Impaginato 49 Adv. Hort. Sci., 2019 33(1): 49-56 DOI: 10.13128/ahs-22863 Ameliorate the cadmium toxicity in Solanum tuberosum L. plants with selenium and silicon application A.O.S. Dorneles 1 (*), A. Soares Pereira 2, G. Possebom 3, C. Peligrinotti Tarouco 4, L.V. Rossato 4, L. Almeri Tabaldi 4 1 Federal University of Pelotas, Postgraduate Program Plant Physiology, 96010-900 Pelotas, Rio Grande do Sul, Brazil. 2 Federal University of Pelotas, Postgraduate Program Family Farm Production Systems, 96010-900 Pelotas, Rio Grande do Sul, Brazil. 3 Federal University of Santa Maria, Postgraduate Program Agricultural Engineering, 97105-900 Santa Maria, Rio Grande do Sul, Brazil. 4 Federal University of Santa Maria, Biology Department, 97105-900 Santa Maria, Rio Grande do Sul, Brazil. Key words: elements beneficial, potato, toxic metals, toxicity. Abstract: The present study aimed to prove the efficiency of Se or Si as reliev- ers of the Cd toxicity in potato plants. Solanum tuberosum plants, Asterix genotype, from in vitro propagation were placed in pots with sand and irrigat- ed with complete nutrient solution for 60 days under six treatments: T1: Control (nutrient solution); T2: 2.5 µM Se; T3: 2.5 mM Si; T4: 50 µM Cd; T5: 50 µM Cd + 2.5 µM Se; T6: 50µM Cd + 2.5 mM Si. The treatments were arranged in completely randomized design, with four replicates for each treatment and six plants per replicate. The plants were collected at 30 and 60 days after application of the treatments. Cadmium was highly toxic in all parameters (dry and fresh weight, plant height, leaf number, leaf area, root and photo- synthetic parameters), in both assessments. However, Se and Si were effec- tive in mitigating Cd toxicity in all parameters, although Si has been shown to be more efficient than Se in dry weight and plant height parameters. Thus, from data obtained in this study, it is clear that the beneficial elements tested have power to ameliorate Cd toxicity. 1. Introduction Plants differ in their ability to absorb, accumulate and tolerate heavy metals, including cadmium (Cd), and toxic levels of heavy metals affect a variety of plant processes (Gupta et al., 2013). Cadmium is one of most toxic heavy metals, having a high mobility in environment (Tang et al., 2015), being absorbed by roots and transported to shoot of many plant species (Shi et al., 2005). Although Cd has no known biological function in plants (Pence et al., 2000; Pereira et al., 2016), it can be easily absorbed and transported by xylem (Lux et al., 2011), since it has an electronic con- (*) Corresponding author: athos_odin@hotmail.com Citation: DORNELES A.O.S., SOARES PEREIRA A., POSSE- BOM G., PELIGRINOTTI TAROUCO C., ROSSATO L.V., ALMERI TABALDI L., 2019 - Ameliorate the cadmium toxicity in Solanum tuberosum L. plants with selenium and silicon application. - Adv. Hort. Sci., 33(1):49-56 Copyright: © 2019 Dorneles A.O.S., Soares Pereira A., Possebom G., Peligrinotti Tarouco C., Rossato L.V., Almeri Tabaldi L. This is an open access, peer reviewed article published by Firenze University Press (http://www.fupress.net/index.php/ahs/) and distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Data Availability Statement: All relevant data are within the paper and its Supporting Information files. Competing Interests: The authors declare no competing interests. Received for publication 9 March 2018 Accepted for publication 4 October 2018 AHS Advances in Horticultural Science Adv. Hort. Sci., 2019 33(1): 49-56 50 figuration and state of zinc-like valence (Nan et al., 2002). This heavy metal is toxic even at low concen- trations, inducing stress responses in plants from 5- 10 μg g-1 soil (While and Brown, 2010). Several stud- ies have reported significant reductions in biomass accumulation in plants exposed to Cd (Farooq et al., 2013; Said et al., 2014; Vaculík et al., 2015), and this inhibition in biomass production can occur in short time of exposure to this heavy metal (Han et al., 2015). In contrast to others toxic heavy metals, Cd present in soil is easily absorbed by plant roots, par- ticularly in acidic soils (Guimarães et al., 2008). The plant development stage and time of exposure to heavy metal affect the absorption and distribution of Cd in different parts of plant (Gonçalves et al., 2009 a). Thus, it is necessary to develop strategies that result in lesser absorption of these toxic elements present in soil by plants, optimizing the use of natural resources and production of safe food, especially when it comes to a