EFFECTS OF SQUIRTING CUCUMBER FRUIT JUICE ON
SEED GERMINATION AND SEEDLING GROWTH IN
RAPE-SEED
Ramazan Beyaz 1,*, Yasin Ozgen2, Irem Poyraz3, Aslinur Cavdar4, Mustafa Yildiz2 1Kırşehir Ahi Evran University, Faculty of Agriculture, Department of Soil Science and Plant Nutrition, Kırşehir
2Ankara University, Faculty of Agriculture, Department of Field Crops, Ankara
3Ankara University, Graduate School of Natural and Applied Sciences, Department of Field Crops, Ankara 4Ankara University, Biotechnology Institute, Ankara
ABSTRACT
The aim of this study was to evaluate the effects of mature squirting cucumber fruit juice (SCFJ) on seed germination and seedling growth in rapeseed cv. "Elvis". Seeds were surface-sterilized with commercial bleach (containing 5% sodium hypochlorite) at a concentration of 40% for 5 min. with continuous stirring and then were washed 6-7 times with sterile distilled water. After sterilization, seeds were imbibed in 50 ml sterile distilled water having SCFJ at different concentrations (0-control, 200, 400, 800 and 1600 µl/L) and rinsed for 72 h at 180 rpm. For each SCFJ concentrations, 250 seeds were put in 50 ml sterile distilled water. After 72 h, grown seedlings were transferred to soil in pots. Five pots having 20 seedlings were set for each SCFJ concentration. 14 days after study initiation, seedling height, root length, leaf width, leaf length, approx. leaf area, and chlorophyll (chl. a, chl. b and total chl.) contents were determined. The highest results in all examined parameters were obtained from 800 µl/L SCFJ concentration. This study showed that SCFJ could be used as a stimulant accelerating seed germination and seedling growth. Putting production material to the market earlier is the guarantee of higher income by providing early harvest.
KEYWORDS:
Seedling growth, germination, fruit juice, squirting cu-cumber
INTRODUCTION
The chemical treatments influence the physiological and biochemical process in seeds and consequently contribute in a faster germination and a greater vegetative growth. Faster germination and seedling growth is a prerequisite for early harvest which means putting agricultural products on market earlier. The most important factor in earliness is the seedling which will be used instead of seed material. Healty and well-grown seedlings
is the guarantee of higher yield. Early supply of agricultural products to market, increasing the number of product harvested from the field in the same year, saving on pesticides, fertilizers and seed material, easiness in controlling of weeds, increasing income of producers are the some of the advantages of using seedlings instead of seed materials.
Squirting cucumber (Ecballium elaterium [L.] A. Rich.) is a plant of the cucumber family that is peculiar to Europe, Asia, and North Africa. In particular, the fruit of the plant consists of elatericin A (cucurbitacin D), elatericin B (cucurbitacin I), β-elaterin (cucurbitacin B), and α-β-elaterin (cucurbitacin E) [1]. In addition, alkaloids, vitamins, fatty acids, flavonoids, sterols, resins, starch, amino acids, and phenolic compounds are among the components of plant [2]. The compounds in the leaves, roots, stem and boll of plants also have antibacterial properties [3,4,5]. As a result, the squirting cucumber is very common in Turkey [6]. In the current study, SCFJ as a biostimulant was evaluated for rapid seed germination and seedling growth in rapeseed.
Rapeseed (Brassica napus ssp. oleifera L.) has been widely used for its oil in the world. Moreover, it has been used as livestock feed, biodiesel source, lamp oil, soap, plastics, rotating crop, heavy metal accumulator, medicinal plant (diuretic, anti-scurvy and anti-inflammatory) [7].
To our knowledge, this is the first report evaluating biostimulant effect of SCFJ on seed germination and seedling growth of rapeseed. In the present study, the effects of SCFJ on seed germination and seedling growth of rapeseed (Brassica napus L. cv. 'Elvis') were investigated.
