Nitrogen concentration in quinoa seeds was significantly influenced by the year, irrigation levels, and vermicompost application, including their triple interaction (P < 0.01). Notably, there was an increase in seed nitrogen concentration in the second year, with varying effects of irrigation and vermicompost levels. Upon examining the interaction effect of year with different irrigation and vermicompost levels, it was observed that as the vermicompost level increased across all irrigation treatments, the nitrogen percentage in quinoa seeds also increased. Additionally, seed nitrogen concentration was higher in low irrigation treatments compared to full irrigation and 125% of the water requirement (Fig. 1).
Seed oil
The study revealed a significant impact of irrigation × vermicompost application on seed oil content (p < 0.001). However, the triple interaction of year × irrigation × vermicompost was not statistically significant (p < 0.001 and p < 0.05). The seed oil content ranged from 2.31% under 50% irrigation stress without vermicompost to 4.2% under 125% irrigation level with 15 tons per hectare of vermicompost. The comparison of means indicated a substantial increase in oil percentage with the use of vermicompost across all tested irrigation levels. Conversely, in treatments without vermicompost, irrigation did not lead to an increase in the oil percentage. The oil percentage in the irrigation treatments at 50% of the plant's water requirement with 10 and 15 tons per hectare of vermicompost was statistically similar at 3.37% and 3.52%, respectively, to the oil percentage in the treatment at 75% of the plant's water requirement with 5 tons per hectare of vermicompost, which was 3.93% (Table 4).
Table 4. Mean comparison of the effect of different levels of irrigation and vermicompost on Fe, Mn and Cu concentration and seed oil.
|
Irrigation (%)
|
Vermicompost
(t ha− 1)
|
Fe concentration
|
Mn concentration
|
Cu concentration
|
|
Seed oil
|
mg kg− 1
|
|
%
|
50
|
0
|
228.04j
|
49.54c
|
1.91d
|
|
2.31d
|
5
|
236.34j
|
53.45bc
|
2.23c
|
|
3bc
|
10
|
244.01i
|
59.42b
|
2.25c
|
|
3.52b
|
15
|
262.66h
|
63.39a
|
2.62bc
|
|
3.37b
|
75
|
0
|
245.67i
|
43.35d
|
2.29c
|
|
2.79c
|
5
|
272.16g
|
48.29c
|
2.26c
|
|
3.59b
|
10
|
290.81f
|
56.25b
|
2.22c
|
|
3.8ab
|
15
|
302.26e
|
47.54b
|
2.25c
|
|
4.2a
|
100
|
0
|
275.01g
|
41.06d
|
3.12b
|
|
2.8c
|
5
|
306.48d
|
42.4d
|
3.29ab
|
|
4a
|
10
|
352.06c
|
43.34d
|
3.25ab
|
|
3.9a
|
15
|
368.75a
|
53.32bc
|
3.82a
|
|
4a
|
125
|
0
|
275.06g
|
4.32d
|
3b
|
|
2.9c
|
5
|
309.03d
|
49.94d
|
2.77c
|
|
3.9a
|
10
|
346.74c
|
55.4b
|
3.26ab
|
|
4.1a
|
15
|
370.07a
|
54.64b
|
3.23ab
|
|
3.8a
|
The same letters indicate statitically insignificant differences (P < 0.05).
|
The findings of numerous studies have demonstrated that water deficiency during seed filling reduces seed yield per unit area by impeding photosynthesis. Water stress during the seed filling stage, particularly when coupled with elevated temperatures, accelerates leaf aging, shortens the seed filling period, decreases average seed weight, and reduces overall yield. This is attributed to the diminished transfer of photosynthetic materials to developing seeds (Piri et al. 2022). The higher seed yield in stress-free conditions can be attributed to the availability of ample water for the plant, which enhances yield components and ultimately boosts seed yield. Sufficient water during plant growth enhances source and sink strength, leading to increased seed yield (Piri et al. 2022).
In addition to its nutrient richness compared to other fertilizers, vermicompost possesses characteristics such as gradual nutrient release, particularly nitrogen, which mitigates nitrate leaching. Calcium and potassium compounds in vermicompost reduce soil acidity and increase soil organic matter, thereby enhancing water and nutrient retention in the soil (Wang et al. 2022). Conversely, augmenting seed yield through increased vermicompost application can be achieved by promoting plant vegetative growth, enhancing reserve material production, improving flower fertility, and increasing thousand-seed weight, ultimately leading to increased seed yield (Arancon et al. 2022). Our results corroborate the earlier findings of (Ranva and Singh 2021; Wang et al. 2022). It appears that in the current experiment, the reduced biological yield in treatments receiving less water than the plant's requirement is due to water stress leading to reduced stem and leaf formation, consequently diminishing the plant's dry matter yield. In simpler terms, as drought stress increases, soil moisture decreases during plant growth, leading to reduced material transfer and limiting the production of photosynthetic materials in plants. The decrease in plant photosynthesis due to stress factors also reduces hydrostatic pressure and material transfer rate, which may explain the decreased biological performance in treatments with low irrigation (González-Teuber et al. 2018).
