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Response of potato (Solanum tuberosum L.) cultivars to drought stress under in vitro and field conditions

Abstract

Background

Potato (Solanum tuberosum L.), the world’s third most important crop, is frequently thought to be sensitive to moderately sensitive to drought, and yield has fallen considerably over consecutive stress periods. Drought produces a wide range of responses in potato, from physiological alterations to variations in growth rates and yield. Knowledge about these responses is essential for getting a full understanding of drought-tolerance mechanism in potato plants which will help in the identification of drought-tolerant cultivars.

Results

A set of 21 commercial potato cultivars representing the genetic diversity in the Middle East countries market were screened for drought tolerance by measuring morpho-physiological traits and tuber production under in vitro and field trials. Cultivars were exposed to drought stress ranging from no drought to 0.1, 0.2 and 0.3 mol L−1 sorbitol in in vitro-based screening and 60, 40 and 20% soil moisture content in field-based screening. Drought stress adversely affected plant growth, yield and cultivars differed for their responses. Shoots and roots fresh weights, root length, surface area of root, no. of roots, no. of leaves, leaf area, plant water content %, K+ content, under in vitro drought treatments and shoots fresh and dry weights, no. of tubers and tuber yield under field drought treatments were examined and all decreased due to drought. The stress tolerance index decreased with increasing drought in examined cultivars; nevertheless, it revealed a degree of tolerance in some of them. Grouping cultivars by cluster analysis for response to drought resulted in: (i) a tolerant group of five cultivars, (ii) a moderately tolerant group of 11 cultivars, and (iii) a sensitive group of five cultivars. Furthermore, stress-related genes, i.e., DRO, ERECTA, ERF, DREB and StMYB were up-regulated in the five cultivars of the tolerant group. Likewise, the stomatal conductance and transpiration explained high correlation with the tuber yield in this group of cultivars.

Conclusion

The diversity in germplasm indicated that potato cultivars can be developed for production under certain degrees of drought. Some cultivars are good candidates to be included in drought-tolerant breeding programs and recommended for cultivation in drought-stricken regions.

Graphical Abstract

Introduction

Plants are subjected to a wide range of stressors as a result of their environment. Abiotic stress is one of the most serious and prevalent agricultural problems, resulting in considerable crop yield losses and jeopardizing long-term crop production [1,2,3,4]. One of the primary problems for the coming decade will be to reduce the impact of abiotic stress on crop development, with a focus on maintaining agricultural production rates in the face of reduced water supply or drought [5].

Potato is one of the world’s most important food crops, alongside wheat, rice and maize. Cultivated potatoes (Solanum tuberosum L.) are sensitive to moderately sensitive to drought, depending on the criteria used for classification [6, 7]. This sensitivity is related to the growth stage of the plant, being more sensitive to drought in the early growth and tuberization periods [8, 9]. Drought stress occurs when soil moisture content is low, relative humidity is low and temperature is high. If drought continues, plants will dry up and production will be adversely affected. In potatoes, drought stress delays emergence, slows plant development, and reduces plant mass weight [10,11,12] as well as dramatically reducing tuber number, size and yield [13,14,15,16].

Under open field conditions, environmental factors may vary from season to season, and a genotype that is successful at one season may fail in another season, although no extensive evaluation of the varieties has been reported [17, 18]. The identification and development of potato stress tolerant cultivars are currently needed as climate change is associated with an increase in global temperature and a decrease in precipitation [19, 20]. A solution to this issue may be the cultivation of cultivars that can withstand abiotic stress while retaining high productivity. The most promising solution for the drought problem is to develop drought-tolerant crops, although in the past this has not been a high priority [8]. Variations in flowering ability, leaf characteristics, plant maturity, and tuber production have all been documented in potatoes [1, 6, 8]. Growing asexual propagating crops on a wide scale creates possibilities for different genes and, as a consequence, selection for desired characteristics [2, 6, 7, 13]. Drought tolerance in plants can be improved through traditional breeding or genetic manipulation techniques. Traditional breeding for drought tolerance has been problematic, since it appears that drought tolerance is a complex trait [21, 22]. However, progress has been made in identifying candidate genes in recent years. The identification of genes involved in drought tolerance opens up new possibilities for identifying tolerant germplasm. Genes associated with drought tolerance, such as deeper rooting (DRO) in potato, LRR receptor-like serine/threonine-protein kinase ERECTA (ERECTA), ethylene response factor (ERF), dehydration responsive element binding (DREB) and Solanum tuberosum MYB (StMYB) have been identified [23,24,25,26,27,28,29].

Potato seed tubers are largely imported from the Netherlands, Ireland, and the United Kingdom into several Middle East countries, including Egypt. This is practiced every summer season to avoid the buildup of viral infection caused by high temperature and high population of insects that spread potato viruses. Many of these countries are located in arid or semi-arid environments, therefore evaluating existing potato cultivars for drought tolerance can assist in sustaining the potato production. Several researchers have suggested in vitro-based screening for potato germplasm with drought tolerance [7, 20, 30, 31] but field-based screening is still required. Although field-based screening for drought-tolerant variations is limited and time demanding, the current study’s main goal is to analyze 21 potato cultivars for drought tolerance in vitro and in the field. It is therefore very important to discuss the genotypic variability of drought tolerance in cultivated potatoes with regard to morpho-physiological parameters, molecular traits and tuber yield under in vitro, and field conditions. The detailed objectives of this study are to understand (1) the extent of genetic variability for drought tolerance among potato cultivars using in vitro studies; (2) the stability of drought tolerance of potato cultivars under field conditions; (3) the regulation of some drought-related genes and transcription factors that contribute to plant response to drought, and (4) the correlation between physiological traits and tuber production under drought stress.

Materials and methods

The present study was carried out at the experimental farm of the Departments of Horticulture and Plant Pathology, Faculty of Agriculture, Minia University, El-Minia, Egypt, laboratory of the Department of Applied Biotechnology, University of Technology and Applied Sciences-Sur, The Sultanate of Oman, and the laboratory of Plant Breeding, Faculty of Agriculture, Iwate University, Morioka, Iwate, Japan.

Plant materials

Twenty-one potato cultivars (Solanum tuberosum L.), Agata, Almond, Anya, Atlantic, Burren, Cara, Champion, Desiree, Diamond, Gala, Gazella, Kennebec, King Edward, Lady Balfour, Lady Rosetta, Marble, Marfona, Maritiema, Mizen, Russet Burbank and Spunta were used for in vitro and field drought response studies. Tuber seeds of cultivars were obtained from Agricultural research center (ARC), Egypt. These cultivars were chosen as they have a wide range of morphology and are widely grown in Middle East countries, including Egypt. The list of cultivars with place of origin, year, maturation, skin color, flesh color and tuber shape is shown in Table 1.

Table 1 List of potato cultivars, their place of origin, year, maturation, skin and flesh color and tuber shape

In vitro experiment

Plant micropropagation

This experiment was carried out at the Applied Biotechnology Department of the University of Technology and Applied Sciences in Sur, Oman. A tissue culture technique was implemented for rapid propagation of the previously mentioned potato cultivars. To collect the explants, sprouted potato tubers were planted in greenhouse in 15 cm pots of ProMix with basic soil composition. In a sequential procedure, shoot tips were cut and washed roughly. Then shoot tips were surface sterilized in a solution of 0.5% (v:v) sodium hypochlorite for 5 min and they were rinsed thoroughly with sterile distilled water. Subsequently, the surface sterilized shoot tips were cultured on agar-solidified (8 g L−1) and sucrose (30 g L−1) Murashige and Skoog (MS) basic medium [32]. The pH was adjusted to 5.7 ± 0.1 before the addition of Agar and subsequent autoclaving at 121 °C and 15 psi for 20 min. Tissue excision, implantation and transfer procedure were performed under sterile conditions. The cultured shoot tips were incubated for four weeks in the incubation room at 16 h photoperiod, 25 ± 2 °C and white cool fluorescent bulbs providing approximately 90 µmol m−2 s−1 PPFD. Subcultures were done using shoot tip.

