3.1. Dry matter
The post-harvest dry matter content in tubers of the tested Satina variety, regardless of factors, averaged 152.0 and 146.9 g kg
-1 for Experiment I and Experiment II, respectively (
Table 1 and
Table 2). The varying dry matter content of tubers of edible varieties is reported by Koch et al. [
38], Mystkowska [
39] and Naeem and Caliskan [
40], among others. According to the authors, the dry matter content of tubers depending on the variety and the cultivation technology and can vary within very large limits from 183 g kg
-1 in the study of Mystkowska [
39] to 350 g kg
-1 in the study of Naeem and Caliskan [
40]. In our study, the application of organic matter had a significant effect on the change in dry matter content (
Table 1 and
Table 2).
The highest dry matter content of 154.2 and 151.7 g kg
-1 was obtained after the application of manure and the lowest of 150.0 and 143.4 g kg
-1 on the control object (for Experiment I and II, respectively) (
Table 1 and
Table 2). The positive effect of natural fertilization in the cultivation of edible potato especially in the context of dry matter content has been reported by many researchers [
5,
41,
42]. The authors indicate that the increase in dry matter content after the application of natural fertilizers is the result of a better supply of nutrients to plants and the gradual availability of nutrients during the growing season. This is closely related to the type and rate of application of these fertilizers. In addition, it was observed that reducing mineral fertilization to 50% resulted in a significant decrease in dry matter content by 7.1% (
Table 1). Manolov et al. [
43] and Bărăscu et al. [
44] report that regardless of variety for dry matter content, the most important thing is to maintain the ratio of fertilizer application rates (N:P:K). Any change in the N:P:K proportion results in a reduction in dry matter content [
45,
46,
47,
48]. However, it is important to remember that a key component of potato production systems is nitrogen management [
45,
49]. Kazimierczak et al. [
5] and Lombardo et al. [
25], report that the use of excessively high doses of nitrogen fertilization in conventional agriculture affects excessive vegetative growth of plants which leads to a decrease in the dry matter content of agricultural crops. The negative effect of high doses of nitrogen fertilization on the dry matter content of potato tubers is also reported by Bombik et al. [
50] and Milroy et al. [
51].
In addition, it was observed that the dry matter content was determined by the interaction of the tested organic material and mineral fertilization. In Experiment I at 100% mineral fertilization rate, a significant increase in dry matter content compared to the control was obtained after the application of manure (9.5%) and stubble intercrop (4.5%). On the other hand, when the NPK rate was reduced to 50%, a significant decrease in dry matter content was obtained only after the application of manure (4.1%) (
Table 1).
After inoculation of soil and plants with soil conditioner, regardless of the factors studied, a significant increase in dry matter content was found, on average by 4.3 and 1.4% for Experiment I and II, respectively, compared to facilities where UGmax was not applied (
Table 1 and
Table 2). The use of a soil conditioner in potato production reduces tuber infestation by pathogens, which contributes to improving the quality of the tubers in terms of nutrients [
52,
53,
54].
There was also a positive interaction between the soil conditioner and organic matter and a statistically significant increase in dry matter was obtained only with manure (8.5%) in Experiment I (
Table 1). On the other hand, in Experiment II after the application of stubble intercrop in the form of peas and manure (
Table 2). Simultaneous application of UGmax and mineral fertilization caused an increase in the dry matter content of potato tubers, while a significant increase was obtained on 100% NPK fertilization (in Experiment I) (
Table 1).
There was a statistically significant effect on dry matter content, of the protection applied regardless of the other experimental factors (
Table 2). The application of protection devoid of fungicides and insecticides caused the greatest decrease in dry matter content by an average of 12.9 and 11.1%, respectively (
Table 2). This is confirmed by the study of Zarzecka et al. [
55], who proved that although there was no significant effect of insecticides on dry matter content, there was a tendency for this component to decrease under their influence compared to the control object. Other results were obtained in the study by Sayuk et al. [
56], who proved that in the variants with fungicide application there is an increase in the dry matter content of tubers by 0.1-0.6%.
