rus
Issue #5/2020
V. N. Zelenkov, V. V. Latushkin, M. I. Ivanova, A. A. Lapin, V. V. Karpachev, A. A. Kosobryukhov, P. A. Vernik, S. V. Gavrilov
Effect of Pulsed Illumination on the Germination of Seeds of Some Vegetable, Oil-Bearing and Medicinal Plants
Effect of Pulsed Illumination on the Germination of Seeds of Some Vegetable, Oil-Bearing and Medicinal Plants
DOI: 10.22184/1993-7296.FRos.2020.14.5.442.460
The results of a comprehensive study of the effect of pulsed light irradiation on seed germination, plant growth and green mass yield are presented. The objects of research included 8 vegetable, medicinal and oil-bearing plants. A hypothesis was put forward that it is necessary to develop differentiated modes of light irradiation of plants (repetition period and pulse duration). It has been found that the effect of pulsed irradiation largely depends on the genetic characteristics of the objects. The most favorable and worst irradiation regimes were found in comparison with dark germination for plant productivity. Chlorophyll fluorescence parameters differ when exposed to pulsed light in the second and millisecond ranges. Analysis of the total antioxidant activity (TAOA) of green mass (microgreens) also showed significant differences depending on the genetic nature of the plant.
The results of a comprehensive study of the effect of pulsed light irradiation on seed germination, plant growth and green mass yield are presented. The objects of research included 8 vegetable, medicinal and oil-bearing plants. A hypothesis was put forward that it is necessary to develop differentiated modes of light irradiation of plants (repetition period and pulse duration). It has been found that the effect of pulsed irradiation largely depends on the genetic characteristics of the objects. The most favorable and worst irradiation regimes were found in comparison with dark germination for plant productivity. Chlorophyll fluorescence parameters differ when exposed to pulsed light in the second and millisecond ranges. Analysis of the total antioxidant activity (TAOA) of green mass (microgreens) also showed significant differences depending on the genetic nature of the plant.
Теги: chlorophyll fluorescence green photonics growth and development leds medicinal plants oilseeds pulsed light seeds vegetables импульсный свет лекарственные растения масличные культуры овощные культуры рост и развитие светодиоды семена фитофотоника флуоресценция хлорофилл
Effect of Pulsed Illumination on the Germination of Seeds of Some Vegetable, Oil-Bearing and Medicinal Plants
V. N. Zelenkov 1, 2, 3, V. V. Latushkin 3, M. I. Ivanova 2, A. A. Lapin 4, V. V. Karpachev 5, A. A. Kosobryukhov 6, P. A. Vernik 3, S. V. Gavrilov 3
FSBSI “All-Russian Research Institute of Medicinal and Aromatic Plants”, Moscow, Russia
All-Russian Scientific Research Institute of Vegetable Growing – branch of FSBSI “Federal Scientific Center of Vegetable Growing”, Vereya, Ramensky District, Moscow region, Russia
ANO “Institute for Development Strategies”, Moscow, Russia
FSBEI HE “Kazan Power Engineering University”, Kazan, Russia
FSBSI “All-Russian Research Institute of Rapeseed”, Lipetsk, Russia
FSBSI “Institute for Fundamental Problems of Biology of RAS”, Pushchino, Moscow region, Russia
The results of a comprehensive study of the effect of pulsed light irradiation on seed germination, plant growth and green mass yield are presented. The objects of research included 8 vegetable, medicinal and oil-bearing plants. A hypothesis was put forward that it is necessary to develop differentiated modes of light irradiation of plants (repetition period and pulse duration). It has been found that the effect of pulsed irradiation largely depends on the genetic characteristics of the objects. The most favorable and worst irradiation regimes were found in comparison with dark germination for plant productivity. Chlorophyll fluorescence parameters differ when exposed to pulsed light in the second and millisecond ranges. Analysis of the total antioxidant activity (TAOA) of green mass (microgreens) also showed significant differences depending on the genetic nature of the plant.
Key words: pulsed light, LEDs, green photonics, vegetables, oilseeds, medicinal plants, seeds, chlorophyll, fluorescence, growth and development
Received on: 28.05.2020
Accepted on: 10.07.2020
Introduction
When artificially growing plants in closed agroecosystems using photoculture, the regulation of lighting parameters becomes especially important [1–2]. Pulsed lighting modes, which allow saving energy, have long attracted the attention of researchers [3]. However, the complexity of the response of the photosynthetic and growth processes of plants to pulsed irradiation did not allow obtaining an unambiguous answer. This conclusion was reached both by the author of a fundamental monograph on photosynthesis, published back in 1954 [4], and by the authors of a newer review in 1980 [5]. Using modern techniques, it was found that reaction centers are capable of absorbing and storing energy from light pulses with a duration of about 100 μs and shorter, and then use it for the transport of electrons in the electron transport chain (ETC) during the dark pause between pulses. It has been suggested that the supply of light with short high-intensity pulses during the periods of activation of light-absorbing complexes and turning off the light during periods of their inactivation will be able to satisfy the energy needs of a number of green crops with relatively low values of the averaged PFD (photon flux density). Data were obtained indicating that light directed to plant sowing in short intense pulses provided a higher quantum efficiency of photosystem II and plant productivity compared to continuous light [6].
However, practical testing of pulsed irradiators in crop production has shown that they can have both stimulating and depressing effects. Practitioners have found that changing the constant illumination mode to a pulsed one (conditions: 2 μs – darkness, 4 μs – light, spectral ratio red light / blue light – RL / BL 2.3:1) at low (160 µmol m–2 s–1) the photon flux density does not cause changes in the rate of photosynthesis and water-retention capacity of leaves. The pulse mode in the variant with volumetric illumination throughout the entire growing season of tomato plants led to a decrease in the growth and development of plants, which ultimately does not allow reaching the levels of productivity obtained with constant illumination. In another study, it was shown that at 400 µmol m–2 s–1, pulsed light inhibited plant growth as compared to continuous radiation, especially when the pulse repetition period was lengthened above 350 µs. However, at an average irradiation level of about 500 µmol m–2 s–1, the dry weight of the crop was higher under pulsed illumination. The nature of the effect of pulsed illumination on plants significantly depended on the duration of the pulse repetition period. With pulsed light with a pulse repetition period of more than 450 μs, the inoculation productivity was higher than with continuous light. However, at values less than 400 μs, pulsed illumination negatively affected plant growth.