In vitro drought treatments

By using the plantlets of the same age, individual stem nodes were cultured in tubes containing 8 mL of MS growth medium, 30 g L−1 sucrose and 8 g L−1 agar supplemented with/without sorbitol of 0.0, 0.1, 0.2 or 0.3 mol L−1 at pH 5.7. The plantlets were allowed to grow for 30 days to test the cultivars drought tolerance or sensitivity. The experimental design was a factorial experiment with three replications in a split block design. Twenty plantlets were evaluated per cultivar per each of the four drought treatments and the experiment was repeated at approximately monthly intervals three times. All subcultures were maintained under 16 h photoperiod, 25 ± 2 °C and white cool fluorescent bulbs providing approximately 90 µmol m−2 s−1 PPFD.

In vitro propagation measurements

When subcultures were 30 days old (without subculturing) and fully grown with stout stems and broad leaves in the control treatment (i.e., MS medium without sorbitol), data were recorded for various morphological and physiological characteristics associated with drought stress treatments. Plantlets were removed from the tubes, and their fresh weight of roots and shoots, and number of leaves/plantlet were measured. The millimeter graph paper was used to estimate leaf area/plantlet. Furthermore root traits, i.e., total root length, surface area of root and number of roots were analyzed by WinRhizo Basic 2009 software (Regent Instruments Canada, Inc.). Meanwhile, plantlets water content (PWC%) was measured according to the previous study [33]. Ions of potassium (K+) content were determined based on the described method [34].

Field experiment

Drought treatments under field conditions

The experiment was conducted at the experimental farms and labs of the Departments of Horticulture and Plant Pathology, Faculty of Agriculture, Minia University, El-Minia, Egypt. Potato seed tubers of the 21 cultivars were planted on September 10 and 15 during the 2016 and 2017 seasons, respectively. Sowing was performed on one side of the row in all plots, each plot consisted of five rows (0.70 m in width and 3.0 m in length) and the planting distance within the row was 20 cm interval. The area of each experimental plot was 10.5 m2 to be considered (1/400 of feddan). Over the two seasons, physical and chemical analyses of soil samples from a depth of 0.0 to 30 cm were performed, and the average findings are shown in Additional file 1: Table S1. Three different drought regimes (60%, 40% and 20% soil moisture content) were examined compared to control (no drought). The drip irrigation system was patched up, and irrigation was undertaken on a daily basis to keep the three drought regimes in place. Soil water content was monitored daily by time domain transmission (Sidney, BC, Canada). Ten plants per plot per cultivar were evaluated for each of the drought regimes and the applied experimental design was randomized complete block design (RCBD) in a split plot with three replications which contained four levels of drought stress and 21 potato cultivars. The main plots concerned the four stress treatments, while the 21 potato cultivars were randomly distributed in the subplots. All other agricultural practices such as fertilization and control of pests and diseases were performed as recommended for the commercial production [35].

Field measurements

At 100–110 days after planting, tubers were harvested and the following horticultural traits were characterized: number of tubers/plant, weight of tubers (g/plant), shoot fresh and dry weights (g).

Drought tolerance analysis

Classification of potato cultivars as a drought tolerant or sensitive was performed using both in vitro and field measurements. The stress tolerance index (STI) for each cultivar was estimated according to the method [36]. STI was calculated as the ratio of the trait performance at 0.1, 0.2, or 0.3 mol L−1 to the trait performance at 0.0 mol L−1 sorbitol for in vitro experiment or at 60%, 40% and 20% soil moisture content to the trait performance at no drought regime for field experiment as described in the following formula:

$${\text{STI}} = \frac{{T_{{\text{s}}} }}{{T_{{\text{p}}} }},$$

where Ts is the trait of cultivar under stress conditions and Tp is the trait of cultivar under normal conditions.

Glasshouse experiments

Expression profile of drought-related genes

The experiment was carried out at the laboratory of Plant Breeding, Faculty of Agriculture, Iwate University, Morioka, Iwate, Japan. Potato seed tubers of examined cultivars were planted pots until harvesting the roots for comparative genes expression experiment. All tubers were planted in commercial soil (Gattirikun N-120; Tokita Seed Co., Saitama, Japan) for two months in controlled climate chambers (Koito, Tokyo, Japan) at 25°/15 °C (day/night) temperatures. The number of plants was adjusted to 1 plant per pot (25 cm up diameter—22.5 cm base diameter—13.5 cm deep), with 10 pots for each cultivar. All other agricultural practices such as irrigation, fertilization and control of pests and diseases were performed as recommended for the commercial production [35]. 45-day-old plants were subjected to drought stress, with no water for 2 weeks. Roots were collected from normal irrigated plants (control) and drought exposed plants after 2, 4, 6, 8 and 10 days. Soil water content was monitored daily by time domain transmission (Sidney, BC, Canada). Poly (A)+ RNA was isolated from the roots using a micro-FastTrack 2.0 mRNA isolation kit (Invitrogen, San Diego, CA, USA) as instructed by the manufacturer. cDNAs were synthesized from 1 μg of mRNA in a total volume of 20 μL containing 1 μL of oligo (dT) primer (0.5 μg μL−1), 4 μL of first-strand buffer (5× concentrated), 2 μL of dithiothreitol (100 mM), 2 μL of dNTPs (10 mM) and 0.2 μL Superscript II (300 unit). The reaction was carried out at 30 °C for 10 min, 42 °C for 50 min, and 70 °C for 10 min. For this study, five distinct drought-related genes; deeper rooting (DRO) in potato, LRR receptor-like serine/threonine-protein kinase ERECTA (ERECTA), ethylene response factor (ERF), dehydration responsive element binding (DREB) and Solanum tuberosum MYB (StMYB) were selected. To evaluate the differential expression of the chosen genes in roots of potato cultivars with different tolerance reaction, RT-PCR was carried out using gene-specific primers (Additional file 1: Table S2). Conditions for the thermal cycling were 94 °C for 2 min; 28 cycles of 94 °C for 20 s, 55 °C for 20 s, and 72 °C for 40 s; and finally, 72 °C for 2 min. The reproducibility of the results was confirmed using samples from three independently grown plants.

Physiological measurements

The experiment was carried out at the laboratory of Plant Breeding, Faculty of Agriculture, Iwate University, Morioka, Iwate, Japan. Potato seed tubers were planted in pots (25 cm up diameter—22.5 cm base diameter—13.5 cm deep) in a controlled chamber at 25°/15 °C (day/night) temperatures. Drought stress (20% soil moisture content) was applied compared to control (no drought). Soil water content was monitored daily by time domain transmission (Sidney, BC, Canada). Ten plants per cultivar per treatment were evaluated and physiological characteristics were examined 60 days after planting. A portable photosynthesis system (LI6400 TX model, LI-COR, Lincoln, NE, USA) was used to determine the water stomatal conductance at night (8:00 pm). Total daily transpiration was measured by using a transparent plastic to cover the soil surface of the pots to avoid loss of evaporation. Total daily transpiration was measured in a period of 24 h by weight difference as previously defined [37] and 100–110 days after planting, tubers were weighed to have tubers fresh weight/plant and examine the correlation between the physiological traits and tuber yield. All experiments and strategy used to identify the drought-tolerant cultivars are shown in Fig. 1.

