
==== Front
Sci Rep
Sci Rep
Scientific Reports
2045-2322
Nature Publishing Group UK London

73093
10.1038/s41598-024-73093-5
Article
Manure-biochar compost mitigates the soil salinity stress in tomato plants by modulating the osmoregulatory mechanism, photosynthetic pigments, and ionic homeostasis
Kamal Mohammed Zia Uddin 12
http://orcid.org/0000-0002-6814-8816
Sarker Umakanta umakanta@bsmrau.edu.bd

3
Roy Siddhartha Kumar 4
Alam Mohammad Saiful 12
Azam Mohammad Golam 5
Miah Md. Yunus 1
Hossain Nazmul 6
Ercisli Sezai 7
Alamri Saud 8
1 https://ror.org/04tgrx733 grid.443108.a 0000 0000 8550 5526 Department of Soil Science, Faculty of Agriculture, Bangabandhu Sheikh Mujibur Rahman Agricultural University (BSMRAU), Gazipur-1706, Bangladesh
2 grid.443108.a 0000 0000 8550 5526 Institute of Climate Change and Environment, BSMRAU, Gazipur, 1706 Bangladesh
3 https://ror.org/04tgrx733 grid.443108.a 0000 0000 8550 5526 Department of Genetics and Plant Breeding, Faculty of Agriculture, Bangabandhu Sheikh Mujibur Rahman Agricultural University (BSMRAU), Gazipur-1706, Bangladesh
4 Khulna Public College, Boyra, Khulna Bangladesh
5 https://ror.org/01n09m616 grid.462060.6 0000 0001 2197 9252 Pulses Research Centre, Bangladesh Agricultural Research Institute, Ishurdi, 6620 Bangladesh
6 https://ror.org/04rswrd78 grid.34421.30 0000 0004 1936 7312 Department of Agronomy, Iowa State University, Iowa, Ames, 50010 USA
7 https://ror.org/03je5c526 grid.411445.1 0000 0001 0775 759X Department of Horticulture, Faculty of Agriculture, Ataturk University, Erzurum, 25240 Türkiye
8 https://ror.org/02f81g417 grid.56302.32 0000 0004 1773 5396 Department of Botany and Microbiology, College of Science, King Saud University, Riyadh, Saudi Arabia
20 9 2024
20 9 2024
2024
14 2192927 1 2024
13 9 2024
© The Author(s) 2024
2024
https://creativecommons.org/licenses/by/4.0/ Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if changes were made. The images or other third party material in this article are included in the article’s Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by/4.0/.
One of the main abiotic stresses that affect plant development and lower agricultural productivity globally is salt in the soil. Organic amendments, such as compost and biochar can mitigate the opposing effects of soil salinity (SS) stress. The purpose of this experiment was to look at how tomato growth and yield on salty soil were affected by mineral fertilization and manure-biochar compost (MBC). Furthermore, the study looked at how biochar (organic amendments) work to help tomato plants that are stressed by salt and also a mechanism by which biochar addresses the salt stress on tomato plants. Tomato yield and vegetative growth were negatively impacted by untreated saline soil, indicating that tomatoes are salt-sensitive. MBC with mineral fertilization increased vegetative growth, biomass yield, fruit yield, chlorophyll, and nutrient contents, Na/K ratio of salt-stressed tomato plants signifies the ameliorating effects on tomato plant growth and yield, under salt stress. Furthermore, the application of MBC with mineral fertilizer decreased H2O2, but increased leaf relative water content (RWC), leaf proline, total soluble sugar, and ascorbic acid content and improved leaf membrane damage, in comparison with untreated plants, in response to salt stress. Among the composting substances, T7 [poultry manure-biochar composting (PBC) (1:2) @ 3 t/ha + soil-based test fertilizer (SBTF)] dose exhibited better-improving effects on salt stress and had maintained an order of T7 > T9 > T8 > T6 in total biomass and fruit yield of tomato. These results suggested that MBC might mitigate the antagonistic effects of salt stress on plant growth and yield of tomatoes by improving osmotic adjustment, antioxidant capacity, nutrient accumulation, protecting photosynthetic pigments, and reducing ROS production and leaf damage in tomato plant leaves.

Supplementary Information

The online version contains supplementary material available at 10.1038/s41598-024-73093-5.

Keywords

Growth
Yield
Organic amendments
Salinity
Tomato
Subject terms

Salt
Abiotic
issue-copyright-statement© Springer Nature Limited 2024
==== Body
pmcIntroduction

One of the worst environmental pressures that seriously jeopardize agricultural sustainability and productivity is salinity1.2. The salinity of the soil is a worldwide issue that affects both developed and developing nations3. Future climate change scenarios are anticipated to worsen the salinity-induced deterioration of soil. These scenarios include increasing sea levels and their impacts on coastal regions, increasing temperatures, and rising evaporation. Soil salinization4 affects approximately 1,128 million hectares of land worldwide, and secondary salinization has added 77 million hectares of land5. Approximately 20% of all farmed and 33% of irrigated agricultural fields globally are affected by soil salinization6. Estimates of the global distribution of regions impacted by salt indicate that 50.8 million hectares are in Europe and 87.6 million hectares are in South Asia7. Comparably, 30% of Bangladesh’s farmland area roughly one million hectares is damaged by salinization to varied degrees8,9. Salinity-induced losses in crop production cause irrigated agriculture to lose USD 27.2 billion annually in economic value. Abiotic stress, such as salinity/drought simultaneously causes ion toxicity, osmotic stress10, and oxidative stress11,12, which affect the physiological and biochemical processes of many plants, including photosynthesis13, energy metabolism, plant water, nutrient uptake imbalance14, protein15, DNA, membrane damage16, etc., and ultimately affect the growth17 and yield of crops18. Osmotic stress in plants changes many enzymatic antioxidants, such as CAT, SOD, GOPX11, non-enzymatic antioxidants, such as carotenoids19,20, betalains21,22, ascorbic acids23,24, betaxanthins25,26, chlorophyll a27, betacyanins28, chlorophyll b29, xanthophylls30, beta-carotene31, phenolic and flavonoids, such as hydroxycinnamic acids32, hydroxybenzoic acids33, flavanols34, flavones35, flavanones36, flavonols37, etc. having high antiradical capacities38 which can manage the adverse impact of abiotic stresses. Under abiotic stress, the plant itself regulates different pathways to increase the application of these antioxidants39 to detoxify the ROS40 and mitigate environmental stress.

Furthermore, SS poses a serious risk to farmland soil production by adversely affecting microbial activity, nutrient availability, and soil characteristics41,42. There is usually a 30–50% yield loss, depending on the SS43. Plants have developed a variety of defense mechanisms to withstand salinity, including ion homeostasis and compartmentalization, osmoprotectant production, antioxidant activation, and the control of several stress-sensitive genes44. Various reclamation techniques, including chemical remediation, phytoremediation, soil water leaching, organic amendment, etc., can be used to improve the congenial environment that plants use to withstand stress. Thus, reducing the salt burden on crop growth in appropriate and economical ways has become a top concern in the development of technologies to guarantee global food security.

In salinity-degraded soil, soil management and agricultural production techniques depend on maintaining a suitable level of soil organic matter and guaranteeing the effective biological cycling of nutrients45. Organic amendments may be considered a sustainable method of lessening the effects of salinity stress on crops since they have both ameliorative effects and enhance the fertility status of saline soils46. Salinity stress can have significant effects on various crops. A high soil salinity level can lead to water stress and ion toxicity, decreasing water availability and increasing the rate of transpiration in plants47. In addition, high soil salinity causes ion homeostasis to be disrupted, resulting in an accumulation of toxic ions such as sodium and chloride in plant tissues48. According to several studies, organic matter’s improved physical, chemical, and biological qualities could help reduce salt stress49,50. However, adding organic matter won’t be able to sustainably improve this deteriorated soil in the tropics because of the natural quick degradation of organic matter and poorly stable chemicals51. Almond shell-derived biochar (ASB) induced changes in soil microbial composition and communities52. The use of more stable carbon compounds, such as carbonized materials, is an alternative to organic additions. Because of its strong Na+ adsorption potential and relatively stable nature, the biomass product known as “biochar,” which is produced through anaerobic or limited oxygen condition pyrolysis, may be able to reduce salinity stress53,54. Because of its increased ability to add Ca2+ and Mg2+, improve the physical characteristics of the soil, including aggregate stability, porosity, saturated hydraulic conductivity, and bulk density, and consequently improve Na+ leakage, biochar becomes an efficient remediation approach in salty soil55. However, due to its expensive and unusual production methods, there is limited data available on the practice of biochar as a viable soil additive for salty soils. For agriculture that is susceptible to salinity stress, appropriate and economical methods of minimizing the negative effects of salt stress on crop growth have thus become crucial. The study reported that biochar-manure compost (poultry manure mixed with biochar (1:3) in conjunction with pyroligneous solution had significant positive effects on salinity reduction in cereals like maize productivity54. However, further research is needed to elucidate the effects of different manure and biochar compost combinations on salt-sensitive vegetables, such as tomatoes. Thus, we hypothesize that composting manure and biochar can reduce salt stress and will be a cost-effective strategy for tomato growers in saline-sensitive locations. Different crops have varying levels of salt tolerance. Some crops, like barley, sugar beet, and spinach, are more salt-tolerant and can withstand higher levels of salinity. On the other hand, crops such as tomatoes, eggplant, beans, and many fruit trees are more sensitive to salt stress56.

The majority of Bangladesh’s salinity-affected areas experience seasonal flooding from salt water, which varies the degree of salinization of the soil. The majority of the land remains fallow during the dry season because of the elevated salinity of the soil and the shortage of high-quality water57. One of the most significant and nutrient-dense foods grown in arid, salinity-degraded agricultural regions maybe tomatoes. The Solanaceae family includes the tomato (Lycopersicon esculentum L.), most of which are cultivated in nearly every part of the globe. It ranks as the second most significant crop among the 24 vegetable crops. Tropical, subtropical, and even temperate regions of the world are where it is primarily farmed58. Due to their flavor and nutritional worth, tomatoes are an essential part of the human diet. Because of its short root structure, high stomatal conductance, and wide diversity of leaf types, the tomato plant is the most vulnerable to salt stress59. If we are to convert a large number of saline soils under circumstances of intensive agriculture and adapt to climate change, we must comprehend the impacts of salt stress and find measures to decrease it. There is scarce information on the effects of salt stress on tomatoes as well as the affordable and practical methods of lowering the effects of salt by organic amendment. This study aims to assess the possibility of mitigating SS stress with the addition of compost made of manure and biochar, as well as the mechanisms involved.

Results

The effects of different fertilizers - biochar compost and organic materials on tomato plants under SS stress were studied, including vegetative development, photosynthetic pigmentation, oxidative damage, and their osmotic adjustment properties, leaf nutrient content, biomass, and fruit yield characteristics.

Growth traits

Tomato plants grown in untreated saline soil had lower plant height (PH) (40.20 cm), number of branches per plant (NBP) (2.89), number of leaves per plant (NLP) (19.17), and leaf area (LA) (49.90 cm2), hence exhibiting significant inhibition of plant growth (Fig. 1). The application of different manures, rice husk biochar, and their composting significantly improved the growth characteristics of salt-stressed tomato plants (Fig. 1).  

Fig. 1 Manure and biochar compost (MBC) application impacts on salt-stressed tomato plant growth (A) PH, (B) NBP, (C) NLP, and (D) LA (cm2). The value indicates the mean (± SD); p < 0.05; n = 9; Different letters indicate significant differences among treatments (p < 0.05) by Tukey’s HSD test. Treatments combinations are T1 = SS; T2 = soil-based test fertilizer (SBTF); T3 = B @ 3 t/ha + SBTF; T4 = P @ 3 t/ha + SBTF; T5 = C @ 3 t/ha + SBTF; T6 = poultry manure-biochar composting (PBC) (1:1) @ 3 t/ha + SBTF; T7 = PBC (1:2) @ 3 t/ha + SBTF; T8 = cow dung-biochar composting (CBC) (1:1) @ 3 t/ha + SBTF; T9 = CBC (1:2) @ 3 t/ha + SBTF.

Treatment T8 [cow dung-biochar composting (CBC) (1:1) @ 3 t/ha + SBTF] showed the highest PH (76.09 cm) and statistically similar data in T3 (B @ 3 t/ha + SBTF), T6 (poultry manure-biochar composting (PBC) (1:1) @ 3 t/ha + SBTF), T7 [PBC (1:2) @ 3 t/ha + SBTF] and T9 [CBC (1:2)) @ 3 t/ha + SBTF] treatment (Fig. 1A). Treatments of T2, T3, T4, T5, T6, T7, T8, and T9 increased PH by 26.83%, 69.92%, 50.50%, 51.00%, 80.27%, 84.34%, 91.26%, and 86.58%, respectively, compared with T1 treatment (Table 1). Manure-biochar compost treatment T9 showed the maximum NBP (7.27), and responses were similar for all organic amendments (Fig. 1B). No significant changes in NBP were found after a single fertilization compared to the control NBP increased by 96.30-151.73% in MBC treatments compared to the control (Table 1). Meanwhile, the T9 treatment showed maximum leaf proliferation i.e. NLP (34.17) and LA (92.77 cm2) (Fig. 1C). Organic amended plants showed more NLP i.e., in T3 (55.65%), T4 (26.05%), T5 (34.87%), T6 (57.39%), T7 (76.61%), T8 (65.22%) and T9 (78.26%) than plots with untreated saline soil (Table 1). Likewise, the result in Fig. 1D revealed that LA of organic amended salt-stressed plants increased in T3 (53.49%), T4 (39.68%), T5 (46.45%), T6 (59.73%), T7 (81.77%), T8 (65.78%) and T9 (85.92%) compare to T1 (Table 1).

Table 1 Percent change in growth, yield, and yield attributes of tomato plants treated with manure and biochar compost in salt-stressed soil compared to untreated control.

Parameters	Treatments	
	T2	T3	T4	T5	T6	T7	T8	T9	
PH	26.83	69.92	50.50	51.00	80.27	84.34	91.26	86.58	
NBP	15.47	90.53	55.88	78.98	105.54	142.49	96.30	151.73	
NLP	12.17	55.65	26.05	34.87	57.39	76.61	65.22	78.26	
LA	13.83	53.49	39.68	46.45	59.73	81.77	65.78	85.92	
NEFC	17.95	64.10	15.38	43.59	94.87	102.05	130.77	128.21	
NFC	6.96	23.41	29.11	18.99	40.38	42.91	56.96	48.73	
NFP	24.83	100.00	47.83	68.50	170.02	186.65	260.97	238.00	
SFW	15.50	24.91	29.24	19.69	23.87	27.58	31.21	28.54	
FWP	44.50	150.49	91.74	103.83	235.40	264.62	374.53	337.69	
FY	44.50	150.49	91.74	103.83	235.40	264.62	374.53	337.69	
TDM	38.34	88.47	56.98	74.94	92.67	121.69	107.22	117.98	
PH = Plant height, NBP = Number of branches per plant, NLP = Number of leaves per plant, LA = Leaf area, NEFC = Effective flower cluster per plant, NFC = Number of fruit per cluster, NFP = Number of fruit per plant, SFW = Single fruit weight, FWP = Fruit weight per plant. FY = Fruit yield, TDM = Total dry mass. Treatments combinations are T 1  = SS; T 2  = SBTF; T 3  = B @ 3 t/ha + SBTF; T 4  = P @ 3 t/ha + SBTF; T 5  = C @ 3 t/ha + SBTF; T 6  = PBC (1:1) @ 3 t/ha + SBTF; T 7  = PBC (1:2) @ 3 t/ha + SBTF; T 8  = CBC (1:1) @ 3 t/ha + SBTF; T 9  = CBC (1:2) @ 3 t/ha + SBTF.