The analysis showed that further studies of pulsed irradiation are needed in order to better understand the mechanisms of its effect on plants. In particular, the reaction of plants in the genetic aspect has not been practically studied, systematic screening by genera, species and varieties of agricultural plants has not been carried out. It is also obvious that the plant response can be associated with a change in pulsed light modes. One of the important points is also the assessment of antioxidant activity as a possible marker of changes in metabolic processes in plants with changes in environmental conditions, including the lighting regime [7, 8]. It is known that when stressors initiate defense reactions, the composition and content of metabolites of the antioxidant metabolome change. Plant resistance to biotic and abiotic stresses may be associated with the accumulation of biologically active substances with antioxidant activity [9, 10].
In previous works [11–18], we studied the effect of pulsed irradiation modes on seed germination of a number of crops (germination, biometric parameters of seedlings and total antioxidant activity). This work continues this line of research. The aim of the research was to study the effect of a pulsed irradiation regime of plants of different species on the parameters of the photosynthetic apparatus, as well as growth processes when they are grown under controlled conditions of the agrobiotechnological system.
Materials and methods
The experiment was carried out in an ISR1.01 synergotron designed by the Institute for Development Strategies (Fig. 1, 2). Seeds were sown in Petri dishes with a mineral wool substrate, 25–50 pcs. in a dish, 3 times repetition. Temperature 24–25 °C. Sowing area included 9 cm Petri dishes with an area of 63.6 cm2.
The objects of research are seeds and sprouts of some vegetable, medicinal and oil-bearing crops: “Yubileiny” radish, salad mustard “Mei Lin”, amaranth “Lipetskiy”, ramtil “Lipchanin”, calendula, caraway seeds, fenugreek and Moldavian snakehead.
Polychrome phyto-lamps (produced by ANO “Institute for Development Strategies”) were used as LED lighting. The ratio of the spectra in all experiments: red 640 nm – 61.6%, blue 440 nm – 23.8%, green 520–530 nm – 6%, far red 740 nm – 7.2%, UV 380 nm – 1.5%. The protocol of the spectral characteristics of the luminaires in the ISR1.01 synergotron is shown in Fig. 3.
Moreover, we studied the option of germinating seeds in the dark, this is the main option prescribed by the GOST requirements for germinating seeds of agricultural crops. The intensity of illumination at the level of the seed surface: peak (during the period of the pulse) 265 μM / m2s, averaged over time, differs in different experiments (see below).
In total, 3 variants of experiments with different modes of pulsed illumination were carried out for 8 agricultural crops of vegetable, oilseed and medicinal uses:
modes of pulsed irradiation 1 / 3 s, 1 / 3 ms – time-averaged irradiation intensity of 66.3 μM / m2 ∙ s;
mode 1 / 2 s – time-averaged irradiation intensity of 88.3 μM / m2 ∙ s;
mode 1 / 1 s – time-averaged irradiation intensity of 132.5 μM / m2 ∙ s,
The 1 / 3 s designation corresponds to the mode: 1 second – duration of light emission by the LED, 3 seconds – duration of the dark period; the 1 / 3 ms designation corresponds to the mode: 1 millisecond – duration of light emission by the LED, 3 milliseconds – duration of the dark period; etc. Lighting in this mode was around the clock, i. e. 24 hours a day.
Germination energy and germination were determined according to GOST 12038-84. The total antioxidant activity (TAOA) was determined by the coulometric method using the electric generation of bromine radicals. The samples were analyzed on an Expert‑006 coulometer (by Econix-Expert LLC, Russia) according to our certified method [19]. The electric generation of bromine radicals was carried out from a 0.2 M solution of potassium bromide in a 0.1 M aqueous solution of sulfuric acid at a constant current of 100.0 mA. 30 ml of the background solution was introduced into the electrolytic cell, and when the indicator current reached a certain value, an aliquot of the water extract of the test sample with a volume of 0.200–0.500 cm3. The determination was carried out at room temperature. The essence of TAOA measurement is that radicals are generated in the measuring cell under the action of an electric current, in this case bromine: , , ; reactive oxygen species: , , , ,; HOBr. When 5 cm3 of aqueous extracts of plant samples are introduced into the measuring cell, they react with radicals, and the device gives out quantitative contents of antioxidants, which are statistically processed and entered into the memory of a personal computer in the form of a table of values. The device was calibrated with an alcohol solution of the Russian standard sample (RSS) of rutin [20] prepared according to the current State Pharmacopoeia, XI edition [21]. The TAOA was expressed in g of a standard sample of rutin (Ru) per 100 g of a sample per dry (d. s.) or completely dry (c. d. s.) sample.
Statistical processing of the obtained results was carried out through the modal value (mode) of 10 determinations [22], the relative error in determining the TAOA of the investigated water samples (E rel.) according to Table 1. was in the range of 1.34–3.25%.
Chlorophyll fluorescence in plant leaves after exposure to pulsed illumination was determined using a PAM fluorometer.
Results and discussion
Experiment 1.
Second and millisecond ranges
of pulsed illumination (1 / 3 s, 1 / 3 ms)
The data given in Fig. 4 show that the effect of pulsed irradiation is highly dependent on the genetic characteristics of the object. Stimulation of seed germination under the influence of pulsed irradiation of 1 / 3 s is typical for crops such as snakehead and calendula, the negative effect of pulsed irradiation is for caraway and mustard. No significant differences were observed for radish, fenugreek, amaranth and ramtil.
Comparison of the second (1 / 3 s) and millisecond (1 / 3 ms) ranges of pulsed irradiation showed that, in general, the second range is better tolerated by plants, especially in amaranth and ramtil crops. For other crops, the germination energy under different irradiation regimes does not differ significantly, and for the snakehead, the effect of stimulation under the influence of millisecond pulsed irradiation (by 10.7%) was noted.