Fig. 1
figure 1

Schematic diagram of the experiments conducted under current research to identify drought-tolerant potato genotype

Statistical analyses

The experimental design was a 21 × 4 (cultivars × drought treatments) in a factorial experiment with three replications in a split block design for in vitro-based screening. Randomized complete block design (RCBD) in a split plot with three replications was used for field-based screening and all obtained data were subjected to the analysis of variance using the combined data from the two seasons. Data obtained from this study were subjected to analysis using SAS, version 9.3 (Cary, NC). Differences among potato cultivars were tested by an analysis of variance (ANOVA), and mean significant differences were tested by the Least Significant Difference (LSD) test at the 0.05 level of significance. The interplay of the characteristics evaluated in vitro and in field conditions, as well as their contributions to heatmaps, was investigated using multivariate data. The traits of the 21 potato cultivars were recorded in the dataset under the studied drought stress levels or regimes. Meanwhile, Clustvis was used to determine the correlation coefficients between each trait.

Results

Several researchers advocated in vitro screening for drought-tolerant potato germplasm; however, few studies examined the relationship between in vitro and field (in vivo) screening. This study looked at the differences in morphological traits and tuber yield production in 21 potato cultivars under in vitro and field-based complementary drought assessment. Furthermore, a series of molecular and physiological experiments were carried out on potato plants grown in a glasshouse and subjected to drought stress. In the meantime, these cultivars have been chosen for this research as they exhibit a high degree of morphology diversity and many of them are also widely cultivated in Egyptian farms and other Middle East regions. Figure 1 shows a schematic diagram of the tests that were conducted.

In vitro experiment

Effects of drought on plantlets grown in vitro

The plantlets development of 21 different potato cultivars was studied in vitro under three different levels of drought stress. Sorbitol was utilized at three doses to alleviate drought: 0.1, 0.2, and 0.3 mol L−1. The water potentials (MPa) of these concentrations of cultural media were − 1.11, − 1.51, and − 1.84, respectively. The drought treatments were compared to 0.0 mol L−1 sorbitol (no-drought), which yielded − 0.79 MPa.

Potato plantlets that were successfully grown in vitro onto MS medium showed significant differences after 30 days of culture. In general, shoots fresh weight (ranging from 0.21 to 1.51 g) and roots fresh weight (ranging from 0.05 to 1.29 g) greatly varied among the examined cultivars under the drought treatments (Table 2). Over the drought treatments, there were significant differences among cultivars for average shoots fresh weight and Burren cv. and Almond cv. gave the heaviest weight. With respect to roots fresh weight, most of the 21 cultivars tested showed sensitive reaction to drought stress treatments as the roots fresh weight significantly decreased however, five cultivars showed different degree of tolerance reaction and could develop roots in response to drought stress levels. Two of them namely, Diamond cv. and Russet Burbank cv., respectively gave higher roots fresh weight (1.11 and 0.84 g, respectively) under drought stress treatment (0.3 mol L−1 sorbitol) than the other cultivars. In Agata cv., Atlantic cv., and Kennebec cv. nevertheless, the roots fresh weight decreased with a rise in stress levels, but the reduction was non-significant under the low (0.1 mol L−1 sorbitol) and intermediate (0.2 mol L−1 sorbitol) drought stress treatments. On the other hand, Anya cv. was the most significantly affected cultivar and gave the lowest roots fresh weigh (0.05 g) at the highest stress level of drought (Table 2).

Table 2 Shoots fresh weight, roots fresh weight, total root length, surface area of root and no. of roots of stem node derived in vitro plantlets of 21 potato (Solanum tuberosum L.) cultivars

Furthermore, other root characteristics such as total root length, root surface area and no. of roots under drought stress were examined using WinRhizo (Table 2). A reduction of 35 to 60% in root length, 42 to 69% in mean root surface area and 37 to 62% in no. of roots due to drought were observed. Drought stress significantly affected most cultivars to grow long roots; however, Agata, Diamond, Kennebec, Russet Burbank and Atlantic cvs, respectively, showed substantial differences with drought treatment (Table 2). Under drought stress (0.3 mol L−1 of sorbitol), Agata cv., had longer root length (99.35 mm) than the other cultivars however, Kennebec cv. gave longer roots (64.51 mm) under low drought treatment (0.1 mol L−1 of sorbitol) compared with 56.24 mm under no drought treatment (0.0 mol L−1 of sorbitol). On the other hand, Almond cv., Marfona cv., Desiree cv. and Gala cv. were significantly sensitive and produced short roots (5.21, 11.23, 11.24 and 11.36 mm, respectively) under drought stress (0.3 mol L−1 of sorbitol). Meanwhile, roots covered narrow surface area with drought stress levels in most cultivars under this investigation. Overall, mean surface area of roots among the cultivars ranged from 0.1 to 1.14 cm. While the root surface area decreased significantly in most of the cultivars, there was a non-significant reduction in Diamond cv., Agata cv., Russet Burbank cv., Atlantic cv. and Kennebec cv. (Table 2). There was also a decrease in the no. of roots from 9.54 to 1.0 due to drought stress. Most of cultivars had significant low no. of roots under stress while the reduction was not significant in Diamond, Atlantic and Kennebec cultivars. Interestingly, Diamond cv. Atlantic cv. and Kennebec cv. developed more roots (9.65, 6.48 and 6.32, respectively) under low drought treatment (0.1 mol L−1 of sorbitol) than the control treatment (8.67, 6.21 and 6.14, respectively). On the other hand, Lady Balfour cv., Maritiema cv., Gala cv. and Marfona cv. had the lowest average roots number in response to drought treatments.

Likewise, cultivars plantlets showed a decrease in the average no. of leaves (from 10.62 to 4.55) and leaf area (from 962.64 to 30.08) due to drought stress treatments; 0.1, 0.2 and 0.3 mol L−1 of sorbitol as shown in Table 3. The number of leaves significantly declined in all cultivars with great reduction in Gala cv., (1.5) although plantlets of Diamond cv. yielded a large number of leaves/plantlet (6.67) under 0.3 mol L−1 of sorbitol. At the same time, Anya, Gala and Almond cultivars had the lowest average leaf area, while Burren and Atlantic cultivars had the highest average leaf area over the drought treatments.

Table 3 Number of leaves/plantlet, leaf area/plantlet, plantlet water content (PWC%), and potassium content (K+) of stem node derived in vitro plantlets of 21 potato (Solanum tuberosum L.) cultivars

Effects of drought on plantlets water content %

The percentage of plantlet water content (PWC%) is one of the stress physiological indices primarily of drought. Variations in PWC % based on cultivar and degree of drought were observed, and it ranged from 20% under high stress level to 95% under no-drought treatment with an approximately fivefold difference (Table 3). PWC % of potato cultivars was low with increasing concentrations of sorbitol compared to control (no drought) but the decrease in PWC was more pronounced in Gala, Anya and Lady Balfour cultivars, respectively, with significant differences. On the other hand, data showed that Diamond, and Agata cultivars did not differ significantly in PWC % with increasing levels of drought stress when they gave the highest PWC % (87% and 84%, respectively) under the highest level of drought.