Yield and yield features

Tomato fruit yield and yield traits were calculated on abandoned salt-stressed (control) plants and various organic amended plants (Table 2; Fig. 2). Yield-promoting parameters of salt-stressed tomato plants were significantly influenced by the application of manure-biochar compost, its raw material plus soil test-based (SBTF) fertilizer (Table 2). Plants grown under salt stress (control treatment) showed the lowest number of effective flower cluster plant−1 (NEFC) (3.25), number of fruit cluster−1 (NFC) (2.57), number of fruit plant−1 (NFP) (8.63), and single fruit weight (SFW) (31.18 g). In the T9 treatment, the maximum NEFC (7.50), NFC (4.13), NFP (31.13), and SFW (40.92 g) were recorded. Notably, NEFC, NFC, NFP, and SFW increased significantly by 94.87–130.77%, 40.38–56.96%, 170.02–260.97%, 23.87–31.21%, respectively under the MBC treatments over the absolute saline soil plants (Table 1). The manure-biochar compost treatments showed better yield traits data than the cow dung, poultry manure, and biochar alone.

Table 2 Manure and biochar compost (MBC) application impacts on yield and yield attributes of salt-stressed tomato plant.

Treatments	NEFC	NFC	NFP	SFW (g)	
T1	3.25 ± 0.25 e	2.63 ± 0.40 d	8.61 ± 1.07 c	31.18 ± 1.05 b	
T2	3.83 ± 0.28 de	2.82 ± 0.16 cd	10.77 ± 0.25 c	36.02 ± 3.28 ab	
T3	5.33 ± 0.58 bcd	3.25 ± 0.25 abcd	17.25 ± 0.90 bc	38.95 ± 2.57 ab	
T4	4.48 ± 0.64 de	3.05 ± 0.32 bcd	13.55 ± 0.83 c	35.82 ± 4.25 ab	
T5	4.67 ± 0.58 cde	3.13 ± 0.31 bcd	14.53 ± 1.26 c	37.32 ± 4.80 ab	
T6	6.33 ± 0.50 abc	3.70 ± 0.35 abc	23.29 ± 0.89 ab	38.63 ± 2.00 ab	
T7	7.50 ± 0.68 a	4.13 ± 0.35 a	31.13 ± 5.23 a	40.92 ± 1.53 a	
T8	6.57 ± 0. 48ab	3.76 ± 0.06 abc	24.72 ± 2.38 ab	39.78 ± 4.12 ab	
T9	7.42 ± 0. 52a	3.92 ± 0.39 ab	29.15 ± 4.60 a	40.08 ± 1.45 ab	
Significance level	< 0.001***	< 0.001***	< 0.001***	< 0.043*	
The value indicates the mean (± SD); p < 0. 05; n  = 9. Different letters indicate significant differences among treatments ( p  < 0.05) by Tukey’s HSD test. Treatments combinations are T 1  = SS; T 2  = SBTF; T 3  = B @ 3 t/ha + SBTF; T 4  = P @ 3 t/ha + SBTF; T 5  = C @ 3 t/ha + SBTF; T 6  = PBC (1:1) @ 3 t/ha + SBTF; T 7  = PBC (1:2) @ 3 t/ha + SBTF; T 8  = CBC (1:1) @ 3 t/ha + SBTF; T 9  = CBC (1:2) @ 3 t/ha + SBTF; . NEFC = Effective flower cluster per plant, NFC = Number of fruit per cluster, NFP = Number of fruit per plant, SFW = Single fruit weight.

No significant changes in tomato plant yield characteristics were observed with absolutely SBTF-based fertilization compared to unamended saline soils.

The data in Fig. 2 showed that the treatments of different organic amendments have a significant positive effect on the fruit weight plant−1 (FWP) and the total fruit yield of salt-stressed tomato plants. The T7 treatment of poultry manure-biochar compost (1:2) recorded the highest FWP (1271.64 g) and the highest total tomato yield (FY) (40.69 t/ha). Meanwhile, tomato fruit yield was destructively affected by salt stress, showing minimum FWP (267.98 g) and total tomato yield (8.58 t/ha) in T1 treated plants (Fig. 2(A) and Fig. 2(B)). Other plants treated with manure-biochar compost showed similar improvements in FWP and total tomato yield and were represented in T6 (898.79 g and 28.76 t/ha, respectively), T8 (977.08 g, and 31.27 t/ha, respectively) and T9 (1172.91 g and 37.53 t/ha, respectively) treatment. The total FY under the MBC treatments increased by 235.40–374.53%, compared to absolute saline soil plants (Table 1). Manure-biochar compost-treated plants produced higher tomato yields than other organic amended plants under salt stress.

Fig. 2 Effects of manure and biochar compost (MBC) on (A) tomato fruit weight plant−1 (g) and (B) tomato yield (t/ha) of tomato plant grown in saline soil. The value indicates the mean (± SD); p < 0.05; n = 9; Different letters indicate significant differences among treatments (p < 0.05) by Tukey’s HSD test. Treatments combinations are T1 = SS; T2 = SBTF; T3 = B @ 3 t/ha + SBTF; T4 = P @ 3 t/ha + SBTF; T5 = C @ 3 t/ha + SBTF; T6 = PBC (1:1) @ 3 t/ha + SBTF; T7 = PBC (1:2) @ 3 t/ha + SBTF; T8 = CBC (1:1) @ 3 t/ha + SBTF; T9 = CBC (1:2) @ 3 t/ha + SBTF.

Biomass yield of tomato plant

Manure-biochar compost and its organic matter had significant effects on above-ground biomass (FAGB), dry above-ground biomass (DAGB), fresh below-ground biomass (FBGB), dry below-ground biomass (DBGB), and total dry mass (TDM) (Table 3). Untreated saline soil showed significantly decreased tomato biomass yield. The organic amended plants showed higher fresh and dry biomass compared to abandoned saline soil (control treatment). The T7-treated tomato plant gave the maximum FAGB, FBGB, DAGB, DBGB, and TDM production (72.73 g, 18.84 g, 15.11 g, 7.61 g and 22.71 g, respectively). Like FAGB, FBGB, DAGB, DBGB, and TDM were produced in T9 (71.15 g, 18.56 g, 15.04 g, 7.30 g, and 22.34 g, respectively), T8 (69.32 g, 17.52 g, 14.05 g, 7.18 g, and 21.23 g, respectively) and T6 (68.20 g, 16.83 g, 13.03 g, 6.71 g, and 19.74 g, respectively) treatment. TDM was found significantly higher by 92.67–121.69%, under the MBC treatments over control plants (Table 1).

Table 3 Manure and biochar compost (MBC) effects on tomato plants cultivated in saline soil in terms of biomass yield.

Treatments	FAGB (g)	FBGB (g)	DAGB (g)	DBGB (g)	TDM (g)	
T1	34.63 ± 1.80 c	9.68 ± 0.36 e	7.89 ± 0.95 e	2.36 ± 0.09 e	10.25 ± 1.04 g	
T2	39.46 ± 3.76 c	12.47 ± 1.15 d	9.86 ± 0.57 de	4.32 ± 0.67 d	14.17 ± 0.46 f	
T3	58.43 ± 1.86 b	15.75 ± 0.70 bc	12.92 ± 0.36 abc	6.39 ± 0.36 abc	19.31 ± 0.35 cd	
T4	51.93 ± 1.40 b	13.99 ± 0.65 cd	10.87 ± 0.90 cd	5.59 ± 0.29 cd	16.46 ± 0.61 ef	
T5	53.65 ± 5.52 b	14.35 ± 0.75 cd	11.83 ± 1.43 bcd	6.09 ± 0.22 bc	17.92 ± 1.21 de	
T6	68.20 ± 2.48 a	16.83 ± 0.62 ab	13.03 ± 0.62 abc	6.71 ± 0.20 cde	19.74 ± 0.80 bcd	
T7	72.73 ± 4.29 a	18,84 ± 1.17 a	15.11 ± 0.25 a	7.61 ± 0.94 a	22.71 ± 1.18 a	
T8	69.32 ± 1.81 a	17.52 ± 0.78 ab	14.05 ± 0.57 ab	7.18 ± 0.17 ab	21.23 ± 0.43abc	
T9	71.15 ± 2.78 a	18.56 ± 0.31 a	15.04 ± 1.06 a	7.30 ± 0.62 ab	22.34 ± 1.76ab	
Significance level	< 0.001***	< 0.001***	< 0.001***	< 0.001***	< 0.001***	
The value indicates the mean (± SD); P ≤ 0.05; n = 9; Different letters indicate significant differences among treatments by Tukey’s HSD test; FAGB = Fresh above-ground biomass, FBGB = Fresh below-ground biomass, DAGB = Dry above-ground biomass, DBGB = Dry below-ground biomass, TDM = Total dry mass; Treatments combinations are T 1  = SS; T 2  = SBTF; T 3  = B @ 3 t/ha + SBTF; T 4  = P @ 3 t/ha + SBTF; T 5  = C @ 3 t/ha + SBTF; T 6  = PBC (1:1) @ 3 t/ha + SBTF; T 7  = PBC (1:2) @ 3 t/ha + SBTF; T 8  = CBC (1:1) @ 3 t/ha + SBTF; T 9  = CBC (1:2) @ 3 t/ha + SBTF.

Photosynthetic pigment content

The application of manure-biochar compost and its raw organic material increased the Chlorophyll (Ch) of salt-stressed tomato plants (Table 4). In comparison between both types of organic amendment applications (alone and compost), manure-biochar compost gave more positive results. SS stress significantly impaired the photosynthetic pigment of tomato plants and the lowest total Ch Ch a, and b contents were found in the T1 treatment (0.73, 0.51, and 0.22 mg/g, respectively). The application of different organic amendments considerably diminished the salt-stress impacts and improved pigmentation. In the T7 treated plant, the maximum total Ch and Ch a content (2.03 and 1.58 mg/g, respectively) were calculated. Likewise, salt-stressed plants in T8 (CBC (1:1) @ 3 t/ha + SBTF) showed the maximum Ch b (0.49 mg/g) content.

Table 4 Effects of manure and biochar compost (MBC) on the chlorophyll content of tomato leaf at 68 days after transplanting (DAT) grown in saline soil.  The value indicates the mean (± SD); p < 0. 05; n  = 9.

Treatments	Ch a
(mg/g)	Ch b
(mg/g)	Total Ch
(mg/g)	
T1	0.51 ± 0.11 f	0.22 ± 0.02 c	0.73 ± 0.13 e	
T2	0.80 ± 0.12 e	0.28 ± 0.03 bc	1.08 ± 0.24 d	
T3	1.30 ± 0.18 bcd	0.40 ± 0.10 ab	1.70 ± 0.23 bc	
T4	1.11 ± 0.07 d	0.45 ± 0.08 a	1.57 ± 0.15 c	
T5	1.18 ± 0.24 cd	0.48 ± 0.05 a	1.66 ± 0.20 c	
T6	1.40 ± 0.04 abc	0.45 ± 0.07 a	1.85 ± 0.10 abc	
T7	1.58 ± 0.05 a	0.45 ± 0.11 a	2.03 ± 0.09 a	
T8	1.53 ± 0.13 ab	0.49 ± 0.16 a	2.02 ± 0.17 ab	
T9	1.51 ± 0. 22 ab	0.46 ± 0.04 abc	1.97 ± 0.22 abc	
Significance level	< 0.001***	˂ 0.021*	< 0.001***	
Different letters indicate significant differences among treatments ( p  < 0.05) by Tukey’s HSD test. Treatments combinations are T 1  = SS; T 2  = SBTF; T 3  = B @ 3 t/ha + SBTF; T 4  = P @ 3 t/ha + SBTF; T 5  = C @ 3 t/ha + SBTF; T 6  = PBC (1:1) @ 3 t/ha + SBTF; T 7  = PBC (1:2) @ 3 t/ha + SBTF; T 8  = CBC (1:1) @ 3 t/ha + SBTF; T 9  = CBC (1:2) @ 3 t/ha + SBTF; Chlorophyll = Ch; chlorophyll a  = Ch a ; chlorophyll b  = Ch b.

Response in osmoregulatory physiological traits

Salt stress considerably declined the RWC of tomato plant leaves, the lowest content in the T1 treatment (63.48%). Adding organic amendment plus SBTF fertilizer increased the RWC (%) of salt-stressed plant leaves, with the superiority of manure-biochar composted treatments (Table 5).

Table 5 Effects of manure and biochar compost (MBC) on RWC, water saturation deficit (WSD), and water retention capacity (WRC) of tomato plants at 65 DAT grown in saline soil. .

Treatments	RWC (%)	WSD (%)	WRC (%)	
T1	63.48 ± 3.95 d	36.52 ± 3.95 a	10.90 ± 2.80 a	
T2	70.74 ± 3.83 cd	29.26 ± 3 0.83 ab	9.54 ± 0.43 ab	
T3	82.96 ± 3.18 ab	17.04 ± 3.18 cd	8.38 ± 0.39 bc	
T4	79.54 ± 3.83 bc	20.46 ± 3.83 bc	8.64 ± 0.43 bc	
T5	82.04 ± 1.97 ab	17.96 ± 1.97 cd	8.43 ± 0.55 bc	
T6	85.72 ± 2.49 ab	14.28 ± 2.49 cd	8.03 ± 0.86 bc	
T7	87.29 ± 3.25 ab	12.71 ± 3.25 d	8.28 ± 0.92 c	
T8	86.69 ± 4.30ab	13.31 ± 4.30 cd	2.38 ± 0.15 c	
T9	88.33 ± 1.95a	11.67 ± 1.95 cd	4.75 ± 0.44 c	
Significance level	< 0.001***	< 0.001***	< 0.001***	
The value indicates the mean (± SD); p < 0. 05; n  = 9. Different letters indicate significant differences among treatments ( p  < 0.05) by Tukey’s HSD test. Treatments combinations are T 1  = SS; T 2  = SBTF; T 3  = B @ 3 t/ha + SBTF; T 4  = P @ 3 t/ha + SBTF; T 5  = C @ 3 t/ha + SBTF; T 6  = PBC (1:1) @ 3 t/ha + SBTF; T 7  = PBC (1:2) @ 3 t/ha + SBTF; T 8  = CBC (1:1) @ 3 t/ha + SBTF; T 9  = CBC (1:2) @ 3 t/ha + SBTF.

