1. Introduction
Iodine is a micronutrient crucial for the proper growth and development of human organisms. Despite wide programmes of the prevention of iodine deficiency diseases, the status of iodine nutrition in some populations still requires effective supplementation [
1,
2]. One of the most popular ways is the usage of iodized table salt. However, the efficiency of that approach can be hardened by major factors, including iodine loss during salt manufacturing and storage [
3], thermal processing [
4], as well as growing awareness of the need to reduce salt consumption due to related health hazards [
5]. Over the years, it has been proposed that the consumption of crop plants with increased content of iodine in edible parts can improve its supply in a daily diet. The in-vivo studies have already confirmed the efficiency of that approach to increase iodine nutrition in human organisms [
6].
For the last decades numerous studies have been conducted evaluating the possibility of increasing iodine content in selected parts of various crop plants; extensive reviews were presented by Medrano-Macias et al. [
7] and Gonzali et al. [
8]. The so-far obtained data indicate that the efficiency of iodine biofortification of crop plants depends on numerous factors such as plant type and species, growth conditions, method of iodine application, and chemical form of that chemical element. A great amount of the studies in the area has been focused on greenhouse cultivation of various plant species, including lettuce [
9,
10], pepper [
11], spinach [
12,
13], and tomato [
14,
15,
16]. Fewer works have studied the efficiency of iodine biofortification when crop plants, such as rice, wheat, corn [
17], pea [
18], carrot [
19], or lettuce [
20], were cultivated in field. The possibility of increasing iodine content in crops grown in field conditions is affected by various factors determining the availability of iodine to plants. Two main approaches can be taken into consideration, namely, soil fertilization and foliar application of iodine.
The main factors affecting the efficiency of soil fertilization with iodine compounds are iodine sorption and leaching within soil as well as plant preference towards particular iodine compound. Soil sorption of iodine depends on its chemical forms and is associated with its fixation with both organic (mainly humic and fulvic acids) and inorganic fractions (ferrous, aluminum, and copper complexes) of the soil [
21]. Some amounts of iodine applied or present in the soil may also be released into the atmosphere due to biological or chemical processes leading to the formation of volatile I
2 or CH
3I [
22,
23,
24]. It is estimated that only up to 10% of the total content of iodine in soil is present in the soil solution, therefore available for plants [
24]. Some works indicate that iodates undergo stronger sorption with soil particles than iodide [
25]. The applicability of foliar spraying for obtaining crop plants with increased content of iodine has been evaluated in a minority of works [
18,
26,
27,
28]. However, it has already been suggested as an efficient method for supplying various forms of that element during cultivation [
20].
Potato is one of the most popular vegetables cultivated in various climatic conditions with the greatest producers including countries from Asia (China, India, Bangladesh and Turkey), Europe (Ukraine, Russian Federation, Poland), as well as United States of America [
29]. It is a major source of carbohydrates (mainly starch), protein, vitamins, and micronutrients in a daily diet of over one billion people around the world [
30]. As its popularity increases in some developing countries, it may be proposed as a convenient crop that may help counteract the deficiency of various micronutrients in the populations. Some reports indicate the possibility of obtaining potato yield with increased iodine content by fertigation in greenhouse cultivation [
31] or applying iodine-enriched nutrient solution [
32]. However, no long-term field studies on the efficiency of iodine biofortification of potato tubers have yet been conducted.
The aim of a three-year field study was to evaluate and compare the efficiency of iodine biofortification of potato by the soil and foliar application of iodine compounds. The study included the description of the influence of various biofortification approaches on yielding and major nutritional quality parameters of potato tubers in order to evaluate the potential applicability of tested methods for the production of biofortified crop.
4. Discussion
The role of iodine in plants has not been widely studied until the current decade. The recent findings have allowed to suggest its potential beneficial role for plant growth and selected physiological processes [
8,
9,
10,
11,
12,
13,
14,
15,
16,
17,
18,
19,
20,
21,
22,
23,
24,
25,
26,
27,
28,
29,
30,
31,
32,
33,
34,
35,
36,
37,
38,
39]. Due to a substantial intensification of the research on iodine in plants it has been revealed that, when applied in properly adjusted doses, iodine compounds are tolerated by plants without decreasing its growth and yielding. However, the effect strongly depends on the cultivated species, method of cultivation as well as iodine application as thoroughly reviewed by Medrano-Macias et al. [
7]. As far as potato cultivation in field is concerned, little information is available in the literature. In the field studies conducted by Mao et al. [
40] soil application of KIO
3 in a dose of 0.59 kg I ha
−1 significantly reduced the tuber yield of potato. Similar observations were noted for potato plants grown in pot experiment and irrigated with iodide and iodate solutions of various concentrations with more detrimental effect exerted by iodide [
31]. On the other hand, the introduction of iodate in a concentration of 5 mg of I dm
−3 had no effect on the tuber yield of potato plants cultivated in hydroponic conditions [
32]. The present three-year study reveals that soil application of KI and foliar application of KIO
3 in doses up to 2.0 kg I ha
−1 did not cause any significant drop in potato yield parameters (evaluated as total yield, percentage share of marketable yield, average weight of a tuber, as well as its dry matter content). That indicates no harmful effect exerted by the tested agronomic approaches on potato plants.