Effects of drought on the potassium content

Potassium (K+) is a vital mineral that influences plant growth and metabolism through a variety of physiological and biochemical mechanisms. Root growth and K+ diffusion rates towards the roots were both confined during drought stress, restricting K+ acquisition. Plant tolerance to drought stress, as well as K+ absorption, may be further negatively affected as a result of the lower K+ concentrations. Plant drought tolerance depends on having sufficient plant K+ levels. In Table 3, treatments with drought resulted in a significant reduction in K+ content of the roots of all the 21 cultivars examined, although the K+ decrease was greater for Mizen, Marfona, Maritiema, Lady Rosetta, Marble, King Edward and Almond cultivars. Overall, K+ content among the cultivars ranged from 16 to 85 mmol kg−1 FW. Potassium was significantly reduced by drought stress in roots started from the treatment with the lowest concentration of 0.1 mol L−1 sorbitol. In Russet Burbank, Kennebec, Diamond, Atlantic and Agata cultivars, respectively, a reduction in K+ was observed in the 0.1 mol L−1 sorbitol treatment, although the decrease was not statistically significant. At high stress level, Diamond cv. showed the highest K+ content (38 mmol kg−1 FW) whereas Mizen cv. had the lowest content (16 mmol kg−1 FW).

Field experiment

Effects of drought on potato plants and yield under field conditions

The drought tolerance assessment in potatoes is usually based on a comparison of plant growth and tuber yield responses under non-drought and drought-growing conditions, whether in the lab or in the field. In addition to the in vitro experiment, this field experiment was assigned to determine the drought tolerance variations and stability of the same 21 potato cultivars. The screening was conducted under three drought regimes (60%, 40% and 20% soil moisture content) in comparison to the control (no drought). Field screening was carried out in two successive seasons, and the drought treatments used for the experiments found mild, moderate, and severe drought conditions.

The combined analysis of the effects of drought stress on the shoots fresh weight and shoots dry weight of plants grown under field conditions of the two growing seasons is shown in Table 4. At harvesting time, fresh and dry weights of shoots decreased as the drought regimes increased and the values ranged from 101.0 to 980.0 g and 15.96 to 77.5 g under 20% soil moisture content and no drought treatment, respectively. In general, the 21 studied potato cultivars showed more shoots fresh dry weights at the control level (no drought) than any of the three different drought regimes (60%, 40% and 20% soil moisture content), although Diamond, Russet Burbank and Agata cultivars outperformed other cultivars at 60% and 40% soil moisture content. The reduction was generally greater in 20% soil moisture content. Diamond cv. had the maximum shoots fresh weight (526.67 g) when Agata cv. had the highest shoots dry weight (41.17 g). Marfona cv., Gazella cv., Desiree cv. and Lady Rosetta cv. showed the greatest decrease in shoots fresh and dry weights when grown in field with elevated drought regimes (Table 4).

Table 4 Shoots fresh weight, shoots dry weight, no. of tubers/plant, and weight of tubers/plant at harvesting time of 21 potato (Solanum tuberosum L.) cultivars

Potato yield was highly more variable due to drought regimes than the growth response as clear in Table 4. The number and the weight of tubers/plant were observed under the three drought regimes and compared to the control (no drought). Tuber’s production declined as a result of drought stress in all studied 21 cultivars, and there were significant variations in the number and the weight of tubers/plant at the higher drought regime among the cultivars. The average number of tubers per plant ranged from 0.13 under high stress regime (20% soil moisture content) to 7.55 under no stress treatment. With the lowest drought level (60% soil moisture content), all cultivars formed tubers and the number was significantly different from the production under normal conditions (no drought) with the exception of Agata, Atlantic, Diamond, Kennebec, and Russet Burbank cultivars which showed no significant difference. When cultivars were subjected to the highest level of drought stress (20% soil moisture content), the number of tubers greatly declined in the most of cultivars, however, Diamond, Agata and Russet Burbank, gave higher number of tubers than the other cultivars (2.4, 2.1, 1.88, respectively). On the other hand, Gazella, King Edward, Gala, Marfona and Anya cultivars, respectively, formed the lowest average number of potato tubers over the stress treatments (Table 4). For tuber yield, in parallel statistical analysis of the combined data indicated that the tuber fresh weight/plant significantly affected by increasing drought regimes from 60 to 20%. The fresh weight of tubers ranged from 15 g/plant under high drought stress to 567.4 g/plant under no drought conditions. At the highest stress level, Diamond cv., Agata cv., Russet Burbank cv. and closely followed by Atlantic cv. and Kennebec cv. had the highest weight of tubers/plant (139.1, 130.3, 125.4, 115.1 and 105.01 g/plant, respectively) while Anya, Mizen, Almond, Marfona, Gazella and Gala cultivars gave the lowest fresh weight (Table 4).

Stress tolerance index (STI) and cluster analysis

In the STI, there was a control (no drought) for each experiment and drought stress levels in in vitro or field-based screening. Values for each cultivar and each drought treatment to that cultivar’s no drought treatment were calculated to give the stress-tolerance index (STI) for root parameters, i.e., roots fresh weight, no. of roots, total root length and surface area of root studied under tissue culture conditions (in vitro) and tuber production, i.e., no. of tubers and tuber yield examined under field conditions (in vivo). Drought tolerance, as expressed by the stress tolerance index (STI) is shown in Table 5.

Table 5 Drought tolerance as expressed by the stress tolerance index (STI) of roots fresh weight, no. of roots, total root length and surface area of root of stem node derived in vitro plantlets grown under three different levels of sorbitol 0.1, 0.2 or 0.3 mol L−1, and no. of tubers, and weight of tubers of plants grown in field under three different regimes of drought 60%, 40% or 20% of moisture content of 21 potato (Solanum tuberosum L.) cultivars

The STIs of roots fresh weight, no. of roots, total root length and root surface area revealed a degree of tolerance to drought stress in some of the cultivars compared to others based on the level of drought and/or trait. The STIs decreased with increasing drought in all examined cultivars. Agata cv. showed the highest STI of roots fresh weight when the plants were subjected to slightly or moderately drought conditions (0.1 and 0.2 mol L−1 sorbitol) however, Diamond cv. had better STI with the highest level of drought (0.3 mol L−1 sorbitol). On the other hand, Diamond, Atlantic, Kennebec, Russet Burbank and Agata cultivars showed remarkable STI of number of roots across all drought levels. For STI of root length, Agata cv. was more tolerant than any other cultivars under intermediate and high drought conditions, when Kennebec cv. had higher STI under the low level of drought stress. When the plants were treated with low drought condition (0.1 mol L−1 sorbitol), Russet Burbank cv. exhibited the largest STI of surface area of root; nevertheless, Diamond cv. exhibited great tolerance with the higher levels of drought (0.2 and 0.3 mol L−1 sorbitol).

Variations in the STIs of number of potato tubers and tubers fresh weight among cultivars were noticed from low to high drought; however, the STI decreased with increasing drought stress under field-based screening (60%, 40% and 20% soil moisture content). Diamond cv. and Agata cv. were more tolerant and had higher STI of tuber numbers across high drought regime. Atlantic cv. preferred slightly drought conditions to give higher STI of tubers weight. But under the moderate and high drought environment Diamond cv. followed by Agata cv. had better STI. Thus, based on the results obtained from both in vitro-based screening and field-based screening, some cultivars were rated as drought-tolerant.