The application of the T9 (CBC (1:2) @ 3 tha−1 + SBTF) treatment showed the highest retrieval of RWC (88.33%), followed by T3 (82.96%), T6 (85.72%), T7 (87.29%) and T8 (86.69%). In contrast, absolute SS treatment T1 showed significant increases in water saturation deficit (WSD) and water retention capacity (WRC) (36.52%, and 10.90%, respectively) (Table 5). In the organic amended T9 treatment, the lowest salinity-induced WSD (11.67%) and WRC were in T8 (2.38%) treatment (Table 5).

The concentration of H2O2, MDA, and electrolyte leakage (EL) in tomato plant leaf

As shown in Table 6, H2O2, malondialdehyde (MDA) content, and electrolyte leakage (EL) in-tomato plant leaves were significantly affected by the SS and application of manure-biochar compost. The abandoned salinity treatment T1 treated plant leaves showed the significantly highest H2O2 and MDA contents (28.87 nmol/g and 15.49 nmol/g, respectively). However, the use of compost made of dung and charcoal significantly decreased their accumulation. The greatest reduction in H2O2 content was observed in the T7 amended plant, while the MDA content was reduced in T9 treated plants. Salt stress caused severe impairment of membrane stability by enhancing the electrolyte leakage (34.56%) in tomato plant leaves (Table 6). This reduction was significantly mitigated by the application of MBC amendment and SBTF-based fertilizer. Minimal electrolyte leakage (13.80%) was calculated for T9 treatment.

Table 6 Impact of manure and biochar compost (MBC) on H2O2, MDA, and electrolyte leakage (EL) of salt-stressed tomato plant leaves at 60 DAT.

Treatments	H2O2
(nmol/g)	MDA
(nmol/g)	EL (%)	
T1	28.87 ± 1.72 a	15.49 ± 1.67 a	34.56 ± 2.15 a	
T2	24.19 ± 3.63 b	13.89 ± 0.85 ab	30.91 ± 1.43 ab	
T3	14.98 ± 1.20 d	7.65 ± 1.20 cd	21.40 ± 0.66 cd	
T4	19.38 ± 0.71 c	12.69 ± 1.70 ab	24.53 ± 2.09 bc	
T5	20.02 ± 1.51 c	11.15 ± 1.86 bc	24.25 ± 1.20 bc	
T6	12.48 ± 1.56 de	6.58 ± 0.91 d	17.51 ± 1.81 de	
T7	10.77 ± 1.15 e	5.64 ± 0.15 d	15.18 ± 2.59 de	
T8	13.76 ± 1.44 de	6.00 ± 0.57 d	17.26 ± 2.48 de	
T9	12.18 ± 1.13 de	4.91 ± 0.62 d	13.80 ± 1.21 e	
Significance level	< 0.001***	< 0.001***	< 0.001***	
The value indicates the mean (± SD); p < 0. 05; n  = 9. Different letters indicate significant differences among treatments ( p  < 0.05) by Tukey’s HSD test. Treatments combinations are T 1  = SS; T 2  = SBTF; T 3  = B @ 3 t/ha + SBTF; T 4  = P @ 3 t/ha + SBTF; T 5  = C @ 3 t/ha + SBTF; T 6  = PBC (1:1) @ 3 t/ha + SBTF; T 7  = PBC (1:2) @ 3 t/ha + SBTF; T 8  = CBC (1:1) @ 3 t/ha + SBTF; T 9  = CBC (1:2) @ 3 t/ha + SBTF.

The statistically identical finding was recorded in treatment T6 (17.51%), T7 (15.18%), and T8 (17.26%). No significant improvement in electrolyte leakage was observed in plants treated with SBTF fertilizer alone.

Leaf proline, ascorbic acid, and soluble sugar content in tomato plant leaves

Proline biosynthesis in tomato plants was activated by salt stress. The lowest proline accumulation (11.85 mg/g) was assessed from SS (T1) tomato plant leaves (Table 7). MBC plus SBTF-based fertilizer-amended tomato plant leaves had lower proline accretion under salt-stressed conditions. T7-treated plant leaves provided the maximum proline accretion (28.16 mg/g). Similar Statistics were obtained in T6, T8 and T9 amended plants (22.32, 21.95, and 26.86 mg/g, respectively). The application of various organic amendments significantly augmented the AA of the leaves of salinity-stressed tomato plants, among which MBC-treated plots performed better (Table 7). The application of the T9 [CBC (1:2) @ 3 t/ha + SBTF] treatment distinctly enhanced the ascorbic acid concentration (17.01 mg/g) of tomato plant leaves. SS (T1 treatment) plant leaves have the minimum amount of AA (8.48 mg/g). Similarly, the TSS content (21.63 mg/g) in tomato plant leaves decreased pointedly in abandoned salt-stress T1 treatment.

Table 7 Effects of manure and charcoal compost (MBC) on the levels of total soluble sugar (TSS), ascorbic acid (AA), and proline in the leaves growing in salty soil.

Treatments	Proline
(mg/g)	Ascorbic acid
(mg/g)	Total soluble sugar
(mg/g)	
T1	11.85 ± 1.78 d	8.48 ± 1.53 d	21.63 ± 1.26 f	
T2	13.09 ± 1.93 d	9.42 ± 1.96 cd	22.74 ± 1.38 ef	
T3	18.36 ± 1.85 cd	13.69 ± 1.36 abc	27.64 ± 2.39 cde	
T4	17.02 ± 1.94 cd	12.60 ± 1.96 abcd	26.44 ± 1.37 def	
T5	16.84 ± 2.15 cd	11.81 ± 1.87 bcd	27.32 ± 2.09 cde	
T6	22.32 ± 3.01 abc	14.67 ± 2.44 ab	30.48 ± 2.40 cd	
T7	28.16 ± 1.57 a	15.71 ± 2.06 ab	36.94 ± 1.82 ab	
T8	21.95 ± 1.99 abc	14.39 ± 0.88 abc	32.02 ± 1.32 bc	
T9	26.86 ± 1.51 ab	17.01 ± 1.88 a	39.66 ± 2.70 a	
Significance level	< 0.001***	< 0.001***	< 0.001***	
The value indicates the mean (± SD); p < 0. 05; n  = 9. Different letters indicate significant differences among treatments ( p  < 0.05) by Tukey’s HSD test. Treatments combinations are T 1  = SS; T 2  = SBTF; T 3  = B @ 3 t/ha + SBTF; T 4  = P @ 3 t/ha + SBTF; T 5  = C @ 3 t/ha + SBTF; T 6  = PBC (1:1) @ 3 t/ha + SBTF; T 7  = PBC (1:2) @ 3 t/ha + SBTF; T 8  = CBC (1:1) @ 3 t/ha + SBTF; T 9  = CBC (1:2) @ 3 t/ha + SBTF.

MBC and its raw organic materials amended salt-stress tomato plant leaves showed a significant decline in total soluble sugar content. Plants grown in T9 (CBC (1:2) @ 3 t/ha + SBTF) amended saline soil produce the highest leaf TSS concentration (39.66 mg/g).

Nutritional status of the tomato

Table 8 displays the results of the effects of MBC and its sole raw materials on the nutrients of tomato leaves under salt stress. Statistically noteworthy variances in the N, P, K, Na content, and K/Na ratio were found in organic amended tomato leaves. The N and P contents were highest in tomato leaves under T7 treatment (18.23 and 4.90 mg/g, respectively). While the T9-treated plant leaves in SS condition gave the maximum K content (14.80 mg/g). Similar leaves K concentrations were found in T6 (13.57 mg/g), T7 (14.74 mg/g) and T8 (13.30 mg/g) amended salt-stressed tomato plants. Untreated SS tomato plants in T1 treatment showed the lowest leaf nutrient contents of N, P, and K (9.29, 2.20, and 7.16 mg/g, respectively). Various organic amendments negatively influence the leaf Na accumulation. The T9 amended SS plant leaves exhibited the minimum Na concentration (6.03 mg/g), while the maximum Na content (12.47 mg/g) in T1 treatment.

Table 8 Manure and biochar compost (MBC) effects on tomato leaf nutritional content at 65 DAT in saline soil. .

Treatments	N
(mg/g)	P
(mg/g)	K
(mg/g)	Na
(mg/g)	Na/K ratio	
T1	9.29 ± 1.38 b	2.20 ± 0.32 c	7.16 ± 1.58 e	17.47 ± 0.60 a	2.54 ± 0.67 a	
T2	11.90 ± 1.57 ab	2.92 ± 0.54 bc	9.00 ± 0.59 de	15.00 ± 0.79 b	1.67 ± 0.79 b	
T3	14.76 ± 2.53 ab	3.80 ± 0.37 abc	11.55 ± 0.44 bc	9.30 ± 1.02 cd	0.80 ± 0.07 c	
T4	13.89 ± 1.64 ab	3.92 ± 0.66 abc	10.51 ± 0.59 cd	10.09 ± 0.31 c	0.96 ± 0.08 c	
T5	14.23 ± 0.97 ab	4.04 ± 0.87 ab	10.30 ± 0.76 cd	10.28 ± 1.10 c	1.00 ± 0.10 bc	
T6	17.34 ± 2.22 a	4.31 ± 0.36 ab	13.57 ± 0.68 ab	8.23 ± 0.23 cde	0.61 ± 0.03 c	
T7	18.23 ± 1.03 a	4.90 ± 0.56 a	14.74 ± 0.40 a	6.35 ± 0.48 e	0.43 ± 0.03 c	
T8	16.97 ± 1.76 a	4.14 ± 0.26 ab	13.30 ± 0.62 ab	7.07 ± 1.01 de	0.53 ± 0.06 c	
T9	17.79 ± 2.61a	4.72 ± 0.39 ab	14.80 ± 0.18 a	6.03 ± 0.60 e	0.41 ± 0.04 c	
Significance level	< 0.001***	< 0.001***	< 0.001***	< 0.001***	< 0.001***	
The value indicates the mean (± SD); p < 0. 05; n  = 9. Different letters indicate significant differences among treatments ( p  < 0.05) by Tukey’s HSD test. Treatments combinations are T 1  = SS; T 2  = SBTF; T 3  = B @ 3 t/ha + SBTF; T 4  = P @ 3 t/ha + SBTF; T 5  = C @ 3 t/ha + SBTF; T 6  = PBC (1:1) @ 3 t/ha + SBTF; T 7  = PBC (1:2) @ 3 t/ha + SBTF; T 8  = CBC (1:1) @ 3 t/ha + SBTF; T 9  = CBC (1:2) @ 3 t/ha + SBTF.

The K/Na ratio of manure-biochar compost and its raw material and/or SBTF-fertilized plants increased compared to abandoned saline soils (Table 8).

Discussion

One of the biggest risks to crop growth is soil salinization, which lowers crop yields significantly all over the world60. The findings demonstrated that tomato plant development and biomass productivity were significantly reduced by soil salinization (Fig. 1; Table 3). This might be because low turgor pressure brought on by salinity prevents cells from expanding, which lowers shoot development and growth. Elevated salt stress has the potential to trigger the synthesis of inhibitors such as abscisic acid and impede the development of growth promoters for plants61. Furthermore, salinity inhibits root system growth, altering its morphology and physiology. These modifications result in altered water and ion absorption, ultimately reducing plant growth62. The decrease in compound leaves and leaf area may be due to the salt-induced osmotic effect, which reduces the availability of water and nutrients to the roots and eventually disturbs the plant tissues. This would lead to a decrease in meristematic tissue activity and cell expansion63. Merely applying chemical fertilizer cannot compensate for the decrease in biomass production brought on by salt. The application of MBC and its organic components reduced the impacts of salt stress and led to a noteworthy increase in the vegetative attributes of tomato plants under SS stress, including PH, NBP, NLP, LA, and biomass, demonstrating its processes of mitigating salt stress (Figs. 1 and 2; Table 3). The growth and biomass output of tomato plants under salt stress were enhanced by the application of organic amendments, which also increased the soil’s organic matter and increased hydraulic conductivity, nutrient availability, soil bulk density, and soil water hold capacity64. Improved ionic balance and stressed plants’ physiological efficiency are guaranteed by increased leaf proliferation in tomato plants grown in salinized soil amendments65. Noteworthy differences among traits were demonstrated for ANOVA of our study which were corroborative to the results of agronomic traits of lentil66, field pea67, mungbean68,69, Zea mays70,71, Oryza sativa72–77, Amaranthus spp78,79.

Under salt stress, tomato plant leaves’ photosynthetic pigments were less effective (Table 4). However, because SS denaturates the enzymes that synthesize chlorophyll content, it produces oxidative stress in chloroplasts and lowers chlorophyll content80. Because of the salt-induced osmotic imbalance in this investigation, SS dramatically decreased leaf RWC (Table 5), which in turn reduced the tomato plants WSD and WUC. However, the application of MBC greatly boosted the leaf RWC. This could be because organic amendments have a stronger ameliorating effect and create ideal soil conditions for higher water absorption under SS. Salt stress positively enhanced the oxidative stress markers MDA and H2O2 in tomato plants (Table 6). High accumulation of H2O2 induces oxidative stress, which disrupts plant physiological processes, leading to growth and yield losses. SS directly affects higher membrane damage, called MDA accumulation, which plays a crucial role in increasing the EL of tomato plant leaves (Table 6). SS increases ROS production, thereby damaging cell membranes and causing a significant increase in EL81. The substantial reduction in EL in MBC-treated tomato plants indicates a better-alleviating effect of compost treatment, which was associated with lower MDA and H2O2 (Table 6)2. Under SS, various organic amendments reduced ROS production, ultimately alleviating MDA accusation by reducing oxidative stress in tomato plants1. Tomato plants regulate and increase osmolyte biosynthesis and maintain Na+ refflux to counteract the toxic effects of salt stress64,82. In the current study, the concentration of proline, soluble sugar, and ascorbic acid in tomato leaves of salt-stressed tomato plants was lower, indicating lower adaptability (Table 7). A key component of plants’ defense mechanisms against salt stress is osmoregulation. Under salt stress, plants create osmotically active solutes (like proline) to promote protein biosynthesis or metabolism26 and balance water potential83 to protect themselves from abiotic stress. To control osmotic pressure, a significant amount of osmolytes, such as soluble carbohydrates and proline, build up in the cytoplasm and other organelles59. The application of MBC resulted in higher proline biosynthesis in the leaves of tomato plants exposed to SS (Table 7), suggesting that balanced osmosis protects their photosynthetic machinery. In addition, amended tomato plants can accumulate larger amounts of proline to scavenge ROS, reduce oxidative damage, and protect cell membranes from the negative effects of salt stress.

Higher proline accumulation in MBC-treated soil may be due to salt-induced increases in N content and metabolism84. Under saline conditions, MBC and its raw materials increased the content of alternative important osmolytes, namely TSS, in the tomato plant leaves (Table 7). A substantial increase in sugar accumulation in amended plants indicates effective protection of tomato plants exposed to salt stress. Total soluble sugar plays key roles in many physiological and biochemical processes, including chlorophyll content, scavenging of ROS, and induction of damaging salt-stress conditions in adaptive pathways85, Studies have shown that proline and ascorbic acid are potent antioxidants that can scavenge various types of ROS and protect cells from oxidative damage86. Salt-stressed tomato plants treated with MBC showed higher ascorbic acid (AsA) accumulation (Table 7). A higher concentration of AsA indicates its better detoxification capacity against salt-induced ROS production.

