Indian Journal of Agricultural Research

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Phytopriming of Vigna mungo (L.) Hepper (Blackgram) Seeds using Aqueous Extracts of Moringa oleifera Lam, Gliricidia sepium (Jacq.) Walp and Pongamia pinnata (L.) Pierre Improves Seedling Emergence, Growth and Yield

Ravindran Krishnasamy1,2, Jagan M. Obbineni2,*
  • 0000-0001-7234-8549, 0000-0001-7716-2778
1School of Biosciences and Technology (SBST), Vellore Institute of Technology, Vellore-632 014, Tamil Nadu, India.
2School of Agricultural Innovations and Advanced Learning (VAIAL), Vellore Institute of Technology, Vellore-632 014, Tamil Nadu, India.

ABSTRACT

Background: Phytopriming can enhance seed germination and seedling emergence. Seeds when primed with plant leaf extracts result in better establishment and growth of field crops.

Methods: This study examines the potential of leaf extracts of Moringa oleifera Lam, Gliricidia sepium (Jacq.) Walp and Pongamia pinnata (L.) Pierre on the growth and yield of black gram through seed hydropriming.

Result: Pot studies revealed a 10% increase in shoot length and 3% increase in fresh weight upon priming with P. pinnata (L.) Pierre leaf extract in comparison to liquid fertilizer treatment. Also, 6% increase in root nodule formation was observed in G. sepium (Jacq.) Walp and P. pinnata (L.) Pierre treated plants. Testing the biochemical parameters revealed a 20% increase in total phenolics, 12% increase in total carbohydrates and 64% increase in chlorophyll pigment in the case of P. pinnata (L.) Pierre leaf extract treated group. The data presented shows that a 4% M. oleifera Lam leaf extract, 10% G. sepium (Jacq.) Walp, or 4% P. pinnata (L.) Pierre leaf extract can be used for improving seedling emergence, plant growth and yield. Among the three, the best overall growth and yield can be obtained by using a 4% P. pinnata (L.) Pierre leaf extract as a seed priming agent.

INTRODUCTION

In recent decades, seed priming has been demonstrated to effectively improve seed germination and support seedling growth. Various methods of seed priming such as biopriming, halopriming, hormonal priming, hydropriming and osmopriming have been used to improve germination (Lutts et al., 2016). Among these techniques, hydropriming is the most commonly followed and effective method (Pandita et al., 2007). Hydropriming involves soaking seeds in an aqueous solvent for 24 hours or overnight prior to sowing in the field. This process facilitates the rapid absorption of water and oxygen by the seeds, triggering a range of biochemical activities. Utilization of storage compounds namely carbohydrates, lipids, proteins and the production of ROS, the activation of enzymes like POX, APX and GH, are among these activities (Hussain et al. 2018). Crucial to successful seed germination, antioxidant enzymes can act as molecular markers for seed vigor (Pagano et al., 2019). A study conducted on V. mungo (L.) Hepper (black gram) demonstrated that the application of Spirulina platensis extract positively influenced the germination of seedlings and their subsequent development (Quang Thinh and Sundareswaran 2018). Ocimum sanctum leaf extracts increased the number of nodules in V. mungo (L.) Hepper. The presence of nodules indicates the plant’s ability to form a symbiotic relationship with rhizobia, which enhances nitrogen fixation and promotes plant growth (Dugesar et al., 2017).  
            
M. oleifera Lam is a widely used plant source, with its leaves and slender fruit also known as drumstick commonly used in food across South Asia. The diverse culinary and medicinal applications of Moringa reflect its significance as a valuable plant resource (Baldisserotto et al., 2018). Its leaf extract, when applied to wheat plants, has shown positive effects on nutrient availability, photosynthesis, root development and overall plant growth and yield (Khan et al., 2017).
       
G. sepium (Jacq.) Walp is also a fast-growing tree species and has the ability to fix nitrogen in the soil (Kaba et al., 2019). It has been mostly used as fodder and for firewood, live fencing, intercropping and shade crops in coffee plantations (Alamu et al., 2023; Beedy et al., 2010). P. pinnata (L.) Pierre is a nitrogen-fixing tropical tree that thrives in nitrogen-deficient and contaminated soils, demonstrating its ability to enrich the soil with essential nutrients (Unnikannan et al., 2013). It has diverse applications in agriculture and biology, with its residual seed oil cake utilized as a soil amendment to improve soil quality and nutrient content and its leaves used as green leaf manure and for pest control (Jathanna et al., 2020).
       
V. mungo (L.) Hepper, known as black gram is a nitrogen-fixing tropical pulse crop, has suffered declining yields due to seed-borne infections and poor seed quality (Prakash et al. 2020). One potential solution to this issue involves utilizing plant extracts as hydropriming agents to enhance germination, seedling vigor and growth. To overcome the poor seed quality and declining yields in black gram, the study conducted to examine the use of Pongamia pinnata leaf extract (PLE), Gliricidia sepium leaf extract (GLE) and Moringa oleifera leaf extract (MLE) as priming agents to improve the seed viability as well as plant growth.

