seaweed has developed as a cure with the capability to spare the biological system, because of its eco-friendliness, reasonableness, accessibility, and active ion expulsion rate. This work summarizes the result of saltiness on the organ of plants, and this biomass is revered because the driving redress material among the accessible materials too grandstand the bio activity part of seaweed extricate disengage and its adequacy as a valuable, on morphological and physiological characteristics of peppermint underneath completely different irrigation water salinity was conducted a factorial experiment supported a totally irregular style with four replications within the Yazd University greenhouse in 2020. Experimental factors included salinity stress at four levels of (0, 2, 4 and 8 dS/m) and, seaweed powder application at four levels of (0, 1, 2 and 4 ppt ). Morphological traits or growth parameters and physiological characteristics (relative water content, photosynthetic pigments, total soluble carbohydrate, proline amino acid) assessed. our results demonstrated that seaweed led to changes in peppermint growth and physiology which improved the plant salt stress response. Examination of interaction effects showed was a significant effect on total chlorophyll, chlorophyll a, chlorophyll b, carotenoids (Photosynthetic pigment) and, total soluble carbohydrate at the level of 1% (p≤0.01), and in examining morphological traits results demonstrated that fresh and dry weight of leaves, aerial components, total fresh and dry biomass and harvest index were significant at the level of 1% (p≤0.01). The use of seaweed at 4 ppt under stress-free conditions resulted in the highest weight of leaves, aerial, and total biomass. The highest relative water content was related to the without salinity stress (65.8%). The highest proline content was obtained at 4 and 8 dS/m salinity levels, which showed a rise of 37.5% compared to the management treatment. In general, the results showed that the employment of 2 and 4 ppt seaweed at high levels of salinity (4 and 8 dS/m) reduced the adverse effects of salinity stress by improved physiological traits, together with chemical change pigments and amino acid content. This analysis grandstands the bio activity part of seaweed growth biomass and its adequacy as a helpful, cheap, and reasonable.
Research Article
Morpho-physiological responses of peppermint (Mentha x piperita L.) by seaweed floair under salinity stress
https://doi.org/10.21203/rs.3.rs-2238761/v1
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Plant production
Photosynthetic pigments
Proline
seaweed
salt stress
Peppermint (Mentha x piperita L.) is an herbaceous, perennial, rhizomatous, and hybrid plant (2n = 48) belonging to Lamiaceae. This plant is a cross between M. aquatic and M. spicata species. This herb isn't as it were a therapeutic plant, but it is additionally a well-known herb commonly utilized all through the world in teas, flavorings, and confections (Malekmohammad et al., 2021). Peppermint is one of the most widely used medicinal plants that the annual production of essential oil in the world reaches about 4000 tons, Peppermint leaves contain 1.2-3.9% (v/w) of basic oils and more than 300 recognized components(52% terpenes and 9% sesquiterpenes) (Motiee and Abdoli, 2021). Menthol, menthol, menthofuran, and 1,8-cineole are the most important constituents of the necessary oil of this plant (Moetamedipoor et al., 2021). Essential oil components and morpho-physiological characteristics were changed under environmental and nutritional factors (Mavandi et al., 2021).
The development and yield of plants worldwide are always limited by numerous biotic environmental stresses (Hassanisaadi et al., 2022). Among abiotic stresses, salinity has caused much damage to plants worldwide (Aghighi Shahverdi et al., 2018).
Salt stress not only diminishes the expected yield of crops, but also affectsmetabolic processes in plants through impairment of cell water potential, ion toxicity, membrane integrity and function, and uptake of indispensable mineral nutrients (Taghizadeh-Mehrjardi et al., 2021).One of the crucial biochemical changes befalling when plants are subjected to salt stress is the accumulation of Proline, which can provoke metabolic disorders such as oxidation of membrane lipids, proteins, and nucleic acids (Sodaiizadeh et al., 2016) and, growth decrease is an adaptation for plant survival under stress conditions (Bernstein, 2019). Salinity stress affects the vegetative and reproductive growth of the plant and therefore reduces the dry weight and yield (Fathi et al., 2020; Yu et al., 2021).