Salt stress exerts a notable inhibitory effect on various physiological processes, including seed germination, lateral root growth, and biomass generation, leading to significant yield loss. A significant decline in the morphological, physiological, and biochemical indices of watermelon seedlings exposed to salt stress87. Salt stress persuaded higher concentration of Na content in tomato plant leaves but lessened other nutrients like N, P, and K contents (Table 8). The application of organic amendments creates provision for larger N, P, and K uptake, and diminishes the salt-induced harmful amount of Na uptake in tomato leaves (Table 8). Osmotic stress is a common secondary stress of NaCl salinity in many plants that can negatively influence many aspects of plant metabolisms88,89. The accumulation of large amounts of Na in plant tissues exposed to saline conditions can have damaging effects on the metabolism of cytoplasm and organelles64. Excess Na causes an imbalance in cellular Na and K homeostasis, often resulting in a low Na/K ratio90. Maintaining a low Na/K ratio in leaves is an important feature of plant salt tolerance91. Under salinity stress, the Na/K ratio increased sharply due to excess Na uptake in the leaves. These results indicate that MBC and other organic amendments can promote nutrient balance and reduce ion toxicity thereby accelerating the vegetative development of plants. One practical method of improving salt tolerance is by high intake of mineral nutrients92. Biochar addition reduces plant sodium uptake in salt-stressed soils due to its high uptake capacity through transient binding of Na+ and release of mineral nutrients (K+, Ca2+, Mg2+ particles) into solution93,94. Therefore, it can be concluded that biochar prevents sodium uptake by plants by releasing nutrients into the soil solution. Biochar might have the potential to release large amounts of Ca and Mg and displace Na+ at soil exchange sites, thereby reducing the availability of Na+ to plants95. Since BC has a larger surface area, CEC, and porosity, the addition of BC hinders the absorption and accumulation of Na+96. BC application increases Ca2+ content, thereby improving SS tolerance by altering cell signaling pathways97. The application of biochar can reduce the Na/K ratio and Ca2+ through the Ca2+ dependent SOS pathway98. Sodium elimination in roots occurs primarily through the plasma membrane Na+/H+ antiporter, a well-known gene (SOS1)96. Overexpression of vacuolar Na+/H+ antiporters increase salt tolerance in plants99. Adding biochar to soil improves electrochemical properties, such as root zeta potential, thereby increasing nutrient uptake by plants100. The suppressing effect of vermicompost as well as biochar might be attributed to the release of toxic compounds such as ammonium and improved soil nutrient and plant growth, leading to plants more tolerant to nematode damage101. In addition, the use of manure- biochar compost has a significant positive effect on improving the growth of microorganisms and the release and uptake of nutrients in saline soil, providing a practical option for alleviating salt stress in tomato plant102. The presence of plant nutrients and ash in the biochar and its large surface area, porous nature, and the ability to act as a medium for microorganisms have been identified as the main reasons for the improvement in soil properties and increase in the absorption of nutrients by plants in soils treated with biochar103. The positive effects of biochar on the interactions between soil-plant-water caused better photosynthetic performance and improved nitrogen and water use efficiency also biochar has the potential to improve the properties of soil, microbial abundance, biological nitrogen fixation, and plant growth. Therefore, it is recommended to use biochar as a soil amendment for long-term carbon sink restoration104. These biochars acted as a nutrient source as well as a suitable cation exchanger due to their large surface area resulting in the availability of nutrients and their higher uptake by plant roots105.Under SS stress, plants display disturbances in several physiological processes, leading to a decrease in tomato production (i.e., SFW and TFWP) (Fig. 2) and yield attributes (i.e., NFCP, NFC) (Table 2). Tomato output is significantly reduced by SS because it damages photosynthetic pigments, results in ionic imbalance and ROS generation, and decreases nutrient uptake, RWC, and membrane integrity 106,107. By altering chlorophyll pigments, salinity prevents the buildup of photoassimilates, which lowers tomato output characteristics and, in turn, lowers fruit weight108. But when MBC and its organic materials were applied, tomato plants under salt stress produced much more, suggesting that salt stress can be lessened (Table 2; Fig. 2). The previous research demonstrates that the use of organic amendment enhanced the productivity and yield characteristics of tomato plants109 by improving the physiological mechanism of osmoregulation, photosynthetic efficiency, nutrient uptake, and antioxidant activities, K uptake, and Na/K reduction. Moreover, organic matter enhances the qualities of the soil and provides plants with greater capacity for water absorption, photosynthetic pigments, and biochemical activity. This increases photosynthesis, which is positively correlated with increased fruit yield characteristics and fruit weight110. In addition to organic amendment, literature have shown that K amendment increased yield, quality, and drought tolerance in soybean111, and mung bean growth, yield, nutrient content, and drought tolerance112. Nano iron oxide accelerates growth, yield, and quality of soybean113.

In summary, biochar is the best organic amendment to act as a source of soil nutrients for improved tomato growth, crop yield, and environmental benefits. SS lowers photosynthetic pigments, nutrient uptake, and leaf water content while increasing H2O2, MDA, and Na accumulation. These effects ultimately lower tomato output and plant growth. According to our findings, under salt stress, MBC and its organic matter can enhance tomato output and plant growth. The salt-treated saline soil (PBC (1:2) @ 3 tha−1 + SBTF) had the highest biomass and tomato fruit production. Based on how much salt stress reduces tomato yield, T7 > T9 > T8 > T6 is the order in which MBC treatments are applied. To decrease leaf oxidative damage, MBC was applied. This resulted in an increase in chlorophyll content, leaves’ RWC, nutrient absorption, Na/K ratio, antioxidant activity, and osmoregulatory characteristics. MBC also lessened the generation of ROS and Na buildup. Consequently, MBC exhibits its tolerance mechanism to salt stress and has a good impact on tomato plants’ growth characteristics, photosynthetic mechanism, and physiological and osmotic adaptation features. Therefore, the combined use of biochar and compost can be beneficial in reducing the detrimental effects of salinity on horticultural crops and increasing tomato productivity in salt-affected soils. Based on our findings, additional research is suggested to explore potential mechanisms involved in the improvement of physiology, growth, and productivity under salt stress.

Methods

Planting materials and systems for growth

The investigation was led in a salinity-affected farmer’s field in Batiaghata Union, Khulna (22° 41΄ 36.1˝ N and 89° 31’56. 0˝ E) from December 2018 to March 2019. The site is located under the AEZ of the Ganges tidal floodplain (AEZ 13). The high-yielding indeterminate tomato (Lycopersicon esculentum L.) cultivar BARI Tomato-9 (Lalima) was used. Tomato seeds were surface sterilized with 70% ethanol solution and 5% sodium hypochlorite solution (NaOCl) for 15 min, and then washed with distilled water. For two days, tomato seeds were grown on damp filter paper in the dark. Then sown in sand-filled trays and grown under the transparent plastic shed for 10 d. At the second-leaf stage, the seedlings were uprooted and placed in plastic bags filled with soil, grown for 20 d, and then transplanted to the experimental field. With soil salinity i.e., ECe (6.62 dS/m), pH (7.52), organic carbon (OC) (0.68%), total nitrogen (TN) (0.09%), available phosphorus (P) (18.40 mg kg−1), and exchangeable Ca, Mg, Na, and K (7.44, 3.72, 18.67, and 0.27 cmol kg−1), respectively, the soil belonged to the silty clay loam textural class (sand 18.10%, silt 45.30%, and clay 36.60%) (Table 9). The average daily temperature, minimum temperature, and relative humidity for the tomato cultivation season were T max = 34.3 °C and T min = 26.2 °C; RH max = 95.1% and RH min = 67.3%, respectively. In accordance with our institution’s, as well as national and international norms and laws, experimental research, laboratory, and field investigations of the effects of salinity stress on organic amendments in tomatoes, including the collecting of tomato seeds, were conducted.

Table 9 Physico-chemical features of the research field soil.

Composition (% (w/w))	pH	ECe	OC	TN	P	K	Ca	Mg	Na	
Sand	Silt	Clay		dS m-1	(%)	(%)	(mg kg-1)	(cmol kg-1)	(cmol kg-1)	(cmol kg-1)	(cmol kg-1)	
18.10	45.3	36.60	7.52	6.62	0.68	0.09	18.40	0.27	7.46	3.72	18.67	
ECe = Electrical conductivity; OC = Organic carbon; TN = Total nitrogen; P = Phosphorus; K = Potassium; Ca = Calcium; Mg = Magnesium; Na = Sodium.

Manure-biochar compost preparation

Biochar used in field experiments was obtained by pyrolysis of rice straw in a two-chamber pyrolysis furnace at 400–500 °C. Locally collected cow dung and poultry manure were kept in ambient conditions for a week to lessen extra moisture. Table 10 displays the chemical characteristics of the additional organic compounds.

Table 10 Chemical properties of different applications of organic substances.

OM	pH	OC
(%)	Total N
(%)	P
(%)	K
(%)	Ca (%)	Mg (%)	Zn
(mg kg−1)	
Cow dung	7.28	22.94	1.13	0.35	0.80	0.97	0.54	142.10	
Poultry manure	8.02	34.20	1.45	0.73	0.97	1.51	0.51	178.10	
Biochar (Rice straw)	7.65	44.03	0.86	0.34	2.84	1.68	0.49	240.10	
PBC (1:1)	7.78	39.12	1.81	0.57	0.98	1.55	0.50	153.50	
PBC (1:2)	7.67	39.72	1.94	0.46	0.95	1.67	0.50	194.30	
CBC (1:1)	7.37	34.49	1.53	0.41	0.87	1.49	0.56	198.10	
CBC (1:2)	7.51	38.66	1.76	0.38	0.86	1.63	0.49	212.43	
OC = Organic carbon; N = Nitrogen; P = Phosphorus; K = Potassium; Ca = Calcium; Mg = Magnesium; Zn = Zinc; PBC = poultry manure-biochar compost; CBC = cow dung-biochar compost.

For compost preparation, manuring substances (M) (cow dung (C) and poultry manure (P)) and biochar (B) were mixed at the ratio of 1:1 (M: BC, v/v) and 1:2 (M: BC, v/v). After 6 weeks cow dung-biochar compost (CBC) and poultry manure-biochar compost (PBC) were ready for field application. The produced MBC is thoroughly mixed before use as a soil amendment.

Experimentation

Abandoned salinized land was treated with different manure-biochar compost and their raw organic substances. Three different organic amendment treatments and their compost mixtures [CBC (1:1; and 1:2 v/v ratio)] and [PBC (1:1; and 1:2 v/v ratio)] plus SBTF dose were evaluated. For comparison, we used reclaimed salinized soil and sole SBTF-based chemical fertilizer-treated plots. The treatments are: T1 = absolute SS; T2 = SBTF dose; T3 = biochar (B) @ 3 t/ha + SBTF; T4 = poultry manure (P) @ 3 t/ha + SBTF; T5 = cow dung (C) @ 3 tha−1 + SBTF; T6 = PBC (1:1) @ 3 t/ha + SBTF; T7 = PBC (1:2) @ 3 t/ha + SBTF; T8 = CBC (1:1) @ 3 t/ha + SBTF; T9 = CBC (1:2) @ 3 t/ha + SBTF. Three replications of a randomized complete block design (RCBD) were employed to order the treatments in the field study. A drainage system of 2.5 × 2.0 m and 0.5 m was built into every plot. On December 5, 2018, four-week-old tomato seedlings were transplanted, with 60 cm × 50 cm spacing between each seedling and two seedlings per hill. The same seedlings were used to repair the gaps one week later. Fertilizers based on SBTF were used to fertilize the crop. computed using Fertilizer Recommendation Guide114 to yield 321 kg of urea, 85 kg of triple superphosphate (TSP), 22 kg of muriate of potash (MP), and 20 kg of gypsum per ha−1. Final land preparation included basal application of one-third of urea, TSP, MP, gypsum, and full volumes of organic amendments. Two top dressing applications of the leftover computed urea were made at 15 and 35 days after transplantation (DAT). We maintained the field capacity (FC) of the soil (moisture content of soil 30.7%) for each pot. Irrigation was provided at 12 h intervals. The moisture loss every day was fulfilled by adding tap water using a moisture meter until the soil moisture level was raised to 30.7%. As we maintained FC (water only in the micropores of the soil), so very minimal water was required every day to maintain FC and there was no leaching or percolation of the potting soil. As a result, the salinity of the soil remains constant throughout the cropping season. The conventional cultivation method used in Bangladesh was followed when applying tomato production to agriculture.

Tomato growth and yield characteristics are measured

During the active flowering period at 65 DAT, several growth parameters [PH (cm), NBP, NLP, and LA] were assessed. Yield characteristics of tomato fruit (NEFC and NFC) were counted in three selected plants and their average value was calculated. Tomato fruit yield was calculated by counting NFP and SFW and weighting all the picked fruits collected at the ripening stage from each plant. Fruits were harvested four (4) times during the harvesting time and that began on 13 March 2019. In the senescence stage, the whole plant, fresh weight (FW), and dry weight (DW) were calculated to obtain biomass yield. To determine the total dry mass (TDM), plants were divided into shoots and roots and air-dried first at room temperature and finally oven-dried at 65 °C for 72 h.

Measurement of pigment content

Using the method outlined by Lichtenthaler et al.115, the total chlorophyll content (mg/g FW) was determined from the completely prolonged top most leaf at the flowering stage (67 DAT). To extract leaf chlorophyll, 20 mg of completely inflated leaf samples were taken, put in tubes covered in aluminum foil with 20 mL of 80% acetone, and left in the dark for the entire night at 4 °C. Following centrifugation, a spectrophotometer was used to measure the absorbance at 645 nm and 663 nm wavelengths.

Measurement of electrolyte leakage and water status

The Sullivan116 method was used to measure the electrolyte loss in the leaves at 67 days after harvest. Leaf samples weighing about 0.5 g were collected and placed in two distinct test tube sets, each holding 10 mL of deionized water. The first group was incubated for ten minutes at room temperature, while the second set spent thirty minutes in a water bath at fifty-five degrees. An EC meter was used to determine the electrical conductivity (EC) of the supernatants from both sets. ECa and ECb are the names of the first and second sets of ECs, respectively. The following formula was used to determine the electrolyte leakage:

Electrolyte leakage (%) = {(ECb - ECa)/ECe} × 100.

The technique of Hayat et al.117 was used to calculate the leaf relative water content (RWC). Samples of fully-grown leaves (at 67 DAT) were collected, and the fresh weight (FW) was measured. To determine the turgid weight (TW), the same leaves were immersed in distilled water for four hours. For 72 h, the leaf samples were oven-dried at 70 °C to obtain a consistent dry weight (DW). The formula used to compute the RWC of leaves was: - RWC (%) = {(FW – DW)/ (TW – DW)} × 100.

As per Sangakkara et al.118, the following formulas were used to compute the water saturation deficit (WSD), water retention capacity (WRC), and water uptake capacity (WUC). Water saturation deficit (WSD) = {(TW – FW)/(TW – DW)} × 100.

Water retention capacity (WRC) = (TW/DW).