Based on the variations of the previous examined growth, physiology and yield traits, clustering for response to drought resulted in three distinct groups: (1) drought-tolerant group consisting of cultivars Russet Burbank, Diamond, Agata, Kennebec, Atlantic; (2) a moderately tolerant group consisting of cultivars Maritiema, Lady Balfour, Mizen, Marfona, Lady Rosetta, Marble, Desiree, Champion, Cara, Burren, and Almond, and (3) a sensitive group consisting of cultivars Anya, Gala, Spunta, King Edward, and Gazella (Fig. 2).

Fig. 2
figure 2

Two-way hierarchal clusters associated with the morphological, physiological traits and tuber yield measured per plantlet/plant of combined data on 21 cultivars under different drought stress treatments in vitro and field conditions. Both in vitro and field experiments were conducted, and traits were recorded under drought conditions; 0.0, 0.1, 0.2 or 0.3 mol L−1 sorbitol for in vitro and 60%, 40% or 20% of moisture content for field studies. PFW plantlet fresh weight, SFW shoots fresh weight, RFW roots fresh weight, R/PFW roots/plantlet fresh weight, S/PFW shoots/plantlet fresh weight, RL root length, SAR surface area of root, NR no. of roots, NL no. of leaves, LAP leaf area/plantlet, PWC plantlet water content%, K+ potassium content in roots, SFW-F shoot fresh weight under field trial, SDW-F shoot dry weight under field trial, NT no. of tubers/plant, WT weight of tubers/plant

Glasshouse experiments

Effect of drought on the expression profile of stress-related genes

The main objective of this experiment was to study the regulation of genes and transcription factors that are involved in the response of plant to drought for example; deeper rooting (DRO) potato, LRR receptor-like serine/threonine-protein kinase ERECTA (ERECTA), ethylene response factor (ERF), dehydration responsive element binding (DREB) and Solanum tuberosum MYB (StMYB). Therefore, drought-related genes were chosen and examined in some of the potato cultivars that differed in their response to drought on the in vitro and field-based screening for example; Atlantic cv., Agata cv., Diamond cv., Kennebec cv., Russet Burbank cv. which showed higher STI, and Anya cv. and Gala cv. which showed lower STI. In this experiment, drought stress was applied to 45-day-old plants by withholding water for 2 weeks, and plant roots were tested at 2, 4, 6, 8, and 10 days. Roots from plants that were routinely watered were utilized as a control. RT-PCR analysis was done to identify the expression profile of the DRO, ERECTA, ERF, DREB and StMYB during this drought experiment using mRNA isolated from roots derived from plants of the seven cultivars (Fig. 3). mRNA accumulation of the studied genes was observable beginning early or late after exposed to drought stress of all cultivars examined. In general, the genes mRNA accumulated to a much greater degree in Atlantic cv., Agata cv., Diamond cv., Kennebec cv., Russet Burbank cv. which produced high plant growth, and yield under stress conditions than in Anya cv. and Gala cv. which produced low plant growth, and yield under stress conditions.

Fig. 3
figure 3

Expression profile analysis of the drought-related genes or transcription regulators by RT-PCR. 45-day-old plants were subjected to drought stress, with no water for 2 weeks. mRNA was extracted from roots of normal irrigated plants (Control) and drought exposed plants after 2, 4, 6, 8 and 10 days. Deeper rooting (DRO) potato, LRR receptor-like serine/threonine-protein kinase ERECTA (ERECTA), ethylene response factor (ERF), dehydration responsive element binding (DREB) and StMYB were examined in Atlantic cv., Agata cv., Diamond cv., Kennebec cv., Russet Burbank cv. and Anya cv. and Gala cv. plants. The findings were verified to be reproducible using samples from three independently grown plants. PCR products were analyzed on a 2% agarose gel

DOR potato had the highest expression at drought day 10. It up-regulated over the drought period of time, however in Diamond cv. and Agata cv. DOR potato started to up-regulate directly after exposed to drought day 2. The ERECTA expression level increased in Atlantic and Agata plants and achieved the highest expression at day 10. Again, in Diamond plants, the expression of ERECTA increased early at the drought day 4. On the other hand, ERECTA had a high level of expression at 6 and 8 days in Kennebec cv. and at 10 days in Russet Burbank cv. ERF expressed abundantly in Atlantic cv., Agata cv., Diamond cv. however, the level of expression was low in Kennebec cv., Russet Burbank cv. and slightly increased over the drought days. In StMYB, the level of expression was higher at 6 days of drought in all cultivars which previously showed higher STI although it showed low level of expression at 10 days. DREP exhibited the highest expression at day 6 among Atlantic cv., Agata cv., Diamond cv., Kennebec cv., Russet Burbank cv., and then, the expression decreased at day 8 in Agata cv., Diamond cv. and Kennebec cv. with slight increase at day 10. On the same time, the DREP level of expression was high early at 2 day of drought in Diamond cv. and Russet Burbank cv. In contrary, in Anya cv. and Gala cv. which showed lower STI, the DOR potato, ERECTA, ERF, StMYB and DREP level of expression was relatively stable or continually decreased over the time of drought (Fig. 3).

Effect of drought on stomatal conductance and transpiration

In this experiment, physiological characteristics, i.e., stomatal conductance and transpiration were examined using 60-day-old plants which subjected to drought stress (20% soil moisture content). Seven potato cultivars that differed in their stress tolerance index were studied, Atlantic cv., Agata cv., Diamond cv., Kennebec cv., Russet Burbank cv. which showed greater STI, and Anya cv. and Gala cv. which showed lower STI. Significant variations were identified for stomatal conductance and transpiration among examined cultivars and drought stress (Fig. 4A, B). Overall, Atlantic cv., Agata cv., Diamond cv., Kennebec cv., Russet Burbank cv. had higher stomatal conductance and transpiration than Anya cv. and Gala cv. The highest values were observed in Agata cv. under no drought conditions (0.028 mol H2O m−2 s−1 and 211.1 g H2O plant−1, respectively) however, Diamond cv. had the highest value under the drought stress (0.019 mol H2O m−2 s−1 and 121.32 g H2O plant−1, respectively). In comparison, Anya cv. and Gala cv. under drought conditions, both stomatal conductance and transpiration values showed a substantial decrease. For the correlation between tuber yield and physiological traits under drought conditions, linear function was fitted as clear in Fig. 4C, D. Likewise, the stomatal conductance and transpiration explained high correlation with the tuber yield in the seven studied cultivars under drought and no drought conditions.

