Legume Research

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Legume Research, volume 44 issue 11 (november 2021) : 1362-1370

Seed Priming Influences the Seed Quality and Activities of Nitrate Assimilation and Anti-Oxidant Enzymes in Pigeon Pea Seedlings

T.N. Tiwari1,*, D.K. Agarwal2
1ICAR-Indian Institute of Pulses Research, Kanpur-208 024, Uttar Pradesh, India.
2ICAR-Indian Institute of Seed Science, Mau-275 103, Uttar Pradesh, India.
  • Submitted16-09-2019|

  • Accepted18-01-2020|

  • First Online 15-05-2020|

  • doi 10.18805/LR-4236

Cite article:- Tiwari T.N., Agarwal D.K. (2021). Seed Priming Influences the Seed Quality and Activities of Nitrate Assimilation and Anti-Oxidant Enzymes in Pigeon Pea Seedlings . Legume Research. 44(11): 1362-1370. doi: 10.18805/LR-4236.
Field experiments were conducted with four levels of seed priming including control and two varieties of pigeon pea at ICAR-Indian Institute of Seed Science, Mau during three consecutive years (2011-12 to 2013-14). One-year-old seeds of pigeon pea varieties (Bahar and Malviya-13) were primed with growth regulator (100 ppm GA3), in-organic salt (0.2% KNO3) and tap water (sanitized) separately for 06 hrs. and sown in field under RBD with 03 replications. Observations were recorded on seed quality parameters, biochemical attributes including nitrate assimilatory enzymes and activities of anti-oxidant enzymes during seedling stage. Seed quality parameters including germination, seedling growth and vigor indices were significantly enhanced through seed priming with GA3 followed by KNO3 and tap water over unprimed control. Biochemical attributes viz; chlorophyll a andb contents, were more influenced with GA3 priming followed by KNO3 and tap water whereas the proline accumulation was reduced with priming treatments and maximum reduction was noted with GA3 followed by KNO3 and tap water. Enhancement in nitrate assimilatory enzymes including nitrate and nitrite reductase activities was more with KNO3 priming followed by GA3 and tap water. Anti-oxidant enzymes activities including Catalase, Peroxidase and Super Oxide Dismutase were also increased significantly by KNO3 priming followed by GA3 and tap water over unprimed control.
Pigeon pea [Cajanus cajan (L.) Millsp.] is one of the most important pulse crops cultivated globally on 6.23 million hectare with total production and productivity of 4.67 million tones and 813 kg/ha, respectively. In India area occupied by pigeon pea is about 4.65 million ha with total production of 3.02 million tons and productivity being 7.00 q/ha (FAO,2013) which is quite low because of several factors including its cultivation in rain fed and marginal lands, use of old and low quality seed by the farmers and several abiotic and biotic stresses affecting during different phases in the entire crop duration. This in turns gives low germination, delayed emergence and sick seedlings that lead to poor yield. Seed priming is a powerful tool in which seed are soaked in water or an osmotic solution allows water imbibition’s and permits early stages of germination but does not permit radical protrusion through seed coat (Heydecker, 1973). Priming with different in-organic salts including KNO3 and MgNO3 and MgSO4 etc. has been reported to improve the germination, speed of  emergence, seedling vigor, growth and yield of different vegetable and field crops viz:, beet root, brinjal, cabbage, cauliflower, carrot, summer squash, tomato, water melon, hot pepper, tobacco, maize, mustard, okra ‘ground nut,  moong bean and pigeon pea by several workers (Durran et al.,1983, Saxena and Singh 1987, Fujikera and Karseen 1992, Broklehurst and Dearman1983, Mauromicale et al., 1994, Min Taigi 2001, Kundu and Basu1981, Pandita et al., 2003. Misra and Sahoo 2003, Thakur and Thakur 2006. Dhedhi et al., 2006 and Tiwari et al., 2013, 2014 and 2018). Considerable evidences exist that repair of proteins and enzymes occurs during imbibition of seeds (McDonalds, 2000). It is known that priming increases the activity of enzymes that counteracts the effect of lipid peroxidation and as a result the free radical scavenging enzymes are increased by increasing the hydration in seeds (Sung and Jeng, 1994). Priming also enhanced the membrane repair in seeds and could be ascribed to evoked activities of several lipid peroxide scavenging enzymes (Chiu et al., 1995). It is also possible that cell damage may happen with increased seed drying (Drew et al., 1997). Maximum beneficial effects of priming can be achieved during the drying phase when enzymes and anti-oxidants are afforded sufficient time to affect repair and physiologically stabilize the seed (Aquila and Tritto, 1991). The information’s on  response of seed priming with KNO3 and GA3 on enhancement of seed quality parameters, activities of nitrate assimilation and anti-oxidant enzymes and some of the biochemical attributes in pigeon pea seedlings are very limited in pigeon pea crop. Hence, keeping the above facts into consideration the present experiment was under taken with the objectives to understand the effect of seed priming with KNO3 and GA3 on germination, vigor and stand establishment of pigeon pea seedlings and its impacts on nitrate assimilation and anti-oxidant enzymes activities.