Where, FW = Fresh weight (mg), DW = Dry weight (mg) and TW = Turgid weight (mg).

Determination of malondialdehyde (MDA) and H2O2concentrations

The H2O2 content (nmol/g FW) in plant samples was determined by the method of Su and Silva119. Fresh plant leaf sample (0.5 g) was extracted with 5 mL of 5% (w/v) trichloric acid (TCA) and centrifuged at 6000 × g for 15 min. The reaction mixture consisted of 1 mL 1.0 M potassium iodide (KI) and 100 µL of 10 mM potassium phosphate buffer (pH 7.0). Then the content of H2O2 was determined by using a spectrophotometer at 390 nm wavelength from a standard curve.

The assay for thiobarbituric acid (TBA) was utilized to ascertain the MDA concentration (nmol/g FW)119. To extract MDA, a 0.5 g leaf sample was taken, homogenized in 5.0 mL of 5% (w/v) trichloroacetic acid (TCA), and centrifuged for 10 min at 12,000 × g. After that, 1 mL of supernatant was added to the reaction mixer, which held 4 mL of 20% TCA with 0.5% TBA, and it was maintained in a water bath at 95 °C for 30 min. To find the MDA content, absorbance measurements were made at 532, 600, and 450 nm following cooling in an ice bath. The following formula was used to determine the tomato leaf’s MDA content: 0.56 × A450–6.45 × (A532 – A600).

Determination of proline, ascorbic acid, and total soluble sugar content

The method outlined by Bates et al.120. was utilized to measure the proline content (mg/g FW) of leaves. Leaf samples (0.5 g) were pulverized in 10 mL of 3% (v/v) sulfosalicylic acid and centrifuged at 10,000 × g for 10 min to extract proline from leaves (67 DAT). Two milliliters of the supernatant (2 mL) were added to a test tube along with two milliliters each of glacial acetic acid and acid-ninhydrin solution. After 30 min of incubation in a water bath at 100 °C, the mixer was cooled in an ice bath. To separate the toluene and aqueous phases, 4 mL of toluene was added to the reaction mixture after it had cooled. It was then let to stand in the dark at room temperature for 20 min. After that, the toluene phase was meticulously collected, and a spectrophotometer was used to measure the absorbance at 520 nm. Analytical proline was used to generate a standard curve that showed the concentration of free proline.

The ascorbic acid (AA) content (mg/g FW) was calculated utilizing the method that Jagota and Dani121 outlined. A 0.5 g leaf sample was weighed, homogenized in 2 mL of 5% trichloroacetic acid (TCA), and centrifuged at 10,000 × g for 15 min at 4 °C to produce the AA extraction. Next, 8 mL of 10% TCA was added to 2 mL of supernatant that had been collected in a test tube. After giving the mixer a good shake, it was put in an ice bath for five minutes, and it was centrifuged once more for five minutes at 3000 × g. 5 mL of the extract was combined with 2 mL of distilled water diluted with 2 mL of Folin-Ciocalteu reagent, and the mixture was left for 10 min to develop the blue color. At 760 nm, the AA content was determined by spectrophotometry.

Leaf total soluble-sugar content (mg/g FW) was determined using a colorimetric method122. Fresh leaf sample was collected and homogenized, then extracted with water at room temperature. Phenol and sulfuric acid are then added to prepare the extract. Total soluble sugar content was measured with a spectrophotometer at 485 nm.

Determination of nutrient contents in leaf

The N, P, K, Ca, and Na contents (mg/g DW) of each plant’s growing third and fourth leaves at the flowering stage (67 DAT) were measured. A 20-mesh sieve was used to grind and homogenize the oven-dried leaf samples in preparation for analysis. The micro Kjeldahl method was used to calculate the sample of leaves’ total nitrogen content. A 0.2 g plant sample was digested with concentrated H2SO4 at 360 °C for two hours while a catalyst mixture (K2SO4: CuSO4.5H2O: Se = 100:10:1) was present to determine the amount of N present. Then the digests were distilled off in an auto distillation unit using 40% NaOH solution and collected in 4% H3BO4; finally titrated with auto burette against 0.02 N (N/50) H2SO4 and calculated123. For the determination of nutrient element concentrations except for nitrogen, 0.5 g ground leaf samples were digested with the di-acid mixture (HNO3: HClO4 = 5:1) heated on a sand bath at 130 °C until colorless fume evolved124. Leaf phosphorus content was determined by the ammonium molybdovanadate method using a spectrophotometer125. Leaf potassium ion (K+), calcium ion (Ca2+), and sodium ion (Na+) contents were assessed with the help of an atomic absorption spectrophotometer (Model No 170 − 30, HITACHI, Japan)126.

Statistical analysis

Trait-wise average dta were analyzed statistically127,128 and biometrically129,130. The analysis of variance (ANOVA) test was run on the experimental data using the SPSS statistical tool (SPSS 16.0, SPSS Inc., USA). Tukey’s HSD Test was utilized to determine the significance of variation across treatments.

Electronic supplementary material

Below is the link to the electronic supplementary material.

Supplementary Material 1

Acknowledgements

The authors would like to extend their sincere appreciation to the Researchers Supporting Project number (RSP2024R194), King Saud University, Riyadh, Saudi Arabia.

Author contributions

M.Z.U.K., and U.S. initiated the research work, conceived the study, and performed the experiments; M.Z.U.K., U.S., S.K.R., and M.S.A. performed statistical analysis; M.Z.U.K., U.S., S.K.R., M.S.A., M.Y.M., N.H., S.E., and S.A. drafted, edited, interpreted data, and prepared the manuscript.

Data availability

Data is provided within the manuscript or supplementary information files.

Declarations

Competing interests

The authors declare no competing interests.

Publisher’s note

Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.
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References