The stimulating effect of pulsed irradiation in the second range of 1 / 3 s was manifested only in calendula (an increase in seed germination energy by 8.9%) and ramtil (an increase in seed germination energy by 4.2%). For other crops, the effect of decreasing germination was noted (the most pronounced for crops was caraway, snakehead, amaranth). The millisecond range of pulsed irradiation of 1 / 3 ms turned out to be unfavorable for amaranth and ramtil; for other crops, sharp differences in comparison with 1 / 3 s were not established both in the germination energy (Fig. 4) and in the seed germination of the studied crops (Fig. 5).
With dark germination, etiolated, elongated plants are formed, photosynthesis does not occur. Under pulse illumination of 1 / 3 s, proportionally developed plants are formed. In the experiment for most of the crops, there was a noticeable lag in height of plants grown in the millisecond range of 1 / 3 ms compared to 1 / 3 s (Fig. 6). Only radishes and fenugreek, and to a lesser extent mustard, in which the sprout height was comparable under different illumination conditions, showed resistance to the adverse effects of pulsed radiation in the millisecond range.
One of the most important criteria for assessing the impact of pulsed irradiation is plant biomass. Despite a significant excess in height of plants in the dark variant, their biomass did not increase in proportion to the growth. For most crops, the maximum biomass was observed under pulsed illumination in the 1 / 3 s mode (Fig. 7). However, the differences between the variants of pulsed illumination are not as pronounced as the differences in plant height.
Probably, under illumination of 1 / 3 ms, plants with a different morphological structure are formed – smaller in height, but not very different in biomass compared to plants grown at 1 / 3 s.
The final economically valuable indicator of the efficiency of plant growing is yield. This indicator also showed significant genetic differences. When grown in the mode of pulsed irradiation in a second range of 1 / 3 s, the yield of green mass (microgreens) of calendula and fenugreek increased twice or more compared to dark germination, caraway, snakehead, radish and ramtil – more by 10–28% (Fig. 8). At the same time, the yield of mustard microgreens decreased by 16%, amaranth – by 26%.
In all the medicinal and essential oil plants studied in the experiment, the yield turned out to be higher when grown in a millisecond mode of pulsed illumination of 1 / 3 ms compared to 1 / 3 s. At the same time, for vegetable and oilseeds, a different pattern is characteristic – the yield of radish and amaranth is almost twice as much at 1 / 3 s, than 1 / 3 ms. The yield of microgreens of ramtil and mustard is also higher with pulsed irradiation in the second range. Thus, the nature of the response of plants to pulsed irradiation in different modes depends primarily on the genetic nature of the object under study.
Analysis of the total antioxidant activity (TAOA) of green mass (microgreens) also showed significant differences depending on the genetic nature of the plant. The millisecond range of pulsed illumination of 1 / 3 ms caused an increase in the TAOA of the microgreens of mustard, amaranth, and snakehead (Fig. 9). The TAOA of ramtil and fenugreek microgreens practically did not change under different modes of pulsed illumination. In caraway, radish and calendula plants, the total antioxidant activity in the 1 / 3 ms mode, on the contrary, decreased.
In the majority of the studied cultures, the TAOA of the aerial part during germination in the dark turned out to be lower than under pulsed illumination. Only ramtil in the dark variant showed a slight increase in the TAOA index.
In the experiment, the fluorescence of chlorophyll in radish and ramtil plants was determined using a PAM fluorimeter. In radish seedlings irradiated in the millisecond range of 1 / 3 ms, the maximum quantum yield increases as compared to 1 / 3 s, while a decrease is observed in the rest of the fluorescence parameters (Fig. 10). In ramtil plants, the maximum quantum yield and non-photochemical quenching increase, while the actual quantum yield and electron transport rate decrease.
Experiment 2.
Pulsed illumination in 1 / 2 s mode
Among the studied cultures, stimulation of seed germination energy under the influence of pulsed irradiation in 1 / 2 s mode was observed only in the snakehead – by 14.7% (Fig. 11). A strong inhibitory effect (a decrease in germination energy by 36.6%) was manifested in caraway seeds, a decrease at the level of 5.55% – ramtil, 4.5% – radish, 3.3% – mustard. Compared with the 1 / 3 s regimen (Fig. 4), there was no stimulation effect in calendula and a decrease in the negative effect in mustard.
The previously noted effect of stimulating the germination energy by the seeds of the snakehead was lost with further germination of the seeds – the germination of seeds under pulsed irradiation decreased by 20% compared with dark germination (Fig. 12). The strong inhibitory effect of pulsed illumination on germination is especially characteristic of caraway seeds (61.3%) and ramtil (36.5%). Seed germination of other crops also decreased after pulsed illumination, but to a lesser extent. Compared with the 1 / 3 s option (Fig. 5), there was no effect of stimulating seed germination in calendula and ramtil.
Similarly to experiment 1, in experiment 2, during dark germination, etiolation and stretching of plants were observed. Under pulsed illumination, normally developed plants of lower height were formed (Fig. 13).
Despite a significant excess in height of plants in the dark variant, the biomass did not increase in proportion to the growth. The effect of pulsed illumination on the mass of the aboveground part of plants, in comparison with dark germination, was most significantly manifested for radishes, calendula, and fenugreek (Fig. 14).
In some crops (radish, calendula and fenugreek), the yield of the aboveground mass – microgreens increased, while in other crops the best yield was observed in the dark germination option (Fig. 15)
In most of the studied crops (mustard, amaranth, ramtil), the total antioxidant activity (TAOA) of the aerial part during germination in the dark turned out to be lower than under pulsed illumination (Fig. 16). Only in the dark version of radish, there was a slight increase in TAOA.
Experiment 3.
Pulsed illumination in 1 / 1 s mode
Similarly to experiment 2, in experiment 3 using pulsed irradiation of 1 / 1 s, the stimulation effect was observed only in the seeds of the snakehead (Fig. 17). For all other crops, especially for caraway seeds, the germination energy in the dark variant was higher.
