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Grain and Oil Yields of Safflower (Carthamus Tinctorius L.) Affected By Water Deficit and Growth Regulators

Grain and Oil Yields of Safflower (Carthamus Tinctorius L.) Affected By Water Deficit and Growth... Agriculture (Poľnohospodárstvo), 67, 2021 (2): 87 − 94 Original paper DOI: 10.2478/agri-2021-0008 GRAIN AND OIL YIELDS OF SAFFLOWER (CARTHAMUS TINCTORIUS L.) AFFECTED BY WATER DEFICIT AND GROWTH REGULATORS ROGHIYEH FARZI-AMINABAD, KAZEM GHASSEMI-GOLEZANI , SAFAR NASRULLAHZADEH University of Tabriz, Tabriz, Iran Farzi-Aminabad, R., Ghassemi-Golezani, K. and Nasrullahzadeh, S. (2021). Grain and oil yields of safflower (Carthamus tinctorius L.) affected by water deficit and growth regulators. Agriculture (Poľnohospodárstvo), 67(2), 87 – 94. In order to evaluate the effects of growth regulators on yield parameters and oil content of safflower (Carthamus tinctorius L.), a field experiment was conducted under different irrigation intervals in 2019. All plots were irrigated regularly until the seedling establishment and thereafter irrigation intervals were applied after 70, 100, 130, and 160 mm evaporation from class A pan, as normal irrigation and mild, moderate, and severe water deficits, respectively. Foliar sprays of water (control), pu- trescine (60 µg/L), and 24-epibrassinolide (25 µg/L) at a rate of 1,000 L/ha were applied slightly before flowering. The results revealed that means of plant biomass, grains per capitol, grains per plant, grain yield, harvest index, oil percentage, and yield were decreased under limited irrigations, but 1,000-seeds weight was only reduced under severe water deficit. However, fo- liar sprays of growth regulators, particularly putrescine, increased grains per plant, grain yield, and harvest index, leading to an improvement in oil yield per unit area under different levels of water supply. These results suggest that foliar application of putrescine is a superior treatment for improving the productivity of safflower plants under normal and stressful conditions. Key words: drought stress, harvest index, epi-brasinollide, plant biomass, putrescine Safflower (Carthamus tinctorius L.) is originated availability is less than evapotranspiration demand. from south Asia and is cultivated in the same regions This can cause a decline in carbon assimilation, that favor the growth and development of wheat and cell growth, and tissue expansion. Many genes, en- barley. It is undoubtedly a crop with great potential zymes, hormones, and metabolites are involved in and compatibility to be grown in a wide range of en- each of these processes (Skirycz & Inze 2010). vironments (Gilbert et al. 2008). Nevertheless, the Putrescine as a polyamine is involved in scav- productivity of this crop could be limited by abiot- enging of the free radicals and regulating osmotic ic stresses such as water deficit. Drought is one of potential under drought stress (Gupta et al. 2013; Li the most detrimental environmental stress for plant et al. 2014). It can stimulate plant growth by act- growth and development (Krouma et al. 2015), that ing as a nitrogen source and influencing cell divi- can affect different physiological and biochemical sion and development (Kandil et al. 2011). There is processes in plants (Mohammadi et al. 2018a). Wa- also a relationship between brassinosteroids (BRs) ter deficit can naturally occur in the field when water concentration and oxidative stress in plants (Bajguz Roghiyeh Farzi-Aminabad, MSc Student, Department of Ecophysiology, Faculty of Agriculture, University of Tabriz, Tabriz, Iran Kazem Ghassemi-Golezani, Prof. (*Corresponding Author), Department of Ecophysiology, Faculty of Agriculture, Universi- ty of Tabriz, Tabriz, Iran. E-mail: golezani@gmail.com Safar Nasrullahzadeh, Assoc. Prof., Department of Ecophysiology, Faculty of Agriculture, University of Tabriz, Tabriz, Iran © 2021 Authors. This is an open access article licensed under the Creative Commons Attribution-NonComercial-NoDerivs License (http://creativecommons.org/licenses/by-nc-nd/4.0/). 87 Agriculture (Poľnohospodárstvo), 67, 2021 (2): 87 − 94 & Hayat 2009). Some reports revealed that exoge- length, spaced 25 cm apart. The seeds were sown th nous application of BRs such as 24-epibrassinolide manually on 14 May 2019 at a depth of 3 – 4 cm, alter the activities of antioxidant enzymes such as with a distance of 10 cm on the rows to achieve catalase, superoxide dismutase, ascorbate perox- a density of 40 plants/m . All plots were irrigated idase, and glutathione peroxidase under stressful after sowing. Subsequent irrigations were carried conditions (Ozdemir et al. 2004). These are a group out according to the treatments up to field capacity of steroidal growth regulators (Bajguz & Piotrows- (FC). Weeds were controlled by hand during plant ka-Niczyporuk 2014) influencing stress tolerance, growth and development. Variants of foliar sprays seed germination, growth, and senescence of plants of growth regulators: water (control), putrescine (Rao et al. 2002). This research was aimed to assess (60 µg/L), and 24-epibrassinolide (25 µg/L) at the impact of exogenously applied putrescine and a solution rate of 1,000 L/ha were applied on plants 24-epibrassinolide on reducing the harmful effects slightly before flowering. At maturity, plants in 1 m of drought stress on field performance of safflower of the middle part of each plot were harvested and plants. the number of grains per capitol, number of grains per plant, 1,000-seeds weight, and grain yield were determined. Aboveground biomass was oven-dried MATERIAL AND METHODS at 75°C for 48 hours and then, plant biomass per unit area was recorded. Harvest index was calculated as: A field experiment was conducted in 2019 at Harvest index = (grain yield/plant biomass) × 100 the Research Farm of the University of Tabriz in Iran, located at 38.05° N, 46.17° E with an altitude A Soxhlet extractor was used to extract oil ac- of 1,360 m above sea level, annual rainfall about cording to the AOCS method (1993). Initially, 263 mm, and mean annual temperature of 13.5°C. 5 grams of grains from each plot were crushed by The annual rainfall during 2017 – 2020 ranged from a milling machine. The filter papers were cut into 200.87 mm up to 380.75 mm. The total rainfall bottom tubes and the shredded samples were poured during experimentation was about 7 mm and the into paper tubes and placed inside the Soxhlet ex- mean temperature ranged 15.7 – 29.8°C. The physi- actor. 