Taking into account the interaction of all experimental factors, the highest dry matter content was characterized by potatoes grown on manure and soil conditioner. It was noted that in Experiment I the increase in dry matter was most favorable after the application of 100% NPK and in Experiment II with full protection (
Table 1 and
Table 2). Obtaining such an effect may have been due to a faster release of compounds and an increase in plant resistance to disease under the influence of microorganisms contained in UGmax [
17,
18,
19].
3.2. Starch
The most important component of dry matter is starch [
57,
58,
59,
60], as evidenced by the significantly positive correlation coefficients between dry matter and starch in Experiments I and II, equal to r = 0.509 and 0.716 (p≥0.05), respectively (
Table 3). In the studies conducted, the starch content averaged from 100.6 g kg
-1 f. m. for Experiment II to 104.0 g kg
-1 f. m. for Experiment I (
Table 1 and
Table 2). Many authors point to the influence of variety, agrotechnical and storage conditions on the content of this component, the value of which varies between 11-21% [
39,
61,
62,
63,
64,
65,
66]. However, it should be remembered that with proper agrotechnology, varieties with lower starch content can yield starch at a comparable or higher level than lower-yielding varieties, but with higher starch content [
67,
68].
Each of the applied factors in the experiments had a significant effect on the starch content of the tubers of the tested cultivar Satina (Experiments I and II) (
Table 1 and
Table 2). The application of organic material generally had a significant effect on the increase in starch content compared to the control. Only in Experiment II was there no significant effect of applied straw on the content of this parameter. Murawska et al. [
69], Koireng et al. [
70] and Demidenko et al. [
60] showed that potato tubers grown on manure had the highest starch content, as in our experiments. In the study of Murawska et al. [
69], the average starch content was higher by an average of 3.8%, while in our study an increase of as much as 14.1% was observed (
Table 1 and
Table 2).
In the study conducted, reducing the NPK fertilization rate by 50% resulted in a significant decrease of 4.4% in the starch content of potato tubers. The decrease in starch content was almost twice that of dry matter. This is due to the fact that most potato traits are genetically determined and subject to high phenotypic variability [
71].
El-Zehery [
72] reports that limitations on mineral fertilization in potato cultivation should be implemented with the simultaneous application of organic fertilization, which is consistent with the results of our study (
Table 1 and
Table 2). The starch content of the tubers of the tested Satina variety was the highest on both 100 and 50% NPK fertilization with manure (
Table 1). It was noted that the starch content on the objects where 50% mineral fertilization and manure were applied was higher compared to the object where 100% NPK fertilization was applied. El-Zehery [
72] obtained the highest quality of potato tubers in terms of starch after applying organic and biological fertilization with reduced levels of mineral fertilization by 25%. Thus, reducing the dose of mineral fertilization supplemented with organic matter has an effect on limiting the reduction of starch content [
72,
73].
As for dry matter content, the application of UGmax resulted in an increase in starch content in potato tubers. This is due to the fact that starch content in potato tubers is closely correlated with starch content (r = 0.51, r = 0.72,
p≤0.05 for Experiment I and II, respectively) (
Table 3). However, it was noted that the increase was significantly higher for starch, with 12.6 and 5.8% for Experiment I and II, respectively (
Table 1 and
Table 2). El Zehery [
72] using biofertilizer obtained an increase in starch content equal to 14.6%, comparable to our study. The positive effect of using biostimulants on the starch content of potato tubers has been reported by many authors [
10,
39,
74]. Such an effect is due to the microorganisms contained in the soil conditioner [
52,
53,
54]. On the other hand, a study by Maciejewski et al. [
75] showed no effect of biostimulant application on the starch content of potatoes, which may be due to the application of biostimulants only in foliar form.
The application of the fertilizer UGmax in Experiment I resulted in a significant increase in starch content on average from 12.2% for the object with stubble intercrop to 15.9% for the object with manure in Experiment I (
Table 1), and from 1.1% after the withdrawal of fungicides to 7.3% after the withdrawal of herbicides in Experiment II (
Table 2). This is due to an increase in plant resistance to diseases and pathogens under the application of biostimulants [
76,
77].