The present study has shown higher efficiency of iodine biofortification of potato tubers after foliar spraying with KIO
3 than soil application with KI. This may be due to numerous factors. When iodine is applied into the soil environment, it undergoes fast and strong sorption, as it is easily bound with soil organic matter, mainly through aromatic rings of humic and fulvic acids [
41]. Additionally iodine may interact with inorganic fraction of the soil including Fe/Al, Cu/Al and Cu/Cr hydroxides as well as Cu(I)-sulphides and Cu(I)-Fe(III) complexes [
42,
43,
44]. The process of iodine desorption is low and dependent on numerous factors including soil pH, Eh, and microbial activity [
41,
42,
43,
44,
45]. The described processes substantially limit the availability of that element for the roots of cultivated plants. Comparative studies have revealed that in soil environment iodide ions are characterized by higher bioavailability for plants than iodate [
9,
10,
11,
12,
13,
14,
15,
16,
17,
18,
19,
20,
21,
22,
23,
24,
25,
26,
27,
28,
29,
30,
31,
32,
33,
34,
35,
36,
37,
38,
39,
40,
41,
42,
43,
44,
45,
46]. Studies conducted by Hong et al. [
25] on three types of soil clearly shown stronger soil sorption of iodate and its lower desorption from these analyzed soils as compared to iodide ions. Furthermore, the difference in iodide and iodate bioavailability was verified by vegetation experiment confirming that exogenous iodate remained in the cultivated soil in a greater amount than iodide [
25]. An additional factor that should be considered is that the plant uptake of iodate is preceded by the process of its chemical reduction by respective iodate or nitrate reductases [
47] or driven by microbial activity [
48].
Foliar application of mineral elements has been proposed as one of the most cost-effective and efficient method of agronomic biofortification of crop plants. The studies on the possibility of improving Zn content in rice conducted within the framework of the HarvestPlus programme showed a higher effect of seed enrichment after spraying of plants as compared to soil application of that element [
49]. The efficiency of foliar application of Fe, Zn, and Se for increasing grain content of these micronutrients was also confirmed for rice [
50]. Foliar spraying with multi-element cocktail containing Se, Zn, and I has substantially increased the content of these elements in the grains of various wheat cultivars grown in various locations [
51]. Foliar spraying of plants with iodine solutions has been less frequently tested in terms of its applicability for iodine biofortification of plants. However, it has been shown to provide beneficial effects even as compared to the introduction of iodine solution the root zone, particularly for leafy vegetables such as lettuce [
16,
17,
18,
19,
20,
21,
22,
23,
24,
25,
26,
52] and alfalfa [
28]. Its applicability for increasing iodine content has already been demonstrated for plum, nectarine, and tomato fruits [
31], carrot roots [
53], and kohlrabi tubers [
54]. What is more, foliar spraying with KIO
3 has been proposed as recommended approach to iodine biofortification of cereals [
17]. The results of the present study revealed that foliar application of KIO
3 turned out more efficient in iodine accumulation in potato tubers as compared to soil application of KI when introduced in the same final iodine dose. In the studies conducted by Caffagni et al. [
31] foliar application of KI was less efficient in terms of iodine biofortification of potato tubers as compared to soil application of KI. However, in that study the highest iodine dose applied foliarly was eight times lower than a dose used for soil fertilization. In the work by Lawson et al. [
52], foliar application of iodine proved less efficient in increasing iodine content in kohlrabi stem tuber as compared to soil application of respective iodine compounds (KI, KIO
3). In that case, the significant factor that may have affected the results could include the morphology of kohlrabi leaves that are characterized by a dense cuticle layer. The presence of such a hydrophobic barrier may have significantly decreased the level of iodine absorption into deeper layers of leaves [
52]. In plants, iodine is transported mainly through xylem [
46] therefore its distribution into the tubers, roots, or fruits is hardened. However, there is growing evidence of phloem mobility of that element [
16,
17]. The results obtained in the present studies, indicating greater efficiency of iodine biofortification of potato through foliar spraying, provide the additional evidence of substantial level of phloem transport of iodine.
Calculations of the average coverage of daily requirements for iodine by the consumption of 100 g of iodine-enriched potatoes substantiate its potential applicability as additional iodine sources in the human diet. It needs to be taken into account that iodine-biofortified vegetables, potato included, are not aimed to be the sole source of that element in the human diet. Therefore, it is even more desirable not to obtain particularly high values of RDA-I coverage in order to avoid the risk of excessive intake of that element. In the studies on iodine biofortification of tomato approximately 36.5% of RDA-I was covered by 100 g of tomato fruits biofortified with KIO
3 [
14]. Studies on foliar application of various forms of I during kohlrabi cultivation led to an increase in iodine content in tubers, allowing approximately 1.5% of RDA-I to be provided by a 100 g portion [
52].
The concentration of starch and soluble sugars in potato tubers is a key parameter regarding its consumption and processing quality. The present study showed that the application of iodine through soil or foliar spraying had relatively small effect on the sugar concentration in potato tubers and the main factor affecting the obtained variation were the weather conditions in individual years of the study. The findings described by various authors do not allow to clearly indicate the influence of exogenous iodine on sugar accumulation in crop plants. Similar observations as those from the present study were found by Smoleń et al. [
27] with field cultivation of carrot. The studies by Kiferle et al. [
15] showed only a small reduction of soluble sugar content in tomato fruits grown in the presence of iodine in the nutrient solution. On the other hand, iodine applied in low-to moderate doses may contribute to a significant increase in sugar accumulation in pepper [
11] and strawberry fruits [
55]. No effect of soil or foliar application of various forms of iodine was noted with respect to the content of soluble sugars in radish [
56].