200 ml of petroleum ether was poured into cal and chemical properties of the farm soil are pre- a round bottom flask and a few pieces of boiling sented in Table 1. stone were thrown into it. The Soxhlet extractor and The experiment was laid out as a split plot based refrigerant were then installed on the flask. Then on randomized complete block design in three rep- the faucet was turned on to allow water to flow into licates with irrigation intervals (I , I , I , and I : irri- 1 2 3 4 the refrigerant. As soon as the first drop of solvent gation after 70, 100, 130, and 160 mm evaporation was distilled and dripped from the refrigerant, the from class A pan, for normal irrigation and mild, time was recorded. Extraction was continued for 5 moderate and severe water deficits, respectively) in hours. The heat source was then turned off to cool main plots and foliar spray of growth regulators in the system, and the vapor was cooled to a liquid. sub-plots. Each plot consisted of 6 rows with 5 m After that, the flask was removed from the clamp. T a b l e 1 Physical and chemical properties of farm soil Total Organic Total Mineral Depth Electrical Absorbable elements Soil pH neutralizing carbon nitrogen components of [cm] conductivity [mg/L] texture material [%] [%] [%] soil [%] P K Ca Fe Zn Mn Clay Silt Sand Sandy 0 – 30 8.0 2.92 10.5 0.37 0.04 loam 4.9 255 0.76 2.6 0.92 3.34 12 14 74 88 Agriculture (Poľnohospodárstvo), 67, 2021 (2): 87 − 94 The samples were then weighed and oil percentage leaf area, and intensity of photosynthesis. Water defi- and yield were determined. cit decreases the water potential of plants, leading The data were analyzed using the model of to stomata closure and reduction in photosynthesis two-factorial analysis of variance (irrigation in- rate, leaf growth (Ozturk 1999), and plant biomass. terval, growth regulators), by MSTATC software; Reduction in plant biomass resulted in decreasing and the means were compared by Duncan multiple the number of grains per plant, 1,000-seeds weight, range test at P ≤ 0.05. The Excel software was used and consequently, grain yield and harvest index of to draw figures. milk thistle (Ghassemi-Golezani et al. 2017). De- creasing plant biomass by water limitation has also been observed in sesame (Eskandari et al. 2009) RESULTS AND DISCUSSION plants. The increment of plant biomass in putres- cine-treated safflower plants was related to further Analysis of variance (Table 2) showed that wa- expansion and persistence of leaf area, providing ter deficit and growth regulators had significant ef- sufficient photosynthates for plant growth (Emadi et fects on plant biomass, number of grains per capi- al. 2013). Application of brassinolide can also en- tol, number of grains per plant, 1,000-seeds weight, hance plant dry matter through increasing relative grain yield, harvest index, and oil yield. The oil water content, and chlorophyll content in soybean percentage was also significantly affected by water (Zhang et al. 2008). limitation. The interaction of irrigation × growth The number of grains per capitol was decreased regulator was significant for grains per plant, grain under water deficit, with no significant differences yield, harvest index, and oil yield. among limited irrigations (Table 3). The highest The plant biomass was deceased with increasing number of grains per capitol was recorded for pu- irrigation intervals (Table 3). Foliar spray of growth trescine treated plants, with no significant difference regulators increased plant biomass of safflower, between growth regulators (Table 3). Foliar appli- with no significant difference between putrescine cation of brassinolide also increased the number of and 24-epibrassinolide treated plants (Table 3). As grains per spike of wheat under normal and limited determined by Ghassemi-Golezani and Afkhami irrigations (Dehghan et al. 2017). This growth reg- (2018), plant biomass reduction in water stressed ulator increased the number of flowers and grain plants is the consequence of decreasing cell growth, yield in mung bean, due to the prevention of flower T a b l e 2 Analysis of variance (mean squares) of grain yield and yield components in safflower affected by water deficit and growth regulators Grains 1,000 Plant Grains per Grain Harvest Oil Oil Source of variation df per seeds biomass capitol yield index percentage yield plant weight Replication 2 4954.40 91.29 799.36 5.04 30.99 4.090 0.029 2.73 ++ ++ ++ + ++ ++ ++ ++ Irrigation (I) 3 117,307.33 77.69 19,586.49 5.70 29,794.93 453.57 30.20 3,191.49 E 6 1,002.59 15.78 460.47 0.78 322.08 15.76 0.19 19.74 ++ ++ ++ + ++ ++ ns ++ Regulators (R) 2 26,393.44 303.32 11,587.68 4.36 15,479.56 70.14 0.017 1,147.94 ns ns + ns ++ + ns ++ I × R 6 142.10 3.28 364.39 0.90 504.02 9.35 0.033 56.80 E 16 362.65 4.17 112.75 0.80 116.38 2.26 0.033 10.63 CV [%] – 3.02 7.12 6.29 3.23 5.79 2.67 0.68 6.47 E , E – errors for main and sub-plots, respectively; CV – coefficient of variation; df – degrees of freedom a b + ++ ns , and – significant at p ≤ 0.05, p ≤ 0.01 and no significant (F-test), respectively 89 Agriculture (Poľnohospodárstvo), 67, 2021 (2): 87−94 abortion and higher content of phenolic compounds the number of grains per plant due to water deficit (Ananthi et al. 2013). Foliar application of brassi- is also reported in chickpea (Ghassemi-Golezani et nolide increases the transport of nutrients within the al. 2008), sesame (Eskandari et al. 2009), and milk plant (Fuji et al. 1991). Brassinolide is involved in thistle (Ghassemi-Golezani et al. 2017). This reduc- physiological and metabolic processes such as pho- tion may be caused by excessive loss of leaves at tosynthesis, nucleic acid synthesis, proline accumu- reproductive stages as reported in sunflower (Rauf lation, and protein production, as well as in the gene 2008). transcription and translation, leading to an increase Increasing irrigation intervals decreased in protein content, including enzymes (Anjum et al. 1,000-seeds weight, but there was no significant 2011). difference among normal irrigation, mild stress, and The number of grains per plant significantly moderate stress (Table 3). Foliar application of both decreased with increasing irrigation interval, with- growth regulators increased 1,000-seeds weight out significant differences between mild and mod- (Table 3). Increasing grain weight by foliar spray of erate stress. Foliar sprays of growth regulators led growth regulators could be the result of from pro- to a significant increase in the number of grains per longed leaf area duration and, hence, a longer grain plant at all levels of water stress. A significantly filling period. Environmental stresses such as water higher value was recorded in the variant with pu- shortage, especially during grain filling can reduce trescine compared to brassinolide only under nor- photosynthesis and remobilization of stored mate- mal irrigation and severe stress (irrigation interval × rials and hence, grain filling duration (Sadeghipour growth regulator interaction). This superiority was 2008). Water stress generally accelerates leaf senes- more pronounced under normal irrigation (Figure cence and shortens grain filling duration (Davis et 1). The decrement in the number of grains per plant al. 1999). Also, water deficit reduced 1,000-seeds under water limitation was associated with the re- weight and grain yield by shortening the grain fill- duced number of grains per capitol (Table 3). Ghas- ing period of wheat (Gooding et al. 2003). Reduced semi-Golezani and Afkhami (2018) found that the grain filling occurs due to decreased segregation of main reason for a decline in the number of grains per assimilates and activities of enzymes involved in plant under water deficit was the reduction of green sucrose and starch biosynthesis (Srivastava & Su- cover percentage and plant biomass. Reduction in prasanna 2015). Application of brassinosteroids also T a b l e 3 Means of plant biomass, number of grains per capitol and 1,000-seeds weight for different irrigation intervals and foliar spray of growth regulators Treatment Plant biomass [g/m ] Grains per capitol 1,000-seeds weight [g] Irrigation a a ab I 765.1 32.9 27.9 b b a I 655.4 27.8 28.4 c b ab I 615.5 27.4 28.1 d b b I 488.6 26.4 26.6 Foliar spray b b b Water (control) 580.0 23.1 27.08 a a a Putrescine 672.0 32.8 28.25 a a a 24-epibrassinolide 641.5 30.1 27.95 Different letters in each column indicate significant difference at p ≤ 0.05 (Duncan test) I , I , I , I – irrigation after 70, 100, 130 and 160-mm evaporation from class A pan, respectively 1 2 3 4 90 Agriculture (Poľnohospodárstvo), 67, 2021 (2): 87 − 94 increased grain weight and; consequently, eventu- hibit membrane lipid peroxidation and reduce the ally grain yield in chickpea (Shahid et al. 2011). production of free radicals. It also prevents the syn- Emam and Doghezloo (2015), reported that water thesis of ethylene and delays the aging of the plant, stress has reduced grain weight and foliar applica- thereby increasing the effective period of grain fill - tion of brassinosteroids has increased grain weight ing and grain yield (Bregoli et al. 2002). in wheat cultivars under different irrigation variants. Foliar application of the growth regulators had Reduction in grain yield under water stress a significantly positive impact on the harvest index (Figure 2) can be attributed to reduced grains per of safflower under all irrigation intervals. However, capitol (Table 3), grains per plant (Figure 1) and putrescine was more effective than 24-epibrasin- 1,000-seeds weight (Table 3). There is a signifi - olide, particularly under normal irrigation (Figure cant positive correlation of 1,000-seeds weight and 3). This reflects the higher impact of growth regula - grain yield of safflower (Yari & Keshtakar 2016). tors on grain yield (Figure 2) than on plant biomass Water limitation during flowering leads to flower (Table 3). The high harvest index is related to the abortion and poor seed set, which can potentially formation of more grains per plant and allocation reduce grain yield of chickpea per unit area (Fang et of more assimilates to the grains, which is strong- al. 2009). The growth regulators, especially putres- ly supported by a previous report on beans (Mu- cine, enhanced the grain yield of safflower under noz-Pera et al. 2006). Increasing irrigation intervals normal and limited irrigations, with a greater impact caused a decline in bean harvest index, while foliar in normal watering (Figure 2). Zafari et al. (2017), application of 24-epibrasinolide enhanced it (Mo- reported that safflower yield was reduced by water hammadi et al. 2018b). This improvement in har- deficit, but it was improved by 24-epibrassinolide vest index could be resulted from the higher effect treatment through enhancing plant metabolism and of 24-epibrasinolide on grain yield, compared to stress tolerance (Talaat & Shawki 2013). Applica- plant biomass. tion of putrescine on drought-stressed wheat plants The oil percentage of safflower grains was de - increases the number of spikes, spike weight; and creased with decreasing water availability (Figure grain yield (Gupta et al. 2012). Polyamines such as 4). The oil content was mainly controlled by the ge- putrescine have antioxidant properties, which in- netic characteristics of a cultivar, but photosynthetic Figure 1. Means of safflower grains per plant for interaction of Figure 2. Means of grain yield for interaction of irrigation irrigation intervals × growth regulators intervals × growth regulators in safflower – I1, I2, I3 and I4 – irrigation after 70, 100, 130 and 160-mm – I1, I2, I3 and I4 – irrigation after 70, 100, 130 and 160-mm evaporation, respectively, evaporation, respectively, – Different letters indicate a significant difference at p ≤ 0.05 – Different letters indicate a significant difference at p ≤ 0.05 (Duncan test). (Duncan test). 91 Agriculture (Poľnohospodárstvo), 67, 2021 (2): 87 −94 Figure 3. Means of harvest index for interaction of irrigation Figure 5. Means of safflower oil yield for interaction of irrigati - intervals × growth regulators in safflower on intervals × growth regulators – I1, I2, I3 and I4 – irrigation after 70, 100, 130 and 160-mm – I1, I2, I3 and I4 – irrigation after 70, 100, 130 and 160-mm evaporation, respectively, evaporation, respectively, – Different letters indicate a significant difference at p ≤ 0.05 – Different letters indicate a significant difference at p ≤ 0.05 (Duncan test). (Duncan test). products as the source of oil synthesis can be great- oxidation of the oil. Increasing irrigation intervals ly reduced by water deficit . This might be also ex- in maize also decreased the percentage of seed oil plained by decreased availability of carbohydrates (Ghassemi-Golezani et al. 2016). for oil synthesis under drought stress (Ashrafi & A significant reduction in oil yield of safflow - Razmju 2014). Mohammadi et al. (2018a) reported er with increasing irrigation interval was the result that a decline in oil percentage of safflower grains of decrements in grain yield (Figure 2) and oil per- in water deficit conditions is related to the reduc - centage (Figure 4), which was also supported by tion of the seeds’ capacity for oil accumulation and previous reports on this crop (Ghassemi-Golezani & Afkhami 2018). Drought stress severely limits the growth and development of plants by affecting different metabolic processes such as CO assimila- tion and oil and protein synthesis (Nasirkhan et al. 2007). Esmaeilian et al. (2012), reported that water stress at the grain filling stage of sunflower caused a decrease in oil content. There is a strong positive correlation between oil yield and grain yield of saf- flower (Omidy Tabrizi et al. 2000). Foliar spray of putrescine was the superior treat- ment on increasing oil yield of safflower under normal and limited irrigations, followed by 24-epi- brassinolide. This advantage was greater under nor- mal irrigation, compared to other irrigation intervals (Figure 5). This superiority in oil yield was directly Figure 4. Changes in mean oil percentage of safflower seeds related to the positive impacts of 24-epibrassinolide under different irrigation intervals and particularly putrescine on grain yield per unit – I1, I2, I3 and I4 – irrigation after 70, 100, 130 and 160-mm evaporation, respectively, area (Figure 2), since oil percentage was not affect- – Different letters indicate a significant difference at p ≤ 0.05 ed by these growth regulators (Table 2). (Duncan test). 92 Agriculture (Poľnohospodárstvo), 67, 2021 (2): 87 −94 Agriculture, 39, 181 – 188. DOI:10.1071/Ea98134. CONCLUSIONS Dehghan, M., Baluchi, H., Yodavi, A.R. and Safikhani, F. (2017). Effect of foliar application of brassinolide on grain yield Water limitation significantly reduced plant bio- and yield components of bread wheat (Triticum aestivum L.) cv. Sirvan under terminal drought stress conditions. mass, grains per capitol, grains per plant, 1,000-grain Iranian Journal of Crop Sciences, 19, 40 – 56. weight, grain yield, and harvest index of safflower, Emadi, M.S., Hasibi, P. and Azimi, A.R. (2013). Effect of foliar application of putrescine and nutrients on grain yield and which were improved by foliar spray of growth reg- quality of two wheat cultivars. Iranian Journal of Crop ulators. Oil percentage and yield of safflower grains Sciences, 15, 247 – 261. were also decreased with decreasing water supply. Emam, Y. Jahani and Doghezloo, M. (2017). Effect of salicylic acid and brassinosteroid on alleviation of water stress in two Application of putrescine and 24-epibrassinolide wheat cultivars. Journal of Crop Production and Processing, noticeably enhanced grains per plant, grain yield 7(1), 127 – 139. DOI:10.18869/acadpub.jcpp.7.1.127. 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Ozdemir, F., Bor, M., Demiral, T. and Turkan, I. (2004). Effects Brassinolide alleviated the adverse effect of water deficits of 24-epibrassinolide on seed germination, seedling growth, on photosynthesis and the antioxidant of soybean (Glycine lipid peroxidation, proline content and anti-oxidative max L.). Plant Growth Regulation, 56, 257 – 264. DOI: system of rice (Oriza sativa L.) under salinity stress. 10.1007/s10725-008-9305-4. Plant Growth Regulation, 42, 203 – 211. DOI:10.1023/ Received: March 29, 2021 B:GROW.0000026509.25995.13. Accepted: May 25, 2021 Ozturk, A. (1999). The effect of drought on the growth and http://www.deepdyve.com/assets/images/DeepDyve-Logo-lg.png Agriculture de Gruyter

Grain and Oil Yields of Safflower (Carthamus Tinctorius L.) Affected By Water Deficit and Growth Regulators

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Abstract