Bio-fertilizers are a collection of preservatives with many beneficial microorganisms or their metabolic products, which are mostly used to provide the plant with the nutrients it needs and create the appropriate physical and chemical conditions for the soil to grow (Kumar et al., 2022).
Seaweed extract contains various amounts of biologically active secondary metabolites, vitamins, thymolic compounds, and precursors of vitamins and amino acids. Moreover, it includes several phytohormones such as auxins, cytokinins, gibberellins, abscisic acid, and brassinosteroids (Stirk et al., 2020 Ghatas et al., 2021).
The cell walls of the seaweed have polysaccharides and proteins that function binding sites for uptake of metal ions (Gupta and Rastogi, 2008). There are some elements dependable for the natural action capability of seaweed cell surface; among these variables are unit accessibility of authoritative bunches for metal ions, the affinity constants of the metal with the practical cluster, the chemical state of those destinations, he quantity of utilitarian bunches within the seaweed, and he coordination variety of the metal ion to be occluded (Mehta and Gaur, 2005).
Saltiness influences plant development through various elements of cellular digestion system, such as: supplement take-up, photosynthesis modification, breath, protein and nucleic corrosive integration, amassing of natural solutes, chemical movement, hormonal adjust and water accessibility(Chiahi et al., 2020).
Iran is one of the best regions globally in terms of climate and geographical location for the cultivation of medicinal plants(Hakimzadeh et al., 2022); however, a large part of the country is under salinity stress (Ardakani and Vahdati, 2018). Despite numerous reports on the effects of salinity stress on medicinal plants of the peppermint family (Liu et al., 2022; Askary et al., 2017; Fathi et al., 2020), the morpho-physiological responses of peppermint to the application of seaweed extract under salinity stress have not been considered in any of the existing reports. Therefore, this study was performed to observe the changes in physiological traits of peppermint (Menthax piperita L.) plants at different levels of seaweed extract and salinity stress.
Experimental materials and treatments
The present work was conducted in the research greenhouse of Yazd University as a factorial experiment based on a completely randomized design (CRD) with four replications. Experimental factors included salinity stress at four levels ( 0, 2, 4, and 8 dS/m) and seaweed at four levels ( 0, 1, 2, and 4 per thousand). The pots were put within the greenhouse conditions (22 to 35 ℃ amid the day and night and 70% relative humidity). The soil dampness of the pots was kept inside the 90% field capacity filled with soil (64 pots), soil characteristics are presented in Table 1. Soil characteristics pH, EC, and organic carbon were estimated (Page1982). Also the concentrations of Fe, Cu, Zn, and Mn were calculable by an atomic absorption photometer (Analytik Jena-330, Germany) with an air-acetylene flame. Then, the readings were placed in Eq. (1) to calculate the concentrations of metals in the soil samples in mg/kg.(Esfandiari and hakimzadeh, 2022).
X = (a × v × b × 100/m) (Eq. 1)
in which X is that the concentration of the component in mg/kg, a is that the spectrophotometer reading, v is that the volume of the extract used, b is that the dilution factor, and m is the pant weight.
Treatments were applied after the plants reached the three -leaf stage, different electrical conductivity was conducted using sodium chloride salt (NaCl) To be sure, the electrical conductivity of the solution was checked by an EC-meter. After five weeks, seaweed was sprayed in four concentrations three times at 2-day intervals. The seaweed extract used was ‘Ascokelp’ (Kimia Koodeh Bahar Company, Iran). Water was used instead of seaweed extract for the control treatment. According to the manufacturer, ‘Ascokelp’ is a soluble extract of the algae Ascophilium nodosum with Laminaria japonica and macro and microelements, an effective and rich combination of natural growth stimulants.
Morphological traits and growth parameters
Morphological parameters, like fresh and dry roots weight and shoots, were weighed with a digital scale with an accuracy of 0.001 g (Sartorius Quintix, Germany), for dry weight measurement, the shoot and root samples were incubated in an oven at 75 °C for 48h.