In terms of seed germination, the results are also similar to those of experiment 2. After seed germination under pulse illumination for 1 / 1 s, a decrease in seed germination of caraway, snakehead, ramtil, calendula and, to a lesser extent, fenugreek, radish and mustard is observed (Fig. 18).
Similar to experiments 1 and 2, in experiment 3 during dark germination, etiolation and stretching of plants were observed. Under pulsed illumination, normally developed plants of lower height were formed (Fig. 19).
The stimulating effect of pulsed illumination of 1 / 1 s in comparison with dark germination was manifested only in calendula and fenugreek (Fig. 20). No stimulating effect of 1 / 1 s pulse illumination on radishes was noted, as in experiment No. 2.
In the same crops (fenugreek and calendula), after germination under pulsed irradiation in the 1 / 1 s mode, the yield of the aboveground mass (microgreens) increased (Fig. 21). For other crops, a decrease in microgreen yield was observed compared to dark germination.
As in experiment 2, in most of the studied crops (mustard, amaranth, ramtil), the total antioxidant activity (TAOA) of the aerial part during germination in the dark turned out to be less than under pulsed illumination or did not change (Fig. 22). Only radish in the dark variant showed a slight increase in the TAOA index.
CONCLUSION
In experiments with 8 vegetable, oil and medicinal crops, it was found that the effect of pulsed irradiation on plant growth largely depends on the genetic characteristics of the object. Stimulation of seed germination under the influence of pulsed irradiation of 1 / 3 s is typical for crops such as snakehead and calendula, and a negative effect was found for caraway and mustard. No significant differences were observed for radish, fenugreek, amaranth and ramtil. The yield of green mass (microgreens) in the 1 / 3 s variant increased twice or more in calendula and fenugreek compared to dark germination, caraway, snakehead, radish and ramtil – more by 10–28%, but decreased by 16% in mustard and by 26% for amaranth. Other studied modes of pulsed irradiation of 1 / 2 s and 1 / 1 s are generally less favorable for plants than 1 / 3 s.
Comparison of the second (1 / 3 s) and millisecond (1 / 3 ms) ranges of pulsed irradiation showed that, in general, the second range is more favorable for plant growth. However, in medicinal plants, the yield of green mass (sprouts), on the contrary, turned out to be higher when grown in a millisecond mode of pulsed illumination of 1 / 3 ms compared to 1 / 3 s. The germination capacity of amaranth and ramtil seeds decreases with irradiation in the millisecond range; it differs slightly in other crops. Chlorophyll fluorescence parameters differ when exposed to pulsed light in the second and millisecond ranges.
Analysis of the total antioxidant activity (TAOA) of green mass (microgreens) also showed significant differences depending on the genetic nature of the plant. Irradiation in the millisecond range of 1 / 3 ms led to an increase in the TAOA of the microgreens of mustard, amaranth, and snakehead. In caraway, radish and calendula plants, the total antioxidant activity in the 1 / 3 ms mode, on the contrary, decreased. The TAOA of ramtil and fenugreek microgreens practically did not change under different modes of pulsed illumination. In most of the studied crops (except for ramtil), the TAOA of the aboveground part during germination in the dark turned out to be lower than under pulsed illumination in all studied modes.
Thus, due to the significant dependence of plant responses to pulsed irradiation on genetic characteristics, it is necessary to develop differentiated modes of irradiation with light radiation (repetition period and pulse duration).
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AUTHORS
V. N. Zelenkov, Candidate of Chemical Sciences, Doctor of Agricultural Sciences, Professor, E‑mail: zelenkov-raen@mail.ru, All-Russian Research Institute of Medicinal and Aromatic Plants, Moscow; All-Russian Scientific Research Institute of Vegetable Growing – branch of the Federal State Budgetary Scientific Institution “Federal Scientific Center for Vegetable Growing”, Vereya, Ramensky District, Moscow region; ANO “Institute for Development Strategies”, Moscow, Russia.
ORCID: 0000-0001-5481-2723
V. V. Latushkin, Candidate of Chemical Sciences, ANO “Institute for Development Strategies”, E‑mail: slavalat@yandex.ru, Moscow, Russia.
ORCID: 0000-0003-1406-8965
M. I. Ivanova, Doctor of Agricultural Sciences, Professor of the Russian Academy of Sciences, E‑mail: ivanova_170@mail.ru, All-Russian Scientific Research Institute of Vegetable Growing – branch of the Federal State Budgetary Scientific Institution “Federal Scientific Center of Vegetable Growing”, Vereya, Ramenskiy district, Moscow region, Russia.
ORCID: 0000-0001-7326-2157
A. A. Lapin, Ph.D., Associate Professor, Kazan Power Engineering University, E‑mail: kgeu-oso@mail.ru, Kazan, Russia.
ORCID: 0000-0001-9142-0403
V. V. Karpachev, Doctor of Agricultural Sciences, Professor, Corresponding Member of RAS, Federal State Budgetary Scientific Institution “All-Russian Research Institute of Rapeseed”, E‑mail: vniirapsa@mail.ru, Lipetsk, Russia.
ORCID: 0000-0002-1141-2065
A. A. Kosobryukhov, Doctor of Biological Sciences, Institute of Fundamental Problems of Biology RAS, E‑mail: kosobr@rambler.ru, Pushchino, Moscow region, Russia.