plant used in food such as pota- toes. Potato (Solanum tuberosum L.) is one of main foods for mankind, consumed by more than one bil- lion people worldwide, due to its composition, gas- tronomic and technological versatility, as well as the low market price of tubers (Coelho et al., 1999; Dorneles et al., 2016). According to Birch et al. (2012), potatoes are third most important crop in world, behind only rice and wheat. The potato is sus- ceptible to Cd, and this sensitivity can be accentuat- ed by level of Cd in soil, time of exposure and cultivar (Gonçalves et al., 2009 b). One of options sought to solve this problem with Cd in plant growth is use of beneficial elements, which when used in low concentrations can alleviate the damaging Cd effects. In this sense, selenium (Se) and silicon (Si) are recognized as beneficial elements for growth of some plants, and can increase the tol- erance of plants to abiotic stresses. These elements are recognized as being capable of mitigating metal toxicity in plants (Wu et al., 2017). However, it is nec- essary to evaluate the potential use of these mitigat- ing elements in the presence of Cd, since some stud- ies show that these may not be effective to reduce the toxicity of this element in some species (Khattab, 2004; Liu et al., 2013). However, Si has been shown to be effective in alleviating Al toxicity in potato plants (Dorneles et al., 2016) and Cd in species such as peanuts (Shi et al., 2010), Chinese cabbage (Song et al., 2009), rice (Tripathi et al., 2013), maize (Lukacová et al., 2013), wheat (Khan et al., 2015) and sunflower (Said et al., 2014). Selenium, however, has stress-alleviating properties that are more focused on biochemical mechanisms activation (Kumar et al., 2012; Feng et al., 2013; Tamaoki and Maruyama- Nakashita, 2017). However, it already has great potential in use to relieve stresses by metals (He et al., 2004; Pezzarossa et al., 2012). However, this ele- ment, to date, has not been tested as potential to relieve the stress caused by Cd toxicity in potato plants. Thus, the aim of this work was to test the pos- sibility of using Se or Si as reliever of Cd toxicity in potato plants. 2. Materials and Methods Plants of Solanum tuberosum L., Asterix genotype, were used for the experiment, which were propagat- ed in vitro from nodal segments for 25 days in MS culture medium (Murashige and Skoog, 1962). After this period, plants were transferred to plas- tic vessels containing 1 plant each and 5 kg of sand, being watered daily with complete nutritive solution. This solution had the following composition (mg L-1): 85.31 N; 7.54 P; 11.54 S; 97.64 Ca; 23.68 Mg; 104.75 K; 176.76 Cl; 0.27 de B; 0.05 Mo; 0.01 Ni; 0.13 Zn; 0.03 Cu; 0.11 Mn and 2.68 Fe (FeSO4/Na-EDTA). After five-day of acclimation period, treatments were applied, which consisted of following combinations: Treatment 1: Complete nutritional solution and absence of Cd, Se and Si; Treatment 2: Complete nutritional solution + 2.5 μM Se; Treatment 3: Complete nutritional solution + 2.5 mM Si; Treatment 4 : C o m p l e t e n u t r i t i o n a l s o l u t i o n + 5 0 μ M C d ; Treatment 5: Complete nutritional solution + 50 μM Cd + 2.5 μM Se; Treatment 6: Complete nutritional solution + 50 μM Cd + 2.5 mM Si. These solutions were applied daily maintaining 80% of the vessel capacity, which was monitored by daily determining the weight of the vessel. The treatments were arranged in a completely randomized design, with four replicates for each treatment and six plants per replicate. The pH of the solutions was adjusted daily (4.5±0.1). Two collects were performed, the first collect being at 30 days of exposure to treatments and the second at 60 days. In both collections, the following evaluations were carried out: - Photosynthetic parameters were evaluated in fourth fully expanded leaf of four plants per repli- cate: the photosynthetic rate (A - μmol CO2 m-2 s-1), the stomatal conductance of water vapors (Gs - mol H2O m-2 s-1), internal CO2 concentration (Ci - μmol Dorneles et al. - Selenium and silicon ameliorate the cadmium toxicity in potato 51 m-2), water use efficiency (WUE - mol CO2 mol H2O- 1), and the transpiration rate (Tr mmol - mmol H2O m-2 s-1) obtained by the ratio between the amount of CO2 fixed by photosynthesis and the internal CO2 concentration. The evaluations were carried out in p e r i o d b e t w e e n 8 a n d 1 1 h w i t h u s e o f I R G A portable meter, brand LI-COR, model LI-6400XT. - Growth parameters: fresh and dry biomass, height, leaf area, number of leaves, and morphological parameters of the root system (length, diameter, volume and number of root branches), according to methodology described by Dorneles et al. (2016). For statistical analysis of data, it was verified the normality of error distribution through Anderson- Darling test and homogeneity of error variances through the Bartlett test (Estatcamp, 2012) for all variables of experiment. The averages were submit- ted to analysis of variance and compared by the Scott-Knott test, with 5% significance, using Sisvar (Ferreira, 2011). The graphic program used was SigmaPlot 12.5. 