Relative water content determination
To assess plant water standing, relative water content (RWC) made up our minds in keeping with the method of Yamasaki and Dillenburg (1999):
RWC (%) = (FW-DW) / (TW-DW) × 100 ( Eq. 2)
where
FW: is that the fresh weight, DW: is that the dry weight, and, TW: is that the turgid weight
Photosynthetic pigments measuring
Lichtenthaler (1987) method was wont to live chlorophyll and carotenoids. In keeping with this method, 0.2 g of plant leaves were washed with distilled water and poured into a mortar, and 5 ml of 80% acetone was additional to that. The samples were then filtered through filter paper and then centrifuged at 1500 rpm for 15 min. Then the photometer (Analytik Jena 210, Germany) was tag with 80% acetone, and also the absorption rate of the samples was read at 663, 645, and 470 nm, and finally, the photosynthetic pigments containing the subsequent formulas were calculated.
where W illustrates the fresh weight of extracted tissue (g), V stands for the final size of the extract in 80% acetone, and OD demonstrates optical density at a specific wavelength.
Total soluble carbohydrate measurement
content was measured based on Irigoyen method. Absorbance of the samples was brows employing a spectrophotometer (Analytik Jena 210, Germany) at 485 nm at room temperature. Using the standard glucose curve, the soluble sugar content make up mind in mg kg-1 dry weight (Irigoyen et al.,1992)
Proline measurement
Proline content was measured based on Bates method. An alcoholic extract was ready from leaves supported the amino acid content. Briefly, and the amount of light absorption was measured at 520 nm with a spectrophotometer (Analytik Jena 210, Germany). Using the standard curve, the amount of proline was t was calculated based on mg g-1 FW of leaf (Bates et al., 1973).
Statistical analysis
All data obtained were analyzed by SAS software package (Statistical Analysis code, 9.2). A completely randomized style was performed to estimate the variance components of the consequences of salt stress, seaweed application, and their interactions. Additionally, the variations between treatments were assessed by Duncan’s only if the ANOVA F-test indicated a significance level of 0.05. (P ≤ 0.05). The Pearson correlation between the attributes was analyzed using SAS 9.4 software. Thecorrelation coefficients of mrpho-physiological traits were investigated using Pearson’s correlation matrix (Table 4 and 5).
Analysis of the variance of interactions of seaweed foliar application on the physiological properties of peppermint under salt stress shows that it has a significant effect on total chlorophyll, chlorophyll a, chlorophyll b, carotenoids (Photosynthetic pigment) and total soluble carbohydrates at the level of 1% (p≤0.01) and had no significant effect on free proline and relative water content table 2.
Also, the interaction effect of salt stress and seaweed on leaf fresh and dry, aerial parts, total fresh and dry biomass, and harvest index were significant at the level of 1% (p≤0.01) and had no significant effect on fresh and dry weight the roots table 3.
Photosynthetic pigment content
salinity stress decreased chlorophyll a, b, and total and carotenoid content, while seaweed foliar application (high concentrations) increased photosynthetic pigments. The highest total chlorophyll was achieved foliar application of 4 ppt seaweed under non-stress conditions (4.33 mg.g-1 FW), and the lowest was non-application of seaweed under 4 and 8 dS.m-1 salinity stress (1.38 and 1.42 mg.g-1 FW, respectively) (Fig. 1A).
The foliar application of seaweed at 1, 2, and 4 ppt concentrations showed the highest chlorophyll a content (3.10, 2.95, and 3.40 mg.g-1 FW), which increased by 43.5%, 40.6%, and 48.5%, increased compared to treatment without use. The lowest chlorophyll- a content (1.17 mg.g-1 FW) was associated with non-application of seaweed under a salinity treatment of 4 dS.m-1 (Fig.1B).
As shown in Figure 2A, application of 2 and 4 ppt seaweed under no salinity conditions showed the highest chlorophyll b content (0.93 and 0.92 mg.g-1 FW, respectively), which decreased by 59.1% and 58.6%. Non application treatment or in addition, the lowest chlorophyll b content was observed in the application and non-application of seaweed under saline or non-saline conditions.