Field experiments were conducted during three consecutive kharif season of 2011-12 to 2013-14 at the research farm of Indian Institute of Seed Science, Kushmaur, Mau (U.P.). One-year-old farmer saved seeds of pigeon pea varieties, Bahar and Malviya-13 were collected from Mau district of U.P. The collected seeds were initially surface sterilized with 0.2% HgCl2 and then primed with tap water (sanitized), 100 ppm GA3 and 0.2% KNO3 separately for 06 hours and the treatments were formulated as Control means no priming (T0), Seed priming with tap water(sanitized) (T1), Seed priming with 0.2% KNO3(T2) Seed priming with100 ppm GA3(T3).
Laboratory experiments (Seedling studies)
Total 1200 seeds were taken for each variety of pigeon pea for germination using top of the paper method as described by ISTA procedures (Anonymous, 1999). After sowing, the germination count was started just after one day and counted up to seventh day and the germination was recorded by counting total number of seeds germinated in each treatment and percent germination was expressed on normal seedling basis. 
Root length was measured from the collar region to the tip of the primary root and mean was calculated and expressed in centimeters similarly shoot length was measured from collar region to the base of uppermost leaf and mean was calculated and expressed in centimeters. The seedlings used for seedling length measurement were used for estimating dry weight. Seedlings were dried in a hot air oven maintained at 80±2°C for 24 hours) and after drying, the weight of 10 dry seedlings was recorded and the mean seedling dry weight was calculated and expressed in milligrams. Vigor of the seeds was assessed based on germination percentage, seedling length and seedling dry weight as suggested by Abdul-Baki and Anderson (1973) and expressed in whole number.
Seedling vigour index I = Germination (%) × Mean seedling length (cm)
Seedling vigour index II = Germination (%) × Mean seedling dry weight (mg)
Observations on biochemical constituents including chlorophyll a, b content and proline content, nitrate assimilatory enzymes like nitrate reductase and nitrite reductase activities and anti-oxidant enzymes including catalase, peroxidase and super oxide dismutase were estimated as per standard procedures mentioned below. Three years’ data were pooled and statistically analyzed using SAS software.
Biochemical studies
The content of chlorophyll was estimated using the method of Arnon (1949) in green leaf tissues and Proline content was estimated in fresh leaves following the procedures mentioned by Bates et al.,1973. Nitrate assimilatory enzymes including nitrate reductase activity and nitrite reductase activity were assayed following the methods of Jaworski (1971) and Ferari and Varner (1971) respectively.
Catalase assay was based on the absorbance of H2O2 at 240 nm in UV-range. A decrease in the absorbance was recorded over a time period as described by Aebi (1984). Peroxidase activity is assayed as increase in optical density due to the oxidation of guaiacol to tetra-guaiacol (Castillo et al., 1984). Superoxide dismutase (SOD) assay was based on the formation of blue coloured formazone by nitro-blue tetrazolium and O2.- radical, which absorbs at 560 nm and the enzyme (SOD) decreases this absorbance due to reduction in the formation of O2.- radical by the enzyme (Dhindsa et al., 1981).
Seed quality parameters
Seed priming of one-year-old pigeon pea seeds with growth regulator (100 ppm GA3), in-organic salt (0.2% KNO3) and tap water separately, significantly enhanced the germination over un-primed control (Table 1a,b,c,d,e,f). Among treatments, seed priming with GA3 displayed maximum seed germination (90.17%) which was at par with KNO3 (85.83%) followed by tap water (80.83%) and differed significantly with each other. The varieties of pigeon pea evaluated responded well to priming treatment and germination per cent was significantly higher in Bahar (90.16%) followed by Malviya -13 (77.50%). Interaction between varieties and treatments were non-significant. Maximum germination (95.00%) was recorded in variety Bahar with GA3. The magnitudes of improvement in germination were 2.97, 9.34 and 14.67% with tap water, KNO3 and GA3 respectively over control. Similarly, other seed quality parameters including root length, shoot length, Seedling dry weight, Vigour index I and Vigour-II also showed the significant improvement through seed priming treatment in both the varieties evaluated. Among the priming agents, GA3@100ppm performed better than KNO3(0.2%) and tap water in respect of all the seed quality parameters studied. The percent improvement over control in root and shoot length, seedling dry weight, vigour IandII was recorded higher in GA3 priming followed by KNO3 and tap water. Variety Bahar displayed significantly higher values of all the seed quality parameters over Malviya-13 (Table 1a to f).