Fig. 4
figure 4

Stomatal conductance (A), transpiration (B), linear function of stomatal conductance and tuber fresh weight (C) and linear function of transpiration and tuber fresh weight (D) under no drought and drought conditions in Atlantic cv., Agata cv., Diamond cv., Kennebec cv., Russet Burbank cv. and Anya cv. and Gala cv. plants. Stomatal conductance was measured at night (8:00 pm) while, total day transpiration was measured in a period of 24 h. Values are means ± SE, (n = 10) and the experiment was repeated three times and significant differences were calculated using LSD test

Discussion

Potato is one of the world’s most significant crops [7, 38, 39]. In Egypt, there are two primary growth seasons for potatoes: summer and fall. Every year, potato tuber seeds are imported from cold climate countries such as the Netherlands, Ireland, and the United Kingdom to be grown in Egypt during the summer season. Eventually, the tubers produced from the summer season will be stored for 3 months at 4 °C to be grown in the fall season, then the fall season production will be available for consumption. Egypt is located in a dry climate; yet, finding drought-tolerant cultivars is one approach for mitigating the negative consequences of drought stress [17, 18, 20]. There are significant variations in potato sensitivity, and individual cultivars’ drought tolerance vary greatly [2, 7, 10]. Consequently, it is very important, in order to assign cultivars that can be grown in high drought regions or integrate drought-tolerant ones in breeding programs, to screen the available potato cultivars for their tolerance to drought [13,14,15]. Several researchers [7, 20, 40] have utilized in vitro assay to identify potato germplasm with drought tolerance; however, no thorough in vitro and field combined assessment of cultivars has been reported. In this study, in vitro and field experiments were used to assess the tolerance of potato cultivars (Solanum tuberosum L.) to distinct degrees of drought stress. The current study included the screening of 21 different potato cultivars which are commonly cultivated in the Middle East, including Egypt.

Significant differences were observed among cultivars for morphological and physiological parameters [41, 42]. A strong link was found in potato cultivars between growth and total plant production under drought conditions. Discussions will therefore be summarized over experiments. In vitro-based screening with stem node culture may be a great way to check and select tolerance for drought [7, 20, 40]. The current study involved different levels of drought stress; no drought, 0.1, 0.2 and 0.3 mol L−1 sorbitol. Root traits were the first morphological traits that were significantly affected by stress. Such findings are not consistent with other studies that the reduction in leaf numbers was the first consequence of drought [7, 43]. On the other hand, the results are consistent with other studies reported for salt tolerance in potato and tomato plants [2, 4].

Under drought stress, five cultivars, Agata, Atlantic, Diamond, Kennebec, and Russet Burbank, produced more root growth than other cultivars. The results were greatly clarified when the five cultivars had the highest root traits at 0.3 mol L−1 of sorbitol under in vitro-based screening, suggesting that there is considerable variation in drought tolerance among cultivars. These findings are in agreement with previous studies [44, 45] which reported that developmental plant response to drought stress is manifested by increased root growth. Similarly, drought stress disrupted cell expansion and elongation, leading to a reduction in the leaf area and plant water content. The morphological consequence of drought was a reduction in leaf size, resulting in reduced photosynthesis and reduced accumulation of dry matter in tubers [7, 10, 14, 43]. At the same time, higher levels of drought contributed to a substantial reduction in the level of K+ roots [46].

Screening in vitro may result in a loss of tolerance of plants or a different reaction under field conditions [17, 47]. To obtain full information, it is necessary to evaluate the tolerance of drought under field conditions. Several studies documented the effects of drought stress on tuberization [47,48,49,50]. In this study, the number and weight of tubers significantly declined in many of the evaluated cultivars. Russet Burbank, Agata, Diamond, Atlantic and Kennebec cultivars gave higher number of tubers than the other cultivars while, Diamond, Atlantic and Agata showed more tolerance and had the highest weight of tubers/plant under the drought regimes. These results confirm that the cultivars with higher stress tolerance index (STI) in tissue culture assay showed similar performance under field trial.

Significant developments in drought tolerance studies have addressed a range of primary genes and expression regulators that manipulate different growth or yield traits [24, 26, 27]. Root morphology and stomata growth genes play a crucial role in soil moisture extraction and preservation, and thus expression profile of some of these genes were analyzed under drought scheme [2, 24, 25, 33]. Any cultivars may be useful contributors in drought tolerance breeding programs if tissue culture and field findings are taken into account [16, 18]. Under drought stress of the current study, Atlantic cv., Agata cv., Diamond cv., Kennebec cv., Russet Burbank cv. displayed higher growth traits and yield than other cultivars and, for example, Anya cv. and Gala cv. had a responsive reaction. Meanwhile, based on the physiological and molecular investigations, these cultivars were chosen to emphasize the distinctions between drought-tolerant and drought-sensitive cultivars. Discussion of these findings found that five genes had variations in expression to which up or down-regulation of gene expression was observed in a drought-tolerant cultivars relative to a drought-sensitive cultivar in drought days. DOR, ERECTA and StMYB demonstrated a similar degree of expression on day 0 then highly up-regulated in drought-tolerant cultivars. ERF and DREP showed differential expression on day 0 but they also up-regulated in drought-tolerant cultivars. On the other hands, all genes exhibited low level of expression starting from day 0 to day 10 of drought stress in Anya cv. and Gala cv., the drought-sensitive cultivars. Surprisingly, these genes began to up-regulate early in Atlantic cv., Agata cv., Diamond cv., Kennebec cv., Russet Burbank cv. right after being subjected to stress. Taken together by roots morphological traits and tuber production which conducted in vitro and field, this might clarify why these cultivars response to drought was higher than that of other cultivars. These findings are consistent with those recognized by DRO and ERECTA as a stimulator of drought tolerance through root trait regulation and transpiration efficiency. DRO was classified as the root system architecture controller by modifying the root growth angle for rice. The profound rooting was seen as benefiting in rice not only for a tolerance to drought, but also for improved yield, nitrogen absorption and cytokinin fluxes from root to shoot in grain [25]. ERECTA is one of the transcription efficiency regulated genes which boost the biomass provided by the unit of water transpired and also connected to the deep root system [23]. ERF, DREB and MYB were reported as both positive and negative regulators of drought reactions in wheat, rice, maize and Arabidopsis [25,26,27,28, 33]. Over the last two decades, researchers have taken important steps in our understanding of the mechanisms involved in adaptation and tolerance to drought stress in some plants such as wheat and rice. Wheat cultivars adapted various drought-tolerance mechanisms, such as deeper root formation, higher biomass accumulation, enhanced stomatal control over transpiration [51], betterment of osmoprotective and antioxidant response [52, 53], and, most importantly, improved coordination of positive and negative gene expression. In rice the mechanisms included increasing chlorophyll content, harvest index, stomatal density and conductance, root thickness and length, waxy or thick leaf coverings, as well as decreasing osmotic potential, transpiration rates, and leaf weight and size [25, 33, 54, 55].

Stomatal closing leads to reduced water potential for leaves, reduced carbon assimilation, oxidation and increased canopy in response to stress [56]. Sustaining improved stomata regulation of transpiration is essential to the battle against inhibition of photosynthesis under drought stress [57]. Additionally, physiological traits, i.e., stomatal conductance and transpiration were performed under drought and normal irrigation. In general, cultivars that withstand drought, i.e., Atlantic cv., Agata cv., Diamond cv., Kennebec cv., Russet Burbank cv. had a higher stomatal activity and transpiration than in a drought-sensitive ones, i.e., Anya cv. and Gala cv. and the stomatal conductance and transpiration were highly correlated with the tuber production. Meanwhile, these cultivars, i.e., Atlantic cv., Agata cv., Diamond cv., Kennebec cv., Russet Burbank cv. were the best suited to drought stress conditions and may be recommended for cultivation or as parenting materials for breeding programs to develop higher yielding cultivars. In order to further understand the underlying mechanisms of drought interference with potato growth and yield, an intermediate experiment with interim tuber development and production assessments may be more insightful and a dedicated more molecular, cellular and physiological analysis is suggested.