ORCID: 0000-0001-7453-3123
P. A. Vernik, ANO “Institute for Development Strategies”, E‑mail: slavalat@yandex.ru, Moscow, Russia.
ORCID: 0000-0001-5850-7654
S. V. Gavrilov, ANO “Institute for Development Strategies”, E‑mail: slavalat@yandex.ru, Moscow, Russia.
ORCID: 0000-0003-2824-9302
Contribution of members of the creative team to the project
Zelenkov V. N. – scientific management, planning, organization of experiments, analysis and evaluation of results, analysis of literature; V. V. Latushkin – planning, setting and conducting experiments and their processing, literature analysis; Ivanova M. I. – planing for vegetable crops, setting up and providing an experiment, analysis of literature on the subject; Lapin A. A. – analysis of the total antioxidant activity of plant samples and their processing, analysis of the literature on the subject; V. V. Karpachev – planning experiments on oilseeds, new crops and providing experiments; A. A. Kosobryukhov – planning, setting up and conducting experiments with a number of cultures to study photosynthesis under the influence of impulse exposure to EMP of visible spectrum, literature analysis; Vernik P. A. – organization of work on the synergotron model 1.01 designed by ANO “Institute for Development Strategies”, planning and support of experiments at the facility; S. V. Gavrilov – measureement of spectral characteristics of LED lamps, instrumental support of the experiment in the agrobiotechnological system “synergotron model 1.01”, maintenance of the parameters of the environment for seed germination and plant growth (temperature, humidity, lighting) in programmed mode and control.
The study was funded by the ANO “Institute for Development Strategies” (Moscow) using its financial and technical capabilities using a synergotron (model 1.01) of its own design.
Conflict of interest
The authors declare that they have no conflicts of interest.
V. N. Zelenkov 1, 2, 3, V. V. Latushkin 3, M. I. Ivanova 2, A. A. Lapin 4, V. V. Karpachev 5, A. A. Kosobryukhov 6, P. A. Vernik 3, S. V. Gavrilov 3
FSBSI “All-Russian Research Institute of Medicinal and Aromatic Plants”, Moscow, Russia
All-Russian Scientific Research Institute of Vegetable Growing – branch of FSBSI “Federal Scientific Center of Vegetable Growing”, Vereya, Ramensky District, Moscow region, Russia
ANO “Institute for Development Strategies”, Moscow, Russia
FSBEI HE “Kazan Power Engineering University”, Kazan, Russia
FSBSI “All-Russian Research Institute of Rapeseed”, Lipetsk, Russia
FSBSI “Institute for Fundamental Problems of Biology of RAS”, Pushchino, Moscow region, Russia
The results of a comprehensive study of the effect of pulsed light irradiation on seed germination, plant growth and green mass yield are presented. The objects of research included 8 vegetable, medicinal and oil-bearing plants. A hypothesis was put forward that it is necessary to develop differentiated modes of light irradiation of plants (repetition period and pulse duration). It has been found that the effect of pulsed irradiation largely depends on the genetic characteristics of the objects. The most favorable and worst irradiation regimes were found in comparison with dark germination for plant productivity. Chlorophyll fluorescence parameters differ when exposed to pulsed light in the second and millisecond ranges. Analysis of the total antioxidant activity (TAOA) of green mass (microgreens) also showed significant differences depending on the genetic nature of the plant.
Key words: pulsed light, LEDs, green photonics, vegetables, oilseeds, medicinal plants, seeds, chlorophyll, fluorescence, growth and development
Received on: 28.05.2020
Accepted on: 10.07.2020
Introduction
When artificially growing plants in closed agroecosystems using photoculture, the regulation of lighting parameters becomes especially important [1–2]. Pulsed lighting modes, which allow saving energy, have long attracted the attention of researchers [3]. However, the complexity of the response of the photosynthetic and growth processes of plants to pulsed irradiation did not allow obtaining an unambiguous answer. This conclusion was reached both by the author of a fundamental monograph on photosynthesis, published back in 1954 [4], and by the authors of a newer review in 1980 [5]. Using modern techniques, it was found that reaction centers are capable of absorbing and storing energy from light pulses with a duration of about 100 μs and shorter, and then use it for the transport of electrons in the electron transport chain (ETC) during the dark pause between pulses. It has been suggested that the supply of light with short high-intensity pulses during the periods of activation of light-absorbing complexes and turning off the light during periods of their inactivation will be able to satisfy the energy needs of a number of green crops with relatively low values of the averaged PFD (photon flux density). Data were obtained indicating that light directed to plant sowing in short intense pulses provided a higher quantum efficiency of photosystem II and plant productivity compared to continuous light [6].
However, practical testing of pulsed irradiators in crop production has shown that they can have both stimulating and depressing effects. Practitioners have found that changing the constant illumination mode to a pulsed one (conditions: 2 μs – darkness, 4 μs – light, spectral ratio red light / blue light – RL / BL 2.3:1) at low (160 µmol m–2 s–1) the photon flux density does not cause changes in the rate of photosynthesis and water-retention capacity of leaves. The pulse mode in the variant with volumetric illumination throughout the entire growing season of tomato plants led to a decrease in the growth and development of plants, which ultimately does not allow reaching the levels of productivity obtained with constant illumination. In another study, it was shown that at 400 µmol m–2 s–1, pulsed light inhibited plant growth as compared to continuous radiation, especially when the pulse repetition period was lengthened above 350 µs. However, at an average irradiation level of about 500 µmol m–2 s–1, the dry weight of the crop was higher under pulsed illumination. The nature of the effect of pulsed illumination on plants significantly depended on the duration of the pulse repetition period. With pulsed light with a pulse repetition period of more than 450 μs, the inoculation productivity was higher than with continuous light. However, at values less than 400 μs, pulsed illumination negatively affected plant growth.
The analysis showed that further studies of pulsed irradiation are needed in order to better understand the mechanisms of its effect on plants. In particular, the reaction of plants in the genetic aspect has not been practically studied, systematic screening by genera, species and varieties of agricultural plants has not been carried out. It is also obvious that the plant response can be associated with a change in pulsed light modes. One of the important points is also the assessment of antioxidant activity as a possible marker of changes in metabolic processes in plants with changes in environmental conditions, including the lighting regime [7, 8]. It is known that when stressors initiate defense reactions, the composition and content of metabolites of the antioxidant metabolome change. Plant resistance to biotic and abiotic stresses may be associated with the accumulation of biologically active substances with antioxidant activity [9, 10].