The carotenoid changes were similar to those of chlorophyll b. In this regard, the highest carotenoid content was associated with the use of 4 ppt seaweed extract with no salt load (9.10 mg.g-1 FW), showing an increase of 57.4% compared to the control treatment. Seaweed application and non-application under 2, 4, and 8 dS.m-1 salinity treatments showed the lowest carotenoid content (Fig. 2B).
Relative water content (RWC)
Based on the results of analysis of variance (Table 2), salinity stress at RWC (p≤0.01) was significant. However, seaweed concentrations and interactions with seaweed salinity were not non-significant for RWC.
With increasing salinity and decreasing water potential, the water absorption in the peppermint plant is reduced, so the plant could not hold the water content of its leaves and withered. In this regard, the highest and lowest RWC (65.8 and 48.8%) were observed at 0 and 8 dS.m-1 salinity treatments, respectively (Table 2). One of the effects of salt stress is the preventing of water uptake and drought stress. And the reason for the decrease in uptake by roots under dry conditioans (Colom and Vazana, 2003).
Total soluble carbohydrates
As shown in Table 2, seaweed application and interaction effects of salinity and seaweed were significant on total soluble carbohydrates (p≤0.05). Seaweed application decreased total soluble carbohydrates. The highest total soluble carbohydrate content (11.25 mg.g-1 DW) was related non-application of seaweed under non-stress conditions. The lowest total soluble carbohydrate content (10.43 mg.g-1 DW) was observed when applying of 1 ppt under non-stress conditions (Fig. 3).
Proline content
The results represented the significant effect of salt stress and seaweed foliar application on proline content (p≤0.01). However, the interaction effects of salinity and foliar application of seaweed were found to be in significant (Table 2). As shown in the mean comparative results (Table 2), the salinity of the irrigation water increased the proline content. The highest proline content was related to 4 and 8 dS.m-1 (0.008 and 0.008 mg.g-1 FW) and the lowest (0.005 mg.g-1 FW) was reached under non-stress conditions. In addition, application seaweed at a concentration of 4 ppt caused the highest proline content (0.008 mg.g-1 FW), and other seaweed concentrations showed the lowest proline content (Table 2).
Fresh and dry root weight
As summarized in Table 3, salinity stress and application of seaweed to the leaves were significant for fresh and dry root weight (p≤0.01). However, interaction effects of salinity and seaweed were not significant for these traits. The salinity was caused by decreasing fresh and dry weight. The highest root fresh weight was related to 0 and 2 dS.m-1 (63.08 and 58.58 g.plant-1), whereas the highest root dry weight was achieved without salinity conditions (21.09 g.plant-1). Severe salinity stress (8 dS.m-1) reduced the mean fresh and dry weight of roots by 37.25 and 48.74%, respectively, compared to the control treatment (Table 3). Applying seaweed extract to the leaves resulted in fresh and dry root weight. The most effective concentration of the seaweed extract was 2 ppt, resulting in the highest fresh and dry root weight (81.04 and 22.8 g.plant-1). The lowest mean fresh and dry root weight was associated with not using the extract (Table 3).
Fresh and dry Leaf weight
As shown in Table 3, the effects of salt stress, seaweed application, and salinity- seaweed interaction effects were significant on fresh and dry leaf weight (p≤0.01). Salinity stress reduced, however, application of seaweed extract to the leaves increased the fresh and dry weight of the leaves. As shown in Figures 4A and B, the highest leaf fresh (23.6 and 24.7 g.plant-1) and dry weight (10.3 and 11.5 g.plant-1) were related to 2 and 4 ppt concentrations of seaweed application under non-stress conditions. Under high salinity, application of 2 ppt seaweed kelp increased leaf fresh and dry weight by 80.8% and 94.4%, respectively. The non-application of seaweed under the high salinity (8 dS.m-1) was caused by the lowest fresh and dry weight of the leaves (3.05 and 0.44 g.plant-1).