Agriculture (Poľnohospodárstvo), 67, 2021 (2): 87 − 94 Original paper DOI: 10.2478/agri-2021-0008 GRAIN AND OIL YIELDS OF SAFFLOWER (CARTHAMUS TINCTORIUS L.) AFFECTED BY WATER DEFICIT AND GROWTH REGULATORS ROGHIYEH FARZI-AMINABAD, KAZEM GHASSEMI-GOLEZANI , SAFAR NASRULLAHZADEH University of Tabriz, Tabriz, Iran Farzi-Aminabad, R., Ghassemi-Golezani, K. and Nasrullahzadeh, S. (2021). Grain and oil yields of safflower (Carthamus tinctorius L.) affected by water deficit and growth regulators. Agriculture (Poľnohospodárstvo), 67(2), 87 – 94. In order to evaluate the effects of growth regulators on yield parameters and oil content of safflower (Carthamus tinctorius L.), a field experiment was conducted under different irrigation intervals in 2019. All plots were irrigated regularly until the seedling establishment and thereafter irrigation intervals were applied after 70, 100, 130, and 160 mm evaporation from class A pan, as normal irrigation and mild, moderate, and severe water deficits, respectively. Foliar sprays of water (control), pu- trescine (60 µg/L), and 24-epibrassinolide (25 µg/L) at a rate of 1,000 L/ha were applied slightly before flowering. The results revealed that means of plant biomass, grains per capitol, grains per plant, grain yield, harvest index, oil percentage, and yield were decreased under limited irrigations, but 1,000-seeds weight was only reduced under severe water deficit. However, fo- liar sprays of growth regulators, particularly putrescine, increased grains per plant, grain yield, and harvest index, leading to an improvement in oil yield per unit area under different levels of water supply. These results suggest that foliar application of putrescine is a superior treatment for improving the productivity of safflower plants under normal and stressful conditions. Key words: drought stress, harvest index, epi-brasinollide, plant biomass, putrescine Safflower (Carthamus tinctorius L.) is originated availability is less than evapotranspiration demand. from south Asia and is cultivated in the same regions This can cause a decline in carbon assimilation, that favor the growth and development of wheat and cell growth, and tissue expansion. Many genes, en- barley. It is undoubtedly a crop with great potential zymes, hormones, and metabolites are involved in and compatibility to be grown in a wide range of en- each of these processes (Skirycz & Inze 2010). vironments (Gilbert et al. 2008). Nevertheless, the Putrescine as a polyamine is involved in scav- productivity of this crop could be limited by abiot- enging of the free radicals and regulating osmotic ic stresses such as water deficit. Drought is one of potential under drought stress (Gupta et al. 2013; Li the most detrimental environmental stress for plant et al. 2014). It can stimulate plant growth by act- growth and development (Krouma et al. 2015), that ing as a nitrogen source and influencing cell divi- can affect different physiological and biochemical sion and development (Kandil et al. 2011). There is processes in plants (Mohammadi et al. 2018a). Wa- also a relationship between brassinosteroids (BRs) ter deficit can naturally occur in the field when water concentration and oxidative stress in plants (Bajguz Roghiyeh Farzi-Aminabad, MSc Student, Department of Ecophysiology, Faculty of Agriculture, University of Tabriz, Tabriz, Iran Kazem Ghassemi-Golezani, Prof. (*Corresponding Author), Department of Ecophysiology, Faculty of Agriculture, Universi- ty of Tabriz, Tabriz, Iran. E-mail: golezani@gmail.com Safar Nasrullahzadeh, Assoc. Prof., Department of Ecophysiology, Faculty of Agriculture, University of Tabriz, Tabriz, Iran © 2021 Authors. This is an open access article licensed under the Creative Commons Attribution-NonComercial-NoDerivs License (http://creativecommons.org/licenses/by-nc-nd/4.0/). 87 Agriculture (Poľnohospodárstvo), 67, 2021 (2): 87 − 94 & Hayat 2009). Some reports revealed that exoge- length, spaced 25 cm apart. The seeds were sown th nous application of BRs such as 24-epibrassinolide manually on 14 May 2019 at a depth of 3 – 4 cm, alter the activities of antioxidant enzymes such as with a distance of 10 cm on the rows to achieve catalase, superoxide dismutase, ascorbate perox- a density of 40 plants/m . All plots were irrigated idase, and glutathione peroxidase under stressful after sowing. Subsequent irrigations were carried conditions (Ozdemir et al. 2004). These are a group out according to the treatments up to field capacity of steroidal growth regulators (Bajguz & Piotrows- (FC). Weeds were controlled by hand during plant ka-Niczyporuk 2014) influencing stress tolerance, growth and development. Variants of foliar sprays seed germination, growth, and senescence of plants of growth regulators: water (control), putrescine (Rao et al. 2002). This research was aimed to assess (60 µg/L), and 24-epibrassinolide (25 µg/L) at the impact of exogenously applied putrescine and a solution rate of 1,000 L/ha were applied on plants 24-epibrassinolide on reducing the harmful effects slightly before flowering. At maturity, plants in 1 m of drought stress on field performance of safflower of the middle part of each plot were harvested and plants. the number of grains per capitol, number of grains per plant, 1,000-seeds weight, and grain yield were determined. Aboveground biomass was oven-dried MATERIAL AND METHODS at 75°C for 48 hours and then, plant biomass per unit area was recorded. Harvest index was calculated as: A field experiment was conducted in 2019 at Harvest index = (grain yield/plant biomass) × 100 the Research Farm of the University of Tabriz in Iran, located at 38.05° N, 46.17° E with an altitude A Soxhlet extractor was used to extract oil ac- of 1,360 m above sea level, annual rainfall about cording to the AOCS method (1993). Initially, 263 mm, and mean annual temperature of 13.5°C. 5 grams of grains from each plot were crushed by The annual rainfall during 2017 – 2020 ranged from a milling machine. The filter papers were cut into 200.87 mm up to 380.75 mm. The total rainfall bottom tubes and the shredded samples were poured during experimentation was about 7 mm and the into paper tubes and placed inside the Soxhlet ex- mean temperature ranged 15.7 – 29.8°C. The physi- actor. 