In previous works [11–18], we studied the effect of pulsed irradiation modes on seed germination of a number of crops (germination, biometric parameters of seedlings and total antioxidant activity). This work continues this line of research. The aim of the research was to study the effect of a pulsed irradiation regime of plants of different species on the parameters of the photosynthetic apparatus, as well as growth processes when they are grown under controlled conditions of the agrobiotechnological system.
Materials and methods
The experiment was carried out in an ISR1.01 synergotron designed by the Institute for Development Strategies (Fig. 1, 2). Seeds were sown in Petri dishes with a mineral wool substrate, 25–50 pcs. in a dish, 3 times repetition. Temperature 24–25 °C. Sowing area included 9 cm Petri dishes with an area of 63.6 cm2.
The objects of research are seeds and sprouts of some vegetable, medicinal and oil-bearing crops: “Yubileiny” radish, salad mustard “Mei Lin”, amaranth “Lipetskiy”, ramtil “Lipchanin”, calendula, caraway seeds, fenugreek and Moldavian snakehead.
Polychrome phyto-lamps (produced by ANO “Institute for Development Strategies”) were used as LED lighting. The ratio of the spectra in all experiments: red 640 nm – 61.6%, blue 440 nm – 23.8%, green 520–530 nm – 6%, far red 740 nm – 7.2%, UV 380 nm – 1.5%. The protocol of the spectral characteristics of the luminaires in the ISR1.01 synergotron is shown in Fig. 3.
Moreover, we studied the option of germinating seeds in the dark, this is the main option prescribed by the GOST requirements for germinating seeds of agricultural crops. The intensity of illumination at the level of the seed surface: peak (during the period of the pulse) 265 μM / m2s, averaged over time, differs in different experiments (see below).
In total, 3 variants of experiments with different modes of pulsed illumination were carried out for 8 agricultural crops of vegetable, oilseed and medicinal uses:
modes of pulsed irradiation 1 / 3 s, 1 / 3 ms – time-averaged irradiation intensity of 66.3 μM / m2 ∙ s;
mode 1 / 2 s – time-averaged irradiation intensity of 88.3 μM / m2 ∙ s;
mode 1 / 1 s – time-averaged irradiation intensity of 132.5 μM / m2 ∙ s,
The 1 / 3 s designation corresponds to the mode: 1 second – duration of light emission by the LED, 3 seconds – duration of the dark period; the 1 / 3 ms designation corresponds to the mode: 1 millisecond – duration of light emission by the LED, 3 milliseconds – duration of the dark period; etc. Lighting in this mode was around the clock, i. e. 24 hours a day.
Germination energy and germination were determined according to GOST 12038-84. The total antioxidant activity (TAOA) was determined by the coulometric method using the electric generation of bromine radicals. The samples were analyzed on an Expert‑006 coulometer (by Econix-Expert LLC, Russia) according to our certified method [19]. The electric generation of bromine radicals was carried out from a 0.2 M solution of potassium bromide in a 0.1 M aqueous solution of sulfuric acid at a constant current of 100.0 mA. 30 ml of the background solution was introduced into the electrolytic cell, and when the indicator current reached a certain value, an aliquot of the water extract of the test sample with a volume of 0.200–0.500 cm3. The determination was carried out at room temperature. The essence of TAOA measurement is that radicals are generated in the measuring cell under the action of an electric current, in this case bromine: , , ; reactive oxygen species: , , , ,; HOBr. When 5 cm3 of aqueous extracts of plant samples are introduced into the measuring cell, they react with radicals, and the device gives out quantitative contents of antioxidants, which are statistically processed and entered into the memory of a personal computer in the form of a table of values. The device was calibrated with an alcohol solution of the Russian standard sample (RSS) of rutin [20] prepared according to the current State Pharmacopoeia, XI edition [21]. The TAOA was expressed in g of a standard sample of rutin (Ru) per 100 g of a sample per dry (d. s.) or completely dry (c. d. s.) sample.
Statistical processing of the obtained results was carried out through the modal value (mode) of 10 determinations [22], the relative error in determining the TAOA of the investigated water samples (E rel.) according to Table 1. was in the range of 1.34–3.25%.
Chlorophyll fluorescence in plant leaves after exposure to pulsed illumination was determined using a PAM fluorometer.
Results and discussion
Experiment 1.
Second and millisecond ranges
of pulsed illumination (1 / 3 s, 1 / 3 ms)
The data given in Fig. 4 show that the effect of pulsed irradiation is highly dependent on the genetic characteristics of the object. Stimulation of seed germination under the influence of pulsed irradiation of 1 / 3 s is typical for crops such as snakehead and calendula, the negative effect of pulsed irradiation is for caraway and mustard. No significant differences were observed for radish, fenugreek, amaranth and ramtil.
Comparison of the second (1 / 3 s) and millisecond (1 / 3 ms) ranges of pulsed irradiation showed that, in general, the second range is better tolerated by plants, especially in amaranth and ramtil crops. For other crops, the germination energy under different irradiation regimes does not differ significantly, and for the snakehead, the effect of stimulation under the influence of millisecond pulsed irradiation (by 10.7%) was noted.
The stimulating effect of pulsed irradiation in the second range of 1 / 3 s was manifested only in calendula (an increase in seed germination energy by 8.9%) and ramtil (an increase in seed germination energy by 4.2%). For other crops, the effect of decreasing germination was noted (the most pronounced for crops was caraway, snakehead, amaranth). The millisecond range of pulsed irradiation of 1 / 3 ms turned out to be unfavorable for amaranth and ramtil; for other crops, sharp differences in comparison with 1 / 3 s were not established both in the germination energy (Fig. 4) and in the seed germination of the studied crops (Fig. 5).
With dark germination, etiolated, elongated plants are formed, photosynthesis does not occur. Under pulse illumination of 1 / 3 s, proportionally developed plants are formed. In the experiment for most of the crops, there was a noticeable lag in height of plants grown in the millisecond range of 1 / 3 ms compared to 1 / 3 s (Fig. 6). Only radishes and fenugreek, and to a lesser extent mustard, in which the sprout height was comparable under different illumination conditions, showed resistance to the adverse effects of pulsed radiation in the millisecond range.