Each protection reduction applied resulted in a significant reduction in starch content compared to the control, which was 3.4, 13.8 and 7.0% after the withdrawal of herbicides, fungicides and insecticides, respectively (
Table 2). The withdrawal of certain herbicide and fungicide protective treatments in potato cultivation leads to a decrease in the nutritional value of potato tubers. Withdrawal of fungicides increases the risk of plant infection with dangerous pathogens [
78]. On the other hand, the withdrawal of herbicides leads to excessive weed infestation of the crop and an increase in the proportion of fine tubers in the yield and, consequently, a decrease in starch content [
79,
80].
In our study, significantly the highest starch content of 112.7 g kg
-1 f. m. was obtained after simultaneous application of a soil conditioner and full protection (Experiment II) (
Table 2). Zarzecka et al. [
59] showed that the highest starch yield compared to the control object was obtained after application of the biostimulant together with plant protection in the form of herbicide. Baranowska [
74] also obtained a 16.3% increase in starch content with the simultaneous application of a biostimulant and herbicides. Kaliyeva et al. [
81] report that the starch content of tubers is also affected by the use of insecticides in potato cultivation. The authors report that the starch content of potato tubers in the control variant ranged from 10.1 to 10.3%, while in the variant with insecticide protection the content ranged from 15.0 to 16.4%. The resulting increase over the control ranged from 4.9 to 6.1% [
81].
3.3. Total and reducing sugars
In the conducted studies, the content of total and reducing sugars was at the level of 4.62 and 1.52 g kg
-1 f. m. for Experiment I and 4.97 and 2.15 g kg
-1 f. m. for Experiment II, respectively (
Table 4 and
Table 5). For nutritional value, higher sugar contents in tubers are desirable, while specific standards are set for potatoes intended for processing [
28,
62,
82].
In the studies conducted, organic matter had a significant effect on the content of total sugars and reducing sugars in Experiments I and II (
Table 4 and
Table 5). The highest content of total sugars was recorded for the control object, and the lowest for the object with manure (5.22 and 3.56 g kg
-1 f. m. for Experiment I and 5.60 and 4.44 g kg
-1 f. m. for Experiment II, respectively) (
Table 4 and
Table 5). Similar trends occurred for reducing sugars in Experiment II (
Table 5). These contents were 2.38 and 1.79 g kg
-1 f. m. for the control and manure objects, respectively (
Table 5). A different relationship was observed for Experiment I, where the highest content of reducing sugars was found for the control object and the lowest for the object for which straw was used, 1.81 and 1.28 g kg
-1 f. m., respectively.
In our study, reducing mineral fertilization to 50% resulted in a significant decrease in the content of reducing sugars only. The decrease in the content of reducing sugars was 0.54 g kg
-1 f. m. (
Table 4). As reported by Mona et al. [
83], AbdEl-Nabi et al. [
84] and Jatav et al. [
85], the content of reducing sugars was significantly lower on sites where reduced mineral fertilization was applied. Similar results were obtained by El-Ghamriny and Saeed [
86] for other plants. This may be due to a decrease in the intensity of polysaccharide hydrolysis processes, the conversion of organic acids into soluble sugars, and a decrease in the solubilization of insoluble starch under reduced mineral fertilization [
47].