3. Results For experimental conditions tested, cadmium (Cd) presented great toxicity to potato plants both at 30 and 60 days of growth. This is evidenced by averages of fresh weight accumulation obtained by plants exposed to Cd, which did not increase after 30 days of cultivation, remaining unchanged up to 60 days (Fig. 1) both being statistically smaller than in the control treatment. Besides, in general, when applied in isolation, Si and Se did not promote greater accu- mulation of fresh and dry weight in comparison to control plants (Figs. 1, 2). However, both elements tested proved to be effective in mitigating Cd toxicity from first collect, inducing a higher accumulation of fresh weight in both tissues, compared to the treat- ment where only Cd was applied. The enhancing effects of Se and Si are more evident in shoot, where plants exposed to Cd combined with Se or Si main- tained some growth in second collect (Fig. 1 A). The roots of potato plants (Fig. 1 B) also show a signifi- cant improvement in presence of both beneficial ele- ments. This probably contributed to growth of shoot, considering that roots are the first tissue to come into contact with Cd. For dry weight, potato plants showed similar behavior to that observed for fresh weight in both tissues (Fig. 2). Cadmium showed to be toxic to both tissues for dry weight, inhibiting this accumulation until last collect. However, Se and Si induced biomass accumulation similar to control plants, even when applied together with Cd. Silicon induced greater accumulation of shoot dry weight when compared to Se (Fig. 2 A). In roots, Cd prevented the dry weight increase, while plants treated with Se and Si present- ed increase in root dry weight in second collect (Fig. 2 B), that is, Se and Si alleviated the toxic effects of cadmium. In addition to affecting the production of fresh and dry biomass, Cd inhibited the production and expansion of leaves, as well as the growth in height of potato plants. However, plants that received Si and Se together with Cd obtained higher means even in comparison to control plants (Figs. 3 A, B). In addi- tion, the number of leaves and height of plants exposed to Cd were higher when applied Si, in com- parison to plants treated with Se as amendment. Both Se and Si were effective in reducing the toxicity of cadmium from first collect, inducing higher height, number of leaves and leaf area than plants with only Fig. 1 - Effect of selenium (2.5 μM) or silicon (2.5 mM) on shoot (A) and roots fresh weight (B) of potato plants grown in presence of cadmium (50 μM) at 30 and 60 days after application of the treatments. Different lowercase letters indicate significant differences between treatments in same collect. Different uppercase letters indicate signifi- cant differences between collects for same treatment. Adv. Hort. Sci., 2019 33(1): 49-56 52 Cd in nutrient solution. There was no statistical difference between treat- ments for root length and root diameter at 30 days, but plants treated with Cd presented root length 45% lower than the control (Table 1). At 60 days, plants exposed to Cd showed root length 16% and root Different lowercase letters indicate significant differences between treatments in same collect. Different upper case letters indicate significant differences between collects for same treatment. Fig. 2 - Effect of selenium (2.5 μM) or silicon (2.5 mM) on shoot (A) and roots dry weight (B) of potato plants grown in cadmium presence (50 μM) at 30 and 60 days after appli- cation of the treatments. Different lowercase letters indi- cate significant differences between treatments in same collect. Different uppercase letters indicate significant differences between collects for same treatment. Fig. 3 - Effect of selenium (2.5 μM) or silicon (2.5 mM) on leaves number (A), plants height (B) and leaf area (C) of potato plants grown in cadmium presence (50 μM) at 30 and 60 days