MATERIALS AND METHODS
Plant material. Seeds of Brassica napus L.
cv. 'Elvis' were used in the study.
Squirting cucumber (Ecballium ela-terium (L.) A. Rich.) fruit juice. Squirting
areas in September. Fruit juice was extracted and then were filtered in order that bigger parts were eliminated. Fruit juice were filter sterilized and kept in the refrigerator at -20ºC.
Surface sterilization of seeds. The seeds
were surface-sterilized with 40% commercial bleach (containing 5.0% sodium hypochlorite) for 5 min and then washed 6-7 times with sterile distilled water (sdH20).
Seed germination and seedling establish-ment. After sterilization, seeds were imbibed in 50
ml sterile distilled water having SCFJ at different concentrations (0-control, 200, 400, 800 and 1600 µl/L) and rinsed for 72 h at 180 rpm. For each SCFJ concentrations, 250 seeds were put in 50 ml sterile distilled water. After 72 h, grown seedlings were transferred to soil in pots. Five pots having 20 seedlings were set for each SCFJ concentration.
Observations. Seed germination percentage
was determined at 48th and 72nd h. At the end of 72nd h, seedlings were transferred to soil in pots. A seed was considered germinated when the emerged radicle reached to 2 mm. Fourteen days later, plant-let and root lengths, leaf width, leaf length, approxi-mately leaf area, chlorophyll (chl. a, chl. b and total chl.) contents in the leaves of seedlings were recorded.
Determination of chlorophyll contents. The
protocol proposed by Curtis and Shetty [8]. was used to determine the chl. a, chl. b and total chl. contents of seedling leaves. By following the proto-col, 50 mg of callus tissue was added into methanol (3 ml) and kept at 23°C in darkness for 2 h to allow the chlorophyll in the green material to dissolve into the methanol. Then, the optic density (OD) of 1.5 ml of the liquid part (the chlorophyll-containing methanol) was determined at 650 and 665 nm using a spectrophotometer (UVmini-1240, Shimadzu, Japan), and the chl. a, chl. b and total chl. quantities were calculated as units of "µg chlorophyll/g of fresh tissue".
Determination of leaf area. Leaf area was
determined using a computer software and a desk-top scanner.
Statistical analysis. Three replicates were
tested and there were 250 seeds per replication for seed germination while 5 replicates each with 20 plantlets were set for seedling growth. All experi-ments were repeated twice. One-way Analysis of Variance (ANOVA) was used to evaluate the effect of SCFJ at different concentrations on seed germi-nation and seedling growth in rapeseed (Brassica napus L.). Data were statistically analyzed by using "IBM SPSS Statistics 22". Means were compared using Duncan's multiple range test at the 1% level
of probability. Data given in percentages were sub-jected to arcsine (√X) transformation before statisti-cal analysis [9].
RESULTS
The effects of different SCFJ concentrations on the seed germination, plantlet height, root length, leaf area, chlorophyll (chl. a, chl. b and total chl.) contents in the leaves of 14-day-old seedlings were shown in Table 1. The stimulative effects of SCFJ were observed at 800 µL/L concentration, and among all examined parameters, the best results were obtained at this concentration. Suppressor ef-fects were observed at 1600 µL/L concentration. The lowest results were recorded in the control group, which contained no SCFJ. The seed germi-nation percentages at 72 h increased at 800 µL/L SCFJ concentration, with the highest being 87.20%; the lowest percentages were observed in the control group at 21.60% (Table 1).
After 14 d, similar results were obtained for seedling height, root length, leaf area and for chlorophyll (chl. a, chl. b, and total chl.) contents in the leaf. The best results for these parameters were 16.3 cm, 11.6 cm, 24.75 cm2, 1491.030 µg
chloro-phyll a/g fresh tissue, 956.358 µg chlorochloro-phyll b/g fresh tissue, and 1008.720 µg total chlorophyll/g fresh tissue, respectively, and were observed for those treated with 800 µL/L concentration. At a 1600 µl/l SCFJ concentration, obtained results in all characters decreased significantly (Fig. 1, Table 1).