The application of UGmax had a significant effect on reducing the content of total and reducing sugars (
Table 4 and
Table 5). Thus, it is necessary to use a soil conditioner in the cultivation of potatoes for the production of refined products. The average decrease in total sugars for Experiment I and II was 13.4 and 6.7% and reducing sugars was 57.6 and 67.8%, respectively (
Table 4 and
Table 5). This is confirmed by the results obtained by Haider et al. [
87]. On the other hand, in the studies by Maciejewski et al. [
75], Zarzecka and Gugała [
88] and Głosek-Sobieraj et al. [
27], the results on the effect of biostimulants on the content of sugars in potato tubers are not so clear. Maciejewski et al. [
75] after foliar application of biostimulants Asahi SL and Atonik Sl obtained both an increase and decrease in the content of reducing sugars in tubers of different varieties, but these differences were not statistically proven. Trawczynski [
89] using one biostimulant found no significant effect of its action on the content of reducing sugars in potatoes during the years of the study. On the other hand, Zarzecka and Gugala [
88] obtained a significant increase in the content of reducing sugars for the Gawin and Honorata varieties, while the authors found no effect of the tested biostimulants in the Bartek variety. Głosek-Sobieraj et al. [
27] using four different biostimulants and five varieties obtained a decrease in the content of reducing sugars for two varieties. However, Karak et al. [
90] using six different biostimulants observed an unambiguous increase in total and reducing sugars. It should be noted, however, that this study involved only one variety. The results discussed here indicate that the sugar content of potatoes is influenced by many factors: variety, environmental conditions, type of formulation, application rate and frequency of application [
28]. Ezzat et al. [
91] and Arafa and Hussien [
92] indicate that the effect of soil conditioners also depends on the dose and type of mineral fertilization. In addition, foliar application of biostimulants affects plant metabolism and improves plant growth within the leaves, which increases the carbohydrate content of these organs. Due to transport, these compounds enter the tuber from where they are partially released into the rhizosphere. Thus, soil microorganisms release various organic substances and increase the availability of nutrients for potato tubers [
91,
93].
In our study in Experiment I, there was a significant interaction of organic matter and soil fertilizer on the content of reducing sugars (
Table 4). In contrast, in Experiment II, the content of total sugars was significantly affected by the simultaneous application of chemical protection with soil fertilizer and organic matter with UGmax (
Table 5). The highest content of reducing sugars was characterized by tubers from the object where organic matter and soil conditioner were not applied, and the lowest by tubers grown using manure with conditioner (
Table 4). In addition, the highest total sugar content was characterized by tubers without the use of fungicides and fertilizer (
Table 5). Starch, sucrose and simple sugars play an important role in the formation of potato tubers, and the mechanism of starch metabolism is the dominant pathway. Sucrose is the main form of carbohydrate transport and in potato tubers it is subject to degradation to reducing sugars, which are the substrate for starch metabolism [
94]. This is confirmed by the negative significant correlation coefficients, obtained in our study, between starch content and total sugars and reducing sugars of r = - 0.569 and r = - 0.407 (’
p ≤ 0.05) for Experiment I and r = - 0.587 r = - 0.449 (
p ≤ 0.05) for Experiment II, respectively (
Table 3).
The study found that, regardless of the pesticide withdrawn, there was generally a significant increase in total sugars and reducing sugars in tubers by an average of 7.3 and 26.4%, respectively, compared to the content obtained in tubers from a facility where full pesticide protection was applied (
Table 5). The greatest increase in total sugars was obtained by withdrawing fungicides, and in reducing sugars by withdrawing insecticides (
Table 5). Such results were caused by the stress induced by improper plant protection against pathogens. Kumar et al. [
95] note that sugars content is influenced by abiotic factors. In addition, the authors state that each genotype requires proper cultivation technology, and stress, regardless of the type, increases sugar accumulation. On the other hand, Zarzecka et al. [
96] and Baranowska and Mystkowska [
97] report that the content of sugars in potato tubers depends on the type of pesticide used. Biotic stresses caused by improper chemical protection intensify the defense response of plants by producing a greater amount of secondary metabolites, the production of which is associated with a change in the sugar balance by plants [
98,
99].