1. Afza U Organic amendments mitigate salinity induced toxic effects in maize by modulating antioxidant defense system, photosynthetic pigments, and ionic homeostasis Notulae Botanicae Horti Agrobotanici Cluj-Napoca 2022 50 2 12735 12735 10.15835/nbha50212735
Afza, U. et al. Organic amendments mitigate salinity induced toxic effects in maize by modulating antioxidant defense system, photosynthetic pigments, and ionic homeostasis. Notulae Botanicae Horti Agrobotanici Cluj-Napoca. 50 (2), 12735–12735. 10.15835/nbha50212735 (2022).
2. Kamal MZU Organic amendments can alleviate the adverse effects of soil salinity on the performance of tomato plant J. Agril Crop Res. 2021 9 7 165 175
Kamal, M. Z. U. et al. Organic amendments can alleviate the adverse effects of soil salinity on the performance of tomato plant. J. Agril Crop Res. 9 (7), 165–175 (2021).
3. Shokat S Großkinsky DK Tackling salinity in sustainable agriculture—what developing countries may learn from approaches of the developed world Sustainability 2019 11 17 4558 10.3390/su11174558
Shokat, S. & Großkinsky, D. K. Tackling salinity in sustainable agriculture—what developing countries may learn from approaches of the developed world. Sustainability. 11 (17), 4558. 10.3390/su11174558 (2019).
4. Mandal S Raju R Kumar A Kumar P Sharma PC Current status of research, technology response and policy need of salt-affected soils in India – a review J. Indian Soc. Coast Agric. Res. 2018 36 2 40 53
Mandal, S., Raju, R., Kumar, A., Kumar, P. & Sharma, P. C. Current status of research, technology response and policy need of salt-affected soils in India – a review. J. Indian Soc. Coast Agric. Res. 36 (2), 40–53 (2018).
5. Li J Soil salinization research in China: advances and prospects J. Geogr. Sci. 2014 24 5 943 960 10.1007/s11442-014-1130-2
Li, J. et al. Soil salinization research in China: advances and prospects. J. Geogr. Sci. 24 (5), 943–960. 10.1007/s11442-014-1130-2 (2014).
6. Shrivastava P Kumar R Soil salinity A serious environmental issue and plant growth promoting bacteria as one of the tools for its alleviation Saudi J. Biol. Sci. 2015 22 2 123 131 10.1016/j.sjbs.2014.12.001 25737642
Shrivastava, P., Kumar, R. & Soil salinity A serious environmental issue and plant growth promoting bacteria as one of the tools for its alleviation. Saudi J. Biol. Sci. 22 (2), 123–131. 10.1016/j.sjbs.2014.12.001 (2015).25737642
7. Khan AA McNeilly T Azhar FM Stress tolerance in crop plants Int. J. Agric. Biol. 2001 3 2 250 255
Khan, A. A., McNeilly, T. & Azhar, F. M. Stress tolerance in crop plants. Int. J. Agric. Biol. 3 (2), 250–255 (2001).
8. Haque SA Salinity problems and crop production in coastal regions of Bangladesh Pakistan J. Bot. 2006 38 5 1359 1365
Haque, S. A. Salinity problems and crop production in coastal regions of Bangladesh. Pakistan J. Bot. 38 (5), 1359–1365 (2006).
9. Uddin MS Khan MSI Talukdar MMR Hossain MI Ullah MH Climate change and salinity in Bangladesh: constraints and management strategy for crop production Rajshahi Univ. Jour Environ. Sci. 2011 1 13 20
Uddin, M. S., Khan, M. S. I., Talukdar, M. M. R., Hossain, M. I. & Ullah, M. H. Climate change and salinity in Bangladesh: constraints and management strategy for crop production. Rajshahi Univ. Jour Environ. Sci. 1, 13–20 (2011).
10. Sarker U Oba S The response of salinity stress-induced A. tricolor to growth, anatomy, physiology, non-enzymatic and enzymatic antioxidants Front. Plant. Sci. 2020 11 559876 10.3389/fpls.2020.559876 33178233
Sarker, U. & Oba, S. The response of salinity stress-induced A. tricolor to growth, anatomy, physiology, non-enzymatic and enzymatic antioxidants. Front. Plant. Sci. 11, 559876. 10.3389/fpls.2020.559876 (2020).33178233
11. Sarker U Oba S Catalase, superoxide dismutase and ascorbate-glutathione cycle enzymes confer drought tolerance of A. Tricolor Sci. Rep. 2018 8 1 16496 10.1038/s41598-018-34944-0 30405159
Sarker, U. & Oba, S. Catalase, superoxide dismutase and ascorbate-glutathione cycle enzymes confer drought tolerance of A. Tricolor. Sci. Rep. 8 (1), 16496. 10.1038/s41598-018-34944-0 (2018).30405159
12. Sarker U Oba S Drought stress effects on growth, ROS markers, compatible solutes, phenolics, flavonoids, and antioxidant activity in Amaranthus tricolor Appl. Biochem. Biotechnol. 2018 186 4 999 1016 10.1007/s12010-018-2784-5 29804177
Sarker, U. & Oba, S. Drought stress effects on growth, ROS markers, compatible solutes, phenolics, flavonoids, and antioxidant activity in Amaranthus tricolor. Appl. Biochem. Biotechnol. 186 (4), 999–1016. 10.1007/s12010-018-2784-5 (2018).29804177
13. Sarker U Oba S Response of nutrients, minerals, antioxidant leaf pigments, vitamins, polyphenol, flavonoid and antioxidant activity in selected amaranth under four Soil Water Content Food Chem. 2018 252 72 83 10.1016/j.foodchem.2018.01.097 29478565
Sarker, U. & Oba, S. Response of nutrients, minerals, antioxidant leaf pigments, vitamins, polyphenol, flavonoid and antioxidant activity in selected amaranth under four Soil Water Content. Food Chem. 252, 72–83. 10.1016/j.foodchem.2018.01.097 (2018).29478565
14. Sarker U Oba S Drought stress enhances nutritional and bioactive compounds, phenolic acids and antioxidant capacity of Amaranthus leafy vegetable BMC Plant. Biol. 2018 18 1 258 10.1186/s12870-018-1484-1 30367616
Sarker, U. & Oba, S. Drought stress enhances nutritional and bioactive compounds, phenolic acids and antioxidant capacity of Amaranthus leafy vegetable. BMC Plant. Biol. 18 (1), 258. 10.1186/s12870-018-1484-1 (2018).30367616
15. Mannan MA Biostimulant red seaweed (Gracilaria tenuistipitata var. Liui) extracts spray improves yield and drought tolerance in soybean Peer J. 2023 11 e15588 10.7717/peerj.15588 37377788
Mannan, M. A. et al. Biostimulant red seaweed (Gracilaria tenuistipitata var. Liui) extracts spray improves yield and drought tolerance in soybean. Peer J. 11, e15588. 10.7717/peerj.15588 (2023).37377788
16. Roychowdhury R Multi-omics pipelines and omics-integration approach to decipher plant’s abiotic stress tolerance responses Genes 2023 14 6 1281 10.3390/genes14061281 37372461
Roychowdhury, R. et al. Multi-omics pipelines and omics-integration approach to decipher plant’s abiotic stress tolerance responses. Genes. 14 (6), 1281. 10.3390/genes14061281 (2023).37372461
17. Fatema MK Assessing morpho-physiological and biochemical markers of soybean for drought tolerance potential Sustainability 2023 15 2 1427 10.3390/su15021427
Fatema, M. K. et al. Assessing morpho-physiological and biochemical markers of soybean for drought tolerance potential. Sustainability. 15 (2), 1427. 10.3390/su15021427 (2023).
18. Sarker U Oba S Salinity Stress Enhances Color Parameters, Bioactive Leaf pigments, vitamins, polyphenols, flavonoids and antioxidant activity in selected Amaranthus Leafy vegetables J. Sci. Food Agric. 2019 99 5 2275 2284 10.1002/jsfa.9423 30324618
Sarker, U. & Oba, S. Salinity Stress Enhances Color Parameters, Bioactive Leaf pigments, vitamins, polyphenols, flavonoids and antioxidant activity in selected Amaranthus Leafy vegetables. J. Sci. Food Agric. 99 (5), 2275–2284. 10.1002/jsfa.9423 (2019).30324618
19. Sarker U Oba S Leaf pigmentation, its profiles and radical scavenging activity in selected Amaranthus tricolor leafy vegetables Sci. Rep. 2020 10 1 18617 10.1038/s41598-020-66376-0 33122663
Sarker, U. & Oba, S. Leaf pigmentation, its profiles and radical scavenging activity in selected Amaranthus tricolor leafy vegetables. Sci. Rep. 10 (1), 18617. 10.1038/s41598-020-66376-0 (2020).33122663
20. Sarker U Nutritional and bioactive properties and antioxidant potential of Amaranthus tricolor, A. Lividus, a viridis, and A. spinosus leafy vegetables Heliyon 2024 10 9 e30453 10.1016/j.heliyon.2024.e30453 38720726
Sarker, U. et al. Nutritional and bioactive properties and antioxidant potential of Amaranthus tricolor, A. Lividus, a viridis, and A. spinosus leafy vegetables. Heliyon. 10 (9), e30453. 10.1016/j.heliyon.2024.e30453 (2024).38720726
21. Sarker U Bioactive phytochemicals and quenching activity of radicals in selected Drought-Resistant Amaranthus tricolor Vegetable Amaranth Antioxidants 2022 11 578 10.3390/antiox11030578 35326227
Sarker, U. et al. Bioactive phytochemicals and quenching activity of radicals in selected Drought-Resistant Amaranthus tricolor Vegetable Amaranth. Antioxidants. 11, 578. 10.3390/antiox11030578 (2022).35326227
22. Sarker U Colorant pigments, nutrients, bioactive components, and antiradical potential of danta leaves (Amaranthus lividus) Antioxidants 2022 11 6 1206 10.3390/antiox11061206 35740102
Sarker, U. et al. Colorant pigments, nutrients, bioactive components, and antiradical potential of danta leaves (Amaranthus lividus). Antioxidants. 11 (6), 1206. 10.3390/antiox11061206 (2022).35740102
23. Sarker, U. & Oba, S. Salinity stress enhances color parameters, bioactive leaf pigments, vitamins, polyphenols, flavonoids and antioxidant activity in selected Amaranthus leafy vegetables. Journal of the Science of Food and Agriculture. 99(5): 2275–2284. doi.10.1002/jsfa.9423 (2019).
24. Tarafder SK Influence of foliar spray and post-harvest treatment on head yield, shelf-life, and physicochemical qualities of Broccoli Front. Nutri 2023 10 1057084 10.3389/fnut.2023.1057084
Tarafder, S. K. et al. Influence of foliar spray and post-harvest treatment on head yield, shelf-life, and physicochemical qualities of Broccoli. Front. Nutri. 10, 1057084. 10.3389/fnut.2023.1057084 (2023).
25. Sarker, U., Islam, M. T. & Oba, S. Salinity stress accelerates nutrients, dietary fiber, minerals, phytochemicals and antioxidant activity in Amaranthus tricolor leaves. PLoS One. 13(11), e0206388. doi.10.1371/journal.pone.0206388. (2018).
26. Sarker, U., Oba, S., Alsanie, W. F. & Gaber, A. Characterization of phytochemicals, nutrients, and antiradical potential in slim amaranth. Antioxidants 11(6), 1089. (2022). 10.3390/antiox11061089
27. Hassan J Color and physiochemical attributes of pointed gourd (Trichosanthes dioica Roxb.) Influenced by modified atmosphere packaging and postharvest treatment during storage Front. Plant. Sci. 2022 13 1016324 10.3389/fpls.2022.1016324 36275589
Hassan, J. et al. Color and physiochemical attributes of pointed gourd (Trichosanthes dioica Roxb.) Influenced by modified atmosphere packaging and postharvest treatment during storage. Front. Plant. Sci. 13, 1016324. 10.3389/fpls.2022.1016324 (2022).36275589
28. Sarker U Oba S Protein, dietary fiber, minerals, antioxidant pigments and phytochemicals, and antioxidant activity in selected red morph Amaranthus leafy vegetable PLoS ONE 2019 14 12 e0222517 10.1371/journal.pone.0222517 31830064
Sarker, U. & Oba, S. Protein, dietary fiber, minerals, antioxidant pigments and phytochemicals, and antioxidant activity in selected red morph Amaranthus leafy vegetable. PLoS ONE. 14 (12), e0222517. 10.1371/journal.pone.0222517 (2019).31830064
29. Sarker U Phytonutrients, colorant pigments, phytochemicals, and antioxidant potential of orphan leafy Amaranthus species Molecules 2022 27 9 2899 10.3390/molecules27092899 35566250
Sarker, U. et al. Phytonutrients, colorant pigments, phytochemicals, and antioxidant potential of orphan leafy Amaranthus species. Molecules. 27 (9), 2899. 10.3390/molecules27092899 (2022).35566250
30. Sarker U Oba S Color attributes, betacyanin, and carotenoid profiles, bioactive components, and radical quenching capacity in selected Amaranthus gangeticus leafy vegetables Sci. Rep. 2021 11 1 11559 10.1038/s41598-021-91157-8 34079029
Sarker, U. & Oba, S. Color attributes, betacyanin, and carotenoid profiles, bioactive components, and radical quenching capacity in selected Amaranthus gangeticus leafy vegetables. Sci. Rep. 11 (1), 11559. 10.1038/s41598-021-91157-8 (2021).34079029
31. Sarker, U. & Ercisli, S. Salt eustress induction in red amaranth (Amaranthus gangeticus) augments nutritional, phenolic acids and antiradical potential of leaves. Antioxidants 11, 2434 doi.10.3390/antiox11122434 (2022).
32. Sarker U Oba S Augmentation of leaf color parameters, pigments, vitamins, phenolic acids, flavonoids and antioxidant activity in selected Amaranthus tricolor under salinity stress Sci. Rep. 2018 8 1 12349 10.1038/s41598-018-30897-6 30120319
Sarker, U. & Oba, S. Augmentation of leaf color parameters, pigments, vitamins, phenolic acids, flavonoids and antioxidant activity in selected Amaranthus tricolor under salinity stress. Sci. Rep. 8 (1), 12349. 10.1038/s41598-018-30897-6 (2018).30120319
33. Sarker U Oba S Antioxidant constituents of three selected red and green color Amaranthus leafy vegetable Sci. Rep. 2019 9 1 18233 10.1038/s41598-019-52033-8 31796754
Sarker, U. & Oba, S. Antioxidant constituents of three selected red and green color Amaranthus leafy vegetable. Sci. Rep. 9 (1), 18233. 10.1038/s41598-019-52033-8 (2019).31796754
34. Sarker U Oba S Nutraceuticals, phytochemicals, and radical quenching ability of selected drought-tolerant advance lines of vegetable amaranth BMC Plant. Biol. 2020 20 1 564 10.1186/s12870-020-02780-y 33317465
Sarker, U. & Oba, S. Nutraceuticals, phytochemicals, and radical quenching ability of selected drought-tolerant advance lines of vegetable amaranth. BMC Plant. Biol. 20 (1), 564. 10.1186/s12870-020-02780-y (2020).33317465
35. Sarker U Oba S Phenolic profiles and antioxidant activities in selected drought-tolerant leafy vegetable amaranth Sci. Rep. 2020 10 1 18287 10.1038/s41598-020-71727-y 33106544
Sarker, U. & Oba, S. Phenolic profiles and antioxidant activities in selected drought-tolerant leafy vegetable amaranth. Sci. Rep. 10 (1), 18287. 10.1038/s41598-020-71727-y (2020).33106544
36. Sarker U Oba S Polyphenol and flavonoid profiles and radical scavenging activity in selected leafy vegetable Amaranthus gangeticus BMC Plant. Biol. 2020 20 1 499 10.1186/s12870-020-02700-0 33138787
Sarker, U. & Oba, S. Polyphenol and flavonoid profiles and radical scavenging activity in selected leafy vegetable Amaranthus gangeticus. BMC Plant. Biol. 20 (1), 499. 10.1186/s12870-020-02700-0 (2020).33138787
37. Sarker U Hossain MN Iqbal MA Oba S Bioactive components and radical scavenging activity in selected advance lines of salt-tolerant vegetable amaranth Front. Nutr. 2020 7 587257 10.3389/fnut.2020.587257 33330589
Sarker, U., Hossain, M. N., Iqbal, M. A. & Oba, S. Bioactive components and radical scavenging activity in selected advance lines of salt-tolerant vegetable amaranth. Front. Nutr. 7, 587257. 10.3389/fnut.2020.587257 (2020).33330589
38. Hossain MN Influence of salinity stress on color parameters, leaf pigmentation, polyphenol and flavonoid contents, and antioxidant activity of Amaranthus lividus Leafy vegetables Molecules 2022 27 6 1821 10.3390/molecules27061821 35335185
Hossain, M. N. et al. Influence of salinity stress on color parameters, leaf pigmentation, polyphenol and flavonoid contents, and antioxidant activity of Amaranthus lividus Leafy vegetables. Molecules. 27 (6), 1821. 10.3390/molecules27061821 (2022).35335185
39. Sarker U Salinity stress ameliorates pigments, minerals, polyphenolic profiles, and antiradical capacity in lalshak Antioxidants 2023 12 1 173 10.3390/antiox12010173 36671036
Sarker, U. et al. Salinity stress ameliorates pigments, minerals, polyphenolic profiles, and antiradical capacity in lalshak. Antioxidants. 12 (1), 173. 10.3390/antiox12010173 (2023).36671036
40. Sarker U Lin Y Oba S Yoshioka Y Hoshikawa K Prospects and potentials of underutilized leafy Amaranths as vegetable use for health-promotion Plant. Physiol. Biochem. 2022 182 104 123 10.1016/j.plaphy.2022.04.011 35487123
Sarker, U., Lin, Y., Oba, S., Yoshioka, Y. & Hoshikawa, K. Prospects and potentials of underutilized leafy Amaranths as vegetable use for health-promotion. Plant. Physiol. Biochem. 182, 104–123. 10.1016/j.plaphy.2022.04.011 (2022).35487123
41. Yang Z Li JL Liu LN Xie Q Sui N Photosynthetic regulation under salt stress and salt-tolerance mechanism of sweet sorghum Front. Plant. Sci. 2020 10 1722 10.3389/fpls.2019.01722 32010174