On the 14th day of the study, the best results of seedling height, root length, and leaf area and of chlorophyll (chl. a, chl. b, and total chl.) contents in the leaf were noticed at 800 µL/L SCFJ concentra-tion, which were much higher than those in the con-trol group. It was also observed that the seedlings grown from the seeds treated with 800 µL/L SCFJ concentration grew faster than the seeds treated with other concentrations (Fig. 1). The highest scores were 16.3 and 11.6 cm for seedling height and root length, respectively. Once again, the low-est results were observed in the control group (Ta-ble 1).
The highest leaf area (24.75 cm2) were
ob-served at 800 µL/L SCFJ, and can be attributed to higher amounts of cell division and higher meta-bolic activities (Table 1). The highest chlorophyll (chl. a, chl. b, and total chl.) contents were observed in seedlings from seeds that were treated with a concentration of 800 µL/L SCFJ and were 1491.030 µg chl. a, 956.358 µg chl. b, and 1008.720 µg total chl./g fresh tissue (Table 1). The results were higher because the seedlings grew very well from seeds treated with this SCFJ concentra-tion and had higher metabolic activity (Fig. 1).
We observed from these results that SCFJ can be used for rapid seed germination and seedling
growth in rapeseed. Higher results obtained from 800 µl/l SCFJ concentration in all parameters ex-amined could be attributed to increased enzymatic activity required for germination and for growth. In addition, when we evaluated the results of the mea-sured parameters from seeds treated with 1600
µL/L SCFJ concentration, seed germination and seedling growth was considerably suppressed. As a result of this study, 800 µL/L SCFJ concentration resulted in the healthiest seedlings and the most seedling growth.
FIGURE 1
In vitro seed germination from rapeseed (Brassica napus L.) seeds kept in sterile distilled water having
dif-ferent mature squirting cucumber fruit juice (SCFJ) concentrations for 48 h and 72 h and seedling growth 14 days after developed seedlings were transferred to soil. Bar = 1 cm.
TABLE 1
Effects of different concentrations of squirting cucumber (Ecballium elaterium [L.] A. Rich.) fruit juice (SCFJ) on seed germination of Brassica napus L. cv. 'Elvis' at 48 and 72 h and on seedling height, root length, leaf area and on chlorophyll (chl. a, chl. b, and total chl.) contents 14 d after planting the seedlings
SCFJ Cont. (µl/L) Germination (%)1 Day 14 48 h 72 h Seedling height (cm) Root length (cm) Leaf area (cm2) Chl. a content (µg/g fresh tissue) Chl. b content (µg/g fresh tissue) Total chl. content (µg/g fresh tissue) 0 9.60± 0.40c 21.60±1. 07c 12.4±0.90 c 8.4±0.44b 11.20±0.35c 706.212±18.51 d 615.264±20.76 e 608.088±19.34 e 200 12.0 0±1.3 3bc 24.80±1. 37c 13.2±0.42bc 10.3±0.71ab 16.00±1.55b 953.388±37.74c 682.896±34.83d 702.312±22.14d 400 28.8 0±1.3 0a 69.60±1. 51b 14.6±0.87 b 10.1±0.60 ab 16.00±0.58b 1173.786±43.4 6b 792.438±15.23 b 825.864±19.15 b 800 16.0 0±1.1 5b 87.20±1. 36a 16.3±0.95 a 11.6±0.66 a 24.75±1.08a 1491.030±82.7 4a 956.358±34.83 a 1008.720±45.2 5a 1600 5.60± 0.40c 7.20±0.6 1d 15.5±0.70 ab 8.5±0.38b 16.00±1.15b 1118.022±37.1 6b 719.172±19.95 c 758.028±30.25 c
Values indicate the average of three repetitions and “±” indicates standard error value of the average.