after application of the treatments. Different lower- case letters indicate significant differences between treatments in same collect. Different upper case letters indicate significant differences between collects for same treatment. Table 1 - Length, diameter, volume and number of branch roots of Solanum tuberosum plants grown in presence of selenium (2.5 μM), silicon (2.5 mM) and cadmium (50 μM) at 30 and 60 days after application of the treatments Collect Treatments Root length (cm) Root diameter (mm) Root volume (cm3) Branch number 30 days Control 798±0.24 Aa 0.35±0.02 Ba 1.64±0.49 Aa 715±68 Aa Se 776±18.3 Aa 0.38±0.04 Ba 0.91±0.25 Ab 394±18 Bb Si 743±9.24 Aa 0.35±0.02 Ba 0.84±0.19 Ab 401±20 Bb Cd 438±3.76 Ba 0.34±0.02 Aa 0.37±0.07 Ac 201±13 Ad Se + Cd 650±4.62 Aa 0.36±0.00 Ba 0.65±0.05 Ab 323±35 Bc Si + Cd 711±3.43 Aa 0.41±0.03 Aa 0.83±0.08 Ab 318±38 Bc 60 days Control 998±0.57 Aa 0.45±0.02 Aa 1.83±0.21 Aa 790±48 Aa Se 884±21.6 Aa 0.45±0.01 Aa 0.85±0.03 Ab 794±58 Aa Si 862±25.2 Aa 0.45±0.02 Aa 0.76±0.08 Ab 790±62 Aa Cd 839±10.11 Ab 0.34±0.02 Ab 0.34±0.02 Ac 253±24 Ac Se + Cd 795±5.50 Aa 0.46±0.00 Aa 0.75±0.04 Ab 423±21 Ab Si + Cd 791±6.11 Aa 0.44±0.02 Aa 0.85±0.07 Ab 452±22 Ab Dorneles et al. - Selenium and silicon ameliorate the cadmium toxicity in potato 53 diameter 25% lower, compared to the control. On the other hand, all treatments presented lower root volume compared to control, but this effect was more significant in the treatment containing only Cd in the growth medium, both at 30 and 60 days of cul- tivation. This same behavior was observed for the number of branches at 30 days. At 60 days, branch numbers was lower in the treatments containing Cd, Se + Cd and Si + Cd, and this effect was more signifi- cant in the treatment containing only Cd in the growth medium. Cadmium showed to be highly toxic, reducing volume of roots and ramifications to less than half of values presented by control plants. However, Si and Se presented an amendment effect on all root variables, mainly for total length and diameter, both at 30 and 60 days of cultivation. In these variables, plants exposed to Cd treated with both Si and Se were statistically the same as control plants. Plants exposed to Cd had photosynthetic rate (A) reduced by 37% compared to control plants at first collect, and 42% at second collect (Table 2). Besides, Cd caused significant reductions in stomatal conduc- tance (GS), internal CO2 concentration (Ci) and tran- spiratory rate (Trmmol) in both collects. The water use efficiency (WUE) in both collects was higher only for the treatment with Se + Cd. However, in both col- lects Si and Se were effective in mitigation of Cd toxi- city for all variables except for WUE (Table 2). 4. Discussion and Conclusions In the present study, Cd promoted a significant reduction in biomass production in potato plants (Fig. 1). The reduction in biomass accumulation caused by Cd may be due, in part, to its effect on the inhibition of nutrient uptake by roots (Cao et al., 2014; Li et al., 2016). The effects of Cd on absorption of nutrients may be due to damage caused by this element in the roots. Since root tissues are first to come into contact with the Cd present in solution, they are also the most affected. Several studies have already reported negative effects of Cd on cellular level of root tissues (Benavides et al., 2005; Lux et al., 2011; Martinka et al., 2014). The data presented in this work show the signifi- cant reduction in root parameters caused by Cd. This behavior may be due to effect of Cd on degradation of membranes and nucleotides of root cells (Han et al., 2015). This degradation of membranes and organelles can be explained by the increase in con- centration of reactive oxygen species (ROS) induced by Cd (Chou et al., 2012; Farooq et al., 2013; Said et al., 2014; Han et al., 2015). It is possible that this effect of Cd on increase of ROS, reported in these studies, may inhibit cell division of roots (Said et al., 2014) In addition, the Cd has affinity for phosphates and some amino acids components of enzymes and Table 2 - Effect of selenium (2.5 μM) or silicon (2.5 mM) on photosynthetic rate (A-μmol CO2 m-2 s-1), stomatal conductance (GS - mol H2O m-2 s-1), internal CO2 concentration (Ci - μmol m-2 s-1), transpiration rate (Trmmol - H2O mmol m-2 s-1),and water use effi- ciency (WUE - CO2 mol H2O mol-1) of Solanum tuberosum plants grown in cadmium presence (50 μM) at 30 and 60 days after application of the treatments Collect Treatment A GS Ci Trmmol WUE 