3.4. Storage
In order to preserve the quality of the tubers and increase their availability throughout the year, long-term storage is necessary. In the tests conducted after 6 months of storage, the dry matter content of the tubers decreased at comparable levels for Experiment I and II, by 3.2 and 3.6%, respectively (
Figure 1 and
Figure 2). A decrease in starch content, whose content is closely related to dry matter content, was also obtained in tubers after long-term storage [
100] (
Figure 3 and
Figure 4). This is confirmed by the highly significant correlation coefficients between dry matter and starch content obtained in our study, amounting to r = 0.667 (
p ≤ 0.05) for Experiment I and r = 0.884 (
p ≤ 0.05) for Experiment II, respectively (
Table 3). It should be noted that the decrease in starch content was almost twice as high in Experiment I compared to Experiment II (
Table 3 and
Table 4). Pandey et al. [
101] and Siddiqui et al. [
102] report that the starch content of potatoes decreases during storage due to the conversion of starch to sugar and its use in respiration. Ozturk and Polat [
3] report a decrease and increase in dry matter and starch content in potato tubers after storage (6 months). The authors, storing 7 varieties under controlled conditions, obtained a decrease in dry matter content of 1.5% on average and an increase in starch content of 3% on average. At the same time, in the case of two varieties they recorded an increase in dry matter content by an average of 8.2% while starch content increased by as much as 17.4%. The authors clearly indicate that potato genotype has the greatest impact on losses in dry matter and starch content. Potato varieties differ in the thickness of the periderm and the amount of deposited suberin, which is a natural barrier to water transport and so different varieties carry out vital processes with different intensity under the same conditions. On the other hand, Sahin et al. [
103] and Pobereżny and Wszelaczyńska [
104] point out that losses of dry matter and starch content are highly dependent on storage time. The authors report that extending the storage period increases losses.
In addition, it was shown that the effect of factors applied during the potato growing season on the dry matter and starch content of tubers after storage was the same as after harvest (
Table 1 and
Table 2,
Figure 1,
Figure 2,
Figure 3 and
Figure 4).
After long-term storage, there was an increase in total sugars and reducing sugars in the tubers. For Experiment I, the increase in total sugars was 8.2% and reducing sugars was 27.6% (
Figure 5 and
Figure 6). In contrast, for Experiment II, an increase of 11.8% in total sugars and 8.6% in reducing sugars was achieved (
Figure 7 and
Figure 8). According to many authors, the accumulation of sugars during long-term storage is mainly due to genetic conditions [
100,
105,
106], so the storage period should be determined taking into account the varietal characteristics of the potato [
107]. A similar view is presented by Alamar et al. [
34] and Wszelaczyńska et al. [
108]. In the study of Wszelaczyńska et al. [
108], a higher increase in sugar content (54.3%) was obtained after 6 months of storage for the Denar variety compared to the Gardena variety (43.6%). Stress factors such as moisture deficiency or temperature changes during storage are also important determinants of sugar content in tubers [
66]. As indicated by Amjad et al. [
105] and Zhang and Zhen-Xiang [
106], low temperatures of 2-4 ℃ can contribute to the accumulation of reducing sugars due to so-called cold-induced sweetening. The sweetening process is a natural process that occurs as a result of tuber aging. It is irreversible and involves cellular breakdown. After cellular breakdown, structural and non-structural carbohydrates are depolymerized by hydrolytic enzymes [
34]. Therefore, maintaining appropriate storage conditions, including temperature, can contribute to reducing weight loss and sweetening of potatoes.
Considering the field factors applied during potato cultivation, it was found that in Experiment II, the effect of these factors on the content of total sugars and reducing sugars in tubers after storage was the same as after harvest (
Table 5,
Figure 7 and
Figure 8). In contrast, in Experiment I, it was shown that the field factors had the same effect only on the content of total sugars after storage (
Table 4,
Figure 5).
The highest increase in total sugars and reducing sugars in tubers after storage in Experiment I was obtained in potatoes after cultivation on manure and straw along with soil conditioner (
Figure 5 and
Figure 6). In contrast, reducing NPK fertilization to 50% resulted in the highest increase in reducing sugars (
Figure 8).
In Experiment II, the withdrawal of insecticides and fungicides resulted in the largest increase in total sugars and, at the same time, the smallest increase in reducing sugars (
Figure 7 and
Figure 8). In contrast, the application of UGmax similarly to that in Experiment I caused an increase in the content of reducing sugars (by 24.4% on average). It should be noted, however, that after potato cultivation without the application of soil conditioner, a slight decrease in the content of reducing sugars was obtained (-0.8%) after storage (
Figure 6).