The values indicated in each column with a different letter were statistically different at a 0.01 level of significance.
1Seed germination percentage represents the ratio of germinated seeds to all seeds in the set.
DISCUSSION
Bio-stimulants impove quantity and quality of yield by giving resistance to plants against abiotic stress factors with their biologically active sub-stances [10, 11]. It was reported that extracts of some plants such as Ascophyllum nodosum,
Kappa-phycus alvarezii, Ulva lactuca, Ulva rigida, Caulerpa scalpelliformis, Caulerpa sertularioides, Sargassum liebmannii, Sargassum plagiophyllum, Sargassum wightii, Turbinaria conoides, Padina tetrastromatica, Padina gymnospora, Dictyota di-chotama, Rosenvigea intricata and Fucus spiralis stimulated seed germination, plant growth and yield of crops [12, 13-19]. Kocira et al. [20] have re-ported that extract of Ecklonia maxima increased white bean (Phaseolus vulgaris L.) yield. In addi-tion, extracts from Sargassum wightii and Kappa-phycus alvarezii increased yields of bread wheat and soybean, respectively [14,15,16]. Nwachukwu and Umechuruba [21] have reported that leaf ex-tracts from basil (Ocimum basilicum L.), bitter leaf (V. amygdalina), neem (Azadirachta indica A. Juss.), and Eastern North American pawpaw (Asim-ina triloba (L.) Dunal) increased seed germ(Asim-ination and seedling emergence of African yam bean (Sphenostylis stenocarpa (A. Rich.) Harms). Ex-tract from garlic (Allium sativum) has stimulated seed germination of tomato (Lycopersicon esculent Mill.), chili (Capsicum annuum L.), eggplant (Solanum melongena L.) and bread wheat (Triticum aestivum L.) [22-24]. It has been reported that sea-weed extracts were involved in stimulating plant growth and yield [25]. Jannin et al. [26] and Fan et al. [27] have reported that seaweed extract in-creased root and shoot growth, and chlorophyll con-tent. In our case, the highest values regarding seedling height, root length and chlorophyll con-tents were obtained SCFJ at a concentration of 800 µl/L. At concentration of 1600 µl/L, seed germina-tion reduced dramatically with parallel to the report of Turkyilmaz-Unal et al. [28]
Although squirting cucumber is a medicinal plant with antibacterial properties [5], it has been observed that the juice of the mature squirting cu-cumber at different concentrations increased the re-generation capacity of wheat, flax, potato, and rape-seed under in vitro conditions [29]. In a study con-ducted for ending dormancy in potato tubers of cv. "Marabel", tubers at 40 to 60 g in weight were rinsed for 6 h at 180 rpm in bottles containing ma-ture SCFJ at different concentrations (0-control, 200, 400, 800 and 1600 µl/L). Results clearly showed that SCFJ had a significant role in ending dormancy in potato tubers [30]. Moreover, it has been determined that SCFJ stimulated the growth of Agrobacterium tumefaciens and increased the fre-quency of gene transfer in tobacco [31]; therefore, this study was aimed to observe and evaluate the ef-fect of SCFJ for seed germination and seedling growth in rapeseed.
The chlorophyll contents in the leaf have been accepted as a standard for measuring the photosyn-thetic capacity of tissues [32-34] and directly influ-ence photosynthesis [35-37]. In addition, it has been determined that low levels of germination and seedling growth observed in rice seedlings under stress was a result of low starch transportation and
-amylase activity [38]. Hsia [39]reported that sup-pression of growth under stress conditions was caused by either suppressed cell division, sup-pressed cell proliferation, or both. The decrease in the growth rate under stress conditions might result from membrane permeability and stress-induced changes in water intake [40, 41]. In the present study, a higher level of seedling growth at 800 µL/L SCFJ concentration can be attributed to increases in water and other constituent absorption from the soil and thus the acceleration of the mechanism by which carbohydrates permeate the epidermis.