Conclusion

The study’s main aim is to assess drought tolerance in 21 potato cultivars in vitro and in the field. Drought stress induced by various concentrations of sorbitol (0.1, 0.2, and 0.3 mol L−1) in vitro and different regimes of soil moisture content (60, 40, and 20%) under field conditions exhibited fairly comparable impacts on the parameters studied. The interactions between the factors examined revealed that: potato cultivars differed significantly in their response to drought stress under in vitro and field conditions. In general, some cultivars were considerably more tolerant to drought stress treatments than others. Plant growth, physiological traits, potato tuber yield were all reduced by stress, and the reduction was much greater at the highest drought stress level (0.3 mol L−1 sorbitol in vitro and 20% soil moisture content under field conditions). There was a high association between in vitro and field trials for plant growth and tuber yield. The regulation of drought-related genes was different among the cultivars which have high and low stress tolerance index. Diamond, Kennebec, Russet Burbank, Atlantic and Agata cultivars are good candidates for inclusion in breeding programs for drought tolerance. Choosing the appropriate cultivar(s) is essential for achieving high quality and economic returns under stress. Because potato is a drought-sensitive adapted species, the best approaches to incorporating its drought-tolerance would be to (1) identify more genes involved in drought-tolerance mechanism, or (2) intercross drought-tolerant cultivars with drought-sensitive adapted germplasm and screen for drought tolerance and/or genes involved in drought tolerance. Research on both fronts is being pursued in our laboratory.

Availability of data and materials

Not applicable.

Abbreviations

STI:

Stress tolerance index

DRO:

Deeper rooting in potato

ERECTA:

LRR receptor-like serine/threonine-protein kinase ERECTA

ERF:

Ethylene response factor

DREB:

Dehydration responsive element binding

StMYB:

Solanum tuberosum MYB

References

  1. Bohnert HJ, Nelson DE, Jensen RG. Adaptations to environmental stresses. Plant Cell. 1995;7(7):1099. https://doi.org/10.1105/tpc.7.7.1099.

    CAS  Article  PubMed  PubMed Central  Google Scholar 

  2. Zaki HEM, Haynes KG. In vitro selection for salinity tolerance in wild potato. In: 99th annual meeting of The Potato Association of America-Portland (PAA), Maine, USA, 2015.

  3. United States Department of Agriculture. Salinity in agriculture. http://www.ars.usda.gov/Aboutus/docs.htm?docid=10201. Accessed 23 Jan 2013.

  4. Zaki HE, Yokoi S. A comparative in vitro study of salt tolerance in cultivated tomato and related wild species. Plant Biotechnol. 2016. https://doi.org/10.5511/plantbiotechnology.16.1006a.

    Article  Google Scholar 

  5. Boyer JS. Plant productivity and environment. Science. 1982;218(4571):443–8. https://doi.org/10.1126/science.218.4571.443.

    CAS  Article  PubMed  Google Scholar 

  6. Mullins E, Milbourne D, Petti C, Doyle-Prestwich BM, Meade C. Potato in the age of biotechnology. Trends Plant Sci. 2006;11(5):254–60. https://doi.org/10.1016/j.tplants.2006.03.002.

    CAS  Article  PubMed  Google Scholar 

  7. Albiski F, Najla S, Sanoubar R, Alkabani N, Murshed R. In vitro screening of potato lines for drought tolerance. Physiol Mol Biol Plants. 2012;18(4):315–21. https://doi.org/10.1007/s12298-012-0127-5.

    CAS  Article  PubMed  PubMed Central  Google Scholar 

  8. Iwama K, Yamaguchi J. Abiotic stresses. In: Gopal J, Paul Khurana SM, editors. Handbook of potato production, improvement and postharvest management. New York: Food Product Press; 2006. p. 231–78.

    Google Scholar 

  9. Levy D, Veilleux RE. Adaptation of potato to high temperatures and salinity—a review. Am J Potato Res. 2007;84(6):487–506. https://doi.org/10.1007/BF02987885.

    Article  Google Scholar 

  10. Deblonde PM, Ledent JF. Effects of moderate drought conditions on green leaf number, stem height, leaf length and tuber yield of potato cultivars. Eur J Agron. 2001;14(1):31–41. https://doi.org/10.1016/S1161-0301(00)00081-2.

    Article  Google Scholar 

  11. Tourneux C, Devaux A, Camacho M, Mamani P, Ledent JF. Effects of water shortage on six potato genotypes in the highlands of Bolivia (I): morphological parameters, growth and yield. Agronomie. 2003;23(2):169–79. https://doi.org/10.1051/agro:2002079.

    Article  Google Scholar 

  12. Lahlou O, Ledent JF. Root mass and depth, stolons and roots formed on stolons in four cultivars of potato under water stress. Eur J Agron. 2005;22(2):159–73. https://doi.org/10.1016/j.eja.2004.02.004.

    Article  Google Scholar 

  13. Hang AN, Miller DE. Yield and physiological responses of potatoes to deficit, high frequency sprinkler irrigation 1. Agron J. 1986;78(3):436–40. https://doi.org/10.2134/agronj1986.00021962007800030008x.

    Article  Google Scholar 

  14. Jefferies RA, Mackerron DK. Responses of potato genotypes to drought. II. Leaf area index, growth and yield. Ann Appl Biol. 1993;122(1):105–12. https://doi.org/10.1111/j.1744-7348.1993.tb04018.x.

    Article  Google Scholar 

  15. Schittenhelm S, Sourell H, Löpmeier FJ. Drought resistance of potato cultivars with contrasting canopy architecture. Eur J Agron. 2006;24(3):193–202. https://doi.org/10.1016/j.eja.2005.05.004.

    Article  Google Scholar 

  16. Hassanpanah D. Evaluation of potato advanced cultivars against water deficit stress under in vitro and in vivo conditions. Biotechnology. 2010;9(2):164–9.

    Article  Google Scholar 

  17. Chen D, Neumann K, Friedel S, Kilian B, Chen M, Altmann T, Klukas C. Dissecting the phenotypic components of crop plant growth and drought responses based on high-throughput image analysis. Plant Cell. 2014;26(12):4636–55. https://doi.org/10.1105/tpc.114.129601.

    CAS  Article  PubMed  PubMed Central  Google Scholar 

  18. Luitel BP, Khatri BB, Choudhary D, Paudel BP, Jung-Sook S, Hur OS, Baek HJ, Cheol KH, Yul RK. Growth and yield characters of potato genotypes grown in drought and irrigated conditions of Nepal. Int J Appl Sci Biotechnol. 2015;3(3):513–9. https://doi.org/10.3126/ijasbt.v3i3.13347.

    Article  Google Scholar 

  19. Lobell DB, Schlenker W, Costa-Roberts J. Climate trends and global crop production since 1980. Science. 2011;333(6042):616–20. https://doi.org/10.1126/science.1204531.

    CAS  Article  PubMed  Google Scholar 

  20. Barra M, Correa J, Salazar E, Sagredo B. Response of potato (Solanum tuberosum L.) germplasm to water stress under in vitro conditions. Am J Potato Res. 2013;90(6):591–606. https://doi.org/10.1007/s12230-013-9333-0.

    Article  Google Scholar 

  21. Anithakumari AM, Dolstra O, Vosman B, Visser RG, van der Linden CG. In vitro screening and QTL analysis for drought tolerance in diploid potato. Euphytica. 2011;181(3):357–69. https://doi.org/10.1007/s10681-011-0446-6.