The presence of vermicompost in the soil enhances soil fertility, storage capacity, and porosity. These compounds gradually release nutrients, making them available to plants over time. Vermicompost fertilizers not only provide essential nutrients for plant growth but also increase water availability, promote vegetative growth, and enhance biomass production (Singer et al. 2007). Vermicompost contains vital elements like nitrogen, phosphorus, potassium, calcium, iron, manganese, copper, zinc, and boron in forms that are readily usable by plants and soluble in water (Wang et al. 2022). These findings align with previous studies (Rani et al. 2020; Demir 2019).
The concentration of nutrients in seeds increases with higher irrigation levels due to improved soil moisture, nutrient availability, and enhanced nutrient absorption by plants. Continuous soil moisture availability under drip irrigation helps dissolve nutrients near the roots, facilitating better absorption at higher irrigation levels (Tang et al. 2023). Nutrient absorption is hindered in water scarcity due to reduced transpiration, disruptions in transport systems, membrane permeability, and decreased root absorption capacity. Lower soil moisture reduces nutrient diffusion to the roots and impairs root system efficiency. Phosphorus fixation in dry conditions contributes to its scarcity in plants (Bardel et al. 2023). Studies have shown reduced phosphorus levels in quinoa seeds under stress compared to control treatments (EL-Tahan et al. 2019).
It seems that vermicompost, rich in nutrients and microorganism activity, facilitates plant access to essential resources like phosphorus by stimulating soil microorganisms and aiding nutrient absorption. Additionally, the compounds in vermicompost acidify the rhizosphere environment, leading to the release of insoluble phosphorus as H+ ions are replaced with calcium ions. Phosphorus, being immobile in soil, is absorbed by roots when they encounter organic or inorganic substances containing its absorbable form (Hanč et al. 2018). Research indicates that potassium concentrations decrease under drought stress due to reduced potassium availability in dry conditions. Higher irrigation levels, by increasing soil moisture content, result in a greater proportion of monovalent ions like potassium in the soil solution compared to divalent ions such as magnesium and calcium. Consequently, plants absorb a higher percentage of potassium at these levels and store it in their seeds. As soil moisture decreases, clay colloids gradually bind potassium more tightly to their surfaces, inhibiting the release of these ions. Furthermore, under water stress, plant growth, including root absorption activity, is hindered, reducing the plant's ability to take up potassium from the clay colloids, resulting in decreased absorption of these elements (Rani et al. 2020). Conversely, the decline in potassium concentration in seeds during drought stress may be attributed to the translocation of these elements from the seed to the root, where potassium serves as an osmotic regulator in such conditions. Potassium ions play a crucial role as a secondary messenger in plants, participating in the transmission of diverse signals, making it a key component in plants' response to drought (Osuagwu et al. 2010). The increase in potassium concentration observed with vermicompost application can be linked to enhanced nutrient cycling and improved physical and biological soil properties. Additionally, vermicompost boosts the population of beneficial soil microorganisms, which in turn secrete organic acids, creating an acidic rhizosphere environment that enhances plant access to potassium (Agegnehu et al. 2016).
The decrease in calcium content in the complete irrigation treatments can be attributed to several factors (Bardel et al. 2023). Firstly, the large water limitations imposed in those treatments may have led to reduced oxygen supply to the roots. This, in turn, can hinder respiration, nutrient absorption, and other root activities, ultimately affecting calcium uptake. In a separate study investigating the impact of different irrigation levels on quinoa seeds, it was found that the concentration of calcium was higher in the low irrigation treatment compared to the full irrigation treatment (Walters et al. 2016). This suggests that reduced water availability may have a positive effect on calcium accumulation in the seeds.
The increase in seed nitrogen concentration with a decrease in soil moisture can be attributed to a concentration effect. When soil moisture decreases, the growth rate of the plant's aerial parts tends to decrease more than the rate of nitrogen absorption (Bardel et al. 2023). As a result, the nitrogen becomes more concentrated in the seeds. In treatments that received more water, one of the main reasons for the decrease in seed nitrogen concentration is related to nitrogen wastage, particularly through leaching or washing away of nitrogen from the soil (Rani et al. 2020). This suggests that excessive watering can lead to the loss of nitrogen, thereby reducing its concentration in the seeds.