From the results, it could be speculated that SCFJ could simply be used for seed germination and rapid seedling growth in Brassica napus L. The highest results in germination percentage at 72 h, chl. a, chl. b and total chl. contents, plantlet height, root length and leaf area at the end of 14 days were obtained from 800 µl/l SCFJ concentration. Higher results obtained from 800 µl/l SCFJ concentration in all examined parameters could be attributed to increased enzymatic activity required for germina-tion and for growth. However, from the examined parameters, it was seen that 1600 µl/l SCFJ concen-tration inhibited seed germination and seedling growth significantly. In our study, 800 µl/l SCFJ concentration resulted in the most healthiest and well grown seedlings.
REFERENCES
[1] Memisoğlu, M., Toker, G. (2002a) Chemical constituents of Ecballium elaterium (L.) A. Rich. FABAD Journal of Pharmaceutical Sci-ences 27, 19-25.
[2] Memisoğlu, M., Toker, G. (2002b) Biological activities and traditional usage of Ecballium elaterium (L.) A. FABAD Journal of Pharma-ceutical Sciences. 27, 157-164.
[3] Oskay, M., Sarı, D. (2007) Antimicrobial screening of some Turkish medicinal plants. Pharmaceutical Biology. 45, 176-181.
[4] Koca, U., Ozçelik, B., Ozgen, S. (2010) Com-parative in vitro activity of medicinal plants Arnebia densiflora and Ecballium elaterium against isolated strains of Klebsiella pneumo-niae. Turkish Journal of Pharmaceutical Sci-ences. 7, 197-204.
[5] Adwan, G., Salameh, Y., Adwan, K. (2011) Ef-fect of ethanolic extract of Ecballium elaterium against Staphylococcus aureus and Candida albicans. Asian Pacific Journal of Tropical Biomedicine. 1, 456-460.
[6] Sezik, E. (1997) Research on the Turkish medicinal plant Ecballium elaterium. Chem-istry of Natural Compounds 33, 541–542. [7] Saeidnia, S., Gohari, A. (2012) Importance of
Jour-nal of MediciJour-nal Plants Research. 6(14), 2700-2703.
[8] Curtis, O., Shetty, K. (1996) Growth medium effects on vitrification, total phenolics, chloro-phyll, and water content of in vitro propagated oregano clones. Acta Horticulturae. 426, 498-503.
[9] Snedecor, G.W., Cochran, W.G. (1967) Statis-tical Methods. 6th ed. Iowa State University Press, Ames, IA.
[10] Calvo, P., Nelson, L., Kloepper, J.W. (2014) Agricultural uses of plant biostimulants. Plant Soil. 383, 3-41.
[11] Bulgari, R., Cocetta, G., Trivellini, A., Vernieri, P., Ferrante, A. (2015) Biostimulants and crop responses: a review. Biological Agri-culture and HortiAgri-culture. 31(1), 1-17.
[12] Senturk, B., Sivritepe, H.O. (2016) NaCl priming alleviates the inhibiting effect of salinity during seedling growth of peas (Pisum sativum L.). Fresen. Environ. Bull. 25, 4202-4208.
[13] Kavipriya, R., Dhanalakshmi, P.K., Jayashree, S., Thangaraju, N. (2011) Seaweed extract as a biostimulant for legume crop, green gram. Journal of Ecobiotechnology. 3, 16-19.
[14] Kumar, G., Sahoo, D. (2011) Effect of seaweed liquid extract on growth and yield of Triticum aestivumvar.Pusa Gold. Journal of Applied Phycology. 23(2), 251-255.
[15] Latique, S., Chernane, H., Mansori, M., El Kaoua, M. (2013) Seaweed liquid fertilizer ef-fect on physiological and biochemical parame-ters of bean plant (Phaesolus vulgaris variety Paulista) under hydroponic system. Euopean. Scientific Journal. 9(30), 174-191.