Yang, Z., Li, J. L., Liu, L. N., Xie, Q. & Sui, N. Photosynthetic regulation under salt stress and salt-tolerance mechanism of sweet sorghum. Front. Plant. Sci. 10, 1722. 10.3389/fpls.2019.01722 (2020).32010174
42. Lobell DB Ortiz-Monasterio JI Gurrola FC Valenzuela L Identification of saline soils with multiyear remote sensing of crop yields Soil. Sci. Soc. Am. J. 2007 71 3 777 783 10.2136/sssaj2006.0306
Lobell, D. B., Ortiz-Monasterio, J. I., Gurrola, F. C. & Valenzuela, L. Identification of saline soils with multiyear remote sensing of crop yields. Soil. Sci. Soc. Am. J. 71 (3), 777–783. 10.2136/sssaj2006.0306 (2007).
43. Khatun M Shuvo MAR Salam MTB Rahman SH Effect of organic amendments on soil salinity and the growth of maize (Zea mays L) Plant. Sci. Today 2019 6 2 106 111 10.14719/pst.2019.6.2.491
Khatun, M., Shuvo, M. A. R., Salam, M. T. B. & Rahman, S. H. Effect of organic amendments on soil salinity and the growth of maize (Zea mays L). Plant. Sci. Today. 6 (2), 106–111. 10.14719/pst.2019.6.2.491 (2019).
44. Gupta B Huang B Mechanism of salinity tolerance in plants: physiological, biochemical, and molecular characterization Int. J. Genomics 2014 701596 18 10.1155/2014/701596
Gupta, B. & Huang, B. Mechanism of salinity tolerance in plants: physiological, biochemical, and molecular characterization. Int. J. Genomics. 701596, 18. 10.1155/2014/701596 (2014).
45. Vanlauwe B Sanginga, N. Integrated soil fertility management: operational definition and consequences for implementation and dissemination Outlook Agric. 2010 39 17 24 10.5367/000000010791169998
Vanlauwe, B. et al. Sanginga, N. Integrated soil fertility management: operational definition and consequences for implementation and dissemination. Outlook Agric. 39, 17–24. 10.5367/000000010791169998 (2010).
46. Melero S Madejón E Ruiz JC Herencia JF Chemical and biochemical properties of clay soil under dryland agriculture system as affected by organic fertilization Eur. J. Agron. 2007 26 3 327 334 10.1016/j.eja.2006.11.004
Melero, S., Madejón, E., Ruiz, J. C. & Herencia, J. F. Chemical and biochemical properties of clay soil under dryland agriculture system as affected by organic fertilization. Eur. J. Agron. 26 (3), 327–334. 10.1016/j.eja.2006.11.004 (2007).
47. Muhammad, M. et al. Soil salinity and drought tolerance: an evaluation of plant growth, productivity, microbial diversity, and amelioration strategies. Plant. Stress, p.100319 (2023).
48. Joshi, S., Nath, J., Singh, A. K., Pareek, A. & Joshi, R. Ion transporters and their regulatory signal transduction mechanisms for salinity tolerance in plants. Physiologia Plantarum, 174(3), p.e13702 (2022).
49. Tejada M Garcia C Gonzalez JL Hernandez MT Use of organic amendment as a strategy for saline soil remediation: influence on the physical, chemical and biological properties of soil Soil. Biol. Biochem. 2006 38 6 1413 1421 10.1016/j.soilbio.2005.10.017
Tejada, M., Garcia, C., Gonzalez, J. L. & Hernandez, M. T. Use of organic amendment as a strategy for saline soil remediation: influence on the physical, chemical and biological properties of soil. Soil. Biol. Biochem. 38 (6), 1413–1421. 10.1016/j.soilbio.2005.10.017 (2006).
50. Walker DJ Bernal MP The effects of olive mill waste compost and poultry manure on the availability and plant uptake of nutrients in highly saline soil Biores Technol. 2008 99 2 396 403 10.1016/j.biortech.2006.12.006
Walker, D. J. & Bernal, M. P. The effects of olive mill waste compost and poultry manure on the availability and plant uptake of nutrients in highly saline soil. Biores Technol. 99 (2), 396–403. 10.1016/j.biortech.2006.12.006 (2008).
51. Bol R Amelung W Friedrich C Ostle N Tracing dung-derived carbon in temperate grassland using 13 C natural abundance measurements Soil. Biol. Biochem. 2000 32 10 1337 1343 10.1016/S0038-0717(00)00022-5
Bol, R., Amelung, W., Friedrich, C. & Ostle, N. Tracing dung-derived carbon in temperate grassland using 13 C natural abundance measurements. Soil. Biol. Biochem. 32 (10), 1337–1343. 10.1016/S0038-0717(00)00022-5 (2000).
52. Ghani MI Almond shell-derived biochar decreased toxic metals bioavailability and uptake by tomato and enhanced the antioxidant system and microbial community Sci. Total Environ. 2024 929 172632 10.1016/j.scitotenv.2024.172632 38653412
Ghani, M. I. et al. Almond shell-derived biochar decreased toxic metals bioavailability and uptake by tomato and enhanced the antioxidant system and microbial community. Sci. Total Environ. 929, 172632. 10.1016/j.scitotenv.2024.172632 (2024). (2024).38653412
53. Akhtar SS Andersen MN Liu F Biochar mitigates salinity stress in potato J. Agron. Crop Sci. 2015 201 5 368 378 10.1111/jac.12132
Akhtar, S. S., Andersen, M. N. & Liu, F. Biochar mitigates salinity stress in potato. J. Agron. Crop Sci. 201 (5), 368–378. 10.1111/jac.12132 (2015a).
54. Lashari MS Biochar–manure compost in conjunction with pyroligneous solution alleviated salt stress and improved leaf bioactivity of maize in a saline soil from central China: a 2-year field experiment J. Sci. Food Agric. 2015 95 6 1321 1327 10.1002/jsfa.6825 25042565
Lashari, M. S. et al. Biochar–manure compost in conjunction with pyroligneous solution alleviated salt stress and improved leaf bioactivity of maize in a saline soil from central China: a 2-year field experiment. J. Sci. Food Agric. 95 (6), 1321–1327. 10.1002/jsfa.6825 (2015).25042565
55. Herath HMSK Camps-Arbestain M Hedley M Effect of biochar on soil physical properties in two contrasting soils: an Alison and an Andisol Geoderma 2013 209 188 197 10.1016/j.geoderma.2013.06.016
Herath, H. M. S. K., Camps-Arbestain, M. & Hedley, M. Effect of biochar on soil physical properties in two contrasting soils: an Alison and an Andisol. Geoderma. 209, 188–197. 10.1016/j.geoderma.2013.06.016 (2013).
56. Bischoff, J. & Werner, H. Salt/Salinity Tolerance of Common Horticultural Crops in South Dakota (Garden and Vegetable/Woody Fruit Crops, 1999).
57. Miah, M. Y., Kamal, M. Z. U., Salam, M. A. & Islam, M. S. Impact of salinity intrusion on agriculture of Southwest Bangladesh. Int. J. Agric. Pol. Res., 8(2), 40–47. doi.10.15739/IJAPR.20.005 (2020).
58. Nicola S Tibaldi G Fontana E Tomato production systems and their application to the tropics Acta Hortic. 2009 821 27 34 10.17660/ActaHortic.2009.821.1
Nicola, S., Tibaldi, G. & Fontana, E. Tomato production systems and their application to the tropics. Acta Hortic. 821, 27–34 (2009).
59. Mohammed HN Mahmud TMM Edaroyati P Deficit irrigation for improving the Postharvest Quality of Lowland Tomato fruits Pertanika J. Trop. Agric. Sci. 2018 41 2 741 758
Mohammed, H. N., Mahmud, T. M. M. & Edaroyati, P. Deficit irrigation for improving the Postharvest Quality of Lowland Tomato fruits. Pertanika J. Trop. Agric. Sci. 41 (2), 741–758 (2018).
60. Savy D Fertilisation with compost mitigates salt stress in tomato by affecting plant metabolomics and nutritional profiles Chem. Biol. Technol. Agric. 2022 9 1 104 10.1186/s40538-022-00373-5
Savy, D. et al. Fertilisation with compost mitigates salt stress in tomato by affecting plant metabolomics and nutritional profiles. Chem. Biol. Technol. Agric. 9 (1), 104. 10.1186/s40538-022-00373-5 (2022).
61. Liu L Silicon effects on biomass carbon and phytolith-occluded carbon in grasslands under high-salinity conditions Front. Plant. Sci. 2020 11 657 10.3389/fpls.2020.00657 32528507
Liu, L. et al. Silicon effects on biomass carbon and phytolith-occluded carbon in grasslands under high-salinity conditions. Front. Plant. Sci. 11, 657. 10.3389/fpls.2020.00657 (2020).32528507
62. Tejera NA Soussi M Lluch C Physiological and nutritional indicators of tolerance to salinity in chickpea plants growing under symbiotic conditions Environ. Exp. Bot. 2006 58 1–3 17 24 10.1016/j.envexpbot.2005.06.007
Tejera, N. A., Soussi, M. & Lluch, C. Physiological and nutritional indicators of tolerance to salinity in chickpea plants growing under symbiotic conditions. Environ. Exp. Bot. 58 (1–3), 17–24. 10.1016/j.envexpbot.2005.06.007 (2006).
63. Ullah N Mitigation the adverse effect of salinity stress on the performance of the tomato crop by exogenous application of Chitosan Bull. Natl. Res. Cent. 2020 44 1 1 11 10.1186/s42269-020-00435-4
Ullah, N. et al. Mitigation the adverse effect of salinity stress on the performance of the tomato crop by exogenous application of Chitosan. Bull. Natl. Res. Cent. 44 (1), 1–11. 10.1186/s42269-020-00435-4 (2020).
64. Khan I Organic amendments improved the Productivity and Bio-fortification of Fine Rice by Improving Physiological Responses and Nutrient Homeostasis under salinity stress Plants 2023 12 8 1644 10.3390/plants12081644 37111867
Khan, I. et al. Organic amendments improved the Productivity and Bio-fortification of Fine Rice by Improving Physiological Responses and Nutrient Homeostasis under salinity stress. Plants. 12 (8), 1644. 10.3390/plants12081644 (2023).37111867
65. Latif HH Mohamed HI Exogenous applications of moringa leaf extract effect on retrotransposon, ultrastructural and biochemical contents of common bean plants under environmental stresses South. Afr. J. Bot. 2016 106 221 231 10.1016/j.sajb.2016.07.010
Latif, H. H. & Mohamed, H. I. Exogenous applications of moringa leaf extract effect on retrotransposon, ultrastructural and biochemical contents of common bean plants under environmental stresses. South. Afr. J. Bot. 106, 221–231. 10.1016/j.sajb.2016.07.010 (2016).
66. Hossain MA Integrating BLUP, AMMI, and GGE models to explore GE Interactions for Adaptability and Stability of Winter Lentils (Lens culinaris Medik) Plants 2023 12 2079 10.3390/plants12112079 37299058
Hossain, M. A. et al. Integrating BLUP, AMMI, and GGE models to explore GE Interactions for Adaptability and Stability of Winter Lentils (Lens culinaris Medik). Plants. 12, 2079. 10.3390/plants12112079 (2023).37299058
67. Azam MG Phenotypic diversity in qualitative and quantitative traits for selection of high yield potential field pea genotypes Sci. Rep. 2024 14 18561 10.1038/s41598-024-69448-7 39122809
Azam, M. G. Phenotypic diversity in qualitative and quantitative traits for selection of high yield potential field pea genotypes. Sci. Rep. 14, 18561. 10.1038/s41598-024-69448-7 (2024).39122809
68. Azam MG Genetic analyses of mungbean [Vigna radiata (L.) Wilczek] breeding traits for selecting superior genotype(s) using multivariate and multi-traits indexing approaches Plants 2023 12 10 1984 10.3390/plants12101984 37653901
Azam, M. G. et al. Genetic analyses of mungbean [Vigna radiata (L.) Wilczek] breeding traits for selecting superior genotype(s) using multivariate and multi-traits indexing approaches. Plants. 12 (10), 1984. 10.3390/plants12101984 (2023).37653901
69. Azam MG Genetic analysis in Grain legumes [Vigna radiata (L.) Wilczek] for yield improvement and identifying Heterotic hybrids Plants 2022 11 1774 10.3390/plants11131774 35807726
Azam, M. G. et al. Genetic analysis in Grain legumes [Vigna radiata (L.) Wilczek] for yield improvement and identifying Heterotic hybrids. Plants. 11, 1774. 10.3390/plants11131774 (2022).35807726
70. Azam MG Sarker U Uddin MS Screening maize (Zea mays L.) genotypes for phosphorus deficiency at the seedling stage Turk. J. Agric. For. 2022 46 6 802 821 10.55730/1300-011X.3044
Azam, M. G., Sarker, U. & Uddin, M. S. Screening maize (Zea mays L.) genotypes for phosphorus deficiency at the seedling stage. Turk. J. Agric. For. 46 (6), 802–821. 10.55730/1300-011X.3044 (2022).
71. Azam MG Sarker U Banik BR Genetic variability of yield and its contributing characters of CIMMYT maize inbreds under drought stress Bangladesh J. Agri Res. 2014 39 3 419 426 10.3329/bjar.v39i3.21985
Azam, M. G., Sarker, U., Banik, B. R. & Maniruzzam & Genetic variability of yield and its contributing characters of CIMMYT maize inbreds under drought stress. Bangladesh J. Agri Res. 39 (3), 419–426. 10.3329/bjar.v39i3.21985 (2014).
72. Hasan MJ Assessment of GGE, AMMI, Regression, and its deviation model to identify stable Rice hybrids in Bangladesh Plants 2022 11 2336 10.3390/plants11182336 36145737
Hasan, M. J. et al. Assessment of GGE, AMMI, Regression, and its deviation model to identify stable Rice hybrids in Bangladesh. Plants. 11, 2336. 10.3390/plants11182336 (2022).36145737
73. Habib MA Climate-smart rice (Oryza sativa L.) genotypes identification using stability analysis, multi-trait selection index, and genotype-environment interaction at different irrigation regimes with adaptation to universal warming Sci. Rep. 2024 14 13836 10.1038/s41598-024-64808-9 38879711
Habib, M. A. et al. Climate-smart rice (Oryza sativa L.) genotypes identification using stability analysis, multi-trait selection index, and genotype-environment interaction at different irrigation regimes with adaptation to universal warming. Sci. Rep. 14, 13836. 10.1038/s41598-024-64808-9 (2024).38879711
74. Faysal, A. S. M. et al. Genetic variability, character association, and path coefficient analysis in transplant Aman rice genotypes. Plants 11(21), 2952. (2022). 10.3390/plants11212952
75. Kulsum, U., Sarker, U. & Rasul, M. G. Genetic variability, heritability and interrelationship in salt-tolerant lines of T. Aman rice. Genetika 54(2), 761–776. (2022). 10.2298/GENSR2202761K
76. Hasan-Ud-Daula M Sarker U Variability, heritability, character association, and path coefficient analysis in advanced breeding lines of rice (Oryza sativa L ) Genetika 2020 52 2 711 726 10.2298/GENSR2002711H
Hasan-Ud-Daula, M. & Sarker, U. Variability, heritability, character association, and path coefficient analysis in advanced breeding lines of rice (Oryza sativa L. ) Genetika. 52 (2), 711–726. 10.2298/GENSR2002711H (2020).
77. Hasan MJ Kulsum MU Majumder RR Sarker U Genotypic variability for Grain Quality attributes in Restorer lines of Hybrid Rice Genetika 2020 52 973 989 10.2298/GENSR2003973H
Hasan, M. J., Kulsum, M. U., Majumder, R. R. & Sarker, U. Genotypic variability for Grain Quality attributes in Restorer lines of Hybrid Rice. Genetika. 52, 973–989. 10.2298/GENSR2003973H (2020).
78. Rashad MMI Sarker U Genetic variations in yield and yield contributing traits of green amaranth Genetika 2020 52 1 393 407 10.2298/GENSR2001393R
Rashad, M. M. I. & Sarker, U. Genetic variations in yield and yield contributing traits of green amaranth. Genetika. 52 (1), 393–407. 10.2298/GENSR2001393R (2020).
79. Sarker U Azam MG Talukder MZA Genetic variation in mineral profiles, yield contributing agronomic traits, and foliage yield of stem Amaranth Genetika 2022 54 1 91 108 10.2298/GENSR2201091S
Sarker, U., Azam, M. G. & Talukder, M. Z. A. Genetic variation in mineral profiles, yield contributing agronomic traits, and foliage yield of stem. Amaranth Genetika. 54 (1), 91–108. 10.2298/GENSR2201091S (2022).
80. Chen T Molecular mechanisms of salinity tolerance in rice Crop J. 2021 9 506 520 10.1016/j.cj.2021.03.005
Chen, T. et al. Molecular mechanisms of salinity tolerance in rice. Crop J. 9, 506–520. 10.1016/j.cj.2021.03.005 (2021).
81. Sultan I Improved salinity tolerance in early growth stage of maize through salicylic acid foliar application Ital. J. Agron. 2021 16 3 1 11 10.4081/IJA.2021.1810
Sultan, I. et al. Improved salinity tolerance in early growth stage of maize through salicylic acid foliar application. Ital. J. Agron. 16 (3), 1–11. 10.4081/IJA.2021.1810 (2021).
82. Akhtar SS Andersen MN Naveed M Zahir ZA Liu F Interactive effect of biochar and plant growth-promoting bacterial endophytes on ameliorating salinity stress in maize Funct. Plant. Biol. 2015 42 8 770 781 10.1071/FP15054 32480720
Akhtar, S. S., Andersen, M. N., Naveed, M., Zahir, Z. A. & Liu, F. Interactive effect of biochar and plant growth-promoting bacterial endophytes on ameliorating salinity stress in maize. Funct. Plant. Biol. 42 (8), 770–781. 10.1071/FP15054 (2015b).32480720
83. Szabados L Savoure A Proline: a multifunctional amino acid Trends Plant. Sci. 2010 15 2 89 97 10.1016/j.tplants.2009.11.009 20036181