Table 1: Effect of seed priming on seed quality parameters in pigeon pea seedlings.

Our results were also confirmed by the findings of Bose and Mishra, (1992) and Bose, 1997 who reported that osmo- priming with different chemicals to seeds enhances the rate of germination and encourages the fast emergence of seedling in field and this might lead in enhancement in subsequent phases of plant growth and finally to better performance of a crop. Results obtained were also supported with the findings of Bose and Mishra (1999 and 2001) who opened that during soaking of seed in Mg (NO3)2 or KNO3 solution the cations Mg++ or K+ and anions NO3- in fluxed in the seeds and showed their carry over effects during vegetative growth period and consequently the plant/crop stand was improved and it was further supported by Bose and Pandey (2003) and Tiwari et al., (2018).
Biochemical attributes
Seed priming with GA3@100 ppm, KNO3(0.2%) and tap water significantly improved the chlorophyll a and b contents in the leaves of pigeon pea varieties over unprimed control, whereas the proline accumulation in leaves was decreased with the priming treatments. The reduction in proline content over unprimed control was maximum with GA3 followed by KNO3 and tap water priming might be due to socking of water which minimizes the level of water stress in primed seed as compare to unprimed seed (Table 2 a,b and c).  

Table 2: Effect of seed priming on Chlorophyll and Proline contents in pigeon pea seedlings.

Nitrate assimilatory enzymes were more influenced by KNO3 priming as compared to GA3 and tap water over un primed control. This indicates the role of priming in the enhancement of activities of nitrate assimilatory enzymes and particularly of KNO3 which showed the highest activity of both the enzymes might be due to No3 salt which is the source of nitrogen and act as substrate for enzyme activity (Table 3 a, b). Nitrate is considered the primary source of nitrogen from the soil and the main function of enzyme nitrate reductase (NR) is to reduce nitrate to nitrite (Beevers and Hageman, 1969). Positive effect of auxin hormone and Ca (NO3)2 on NRA was due to its possible role in the activation of the inactive nitrate reductase protein and prevention of enzyme degradation by proteolysis. This might also be involved in the enhancement of enzyme synthesis or its maintenance in the active form and thus, has a protective role on nitrate reductase activity (Richard and Stanely, 1981). The stimulated NR activity in seed priming treatments compared to control plants might be due to enhanced nitrogen uptake by plants Muthuchelian et al., (1994). Inclusion of nitrate and in-organic ions like Cu2+, K+, Mn2+, Mg2+ and Zn2+ in assay medium has also been reported to increase the nitrate reductase activity in roots and leaves of maize under salinity (100 mM NaCl) (Khan and Srivastava, 2000). Beneficial effects of priming have been associated with various biochemical, cellular and molecular events including synthesis of DNA and proteins (Srivastava et al., 2010). Metabolic repair processes and build-up of germination metabolites or osmotic adjustments during priming are due of germination enhancement techniques (Mukhtar et al., 2013). These are the possible reasons for the observed enhancement in seedlings growth, improvement in composition of chlorophyll and enhanced activity of nitrate assimilatory enzymes. Seed priming with beta amino butyric acid (BABA) in rice seeds in varied abiotic stresses (NaCl/PEG-6000) leads to increased seedling growth, photosynthetic pigment chlorophyll, photosystem activities and mitochondrial activities in rice seedlings. Moreover, BABA priming significantly reduced malondialdehyde content in the seedlings and also resulted in accumulation of proline in NaCl tolerant variety. It also enhanced the activities of nitrate reductase and antioxidant enzymes and thus improved the drought and salinity stress in rice varieties (Jisha and Puthur, 2016).