[16] Rathore, S.S., Chaudhary, D.R., Boricha, G.N., Ghosh, A., Bhatt, B.P., Zodape, S.T., Patolia J.S. (2009) Effect of seaweed extract on the growth, yield and nutrient uptake of soybean (Glycine max) under rainfed conditions. South African Journal of Botany. 75(2), 351-355. [17] Raverkar, K.P, Pareek, N., Chandra, R.,
Chauhan, S., Zodape, S.Y., Ghosh, A. (2016) Impact of foliar application of seaweed saps on yield, nodulation and nutritional quality in green gram (Vigna radiata L.). Legume Re-search. 39(2), 315-318.
[18] Thırumaran, G., Arumugam, M., Arumugam, R., Anantharaman, P. (2009) Effect of Seaweed Liquid Fertilizer on growth and pigment con-centration of Abelmoschus esculentus medikus. American-Eurasian Journal of Agronomy. 2(2), 57-66.
[19] Zodape, S.T., Mukhopadhyay, S., Eswaran, K., Reddy, M.P., Chikara, J. (2010) Enhanced yield and nutritional quality in green gram (Phaseolus radiata L.) treated with seaweed (Kappaphycus alvarezii) extract. Journal of Scentific and Industrial Research. 69, 468-471.
[20] Kocira, A., Kornas, R., Koc?ra, S. (2013) Ef-fect assessment of Kelpak SL on the bean yield (Phaseolus vulgaris L.). Journal of Central Eu-ropean Agriculture 14, 67-76.
[21] Nwachukwu, E.O., Umechuruba, C.I. (2001) Antifungal activities of some leaf extracts on seed-borne fungi of African yam, bean seed germination and seedling emergence. Journal of Applied Sciences and Environmental Man-agement. 5(1), 29-32.
[22] Arzoo, K., Biswas, S.K. (2013) Effect of plant extracts as seed treatments on growth parame-ters, seedlings mortality and biochemical changes in tomato. International Journal of Bio-resource and Stress Management. 4, 47-53. [23] Islam, M.T., Faruq, A.N. (2012) Effect of some medicinal plant extracts on damping-off dis-ease of winter vegetable. World Applied Sci-ences Journal. 17, 1498–1503.
[24] Perello, A., Gruhlke, M., Slusarenko, A.J. (2013) Effect of garlic extract on seed germina-tion, seedling health, and vigour of pathogen-infested wheat. Journal of Plant Protection Re-search. 53, 317–323.
[25] Hidangmayum, A., Sharma, R. (2017) Effect of different concentration of commercial seaweed liquid extract of Ascophyllum nodosum as a plant bio stimulant on growth, yield and bio-chemical constituents of onion (Allium cepa L.). Journal of Pharmacognosy and Phyto-chemistry. 6(4), 658-663.
[26] Jannin, L., Arkoun, M., Etıenne, P., Laîné, P., Goux, D., Garnıca, M., Fuentes, M., San Francısco, S., Baigorrı, R., Crus, F., Houdusse, F., Garcıa-Mına, J.M., Yvın, J.C., Ourry, A. (2013) Brassica napus growth is promoted by Ascophyllum nodosum (L.) Le Jol. seaweed ex-tract: microarray analysis and physiological characterization of N, C, and S metabolisms. Journal of Plant Growth Regulation. 32, 31-52. [27] Fan, D., Hodges, D.M., Crıtchley, A.T.,
Prıthıvıraj, B. (2013) A commercial extract of brown macroalga (Ascophyllum nodosum) af-fects yield and the nutritional quality of Spinach in vitro. Communications in Soil Sci-ence and Plant Analysis. 44, 1873-1884. [28] Turkyilmaz-Unal, B., Esiz-Dereboylu, A.,
Gu-vensen, A., Tort, N., Ozturk, M. (2018) Effects of the extract from different plant parts of Fer-ula communis spp. Communis on the seed ger-mination and seedling growth of barley and cu-cumber. Fresen. Environ. Bull. 27, 3375-3381. [29] Yildiz, M., Mirici, S., Ozcan, F.S. (2016)
Uti-lization Possibilities from Gamma Radiation, Magnetic Field and Squirting Cucumber (Ecballium elaterium (L.) A. Rich.) Fruit Juice in Increasing Adventitious Shoot Regeneration and Agrobacterium tumefaciens-mediated Gene Transfer Frequencies in Some Field Crops. Research Project No: 103O280, The
Scientific and Technological Research Council of Turkey TUBİTAK. Ankara, Turkey.