Szabados, L. & Savoure, A. Proline: a multifunctional amino acid. Trends Plant. Sci. 15 (2), 89–97. 10.1016/j.tplants.2009.11.009 (2010).20036181
84. Zhang G Yan Z Wang Y Feng Y Yuan Q Exogenous proline improves the growth and yield of lettuce with low potassium content Sci. Hortic. 2020 271 109469 10.1016/j.scienta.2020.109469
Zhang, G., Yan, Z., Wang, Y., Feng, Y. & Yuan, Q. Exogenous proline improves the growth and yield of lettuce with low potassium content. Sci. Hortic. 271, 109469. 10.1016/j.scienta.2020.109469 (2020).
85. Bolouri-Moghaddam MR Le Roy K Xiang L Rolland F Van den Ende W Sugar signalling and antioxidant network connections in plant cells FEBS J. 2010 277 9 2022 2037 10.1111/j.1742-4658.2010.07633.x 20412056
Bolouri-Moghaddam, M. R., Le Roy, K., Xiang, L., Rolland, F. & Van den Ende, W. Sugar signalling and antioxidant network connections in plant cells. FEBS J. 277 (9), 2022–2037. 10.1111/j.1742-4658.2010.07633.x (2010).20412056
86. Zhang G Exogenous application of Chitosan alleviates salinity stress in lettuce (Lactuca sativa L) Horticulturae 2021 7 10 342 10.3390/horticulturae7100342
Zhang, G. et al. Exogenous application of Chitosan alleviates salinity stress in lettuce (Lactuca sativa L). Horticulturae. 7 (10), 342. 10.3390/horticulturae7100342 (2021).
87. Ghani, M. I. et al. Potential of melatonin and Trichoderma harzianum inoculation in ameliorating salt toxicity in watermelon: Insights into antioxidant system, leaf ultrastructure, and gene regulation. Plant Physiology and Biochemistry, 211, p.108639
88. Souri, M.K. and Tohidloo, G. Effectiveness of different methods of salicylic acid application on growth characteristics of tomato seedlings under salinity. Chemical and Biological Technologies in Agriculture, 6(1), 26 (2019).
89. Acosta-Motos J Plant responses to salt stress: adaptive mechanisms Agronomy 2017 7 1 18 10.3390/agronomy7010018
Acosta-Motos, J. et al. Plant responses to salt stress: adaptive mechanisms. Agronomy. 7 (1), 18 (2017).
90. Mekawy AMM Growth, physiological adaptation, and gene expression analysis of two Egyptian rice cultivars under salt stress Plant. Physiol. Biochem. 2015 87 17 25 10.1016/j.plaphy.2014.12.007 25532120
Mekawy, A. M. M. et al. Growth, physiological adaptation, and gene expression analysis of two Egyptian rice cultivars under salt stress. Plant. Physiol. Biochem. 87, 17–25. 10.1016/j.plaphy.2014.12.007 (2015).25532120
91. Hernández-Hernández H Effects of chitosan–PVA and Cu nanoparticles on the growth and antioxidant capacity of tomato under saline stress Molecules 2018 23 1 178 10.3390/molecules23010178 29337864
Hernández-Hernández, H. et al. Effects of chitosan–PVA and Cu nanoparticles on the growth and antioxidant capacity of tomato under saline stress. Molecules. 23 (1), 178. 10.3390/molecules23010178 (2018).29337864
92. Younas HS Abid M Shaaban M Ashraf M Influence of silicon and chitosan on growth and physiological attributes of maize in a saline field Physiol. Mol. Biol. Plants 2021 27 387 397 10.1007/s12298-021-00940-4 33707876
Younas, H. S., Abid, M., Shaaban, M. & Ashraf, M. Influence of silicon and chitosan on growth and physiological attributes of maize in a saline field. Physiol. Mol. Biol. Plants. 27, 387–397. 10.1007/s12298-021-00940-4 (2021).33707876
93. Akhtar SS Andersen MN Liu FL Residual effects of biochar on improving growth, physiology and yield of wheat under salt stress Agric. Water Manage. 2015 158 61 68 10.1016/j.agwat.2015.04.010
Akhtar, S. S., Andersen, M. N. & Liu, F. L. Residual effects of biochar on improving growth, physiology and yield of wheat under salt stress. Agric. Water Manage. 158, 61–68 (2015).
94. Melas GB Ortiz O Alacañiz JM Can biochar protect labile organic matter against mineralization in soil? Pedosphere 2017 27 822 831 10.1016/S1002-0160(17)60421-1
Melas, G. B., Ortiz, O. & Alacañiz, J. M. Can biochar protect labile organic matter against mineralization in soil? Pedosphere. 27, 822–831 (2017).
95. Huang K Soil acidification and salinity: the importance of biochar application to agricultural soils Front. Plant. Sci. 2023 14 1206820 10.3389/fpls.2023.1206820 37780526
Huang, K. et al. Soil acidification and salinity: the importance of biochar application to agricultural soils. Front. Plant. Sci. 14, 1206820. 10.3389/fpls.2023.1206820 (2023).37780526
96. Wu Y The critical role of biochar to mitigate the adverse impacts of drought and salinity stress in plants Front. Plant. Sci. 2023 14 1163451 10.3389/fpls.2023.1163451 37223815
Wu, Y. et al. The critical role of biochar to mitigate the adverse impacts of drought and salinity stress in plants. Front. Plant. Sci. 14, 1163451. 10.3389/fpls.2023.1163451 (2023).37223815
97. Qin H Rice os DOF 15 contributes to ethylene-inhibited primary root elongation under salt stress New. Phytol 2019 223 798 813 10.1111/nph.15824 30924949
Qin, H. et al. Rice os DOF 15 contributes to ethylene-inhibited primary root elongation under salt stress. New. Phytol. 223, 798–813. 10.1111/nph.15824 (2019).30924949
98. Farouk S Al-Huqail AA El-Gamal SM Potential role of Biochar and Silicon in improving physio-biochemical and Yield Characteristics of Borage Plants under different irrigation regimes Plants 2023 12 8 1605 10.3390/plants12081605 37111829
Farouk, S., Al-Huqail, A. A. & El-Gamal, S. M. Potential role of Biochar and Silicon in improving physio-biochemical and Yield Characteristics of Borage Plants under different irrigation regimes. Plants. 12 (8), 1605. 10.3390/plants12081605 (2023).37111829
99. Athar HUR Salt stress proteins in plants: an overview Front. Plant. Sci. 2023 13 999058 10.3389/fpls.2022.999058
Athar, H. U. R. et al. Salt stress proteins in plants: an overview. Front. Plant. Sci. 13, 999058. 10.3389/fpls.2022.999058 (2023).
100. Farhangi-Abriz S Ghassemi-Golezani K Improving electrochemical characteristics of plant roots by biochar is an efficient mechanism in increasing cations uptake by plants Chemosphere 2023 313 137365 10.1016/j.chemosphere.2022.137365 36427572
Farhangi-Abriz, S. & Ghassemi-Golezani, K. Improving electrochemical characteristics of plant roots by biochar is an efficient mechanism in increasing cations uptake by plants. Chemosphere. 313, 137365. 10.1016/j.chemosphere.2022.137365 (2023).36427572
101. Ebrahimi M Mousavi A Souri MK Sahebani N Can vermicompost and biochar control Meloidogyne Javanica on eggplant? Nematology 2021 1 1 12
Ebrahimi, M., Mousavi, A., Souri, M. K. & Sahebani, N. Can vermicompost and biochar control Meloidogyne Javanica on eggplant? Nematology. 1, 1–12 (2021).
102. Lu H Changes in soil microbial community structure and enzyme activity with amendment of biochar-manure compost and pyroligneous solution in a saline soil from Central China Eur. J. Soil. Biol. 2015 70 67 76 10.1016/j.ejsobi.2015.07.005
Lu, H. et al. Changes in soil microbial community structure and enzyme activity with amendment of biochar-manure compost and pyroligneous solution in a saline soil from Central China. Eur. J. Soil. Biol. 70, 67–76. 10.1016/j.ejsobi.2015.07.005 (2015).
103. Nigussie A Kissi E Misganaw M Ambaw G Effect of biochar application on soil properties and nutrient uptake of Lettuces (Lactuca sativa) grown in chromium polluted soils American-Eurasian J. Agric. Environ. Sci. 2012 12 3 369 376
Nigussie, A., Kissi, E., Misganaw, M. & Ambaw, G. Effect of biochar application on soil properties and nutrient uptake of Lettuces (Lactuca sativa) grown in chromium polluted soils. American-Eurasian J. Agric. Environ. Sci. 12 (3), 369–376 (2012).
104. Rawat J Saxena J Sanwal P Biochar: A Sustainable Approach for improving Plant Growth and Soil Properties IntechOpen 2019 10.5772/intechopen.82151
Rawat, J., Saxena, J. & Sanwal, P. Biochar: A Sustainable Approach for improving Plant Growth and Soil Properties. IntechOpen. 10.5772/intechopen.82151 (2019).
105. Ebrahimi M Mousavi A Souri MK Sahebani N Biochar and Vermicompost improve growth and physiological traits of eggplant (Solanum melongena L.) under deficit irrigation Che Bio Tech. Agri 2021 8 1 1 14
Ebrahimi, M., Mousavi, A., Souri, M. K. & Sahebani, N. Biochar and Vermicompost improve growth and physiological traits of eggplant (Solanum melongena L.) under deficit irrigation. Che Bio Tech. Agri. 8 (1), 1–14 (2021).
106. Munns R Tester M Mechanisms of salinity tolerance Annu. Rev. Plant. Biol. 2008 59 651 681 10.1146/annurev.arplant.59.032607.092911 18444910
Munns, R. & Tester, M. Mechanisms of salinity tolerance. Annu. Rev. Plant. Biol. 59, 651–681. 10.1146/annurev.arplant.59.032607.092911 (2008). (2008).18444910
107. Abbas A Synergistic use of biochar and acidified manure for improving growth of maize in chromium contaminated soil Int. J. Phytorem. 2020 22 1 52 61 10.1080/15226514.2019.1644286
Abbas, A. et al. Synergistic use of biochar and acidified manure for improving growth of maize in chromium contaminated soil. Int. J. Phytorem. 22 (1), 52–61. 10.1080/15226514.2019.1644286 (2020).
108. El-Beltagi HS Mohamed HI Sofy Role of ascorbic acid, glutathione, and proline applied as singly or in sequence combination in improving chickpea plant through physiological change and antioxidant defense under different levels of irrigation intervals Molecules 2020 25 7 1702 10.3390/molecules25071702 32276352
El-Beltagi, H. S., Mohamed, H. I. & Sofy Role of ascorbic acid, glutathione, and proline applied as singly or in sequence combination in improving chickpea plant through physiological change and antioxidant defense under different levels of irrigation intervals. Molecules. 25 (7), 1702. 10.3390/molecules25071702 (2020).32276352
109. Al-Ashkar I Alderfasi A El-Hendawy S Al-Suhaibani N El-Kafafi S Seleiman MF detecting salt tolerance in doubled haploid wheat lines Agronomy 2019 9 4 211 10.3390/agronomy9040211
Al-Ashkar, I., Alderfasi, A., El-Hendawy, S., Al-Suhaibani, N. & El-Kafafi, S. Seleiman MF detecting salt tolerance in doubled haploid wheat lines. Agronomy. 9 (4), 211. 10.3390/agronomy9040211 (2019).
110. El-Tantawy EM Behavior of tomato plants as affected by spraying with chitosan and aminofort as natural stimulator substances under application of soil organic amendments Pakistan J. Biol. Sci. 2009 12 1164 1173 10.3923/pjbs.2009.1164.1173
El-Tantawy, E. M. Behavior of tomato plants as affected by spraying with chitosan and aminofort as natural stimulator substances under application of soil organic amendments. Pakistan J. Biol. Sci. 12, 1164–1173. 10.3923/pjbs.2009.1164.1173 (2009).
111. Mamun MAA Application of Potassium after Waterlogging improves Quality and Productivity of soybean seeds Life 2022 12 1816 10.3390/life12111816 36362971
Mamun, M. A. A. et al. Application of Potassium after Waterlogging improves Quality and Productivity of soybean seeds. Life. 12, 1816. 10.3390/life12111816 (2022).36362971
112. Islam, M. R. et al. Potassium augments growth, yield, nutrient content, and drought tolerance in mung bean (Vigna radiata L. Wilczek.). Sci Rep 14, 9378 doi.10.1038/s41598-024-60129-z (2024).
113. Dola DB Nano-iron oxide accelerates growth, yield, and quality of Glycine max seed in water deficits Front. Plant. Sci. 2022 13 992535 10.3389/fpls.2022.992535 36160973
Dola, D. B. et al. Nano-iron oxide accelerates growth, yield, and quality of Glycine max seed in water deficits. Front. Plant. Sci. 13, 992535. 10.3389/fpls.2022.992535 (2022).36160973
114. FRG Fertilizer Recommendation Guide. BARC (Bangladesh Agricultural Research Council), Farmgate, Dhaka-1215. (2012). (2012).
115. Lichtenthaler HK Chlorophylls, and carotenoids: pigments of photosynthetic biomembranes Meth Enzymol. 1987 148 350 382 10.1016/0076-6879(87)48036-1
Lichtenthaler, H. K. Chlorophylls, and carotenoids: pigments of photosynthetic biomembranes. Meth Enzymol. 148, 350–382. 10.1016/0076-6879(87)48036-1 (1987).
116. Sullivan, C. Y. Selecting for drought and heat resistance in grain sorghum. Stress Physiol. Crop Plants, 263–281. (1979).
117. Hayat S Ali B Hasan SA Ahmad A Brassinosteroid enhanced the level of antioxidants under cadmium stress in Brassica juncea Environ. Exp. Bot. 2007 60 33 41 10.1016/j.envexpbot.2006.06.002
Hayat, S., Ali, B., Hasan, S. A. & Ahmad, A. Brassinosteroid enhanced the level of antioxidants under cadmium stress in Brassica juncea. Environ. Exp. Bot. 60, 33–41. 10.1016/j.envexpbot.2006.06.002 (2007).
118. Sangakkara HR Hartwig UA Nosberger J Response of root branching and shoot water potential of Phaseolus vulgaris L. to soil moisture and fertilizer potassium J. Agron. Crop Sci. 1996 177 165 173 10.1111/j.1439-037X.1996.tb00234.x
Sangakkara, H. R., Hartwig, U. A. & Nosberger, J. Response of root branching and shoot water potential of Phaseolus vulgaris L. to soil moisture and fertilizer potassium. J. Agron. Crop Sci. 177, 165–173. 10.1111/j.1439-037X.1996.tb00234.x (1996).
119. Su MS Silva J Antioxidant activity, anthocyanins, and phenolics of rabbiteye blueberry (Vaccinium ashei) by-products as affected by fermentation Food Chem. 2006 97 447 451 10.1016/j.foodchem.2005.05.023
Su, M. S. & Silva, J. Antioxidant activity, anthocyanins, and phenolics of rabbiteye blueberry (Vaccinium ashei) by-products as affected by fermentation. Food Chem. 97, 447–451. 10.1016/j.foodchem.2005.05.023 (2006).
120. Bates LS Waldem RP Teare ID Rapid determination of free proline for water-stress studies Plant. Soil. 1973 39 205 207 10.1007/BF00018060
Bates, L. S., Waldem, R. P. & Teare, I. D. Rapid determination of free proline for water-stress studies. Plant. Soil. 39, 205–207 (1973).
121. Jagota SK Dani HM A new colorimetric technique for the estimation of vitamin C using Folin phenol reagent Anal. Biochem. 1982 127 1 178 182 10.1016/0003-2697(82)90162-2 7165085
Jagota, S. K. & Dani, H. M. A new colorimetric technique for the estimation of vitamin C using Folin phenol reagent. Anal. Biochem. 127 (1), 178–182. 10.1016/0003-2697(82)90162-2 (1982).7165085
122. Dey, P. Oligosaccharides. Methods Plant Biochem. Carbohydr., 2, 189–218. (1990). 10.1016/B978-0-12-461012-5.50011-2
123. Bremner JM Nitrogen availability indexes. Methods of Soil Analysis: part 2 Chem Microbiol. Prop. 1965 9 1324 1345 10.2134/agronmonogr9.2.c37
Bremner, J. M. Nitrogen availability indexes. Methods of Soil Analysis: part 2 Chem. Microbiol. Prop. 9, 1324–1345. 10.2134/agronmonogr9.2.c37 (1965).
124. Piper, C. S. Soil and Plant Analysis (Han’s Publication, 1966).
125. Jackson, M. L. Soil Chemical Analysis. Prentice Hall of India Pvt144–197 (Ltd, 1967).
126. Page, A. I., Miller, R. H. & Keeny, D. R. Methods of Soil Analysis. Part II. Chemical and Microbiological Methods 2nd edn 225–246 (Am. Soc. Agron. Madison, WI, 1982).
127. Rahman MM Combining ability analysis and marker-based prediction of heterosis in yield reveal prominent heterotic combinations from diallel population of rice Agron 2022 12 8 1797 10.3390/agronomy12081797
Rahman, M. M. et al. Combining ability analysis and marker-based prediction of heterosis in yield reveal prominent heterotic combinations from diallel population of rice. Agron. 12 (8), 1797. 10.3390/agronomy12081797 (2022).
128. Prodhan MM Foliar application of GA3 stimulates seed production in cauliflower Agron 2022 12 6 1394 10.3390/agronomy12061394
Prodhan, M. M. et al. Foliar application of GA3 stimulates seed production in cauliflower. Agron. 12 (6), 1394. 10.3390/agronomy12061394 (2022).
129. Azad AK Evaluation of combining ability and heterosis of popular restorer and male sterile lines for the development of superior rice hybrids Agron 2022 12 4 965 10.3390/agronomy12040965
Azad, A. K. et al. Evaluation of combining ability and heterosis of popular restorer and male sterile lines for the development of superior rice hybrids. Agron. 12 (4), 965. 10.3390/agronomy12040965 (2022).
130. Jahan, N. et al. Evaluation of yield attributes and bioactive phytochemicals of twenty amaranth genotypes of Bengal floodplain. Heliyon 9(9), e19644, doi.10.1016/j.heliyon.2023.e19644 (2023).