MATERIALS AND METHODS

The experiment was conducted from October 2022 to December 2022 at the research cum instructional farm of Vellore Institute of Technology, Vellore, Tamil Nadu, India.
 
Materials
 
Leaves of M. oleifera Lam, G. sepium (Jacq.) Walp and P. pinnata (L.) Pierre were collected from various locations in Vellore, India. All the reagents and chemicals were obtained from HiMedia Laboratories Pvt. Ltd and Sisco Research Laboratories (India).
 
Extraction process
 
The leaves were washed, dried and ground before being extracted with distilled water. The leaf samples were combined with distilled water (1:10 w/v). The solution was then placed in a water bath for 20 mins with a set temperature of 60°C (Krishnaraj et al., 2012). The solution was allowed to cool down to room temperature and filtered. The filtered extract was stored in an airtight container at 4oC. For priming, different concentrations of the extract in distilled water were prepared by dilution (2%, 4%, 6%, 8% and 10%) (Radwan et al., 2019).
 
Seed priming test
 
Surface sterilization of V. mungo (L.) Hepper seeds was done by gentle agitation in 4% sodium hypochlorite solution for 10 mins. Then, the seeds were soaked for 3 hrs in the aforementioned concentrations of the different leaf extracts. A control condition was also setup by soaking seeds in distilled water. To carry out the seed germination test, a sterile Petri dish was taken and two sterile filter papers were placed on it. Distilled water (~10-15 mL) was added to the dish to keep the filter papers moist. Twelve seeds were then placed per plate and kept at room temperature. After 10 days, seedling growth parameters were measured. The experiment was conducted in triplicate and repeated twice to ensure reliable results and account for any experimental variations (Ismail et al., 2016).
 
Pot studies
 
The pot studies were conducted under net house in Vellore, India. Polyvinyl chloride pots (height - 25 cms and outer diameter - 30 cms) were used. A total of 15 pots were filled with 10 kg of dehydrated and sieved red soil (pH 7.1 and EC 237 mSm-1) and split into 5 groups with 3 pots in each group. Pots in each group were labeled as group A to E. The labels referred to the different treatments given viz., group A for distilled water only, group B for liquid fertilizer (N: P: K 19:19:19), group C for undiluted MLE, group D for undiluted GLE and group E for undiluted PLE.
       
Surface sterilized V. mungo (L.) Hepper (blackgram) seeds were soaked for 1 hr in the respective solutions. Eight seeds that were soaked in the respective solutions were sown in each pot. Thinning was done after ten days and only two plants (showing uniform growth) were retained per pot. 1 lit of tap water was applied to each pot three times a week to maintain the soil moisture. 20 mL of the respective solution i.e., distilled water for group A, 4% liquid fertilizer (N: P: K 19:19:19) for group B, undiluted MLE for group C, undiluted GLE for group D and undiluted PLE for group E was given per plant by foliar application on the 15th, 30th and 45th day after sowing (Karthik et al., 2020).
 
Measurement of physiological parameters
 
On the 70th day after sowing, two plants from each group were harvested, washed and analyzed for root nodules, root and shoot length, fresh and dry weight, grain weight and number of pods per plant.
 
Measurement of biochemical parameters
 
On the 47th day leaf samples were collected and stored at -80oC. For the measurement of biochemical parameters samples were prepared for each assay as per previously reported protocol (Karthik et al., 2020). The total carbohydrate in the sample was estimated using phenol-sulphuric acid method (Chow and Landhäusser 2004). Using Lowry’s method, total protein was estimated (Lowry et al., 1951). For the estimation of total amino acid, ninhydrin method was used (Yemm et al., 1955). Folin Ciocalteau reagent was used to estimate the total phenol (Rekha et al., 2012). Photosynthetic pigments were estimated using a previously published method (Sarkar et al., 2018).
 

RESULTS AND DISCUSSION

Effect of hydropriming using leaf extracts on seedling growth
 
Leaf extracts at different concentrations were used for hydropriming the Vigna mungo (L.) Hepper (blackgram) seeds. Seeds hydroprimed with 4% MLE displayed the longest root length of 7.91±0.23 cm and the longest shoot length of 16.47±0.36 cm was observed in the 10% extract group (Fig 1A). Seeds hydroprimed with 10% GLE exhibited the longest root length of 7.99±0.25 cm, whereas seeds primed with a 6% extract had the longest shoot length of 13.74±0.24 cm (Fig 1B). Seeds hydroprimed with 2% PLE showed the longest root length (7.26±0.38 cm) whilst the longest shoot length was observed in the case of 4% extract (14.03±0.31 cm) (Fig 1C).
       