30 days Control 9.47±0.13 Ba 0.20±0.07 Aa 288±0.48 Aa 9.47±0.13 Ba 192±4.59 Bb Se 8.39±0.32 Bb 0.16±0.04 Aa 295±11.8 Ba 8.39±0.32 Bb 178±11.8 Bb Si 8.92±0.31 Bb 0.04±0.00 Ac 272±2.44 Bb 8.92±0.31 Bb 200±20.3 Bb Cd 5.92±0.27 Bc 0.10±0.00 Ab 257±8.54 Ac 5.92±0.27 Bc 228±8.63 Ab Se + Cd 8.73±0.07 Bb 0.06±0.03 Ac 297±5.57 Aa 8.73±0.07 Bb 308±3.05 Aa Si + Cd 8.26±0.03 Bb 0.10±0.00 Ab 255±9.29 Ac 8.26±0.03 Bb 236±12.6 Ab 60 days Control 12.5±0.14 Aa 0.20±0.03 Aa 289±0.24 Aa 12.5±0.14 Aa 205±0.25 Ab Se 10.3±0.32 Ab 0.14±0.01 Aa 268±14.6 Ab 10.3±0.32 Ab 215±0.24 Ab Si 10.1±0.06 Ab 0.06±0.00 Ab 298±5.53 Aa 10.1±0.06 Ab 236±0.02 Ab Cd 7.41±0.31 Ad 0.04±0.00 Bc 269±8.13 Ab 7.41±0.31 Ad 220±0.26 Ab Se + Cd 9.08±0.72 Ac 0.08±0.00 Ab 257±8.54 Bc 9.08±0.72 Ac 298±0.14 Aa Si + Cd 8.98±0.61 Ac 0.10±0.00 Ab 255±9.29 Ac 8.98±0.61 Ac 245±0.22 Ab Different lowercase letters indicate significant differences between treatments in same collect. Different upper case letters indicate significant differences between collects for same treatment. 54 Adv. Hort. Sci., 2019 33(1): 49-56 proteins, which leads, in addition to damage to mem- branes, genetic damage and can disrupt oxidative phosphorylation in exposed tissues (Hasanuzzaman and Fujita, 2012; Nahar et al., 2016). The present study also shows the effect of Cd under photosynthetic parameters, in which there was a significant reduction in plants exposed to this heavy metal. This reduction in photosynthetic para- meters may be a consequence of the degradation of chlorophylls caused by Cd (López-Millán et al., 2009). Furthermore, Cd causes disorder in arrangement of grana and thylakoids, which limits the efficiency of most photosynthetic parameters (Han et al., 2015; Bayçu et al., 2016). The potato plants used in present study, treated with Si or Se as amendments of Cd toxicity, showed higher biomass production than plants exposed to only Cd. In addition, Se and Si significantly reduced the damage caused by Cd in photosynthetic parame- ters. These results of biomass and photosynthetic rate are related, since the beneficial effects of Si and Si on the Cd toxicity in photosynthetic parameters are expressed, to a greater or lesser degree, in higher production of biomass by plants. Each element used as amendment in this work (Se or Si) act in different ways in the plant. Silicon have been proven to be effective in easing stress by Cd in different species such as cucumber (Feng et al., 2010), corn (Malčovská et al., 2014; Vaculík et al., 2015) and cotton (Farooq et al., 2016). This effect of Si may be due to its deposition on cell wall which increases its plasticity and elasticity (Vaculík et al., 2009). The Si deposited on cell wall of the plants may increase Cd retention in apoplast (Lukačová et al., 2013; Vaculík et al., 2015), which may reduce the availability and translocation of Cd. Due to these cell wall modulations, Si can inhibit Cd uptake (Liu et al., 2013). These effects of Si can explain the more evi- dent amelioration of Cd in shoot than in roots for dry weight in potato plants (Fig. 2). In addition, silicon has photoprotective properties when deposited on leaves (Tripathi et al., 2017), which may explain the mitigation of Cd toxicity in gas exchange parameters. There is a chance that Se also has the property of forming complexes with toxic or heavy metals, but were only found complexes of Se-mercury (Hg) in plants (Said et al., 2014). However, there are reports that selenium reduces Cd accumulation in tissues (Lactuca sativa L.) (He et al., 2004). While Si is recog- nized for increasing the enzymatic antioxidants activi- ty (Debona et al., 2017), Se has the property of induc- ing resistance by activating routes of synthesis of hormones linked to stress response and antioxidant activities (Freeman et al., 2010; Feng et al., 2013; Tamaoki and Maruyama-Nakashita, 2017). Thus, the mechanism of mitigation of Se may be more directly related to antioxidant system. Many studies have reported this effect of Se under plant antioxidant enzymes, as well as its effect on direct removal of ROS (Cartes et al., 2010; Zembala et al., 2010). In pre- sent study, the Cd toxicity was reduced in most of evaluated parameters, mainly under photosynthetic parameters. The possible effect of Se under enzyme activity may help to explain its most evident effect under these parameters. Selenium was more effec- tive in reducing the effects of Cd on photosynthetic parameters than for other parameters evaluated. 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