ accumulated in the root system, K⁺ and Ca²⁺ were significantly lost; the photosynthetic charge per unit decreased significantly, the PSⅡ reaction center was partially inactivated, and the donor side OEC and the electron acceptor on the acceptor side were damaged. Salix alba can answer to salt stress by intercepting Na⁺ in the root system, improving the selective absorption of Thousand⁺ and Ca²⁺ and the ground transportation capacity, and increasing the breakthrough ratio used for heat dissipation, indicating that Salix willow has some tolerance to salt stress environments.

Supporting data

Acknowledgments

We limited sincere gratitude to all the authors involved in this study.

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