From Fig 1 it is evident that hydropriming with MLE, GLE and PLE has led to better seedling growth and it can be concluded that 4% MLE, 10% GLE and 4% PLE are effective in promoting seedling growth.

Fig 1: Effect of hydropriming using varying concentrations of A. MLE B. GLE and C. PLE on growth of V. mungo (L.) Hepper (blackgram) seedlings as estimated by measuring the seedling height.


 
Effect of hydropriming using leaf extracts on plant growth and yield
 
Pot experiments were conducted to examine the effect of the different leaf extracts on the growth characteristics and yield of the blackgram. Experiments showed that plants treated with commercial fertilizer (19:19:19) showed the highest root length (39.67±1.05 cm) (Fig 2A). On the other hand, PLE treated plants showed the highest shoot length (42.35±1.53 cm) (Fig 2B). 

Fig 2: Effect of hydropriming using liquid fertilizer (N: P: K 19:19:19), MLE, GLE and PLE on A. Root Length and B. Shoot Length of V. mungo (L.) Hepper (blackgram) plants.


       
Plant biomass analysis revealed that the fresh weight of PLE treated group was the highest (50.09±0.71 g) and that of the untreated plant was lowest (30.31±1.33 g) (Fig 3A). The dry weight of plants treated with liquid fertilizer was highest (23.14±0.89 g) and that in the untreated group was lowest (10.50±0.31 g) (Fig 3B). The morphological differences between the plants of different treatment groups, viz., control, liquid fertilizer, MLE, GLE and PLE can be seen in Fig 4. 

Fig 3: Increase in A. fresh weight B. dry weight of V. mungo (L.) Hepper (blackgram) plants upon hydropriming using liquid fertilizer (N: P: K 19:19:19), MLE, GLE and PLE.



Fig 4: Morphological differences in V. mungo (L.) Hepper (blackgram) plants hydroprimed using A. water, B. liquid fertilizer (N: P: K 19:19:19), C. MLE, D. GLE and E. PLE.


       
Other growth characteristics such as root nodules per plant and yield characteristics such as pods per plant as well as grain weight were measured. Treatment with PLE resulted in the highest number of well-developed root nodules per plant followed by GLE treated group, liquid fertilizer treated group and MLE treated group (Fig 5A). The grain weight and number of pods per plant were highest in liquid fertilizer treated group. Plants treated with PLE also showed a significantly high grain weight and number of pods per plant (Fig 5B and Fig 5C).

Fig 5: Growth and yield characteristics of V. mungo (L.) Hepper (blackgram) plants on hydropriming using liquid fertilizer (N: P: K 19:19:19), MLE, GLE and PLE. A. Root nodules per plant. B. Pods per plant. C. Grain weight.


   
Assessment of plant metabolite levels upon hydropriming
 
The total protein, carbohydrate, phenol, chlorophyll and carotenoid were measured using biochemical assays. Total protein concentration was found to be highest in MLE treated group (0.536±0.0153 mg/ml) (Fig 6A). Carbohydrate content was highest in the plants treated with PLE (0.5915±0.043 mg/ml) (Fig 6B). Phenol content was highest in the plants treated with PLE (0.3668±0.021 mg/ml) (Fig 6C).

Fig 6: Concentration of A. Protein B. Carbohydrate and C. Phenol in Vigna mungo (L.) Hepper (blackgram) plants on hydropriming using liquid fertilizer (N: P: K 19:19:19), MLE, GLE and PLE.



The photosynthetic pigment chlorophyll a was highest in the plants treated with PLE (0.6536±0.018 mg/ml) (Fig 7A). Chlorophyll b content was highest in plants treated with liquid fertilizer (0.4923±0.051 mg/ml) (Fig 7B). Carotenoid content in the liquid fertilizer group was highest at 0.0537±0.003 mg/ml (Fig 7C).

Fig 7: Estimation of photosynthetic pigments A. chlorophyll a B. chlorophyll b and C. carotenoid in V. mungo (L.) Hepper (blackgram) plants on hydropriming using liquid fertilizer (N: P: K 19:19:19), MLE, GLE and PLE.