Fresh and dry aerial weight
The results represented the significant effect of salinity stress, foliar application of seaweed, and salinity stress × seaweed application on aerial fresh and dry weight (p≤0.01). The results showed that a low salinity level did not significantly reduce the mean aerial fresh and dry weight (Table 3). Results showed that seaweed application increased fresh and dry aerial weight under stress and non-stress conditions. Seaweed application (2 and 4 ppt values) under non-stress conditions showed the highest aerial fresh and dry weight. Application of 2 ppt seaweed under high salinity conditions (8 dS.m-1) showed 66.2% and 75.8% increasing aerial fresh and dry weight, respectively, compared to no seaweed no application at the same salinity. The lowest fresh and dry aerial weights (8.52 and 2.98 g.plant-1) were associated with non-application of seaweed under high salinity stress (Figures 5A and B).
Fresh and dry biomass weight
The results represented the significant effect of salinity stress, foliar application of seaweed, and salinity stress × seaweed application on biomass fresh and dry weight (p≤0.01) (Table 3). Seaweed extract resulted in an increase in fresh and dry biomass weight under salt stress and non-salt stress conditions. As shown in Figure 6, the highest biomass fresh and dry weights were associated with the application of 2 ppt of seaweed under non-stress conditions (124.9 and 41.97 g.plant-1, respectively). Application of 2 ppt seaweed extract under high salinity conditions (8 dS.m-1) resulted in 79.6 and 80.2% increases in fresh and dry biomass weight compared to no application of seaweed. The lowest means of these traits (20.02 and 6.4 g.plant-1) were related to non-application of seaweed at high salinity (8 dS.m-1).
Correlation coefficients traits
Under the seaweed foliar application, all the morphological traits were positively correlated, coefficients correlated were significantly affected (p<0.01) (Table 4).
Total chlorophyll were positively correlated with chlorophyll a, chlorophyll b, and carotenoids with coefficient correlation (0.98, 0.96 and 0.61) respectively, significant at (p<0.01). As well as chlorophyll a with chlorophyll b and carotenoids with coefficient correlation (0.89 and 0.73) respectively, significant at (p<0.01). According to the results in Table 5, there was a negative significant correlation between proline trait with all physiological traits, except total soluble carbohydrate and carotenoids. The highest correlation coefficient was associated with RWC% with correlation coefficients of (- 0.62) at the p≤0.01 level (Table 5).
Salt stress is one of the most limiting factors in the production of such crop plants (kiiani, et al., 2021). Salinity stress was found to cause a significant reduction in growth and morphological traits Findings showed that salt stress, particularly at higher levels (4 and 8 dS.m-1), result in a decrease in peppermint's growth and morphological traits, as well as the fresh and dry weight of leaves, aerial, roots, and total biomass. Alternative researchers have expressed that salinity stress usually result in a decrease in growth and morphological characteristics (Aghighi Shahverdi et al., 2019; Fathizad et al., 2022; Shahverdi et al., 2019 Askary et al. 2017).
Plants reduce their leaf area through environmental stresses like salinity and drought and diminish leaf dry weight per unit area. This tolerance mechanism minimizes the production of photosynthetic material and, ultimately, the biomass dry weight. which may be a part of this decrease due to lower water potential within the cell that causes stomata closure and limit carbon dioxide fixation. Content Chlorophyll is one of the important factors in maintaining the capacity of photosynthesis and determining the rate of photosynthesis in plant (Rossi et al., 2016).
Due to the deficiency of photosynthetic materials, the transfer of these materials to the procreative organs is reduced. As a result, plants' decrease in vegetative and procreative performance is ascertained. The decrease in dry weight below stress conditions may well be to the reduction of state pressure caused by the decline of plant leaf area and additionally the decrease in photosynthetic rate due to biochemical limitations as a result of dehydration or a rise in sodium ions. Baniasadi et al. (2015) reportable that with increasing salinity, the fresh and dry weight of the plant decreases, that is in line with the findings of this study.