Table 3: Effect of seed priming on nitrate assimilation and anti-oxidant enzymes activities in pigeon pea seedlings.

Anti- Oxident enzymes
The anti-oxidant enzymes including catalase, peroxidase and super oxide dismutase were assayed using the respective standard methods mentioned in the text in growing seedlings. The observations recorded indicated that seed priming with GA3@100 ppm, KNo3 (0.2%) and tap water significantly enhanced the enzymes activity of catalase, peroxidase and super oxide dismutase over unprimed control. Among the priming agents used, KNO3 enhanced relatively more activity of catalase, peroxidase and super oxide dismutase followed by GA3@100ppm and tap water (Table 4 a, b and c). In Varieties, Bahar displayed significantly higher values of enzymes activities including catalase, peroxidase and super oxide dismutase. Enhancement in anti-oxidant enzymes activity through osmo- priming with PEG socked for 24 h in okra seeds has been reported by Chawla et al., (2014) and priming of rice seeds with distilled water, Mg (NO3) 2 salt, Kinetin and salicylic acid were also found to enhance the activity of SOD, NR, Ascorbate peroxidase, proline and protein content over non primed control under timely and late sown conditions (Kumar et al., 2018). These findings confirm our results in case of pigeon pea where the priming agents used are also of same nature. Further, Priming has been used to improve the performance of germination at the field and potassium nitrate (KNO3) is a promising compound for the purpose. The nitrate (NO3) could be absorbed, being used in the metabolism of the embryo, through the enzyme nitrate reductase (NR). Besides, the priming could also activate the response of the antioxidant system, becoming the primed seeds more prepared for possible stresses. The results obtained showed an increase in the NR activity, as well as in the antioxidant enzymes and seems to be the beneficial effects of priming with KNO3 in pigeon pea seeds (Lara et al., 2014). GA3 priming of alfalfa ‘Bami’ seeds resulted in increased germination percentage and seedlings growth, decreased lipid peroxidation and enhanced antioxidant defences and may be an efficient method to overcome seed germination problems and to improve seedlings growth in the field (Younesi and Moradi, 2014). Osmo-priming strengthens the antioxidant system and increases seed germination potential, resulting in an increased stress tolerance in germinating seeds. Osmo-priming-mediated promotive effect on stress tolerance, however, may diminish in relatively older (e.g. ~5-week) seedlings (Chen and Arora, 2011). In contrary to above findings, Bhattacharya et al., (2019) reported the deteriorative changes in enzyme activity of dehydrogenase, Amylase. Catalase and peroxidase in invigorated soybean seeds.

Table 4: Effect of seed priming on anti-oxidant enzymes activities in pigeon pea seedlings.

Seed priming with beta amino butyric acid(BABA) in rice seeds in varied abiotic stresses (NaCl/PEG-6000) also enhanced the activities of nitrate reductase and antioxidant enzymes and thus improved the drought and salinity stress in rice varieties (Jisha and Puthur, 2016). Priming potential of AGE (aqueous garlic extracts), SA and MeJA to enhance seed germination and early seedling growth in eggplant and the effects were obvious in various morphological and physiological traits. Seed priming significantly altered the antioxidant enzymes activities such as superoxide dismutase (SOD) and peroxidase (POD) with alteration in the reactive oxygen species (ROS). Interestingly, priming duration also affected the bioactivity of these chemicals because seed priming with 300 µg mL-1 AGE for 4 h had a positive influence, however, prolonged exposure to the same concentration inhibited the seed germination process and induced oxidative stress on the seedlings with elevated levels of malondialdehyde (MDA) content. AGE (aqueous garlic extracts) seed priming as a bio-stimulant to enhance seed germination and early seedling growth in eggplant and the results hence lay the foundation for the preparation of garlic-based compounds to improve vegetables production under plastic tunnels and greenhouse production units (Ali et al., 2019).
In the light of above findings, it may be concluded that the observed improvement in present study with respect to germination, seedling growth, biochemical attributes and enzymatic activities are the results of seed priming with growth regulator GA3 and osmo-priming with KNO3 and might be considered as an effective and cheapest tool for the improvement of pigeon pea germination and crop stand establishment.

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