       
Seedling and plant growth can be enhanced by using several plant leaf extracts and organic compounds as priming agents (Duary et al. 2022). Poor seed quality and lower germination rates can be overcome by hydropriming with plant leaf extracts. In the current study, we have assessed the effect of hydropriming using aqueous leaf extracts of M. oleifera Lam, G. sepium (Jacq.) Walp and P. pinnata (L.) Pierre on the growth and yield of V. mungo (L.) Hepper (blackgram). Biomass analysis revealed significant growth enhancement in the case of seeds hydroprimed with different plant extracts. Seeds hydroprimed with 4% MLE showed a 21% increase, 10% GLE showed a 20% and 4% PLE showed the highest increase of 27% in seedling height. In pot trials, growth parameters such as dry weight and fresh weight as well as root and shoot length showed a significant increase. Notably, GLE treated plants exhibited a 3% increase in shoot length and PLE treated plants exhibited a 10% increase in shoot length as well as 3% increase in fresh weight in comparison to liquid fertilizer-treated group. The improved emergence of the seedlings as well as increase in plant growth can be attributed to the several plant nutrients, i.e., N, P, K and Ca present in MLE, GLE and PLE (Alamu et al., 2023; Foidl et al., 2001; Islam et al., 2021). The stimulatory effect may also be due to the presence of several bioactive compounds in the extracts (ElSayed et al., 2022). Another notable observation is a 6% increase in root nodules per plant in GLE and PLE treated plants. This may be attributed to flavones which are vital in mediating the molecular dialogue between rhizobia and legume plants. This symbiosis allows legume plants to access a significant source of nitrogen through biological nitrogen fixation, contributing to their growth, development and ecological adaptation (Zhang et al., 2009). GLE and PLE have been reported to contain many flavones thus playing a role in a multitude of applications (Jurd and Manners,1977; Marzouk et al., 2007). Biochemical attributes such as total protein, phenol and carbohydrates also showed a significant increase upon hydropriming. Treatment with MLE showed a 25% increase in total protein and treatment with PLE resulted in a 12% increase in total carbohydrates as well as 20% increase in total phenol. MLE, GLE and PLE treatments resulted in a significant increase in chlorophyll a by 30%, 49% and 64% respectively. An increase in chlorophyll pigments would facilitate an improvement in the water use efficiency as well as gas exchange attributes. There is also an increase in photosynthetic and respiration activities. Previous studies have reported that seeds pelleted with pungam leaf powder showed increased chlorophyll (Prakash et al. 2021) and exogenous application of MLE, improved chlorophyll pigments (Rashid et al. 2021).
       
Overall, the PLE treated group exhibited higher shoot length, fresh weight and other physiological parameters. The extract is known to have an active compound myo-inositol which acts as a signaling molecule involved in various physiological processes (Sharma et al., 2020). GLE and MLE have an active compound called quercetin which plays a pivotal role in seed germination, photosynthesis and pollen development. Another flavonoid called rutin is also present in these extracts. It regulates the biosynthesis of growth hormones such as cytokinin, auxin and gibberellins (Gorni et al., 2022). MLE is also rich in vitamins, alkaloids and phytohormones which regulate the plant growth (Karthiga et al., 2022). Hydropriming of seeds with plant leaf extracts holds great promise and can be particularly beneficial for farming systems using low external agricultural inputs and to a large extent depending on on-farm resources (Carrillo-Reche  et al. 2018). MLE at 4%, GLE at 10% and PLE at 4% can be used by the farmer as a priming agent for V. mungo (L.) Hepper (blackgram) seeds to enhance growth and productivity with the best results obtained using PLE.

CONCLUSION

Vigna mungo (L.) Hepper (blackgram) seeds hydroprimed with 4% M. oleifera Lam extract, 10% G. sepium (Jacq.) Walp extract and 4% P. pinnata (L.) Pierre leaf extract showed significant improvement in seedling growth and development. Subsequent pot trials revealed that all tested leaf extracts exhibited notable benefits in terms of yield attributes and plant growth parameters. Among the different extracts, P. pinnata (L.) Pierre leaf extract displayed the most pronounced effects, resulting in superior plant growth and increased grain weight. Based on these findings, it can be concluded that the application of these leaf extracts as an alternative to commercial fertilizers holds promise for sustainable agricultural practices. By harnessing the natural growth-promoting properties of these leaf extracts, farmers can potentially reduce their reliance on synthetic chemical fertilizers, mitigating their negative environmental impact. Further investigations are warranted to gain an in-depth understanding of the underlying mechanisms behind the beneficial effects of these leaf extracts. Field trials encompassing a broader range of crop species and environmental conditions would provide valuable insights.

ACKNOWLEDGEMENT

This work was funded by VIT under the rGEMS grant and the Science and Engineering Research Board (SERB), Govt. of India, vide grants TAR/2022/000312 and SRG/2020/001690.
 
Disclaimers
 
The views and conclusions expressed in this article are solely those of the authors and do not necessarily represent the views of their affiliated institutions. The authors are responsible for the accuracy and completeness of the information provided, but do not accept any liability for any direct or indirect losses resulting from the use of this content.

CONFLICT OF INTEREST

The funders had no role in the design of the study; in the collection, analyses, or interpretation of data; in the writing of the manuscript; or in the decision to publish the results. The authors declare no conflict of interest.

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