The results of Jiang et al. (2018) indicated nano dioxide treatment Titanium formation of chlorophyll, Rubisco enzyme activity and it increases the rate of photosynthesis in the plant and foliar spraying with nanopotassium increases chlorophylls a and b were compared to the control.
The findings of this study showed that the utilization of various concentrations of seaweed extract led to a rise in growth and yield traits. The best growth and yield attributes were achieved within the application of 2 ppt of seaweed that had higher results and caused a significant increase in these traits compared to the management treatment (without foliar application of seaweed extract). Seaweed extract because of the distance of development hormones such as auxin and cytokinin, components such as nitrogen, Fe, Zn, Cu, Co, Mo, Mn, Ni, vitamins, and amino acids, encompasses a advantageous impact on development and resistance to saltiness or dry spell conditions. In this regard, Guerreiro et al. (2017) reported that the presence of important compounds in seaweed extract will increase the expansion growth and production of crops. Several researchers have expressed that the utilization of seaweed has positive and increasing effects on plants' growth and yield traits (Chitra and Sreeja, 2013; Selvam and Sirvakumar, 2014).
Researchers declared that the presence of growth regulators like plant hormone, etc. within the composition of seaweed extract results in enlarged longitudinal growth and plant yield. During this regard, Rathore et al. (2009) reported that seaweed biological fertilizers acted equally to plant growth regulators not solely because of their presence of nitrogen, potassium, and phosphorus but also due to the presence of trace elements and secondary metabolites, caused improve plant growth.
Gireesh et al. (2011) reported that the use of seaweed extract increased growth, physiological, and yield traits in bean plants. It appears that the distinction within the desired concentrations for each plant is because of the plant growth species, structure, and morphology of the plant. De-Carvalho et al. (2014)declared that the use of seaweed biological chemical because of macro-and micro-elements and growth hormones results in increased photosynthetic levels and therefore will increase vegetative and reproductive growth and plant yield.
The presence of organic molecules like organic acids, methionine, and polyamines within the composition of seaweed united of the explanations for the positive effects of this type of fertilizer on plant yield and growth. The uptake of minerals by binding to those molecules and forming chelates becomes easier and ultimately improves growth and yield. It appears that the growth of plants in saline environmental conditions result in nutritional imbalance, during which the addition of macro and micro nutrients can improve the growth standing of plants and increase plant tolerance equal to environmental stress.
The present study results showed that the fresh and dry weight of roots in treatment 2 ppt seaweed extracts increased significantly. It looks that higher concentrations of biological fertilizers don't completely have an effect on yield and growth; in different words, higher concentrations are toxic to peppermint and have adverse effects on growth and yield indices. During this regard, Rathore et al. (2009) concluded that biological fertilizers are useful due to containing macro and micro elements, amino and fatty acids, growth regulators, and other elements. The use of high concentrations leads to nutritional imbalance and toxicity to the peppermint plant and negatively affects growth. However, using the optimum concentration will increases the growth and yield by increasing the speed and duration of photosynthesis and also the transfer potency of photosynthetic materials (Mukherjee et al., 2020). Researchers have joined the enhancing effects of seaweed extract to the presence of assorted organic compounds likewise because the presence of phytohormones, particularly cytokines (Yao et al., 2020). These phytohormones area unit concerned in nutritionary metabolism within in the plant's reproductive organs. In other words, the use of seaweed extract will increases the quality of proteins from root to organ and will increases cytokine synthesis, resulting in a rise in growth and yield indices (Yao et al., 2020; Mukherjee et al., 2020; Fathi et al. (2020) reported that photosynthetic processes are one in all the foremost necessary cellular reactions that are strongly affected by salinity stress. Disruption of this process directly reduces carbon stabilization and biomass production in plants.