200 ml of petroleum ether was poured into cal and chemical properties of the farm soil are pre- a round bottom flask and a few pieces of boiling sented in Table 1. stone were thrown into it. The Soxhlet extractor and The experiment was laid out as a split plot based refrigerant were then installed on the flask. Then on randomized complete block design in three rep- the faucet was turned on to allow water to flow into licates with irrigation intervals (I , I , I , and I : irri- 1 2 3 4 the refrigerant. As soon as the first drop of solvent gation after 70, 100, 130, and 160 mm evaporation was distilled and dripped from the refrigerant, the from class A pan, for normal irrigation and mild, time was recorded. Extraction was continued for 5 moderate and severe water deficits, respectively) in hours. The heat source was then turned off to cool main plots and foliar spray of growth regulators in the system, and the vapor was cooled to a liquid. sub-plots. Each plot consisted of 6 rows with 5 m After that, the flask was removed from the clamp. T a b l e 1 Physical and chemical properties of farm soil Total Organic Total Mineral Depth Electrical Absorbable elements Soil pH neutralizing carbon nitrogen components of [cm] conductivity [mg/L] texture material [%] [%] [%] soil [%] P K Ca Fe Zn Mn Clay Silt Sand Sandy 0 – 30 8.0 2.92 10.5 0.37 0.04 loam 4.9 255 0.76 2.6 0.92 3.34 12 14 74 88 Agriculture (Poľnohospodárstvo), 67, 2021 (2): 87 − 94 The samples were then weighed and oil percentage leaf area, and intensity of photosynthesis. Water defi- and yield were determined. cit decreases the water potential of plants, leading The data were analyzed using the model of to stomata closure and reduction in photosynthesis two-factorial analysis of variance (irrigation in- rate, leaf growth (Ozturk 1999), and plant biomass. terval, growth regulators), by MSTATC software; Reduction in plant biomass resulted in decreasing and the means were compared by Duncan multiple the number of grains per plant, 1,000-seeds weight, range test at P ≤ 0.05. The Excel software was used and consequently, grain yield and harvest index of to draw figures. milk thistle (Ghassemi-Golezani et al. 2017). De- creasing plant biomass by water limitation has also been observed in sesame (Eskandari et al. 2009) RESULTS AND DISCUSSION plants. The increment of plant biomass in putres- cine-treated safflower plants was related to further Analysis of variance (Table 2) showed that wa- expansion and persistence of leaf area, providing ter deficit and growth regulators had significant ef- sufficient photosynthates for plant growth (Emadi et fects on plant biomass, number of grains per capi- al. 2013). Application of brassinolide can also en- tol, number of grains per plant, 1,000-seeds weight, hance plant dry matter through increasing relative grain yield, harvest index, and oil yield. The oil water content, and chlorophyll content in soybean percentage was also significantly affected by water (Zhang et al. 2008). limitation. The interaction of irrigation × growth The number of grains per capitol was decreased regulator was significant for grains per plant, grain under water deficit, with no significant differences yield, harvest index, and oil yield. among limited irrigations (Table 3). The highest The plant biomass was deceased with increasing number of grains per capitol was recorded for pu- irrigation intervals (Table 3). Foliar spray of growth trescine treated plants, with no significant difference regulators increased plant biomass of safflower, between growth regulators (Table 3). Foliar appli- with no significant difference between putrescine cation of brassinolide also increased the number of and 24-epibrassinolide treated plants (Table 3). As grains per spike of wheat under normal and limited determined by Ghassemi-Golezani and Afkhami irrigations (Dehghan et al. 2017). This growth reg- (2018), plant biomass reduction in water stressed ulator increased the number of flowers and grain plants is the consequence of decreasing cell growth, yield in mung bean, due to the prevention of flower T a b l e 2 Analysis of variance (mean squares) of grain yield and yield components in safflower affected by water deficit and growth regulators Grains 1,000 Plant Grains per Grain Harvest Oil Oil Source of variation df per seeds biomass capitol yield index percentage yield plant weight Replication 2 4954.40 91.29 799.36 5.04 30.99 4.090 0.029 2.73 ++ ++ ++ + ++ ++ ++ ++ Irrigation (I) 3 117,307.33 77.69 19,586.49 5.70 29,794.93 453.57 30.20 3,191.49 E 6 1,002.59 15.78 460.47 0.78 322.08 15.76 0.19 19.74 ++ ++ ++ + ++ ++ ns ++ Regulators (R) 2 26,393.44 303.32 11,587.68 4.36 15,479.56 70.14 0.017 1,147.94 ns ns + ns ++ + ns ++ I × R 6 142.10 3.28 364.39 0.90 504.02 9.35 0.033 56.80 E 16 362.65 4.17 112.75 0.80 116.38 2.26 0.033 10.63 CV [%] – 3.02 7.12 6.29 3.23 5.79 2.67 0.68 6.47 E , E – errors for main and sub-plots, respectively; CV – coefficient of variation; df – degrees of freedom a b + ++ ns , and – significant at p ≤ 0.05, p ≤ 0.01 and no significant (F-test), respectively 89 Agriculture (Poľnohospodárstvo), 67, 2021 (2): 87−94 abortion and higher content of phenolic compounds the number of grains per plant due to water deficit (Ananthi et al. 2013). Foliar application of brassi- is also reported in chickpea (Ghassemi-Golezani et nolide increases the