One of the most important criteria for assessing the impact of pulsed irradiation is plant biomass. Despite a significant excess in height of plants in the dark variant, their biomass did not increase in proportion to the growth. For most crops, the maximum biomass was observed under pulsed illumination in the 1 / 3 s mode (Fig. 7). However, the differences between the variants of pulsed illumination are not as pronounced as the differences in plant height.
Probably, under illumination of 1 / 3 ms, plants with a different morphological structure are formed – smaller in height, but not very different in biomass compared to plants grown at 1 / 3 s.
The final economically valuable indicator of the efficiency of plant growing is yield. This indicator also showed significant genetic differences. When grown in the mode of pulsed irradiation in a second range of 1 / 3 s, the yield of green mass (microgreens) of calendula and fenugreek increased twice or more compared to dark germination, caraway, snakehead, radish and ramtil – more by 10–28% (Fig. 8). At the same time, the yield of mustard microgreens decreased by 16%, amaranth – by 26%.
In all the medicinal and essential oil plants studied in the experiment, the yield turned out to be higher when grown in a millisecond mode of pulsed illumination of 1 / 3 ms compared to 1 / 3 s. At the same time, for vegetable and oilseeds, a different pattern is characteristic – the yield of radish and amaranth is almost twice as much at 1 / 3 s, than 1 / 3 ms. The yield of microgreens of ramtil and mustard is also higher with pulsed irradiation in the second range. Thus, the nature of the response of plants to pulsed irradiation in different modes depends primarily on the genetic nature of the object under study.
Analysis of the total antioxidant activity (TAOA) of green mass (microgreens) also showed significant differences depending on the genetic nature of the plant. The millisecond range of pulsed illumination of 1 / 3 ms caused an increase in the TAOA of the microgreens of mustard, amaranth, and snakehead (Fig. 9). The TAOA of ramtil and fenugreek microgreens practically did not change under different modes of pulsed illumination. In caraway, radish and calendula plants, the total antioxidant activity in the 1 / 3 ms mode, on the contrary, decreased.
In the majority of the studied cultures, the TAOA of the aerial part during germination in the dark turned out to be lower than under pulsed illumination. Only ramtil in the dark variant showed a slight increase in the TAOA index.
In the experiment, the fluorescence of chlorophyll in radish and ramtil plants was determined using a PAM fluorimeter. In radish seedlings irradiated in the millisecond range of 1 / 3 ms, the maximum quantum yield increases as compared to 1 / 3 s, while a decrease is observed in the rest of the fluorescence parameters (Fig. 10). In ramtil plants, the maximum quantum yield and non-photochemical quenching increase, while the actual quantum yield and electron transport rate decrease.
Experiment 2.
Pulsed illumination in 1 / 2 s mode
Among the studied cultures, stimulation of seed germination energy under the influence of pulsed irradiation in 1 / 2 s mode was observed only in the snakehead – by 14.7% (Fig. 11). A strong inhibitory effect (a decrease in germination energy by 36.6%) was manifested in caraway seeds, a decrease at the level of 5.55% – ramtil, 4.5% – radish, 3.3% – mustard. Compared with the 1 / 3 s regimen (Fig. 4), there was no stimulation effect in calendula and a decrease in the negative effect in mustard.
The previously noted effect of stimulating the germination energy by the seeds of the snakehead was lost with further germination of the seeds – the germination of seeds under pulsed irradiation decreased by 20% compared with dark germination (Fig. 12). The strong inhibitory effect of pulsed illumination on germination is especially characteristic of caraway seeds (61.3%) and ramtil (36.5%). Seed germination of other crops also decreased after pulsed illumination, but to a lesser extent. Compared with the 1 / 3 s option (Fig. 5), there was no effect of stimulating seed germination in calendula and ramtil.
Similarly to experiment 1, in experiment 2, during dark germination, etiolation and stretching of plants were observed. Under pulsed illumination, normally developed plants of lower height were formed (Fig. 13).
Despite a significant excess in height of plants in the dark variant, the biomass did not increase in proportion to the growth. The effect of pulsed illumination on the mass of the aboveground part of plants, in comparison with dark germination, was most significantly manifested for radishes, calendula, and fenugreek (Fig. 14).
In some crops (radish, calendula and fenugreek), the yield of the aboveground mass – microgreens increased, while in other crops the best yield was observed in the dark germination option (Fig. 15)
In most of the studied crops (mustard, amaranth, ramtil), the total antioxidant activity (TAOA) of the aerial part during germination in the dark turned out to be lower than under pulsed illumination (Fig. 16). Only in the dark version of radish, there was a slight increase in TAOA.
Experiment 3.
Pulsed illumination in 1 / 1 s mode
Similarly to experiment 2, in experiment 3 using pulsed irradiation of 1 / 1 s, the stimulation effect was observed only in the seeds of the snakehead (Fig. 17). For all other crops, especially for caraway seeds, the germination energy in the dark variant was higher.
In terms of seed germination, the results are also similar to those of experiment 2. After seed germination under pulse illumination for 1 / 1 s, a decrease in seed germination of caraway, snakehead, ramtil, calendula and, to a lesser extent, fenugreek, radish and mustard is observed (Fig. 18).
Similar to experiments 1 and 2, in experiment 3 during dark germination, etiolation and stretching of plants were observed. Under pulsed illumination, normally developed plants of lower height were formed (Fig. 19).
The stimulating effect of pulsed illumination of 1 / 1 s in comparison with dark germination was manifested only in calendula and fenugreek (Fig. 20). No stimulating effect of 1 / 1 s pulse illumination on radishes was noted, as in experiment No. 2.
In the same crops (fenugreek and calendula), after germination under pulsed irradiation in the 1 / 1 s mode, the yield of the aboveground mass (microgreens) increased (Fig. 21). For other crops, a decrease in microgreen yield was observed compared to dark germination.