The present study results showed that salinity led to a significant reduction in photosynthetic pigments including chlorophyll a, b, and total and carotenoids, the results of correlation between traits showed that photosynthetic pigments have a positive and significant correlation with growth and yield attributes. In other words, one of the reasons for the decrease in the growth and yield traits could be the decrease in the photosynthetic pigment content under salinity stress. Salinity leads to the accumulation of sodium ions in plant organs, while the concentration of potassium, calcium, and magnesium ions decreases, thereby reducing photosynthesis and the photosynthetic pigments content (Shahverdi et al., 2019).
In the molecular structure of photosynthetic pigment (chlorophyll), magnesium plays a critical role, and magnesium deficiency under salinity stress is one of the reasons for the decrease in photosynthetic pigment content. This decrease in photosynthetic pigments may be due to the formation of protein enzymes like chlorophylla, that may be a reaction to chlorophyll depletion or damage to the photosynthetic process (Hussain et al., 2021; Fathi et al., 2020). Additionally, the decrease in photosynthetic pigment content is because of the inhibition of salt ions from the re-biosynthesis of proteins and their destructive effect on chloroplast structure (Mpunitha et al., 2022; Shahverdi et al., 2020).
Salinity led to a significant reduction in RWC. During this regard, researchers expressed that reducing RWC in plants under salinity stress results from high salt concentrations within the external environment, makining osmotic potential and dehydration at the cellular level (Fathi et al., 2020).
The highest total soluble carbohydrate content was related non-application of seaweed under non-stress conditions. Many environmental stress conditions affect the metabolism of sugars and the distribution of photosynthetic materials in growing plants.Researchers have reported a high correlation between the accumulation of soluble sugars (sucrose, glucose and fructose) and drought tolerance in plants (Hoekstra and Buitink, 2001)
Accumulation of compatible soluble substances (osmolite) such as proline amino acids and carbohydrates plays a vital role in maintaining plant osmotic balance (Feng et al., 2022). Shahverdi et al. 019; Fathi et al. 2020) reported that osmolites such as proline, soluble sugars and glycine betaine are synthesized, that additionally to officious with osmotic regulation and helping water absorption, play protective roles on membranes and cell macromolecules, and in sure conditions. Fan et al. (2011) according that the applying of protoctist liquid fertilizer caused a rise in proline amino acid content.This osmolite is that the most stable amino acid that has resisted oxidative hydrolysis and has the most negligible inhibitory effect on cell growth. Proline has also been introduced as a carbon and nitrogen source to repair the effects of stress, modulate the oxidation potential, and regenerate cells (Afshari et al., 2022). the main factors affecting salinity resistance are proline and soluble carbohydrate accumulation, enzyme activity, and photosystem efficiency ( Shahverdi et al., 2019). Increasing the extent of salinity stress in several treatments caused a rise in antioxidant enzymes in tomato plants (Dehnadi et al., 2020).salinity stresses act as plant growth inhibiting factors. They'll amendment the physiological characteristics and limit the production of crops in return that Seaweed extract has positive effects on plant growth, increasing tolerance to pests, diseases, and abiotic stresses. Seaweed extract conjointly will increases antioxidant content.
Soil and water salinity is one among the largest problems and the most important environmental stresses, especially in arid and semi-arid regions, that end up in reduced yield and production in plants. Numerous methods are projected to extend salinity tolerance and increase or protect the growth and yield of plants in these conditions, one among that is that the use of seaweed had positive and increasing effects on plants' growth and yield traits. The results showed that salinity, especially at higher levels (4 and 8 dS/m), decreased chlorophyll a, b, and total chlorophyll, in addition as carotenoids and RWC at higher levels of salinity. However, arise in proline was discovered at the same time with a rise in salt concentration within the growth medium.
The foliar application of seaweed increased growth and yield parameters in peppermint the foremost fascinating concentration of seaweed extract in line with the aid of the studied traits was 2 ppt, ofcourse, a degree of 4 ppt conjointly had acceptable results, however considering the value and several factors, selecting a degree of 2 ppt was desirable. Seaweed extract improved and increased photosynthetic pigments and proline amino acid content, by rising some physiological traits, the seaweed extract, together with increasing the chlorophyll and proline content, led to an increase in resistance plant underneath salinity stress. In general, the results showed that using seaweed extract under salinity stress in peppermint (Mentha piperita L.) improves physiological characteristic and salt tolerance in this plant.