transport of nutrients within the al. 2008), sesame (Eskandari et al. 2009), and milk plant (Fuji et al. 1991). Brassinolide is involved in thistle (Ghassemi-Golezani et al. 2017). This reduc- physiological and metabolic processes such as pho- tion may be caused by excessive loss of leaves at tosynthesis, nucleic acid synthesis, proline accumu- reproductive stages as reported in sunflower (Rauf lation, and protein production, as well as in the gene 2008). transcription and translation, leading to an increase Increasing irrigation intervals decreased in protein content, including enzymes (Anjum et al. 1,000-seeds weight, but there was no significant 2011). difference among normal irrigation, mild stress, and The number of grains per plant significantly moderate stress (Table 3). Foliar application of both decreased with increasing irrigation interval, with- growth regulators increased 1,000-seeds weight out significant differences between mild and mod- (Table 3). Increasing grain weight by foliar spray of erate stress. Foliar sprays of growth regulators led growth regulators could be the result of from pro- to a significant increase in the number of grains per longed leaf area duration and, hence, a longer grain plant at all levels of water stress. A significantly filling period. Environmental stresses such as water higher value was recorded in the variant with pu- shortage, especially during grain filling can reduce trescine compared to brassinolide only under nor- photosynthesis and remobilization of stored mate- mal irrigation and severe stress (irrigation interval × rials and hence, grain filling duration (Sadeghipour growth regulator interaction). This superiority was 2008). Water stress generally accelerates leaf senes- more pronounced under normal irrigation (Figure cence and shortens grain filling duration (Davis et 1). The decrement in the number of grains per plant al. 1999). Also, water deficit reduced 1,000-seeds under water limitation was associated with the re- weight and grain yield by shortening the grain fill- duced number of grains per capitol (Table 3). Ghas- ing period of wheat (Gooding et al. 2003). Reduced semi-Golezani and Afkhami (2018) found that the grain filling occurs due to decreased segregation of main reason for a decline in the number of grains per assimilates and activities of enzymes involved in plant under water deficit was the reduction of green sucrose and starch biosynthesis (Srivastava & Su- cover percentage and plant biomass. Reduction in prasanna 2015). Application of brassinosteroids also T a b l e 3 Means of plant biomass, number of grains per capitol and 1,000-seeds weight for different irrigation intervals and foliar spray of growth regulators Treatment Plant biomass [g/m ] Grains per capitol 1,000-seeds weight [g] Irrigation a a ab I 765.1 32.9 27.9 b b a I 655.4 27.8 28.4 c b ab I 615.5 27.4 28.1 d b b I 488.6 26.4 26.6 Foliar spray b b b Water (control) 580.0 23.1 27.08 a a a Putrescine 672.0 32.8 28.25 a a a 24-epibrassinolide 641.5 30.1 27.95 Different letters in each column indicate significant difference at p ≤ 0.05 (Duncan test) I , I , I , I – irrigation after 70, 100, 130 and 160-mm evaporation from class A pan, respectively 1 2 3 4 90 Agriculture (Poľnohospodárstvo), 67, 2021 (2): 87 − 94 increased grain weight and; consequently, eventu- hibit membrane lipid peroxidation and reduce the ally grain yield in chickpea (Shahid et al. 2011). production of free radicals. It also prevents the syn- Emam and Doghezloo (2015), reported that water thesis of ethylene and delays the aging of the plant, stress has reduced grain weight and foliar applica- thereby increasing the effective period of grain fill - tion of brassinosteroids has increased grain weight ing and grain yield (Bregoli et al. 2002). in wheat cultivars under different irrigation variants. Foliar application of the growth regulators had Reduction in grain yield under water stress a significantly positive impact on the harvest index (Figure 2) can be attributed to reduced grains per of safflower under all irrigation intervals. However, capitol (Table 3), grains per plant (Figure 1) and putrescine was more effective than 24-epibrasin- 1,000-seeds weight (Table 3). There is a signifi - olide, particularly under normal irrigation (Figure cant positive correlation of 1,000-seeds weight and 3). This reflects the higher impact of growth regula - grain yield of safflower (Yari & Keshtakar 2016). tors on grain yield (Figure 2) than on plant biomass Water limitation during flowering leads to flower (Table 3). The high harvest index is related to the abortion and poor seed set, which can potentially formation of more grains per plant and allocation reduce grain yield of chickpea per unit area (Fang et of more assimilates to the grains, which is strong- al. 2009). The growth regulators, especially putres- ly supported by a previous report on beans (Mu- cine, enhanced the grain yield of safflower under noz-Pera et al. 2006). Increasing irrigation intervals normal and limited irrigations, with a greater impact caused a decline in bean harvest index, while foliar in normal watering (Figure 2). Zafari et al. (2017), application of 24-epibrasinolide enhanced it (Mo- reported that safflower yield was reduced by water hammadi et al. 2018b). This improvement in har- deficit, but it was improved by 24-epibrassinolide vest index could be resulted from the higher effect treatment through enhancing plant metabolism and of 24-epibrasinolide on grain yield, compared to stress tolerance (Talaat & Shawki 2013). Applica- plant biomass. tion of putrescine on drought-stressed wheat plants The oil percentage of safflower grains was de - increases the number of spikes, spike weight; and creased with decreasing water availability (Figure grain yield (Gupta et al. 2012). Polyamines such as 4). The oil content was mainly controlled by the ge- putrescine have antioxidant properties, which in- netic characteristics of a cultivar, but photosynthetic Figure 1. Means of safflower grains per plant for interaction of Figure 2. Means of grain yield for interaction of irrigation irrigation intervals × growth regulators intervals × growth regulators in safflower – I1, I2, I3 and I4 – irrigation after 70, 100, 130 and 160-mm – I1, I2, I3 and I4 – irrigation after 70, 100, 130 and 160-mm evaporation, respectively, evaporation, respectively, – Different letters indicate a significant difference at p ≤ 0.05 – Different letters indicate a significant difference at p ≤ 0.05 (Duncan test). (Duncan test). 