In agriculture, when there is sufficient water available in the field during the growth period, plants do not experience water shortage during the physiological ripening stage. Irrigation improves the plant's condition by enhancing photosynthesis and enabling nitrogen absorption to continue throughout the growth period (Bole and Dubetz 1986). Organic fertilizers, such as vermicompost, contain high levels of organic compounds and serve as rich sources of nutrients, particularly nitrogen. Incorporating vermicompost into the soil not only increases the supply of essential nutrients for plants but also enhances soil physical conditions and vital processes. This creates an optimal substrate for root growth, promotes aerial organ development, and boosts nitrogen absorption (Arancón et al. 2022).
In stressful conditions, the accumulation of sodium in various plant tissues, especially seeds, is a result of increased root absorption and xylem-to-leaf transport. This mechanism establishes osmotic balance in the plant, facilitating enhanced water uptake (EL-Tahan et al. 2019).
Vermicompost is a rich source of nutrients, particularly nitrogen, that can enhance plant growth and performance. Studies have shown that the use of vermicompost fertilizer can increase the absorption and transfer of sodium to plant seeds, which helps establish osmotic balance and facilitates water uptake (Gutierrez-Miceli et al. 2007). However, the absorption of zinc is primarily through active absorption mechanisms, and drought stress can limit the transfer of this element to the seed, resulting in decreased zinc concentration. Additionally, drought stress can reduce the amount of active absorption of elements and the production of photosynthetic substances, leading to decreased absorption of elements and reduced growth (Marschner 2012; Osborne et al. 2022).
The high percentage of humic substances in vermicompost, with their low molecular weight, plays a crucial role in stabilizing cell membranes and enhancing the absorption of various essential nutrients, including zinc (Li et al. 2017). Furthermore, the decomposition of organic fertilizers, particularly vermicompost, leads to a decrease in soil pH and the formation of zinc chelates through the addition of organic compounds, effectively increasing the availability of zinc. The presence of short-chain organic matter in the soil enhances the mobility and solubility of zinc, making it more readily available to plants through bonding with zinc (Marschner 2012).
Low soil moisture levels have been found to reduce the absorption of iron by plant roots, which justifies the decrease in iron concentration in drought stress treatments (EL-Tahan et al. 2019). The characteristics of soil organic matter play a role in the availability of low-use nutrients in the soil. As organic matter decomposes, the nutrients it contains are released. This decomposition process also leads to a decrease in soil pH, resulting in increased concentrations of nutrients, particularly low-use elements like iron, in the soil. Research has shown that the addition of vermicompost to the soil can enhance the solubility of iron by forming a complex with it, thereby preventing its precipitation (Li et al. 2017). Other studies have reported a significant decrease in the concentration of iron in quinoa seeds under drought stress (Aly et al. 2018; EL-Tahan et al. 2019).
An increase in manganese concentration in seeds under drought stress may be attributed to the reduction in biomass caused by water scarcity, leading to a higher accumulation of this element in seeds. Our study suggests that under well-irrigated conditions, an increase in iron absorption could potentially lead to a decrease in manganese concentration and uptake by plants due to their antagonistic relationship (Yanga et al. 2015). Drought conditions can limit root access to cations, resulting in reduced copper concentration in seeds. Furthermore, decreased soil moisture levels can restrict plant access to low-demand nutrients such as iron, zinc, copper, and manganese. Consequently, increased drought stress can reduce copper absorption, leading to lower concentrations in leaves and seeds (Arndt et al. 2021). Additionally, research findings indicate that the use of vermicompost not only enhances soil's absorbable copper levels but also improves soil fertility and plant nutrition by promoting nutrient cycling (Li et al. 2017).
The oil percentage in seeds is a quantitative trait controlled by multiple genes, and the decrease in oil percentage under drought stress can be attributed to the damage of numerous genes regulating seed oil content (Sincik et al. 2013). The findings of our study align with previous research in this area, indicating that water stress leads to a reduction in oil percentage due to the impact of stress on the seed's ability to accumulate oil. Drought stress negatively affects seed oil percentage by hampering the seed's capacity to absorb assimilates and convert them into oil (Petcu and Stanciu 2021). Adequate water availability during plant growth appears to promote an increase in seed weight and oil storage. Conversely, the decrease in seed oil content under drought stress may result from a shortened seed filling period and reduced seed weight. Furthermore, the limited availability of carbohydrates for oil synthesis due to drought stress contributes to the decrease in oil percentage (Si et al. 2003).
The oil percentage in seeds is a heritable trait influenced by genetic factors, with the production of oil being derived from compounds generated during photosynthesis. Enhancing plant growth conditions can potentially elevate the oil percentage in seeds (Singer et al. 2007). Utilizing vermicompost fertilizer can create more favorable conditions for root and organ growth, leading to improved water and nutrient uptake. Additionally, vermicompost can help balance soil acidity and enhance nutrient absorption, ultimately resulting in an increase in seed oil percentage (Wang et al. 2022).