Conflict of interest
The authors declare no conflict of interest
Ethical Approval
The authors have thoroughly observed ethical issues and no data from the study has been or will be published separately elsewhere.
Funding
This work was supported by Yazd university. The authors would like to gratitude all people who contributed to this research.
Competing Interests
The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.
Author Contribution Statement
M. H: conceived and designed the research. Conceptualization.Formal analysis. Writing-Review & Editing. Supervision. H. J.K: Formal analysis. The experiments were conducted by H.J.
H. S: data analysis was performed by H.S. M. E: Data curation. Resources. wrote down the draft of the manuscript with subsequent contributions from all the authors. All authors contributed to the article and approved the submitted version.
Data Availability Statement
The data that support the findings of this study are available from the corresponding author, upon reasonable request.
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Table 1: Physico-chemical characteristics of experimental soil at depths of 0 to 30 cm
|
Soil texture |
Nitrogen (mg.kg-1) |
Phosphorous (mg.kg-1) |
Potassium (mg.kg-1) |
Zinc extractable with DTPA (ppm of soil) |
Copper extractable with DTPA (ppm of soil) |
Manganese extractable with DTPA (ppm of soil) |
Iron extractable with DTPA (ppm of soil) |
Electrical conductivity (dS.m-1) |
pH |
Organic carbon (%) |
|
Sandy loam |
1200 |
7.4 |
250 |
0.59 |
0.27 |
2.17 |
1.42 |
2.8 |
6.93 |
1.7 |
Table 2: Analysis of variance of the influence of seaweed application on the physiological properties of peppermint under salinity stress
|
MS |
||||||||
|
S.O.V |
df |
Total chlorophyll |
Chlorophyll a |
Chlorophyll b |
Carotenoids |
RWC |
Total soluble carbohydrate |
Proline |
|
Salinity (S) |
3 |
8.56** |
5.26** |
0.40** |
27.05** |
730.17** |
0.079ns |
0.0000189** |
|
Seaweed (SW) |
3 |
1.87** |
1.09** |
0.15** |
7.34** |
112.67ns |
0.29* |
0.0000064** |
|
S × SW |
9 |
0.48** |
0.30** |
0.59** |
2.82** |
243.99ns |
0.16* |
0.0009641ns |
|
Error |
32 |
0.104 |
0.084 |
0.012 |
0.86 |
118.47 |
0.075 |
0.0006323 |
|
CV (%) |
- |
14.41 |
15.66 |
22.72 |
19.77 |
19.02 |
2.52 |
3.50 |
|
*, ** and ns: Significant at P < 0.05, P < 0.01 and insignificant, respectively. |
||||||||
Table 3: Analysis of variance of the influence of seaweed application on the morphology properties of peppermint under salinity stress.
|
MS |
|||||||||
|
S.O.V |
df |
Leaf fresh weight |
Leaf dry weight |
Aerial fresh weight |
Aerial dry weight |
Root fresh weight |
Root dry weight |
Biomass fresh weight |
Biomass dry weight |
|
Salinity (S) |
3 |
217.4** |
38.16** |
302.92** |
60.08** |
1467.84** |
228.42** |
4927.24** |
824.32** |
|
Seaweed (SW) |
3 |
217.6** |
58.37** |
461.37** |
108.15** |
7299.53** |
400.00** |
14311.26** |
1332.44** |
|
S × SW |
9 |
62.63** |
14.29** |
112.51** |
30.29** |
83.99ns |
6.78ns |
537.39** |
94.98** |
|
Error |
32 |
10.94 |
2.35 |
22.92 |
4.64 |
73.96 |
14.87 |
155.66 |
28.37 |
|
CV (%) |
- |
24.48 |
26.25 |
21.32 |
22.62 |
16.66 |
25.22 |
14.24 |
17.36 |
|
**and ns: Significant at P < 0.01 and insignificant, respectively. |
|||||||||
No competing interests reported.