91 Agriculture (Poľnohospodárstvo), 67, 2021 (2): 87 −94 Figure 3. Means of harvest index for interaction of irrigation Figure 5. Means of safflower oil yield for interaction of irrigati - intervals × growth regulators in safflower on intervals × growth regulators – I1, I2, I3 and I4 – irrigation after 70, 100, 130 and 160-mm – I1, I2, I3 and I4 – irrigation after 70, 100, 130 and 160-mm evaporation, respectively, evaporation, respectively, – Different letters indicate a significant difference at p ≤ 0.05 – Different letters indicate a significant difference at p ≤ 0.05 (Duncan test). (Duncan test). products as the source of oil synthesis can be great- oxidation of the oil. Increasing irrigation intervals ly reduced by water deficit . This might be also ex- in maize also decreased the percentage of seed oil plained by decreased availability of carbohydrates (Ghassemi-Golezani et al. 2016). for oil synthesis under drought stress (Ashrafi & A significant reduction in oil yield of safflow - Razmju 2014). Mohammadi et al. (2018a) reported er with increasing irrigation interval was the result that a decline in oil percentage of safflower grains of decrements in grain yield (Figure 2) and oil per- in water deficit conditions is related to the reduc - centage (Figure 4), which was also supported by tion of the seeds’ capacity for oil accumulation and previous reports on this crop (Ghassemi-Golezani & Afkhami 2018). Drought stress severely limits the growth and development of plants by affecting different metabolic processes such as CO assimila- tion and oil and protein synthesis (Nasirkhan et al. 2007). Esmaeilian et al. (2012), reported that water stress at the grain filling stage of sunflower caused a decrease in oil content. There is a strong positive correlation between oil yield and grain yield of saf- flower (Omidy Tabrizi et al. 2000). Foliar spray of putrescine was the superior treat- ment on increasing oil yield of safflower under normal and limited irrigations, followed by 24-epi- brassinolide. This advantage was greater under nor- mal irrigation, compared to other irrigation intervals (Figure 5). This superiority in oil yield was directly Figure 4. Changes in mean oil percentage of safflower seeds related to the positive impacts of 24-epibrassinolide under different irrigation intervals and particularly putrescine on grain yield per unit – I1, I2, I3 and I4 – irrigation after 70, 100, 130 and 160-mm evaporation, respectively, area (Figure 2), since oil percentage was not affect- – Different letters indicate a significant difference at p ≤ 0.05 ed by these growth regulators (Table 2). (Duncan test). 92 Agriculture (Poľnohospodárstvo), 67, 2021 (2): 87 −94 Agriculture, 39, 181 – 188. DOI:10.1071/Ea98134. CONCLUSIONS Dehghan, M., Baluchi, H., Yodavi, A.R. and Safikhani, F. (2017). Effect of foliar application of brassinolide on grain yield Water limitation significantly reduced plant bio- and yield components of bread wheat (Triticum aestivum L.) cv. Sirvan under terminal drought stress conditions. mass, grains per capitol, grains per plant, 1,000-grain Iranian Journal of Crop Sciences, 19, 40 – 56. weight, grain yield, and harvest index of safflower, Emadi, M.S., Hasibi, P. and Azimi, A.R. (2013). Effect of foliar application of putrescine and nutrients on grain yield and which were improved by foliar spray of growth reg- quality of two wheat cultivars. Iranian Journal of Crop ulators. Oil percentage and yield of safflower grains Sciences, 15, 247 – 261. were also decreased with decreasing water supply. Emam, Y. Jahani and Doghezloo, M. (2017). Effect of salicylic acid and brassinosteroid on alleviation of water stress in two Application of putrescine and 24-epibrassinolide wheat cultivars. Journal of Crop Production and Processing, noticeably enhanced grains per plant, grain yield 7(1), 127 – 139. DOI:10.18869/acadpub.jcpp.7.1.127. Eskandari, H., Zehtab-Salmasi, S., Ghassemi-Golezani, K. and harvest index, leading to an improvement in and Gharineh, M.H. (2009). Effects of water limitation on oil yield per unit area under all irrigation intervals, grain and oil yields of sesame cultivars. Journal of Food especially under normal irrigation. Foliar spray of Agriculture and Environmental Science, 7, 339 – 342. Esmaeilian, Y., Sirousmehr, A.R., Asghripour, M.R. and Amiri, putrescine was the superior treatment for improving E. (2012). Comparison of sole and combined nutrient these traits. Therefore, application of 24-epibrass- application on yield and biochemical composition of sunflower under water stress. International Journal of inolide and especially putrescine can considerably Applied Science and Technology, 2, 214 – 220. enhance safflower plant productivity under normal Fang, X., Turner, N.C., Yan, G., Li, F. and Siddique, KH.M. and limited irrigations. (2009). 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Journal

Agriculture – de Gruyter

Published: Jul 1, 2021

Keywords: drought stress; harvest index; epi-brasinollide; plant biomass; putrescine

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