[30] Yildiz, M., Beyaz, R., Gursoy, M., Aycan, M., Koc, Y., Kayan, M. (2017) Seed Dormancy. In: Advances in Seed Biology, Jose C. Jimenez-Lopez (Ed.), ISBN 978-953-51-3622-4, In-Tech.
[31] Ozcan, S.F,. Yildiz, M., Ahmed, H.A.A., Aasim, M. (2015) Effects of squirting cucum-ber (Ecballium elaterium) fruit juice on Agrobacterium tumefaciens-mediated transfor-mation of plants. Turkish Journal of Biology. 39, 790-799.
[32] Pal, R., Laloraya, M.M. (1972) Effect of cal-cium levels on chlorophyll synthesis in peanut and linseed plants. Biochemie und Physiologie Pflanezen. 163, 443-449.
[33] Wright, G.C., Nageswara, R.R.C., Farquhar, G.D. (1994) Water use efficiency and carbon isotope discrimination in peanut under water deficit conditions. Crop Science. 34, 92-97. [34] Nageswara, R.R.C., Talwar, H.S., Wright, G.C.
(2001) Rapid assessment of specific leaf area and leaf nitrogen in peanut (Arachis hypogaea L.) using chlorophyll meter. Journal of Agron-omy and Crop Science. 189, 175-182.
[35] Rensburg, L.V., Kruger, G.H.J. (1994) Evalua-tion of components of oxidative stres metabo-lism for use in selection of drought tolerant cul-tivars of Nicotiana tabacum L. Journal of Plant Physiology. 143, 730-737.
[36] Kyparıssıs, A., Manetas, Y. (1995) Summer survival of leaves in a soft-leaved shrub (Phlomis fruticosa L., Labiatae) under Mediter-ranean field conditions: Avoidance of photoin-hibitory damage through decreased chlorophyll contents. Journal of Experimental Botany. 46, 1825-1831.
[37] Jagtap, V., Bhargava, S., Sterb, P., Feıerabend, J. (1988) Comparative effect of water, heat and light stresses on photosynthetic reactions in Sorghum bicolor (L.). Moench. Journal of Ex-perimental Botany. 49, 1715-1721.
[38] Lin, C.C., Kao, C.H. (1995) NaCl stress in rice seedlings: Starch mobilization and the influ-ence of GA3 on seedling growth. Botanical
Bulletin Academia Sinica 36, 169-173.
[39] Hsia, T.C. (1973) Plant responses to water stress. Annual Review of Plant Physiology. 24, 219-270.
[40] Ilan, I. (1971) Evidence for hormonal regula-tion of the selectivity of ion uptake by plant cells. Physiologia Plantarum. 25, 230-233. [41] Karmoker, J.L., Van Stevenınck, F.M. (1979)
The effect of abscisic acid on the uptake and distribution of ions in intact seedlings of Phaseolus vulgaris cv. Redland Pioneer. Physi-ologia Plantarum. 45, 453-459.
Received: 23.09.2019
Accepted: 18.10.2020
CORRESPONDING AUTHOR Ramazan Beyaz
Kırşehir Ahi Evran University, Faculty of Agriculture,
Department of Soil Science and Plant Nutrition, Kırşehir