As in experiment 2, in most of the studied crops (mustard, amaranth, ramtil), the total antioxidant activity (TAOA) of the aerial part during germination in the dark turned out to be less than under pulsed illumination or did not change (Fig. 22). Only radish in the dark variant showed a slight increase in the TAOA index.
CONCLUSION
In experiments with 8 vegetable, oil and medicinal crops, it was found that the effect of pulsed irradiation on plant growth largely depends on the genetic characteristics of the object. Stimulation of seed germination under the influence of pulsed irradiation of 1 / 3 s is typical for crops such as snakehead and calendula, and a negative effect was found for caraway and mustard. No significant differences were observed for radish, fenugreek, amaranth and ramtil. The yield of green mass (microgreens) in the 1 / 3 s variant increased twice or more in calendula and fenugreek compared to dark germination, caraway, snakehead, radish and ramtil – more by 10–28%, but decreased by 16% in mustard and by 26% for amaranth. Other studied modes of pulsed irradiation of 1 / 2 s and 1 / 1 s are generally less favorable for plants than 1 / 3 s.
Comparison of the second (1 / 3 s) and millisecond (1 / 3 ms) ranges of pulsed irradiation showed that, in general, the second range is more favorable for plant growth. However, in medicinal plants, the yield of green mass (sprouts), on the contrary, turned out to be higher when grown in a millisecond mode of pulsed illumination of 1 / 3 ms compared to 1 / 3 s. The germination capacity of amaranth and ramtil seeds decreases with irradiation in the millisecond range; it differs slightly in other crops. Chlorophyll fluorescence parameters differ when exposed to pulsed light in the second and millisecond ranges.
Analysis of the total antioxidant activity (TAOA) of green mass (microgreens) also showed significant differences depending on the genetic nature of the plant. Irradiation in the millisecond range of 1 / 3 ms led to an increase in the TAOA of the microgreens of mustard, amaranth, and snakehead. In caraway, radish and calendula plants, the total antioxidant activity in the 1 / 3 ms mode, on the contrary, decreased. The TAOA of ramtil and fenugreek microgreens practically did not change under different modes of pulsed illumination. In most of the studied crops (except for ramtil), the TAOA of the aboveground part during germination in the dark turned out to be lower than under pulsed illumination in all studied modes.
Thus, due to the significant dependence of plant responses to pulsed irradiation on genetic characteristics, it is necessary to develop differentiated modes of irradiation with light radiation (repetition period and pulse duration).
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AUTHORS
V. N. Zelenkov, Candidate of Chemical Sciences, Doctor of Agricultural Sciences, Professor, E‑mail: zelenkov-raen@mail.ru, All-Russian Research Institute of Medicinal and Aromatic Plants, Moscow; All-Russian Scientific Research Institute of Vegetable Growing – branch of the Federal State Budgetary Scientific Institution “Federal Scientific Center for Vegetable Growing”, Vereya, Ramensky District, Moscow region; ANO “Institute for Development Strategies”, Moscow, Russia.
ORCID: 0000-0001-5481-2723
V. V. Latushkin, Candidate of Chemical Sciences, ANO “Institute for Development Strategies”, E‑mail: slavalat@yandex.ru, Moscow, Russia.
ORCID: 0000-0003-1406-8965
M. I. Ivanova, Doctor of Agricultural Sciences, Professor of the Russian Academy of Sciences, E‑mail: ivanova_170@mail.ru, All-Russian Scientific Research Institute of Vegetable Growing – branch of the Federal State Budgetary Scientific Institution “Federal Scientific Center of Vegetable Growing”, Vereya, Ramenskiy district, Moscow region, Russia.
ORCID: 0000-0001-7326-2157
A. A. Lapin, Ph.D., Associate Professor, Kazan Power Engineering University, E‑mail: kgeu-oso@mail.ru, Kazan, Russia.
ORCID: 0000-0001-9142-0403
V. V. Karpachev, Doctor of Agricultural Sciences, Professor, Corresponding Member of RAS, Federal State Budgetary Scientific Institution “All-Russian Research Institute of Rapeseed”, E‑mail: vniirapsa@mail.ru, Lipetsk, Russia.
ORCID: 0000-0002-1141-2065
A. A. Kosobryukhov, Doctor of Biological Sciences, Institute of Fundamental Problems of Biology RAS, E‑mail: kosobr@rambler.ru, Pushchino, Moscow region, Russia.
ORCID: 0000-0001-7453-3123
P. A. Vernik, ANO “Institute for Development Strategies”, E‑mail: slavalat@yandex.ru, Moscow, Russia.
ORCID: 0000-0001-5850-7654
S. V. Gavrilov, ANO “Institute for Development Strategies”, E‑mail: slavalat@yandex.ru, Moscow, Russia.
ORCID: 0000-0003-2824-9302
Contribution of members of the creative team to the project
Zelenkov V. N. – scientific management, planning, organization of experiments, analysis and evaluation of results, analysis of literature; V. V. Latushkin – planning, setting and conducting experiments and their processing, literature analysis; Ivanova M. I. – planing for vegetable crops, setting up and providing an experiment, analysis of literature on the subject; Lapin A. A. – analysis of the total antioxidant activity of plant samples and their processing, analysis of the literature on the subject; V. V. Karpachev – planning experiments on oilseeds, new crops and providing experiments; A. A. Kosobryukhov – planning, setting up and conducting experiments with a number of cultures to study photosynthesis under the influence of impulse exposure to EMP of visible spectrum, literature analysis; Vernik P. A. – organization of work on the synergotron model 1.01 designed by ANO “Institute for Development Strategies”, planning and support of experiments at the facility; S. V. Gavrilov – measureement of spectral characteristics of LED lamps, instrumental support of the experiment in the agrobiotechnological system “synergotron model 1.01”, maintenance of the parameters of the environment for seed germination and plant growth (temperature, humidity, lighting) in programmed mode and control.
The study was funded by the ANO “Institute for Development Strategies” (Moscow) using its financial and technical capabilities using a synergotron (model 1.01) of its own design.
Conflict of interest
The authors declare that they have no conflicts of interest.
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