    Article  Google Scholar 

  22. Anithakumari AM, Nataraja KN, Visser RG, van der Linden CG. Genetic dissection of drought tolerance and recovery potential by quantitative trait locus mapping of a diploid potato population. Mol Breed. 2012;30(3):1413–29. https://doi.org/10.1007/s11032-012-9728-5.

    CAS  Article  PubMed  PubMed Central  Google Scholar 

  23. Condon AG, Richards RA, Rebetzke GJ, Farquhar GD. Breeding for high water-use efficiency. J Exp Bot. 2004;55(407):2447–60. https://doi.org/10.1093/jxb/erh277.

    CAS  Article  PubMed  Google Scholar 

  24. López-Maury L, Marguerat S, Bähler J. Tuning gene expression to changing environments: from rapid responses to evolutionary adaptation. Nat Rev Genet. 2008;9(8):583–93. https://doi.org/10.1038/nrg2398.

    CAS  Article  PubMed  Google Scholar 

  25. Arai-Sanoh Y, Takai T, Yoshinaga S, Nakano H, Kojima M, Sakakibara H, Kondo M, Uga Y. Deep rooting conferred by DEEPER ROOTING 1 enhances rice yield in paddy fields. Sci Rep. 2014;4(1):1–6. https://doi.org/10.1038/srep05563.

    CAS  Article  Google Scholar 

  26. Gahlaut V, Jaiswal V, Kumar A, Gupta PK. Transcription factors involved in drought tolerance and their possible role in developing drought tolerant cultivars with emphasis on wheat (Triticum aestivum L.). Theor Appl Genet. 2016;129(11):2019–42. https://doi.org/10.1007/s00122-016-2794-z.

    CAS  Article  PubMed  Google Scholar 

  27. Joshi R, Wani SH, Singh B, Bohra A, Dar ZA, Lone AA, Pareek A, Singla-Pareek SL. Transcription factors and plants response to drought stress: current understanding and future directions. Front Plant Sci. 2016;14(7):1029. https://doi.org/10.3389/fpls.2016.01029.

    Article  Google Scholar 

  28. Kulkarni M, Soolanayakanahally R, Ogawa S, Uga Y, Selvaraj MG, Kagale S. Drought response in wheat: key genes and regulatory mechanisms controlling root system architecture and transpiration efficiency. Front Chem. 2017;5(5):106. https://doi.org/10.3389/fchem.2017.00106.

    Article  PubMed  PubMed Central  Google Scholar 

  29. Pieczynski M, Wyrzykowska A, Milanowska K, Boguszewska-Mankowska D, Zagdanska B, Karlowski W, Jarmolowski A, Szweykowska-Kulinska Z. Genomewide identification of genes involved in the potato response to drought indicates functional evolutionary conservation with Arabidopsis plants. Plant Biotechnol J. 2018;16(2):603–14. https://doi.org/10.1111/pbi.12800.

    CAS  Article  PubMed  Google Scholar 

  30. Hamooh BT, Sattar FA, Wellman G, Mousa MAA. Metabolomic and biochemical analysis of two potato (Solanum tuberosum L.) cultivars exposed to in vitro osmotic and salt stresses. Plants. 2021;10:98. https://doi.org/10.3390/plants10010098.

    CAS  Article  PubMed  PubMed Central  Google Scholar 

  31. Sattar FA, Hamooh BT, Wellman G, Ali MA, Shah SH, Anwar Y, Mousa MAA. Growth and biochemical responses of potato cultivars under In Vitro lithium chloride and mannitol simulated salinity and drought stress. Plants. 2021;10:924. https://doi.org/10.3390/plants10050924.

    CAS  Article  PubMed  PubMed Central  Google Scholar 

  32. Murashige T, Skoog F. A revised medium for rapid growth and bio assays with tobacco tissue cultures. Physiol Plant. 1962;15(3):473–97. https://doi.org/10.1111/j.1399-3054.1962.tb08052.x.

    CAS  Article  Google Scholar 

  33. Guo Q, Zhang J, Gao Q, Xing S, Li F, Wang W. Drought tolerance through overexpression of monoubiquitin in transgenic tobacco. J Plant Physiol. 2008;165(16):1745–55. https://doi.org/10.1016/j.jplph.2007.10.002.

    CAS  Article  PubMed  Google Scholar 

  34. Cano EA, Pérez-Alfocea F, Moreno V, Caro M, Bolarín MC. Evaluation of salt tolerance in cultivated and wild tomato species through in vitro shoot apex culture. Plant Cell Tissue Organ Cult. 1998;53(1):19–26. https://doi.org/10.1023/A:1006017001146.

    Article  Google Scholar 

  35. Ministry of Agriculture and Land Reclamation, Agriculture Research Center (ARC). Potato production. 2003 no. 813. http://www.vercon.sci.eg/.

  36. Reddy PJ, Vaidyanath K. Note on salt tolerance of some rice varieties of Andhra Pradesh during germination and early seedling growth. Indian J Agric Sci. 1982;52:472–4.

    Google Scholar 

  37. Ramírez DA, Yactayo W, Rolando JL, Quiroz R. Preliminary evidence of nocturnal transpiration and stomatal conductance in potato and their interaction with drought and yield. Am J Potato Res. 2018;95(2):139–43. https://doi.org/10.1007/s12230-018-9652-2.

    Article  Google Scholar 

  38. FAO. World crop production statistics. http://faostat.fao.org. Accessed 10 Sept 2016.

  39. Tican A, Cioloca M, Chiru N, Bădărău C. Behavior of different potato varieties by simulating in vitro of hydric stress conditions. Agron Ser Sci Res/Lucrari Stiintifice Seria Agronomie. 2016;59(1):97–102.

    Google Scholar 

  40. Gopal J, Iwama K. In vitro screening of potato against water-stress mediated through sorbitol and polyethylene glycol. Plant Cell Rep. 2007;26(5):693–700. https://doi.org/10.1007/s00299-006-0275-6.

    CAS  Article  PubMed  Google Scholar 

  41. Gopal J, Iwama K, Jitsuyama Y. Effect of water stress mediated through agar on in vitro growth of potato. In Vitro Cell Dev Biol-Plant. 2008;44(3):221–8. https://doi.org/10.1007/s11627-007-9102-1.

    Article  Google Scholar 

  42. Haynes KG, Zaki HE, Christensen CT, Ogden E, Rowland LJ, Kramer M, Zotarelli L. High levels of heterozygosity found for 15 SSR loci in Solanum chacoense. Am J Potato Res. 2017;94(6):638–46. https://doi.org/10.1007/s12230-017-9602-4.

    CAS  Article  Google Scholar 

  43. Frensch J. Primary responses of root and leaf elongation to water deficits in the atmosphere and soil solution. J Exp Bot. 1997;48(5):985–99. https://doi.org/10.1093/jxb/48.5.985.

    CAS  Article  Google Scholar 

  44. Yamaguchi M, Sharp RE. Complexity and coordination of root growth at low water potentials: recent advances from transcriptomic and proteomic analyses. Plant Cell Environ. 2010;33(4):590–603. https://doi.org/10.1111/j.1365-3040.2009.02064.x.

    CAS  Article  PubMed  Google Scholar 

  45. Xu W, Jia L, Shi W, Liang J, Zhou F, Li Q, Zhang J. Abscisic acid accumulation modulates auxin transport in the root tip to enhance proton secretion for maintaining root growth under moderate water stress. New Phytol. 2013;197(1):139–50. https://doi.org/10.1111/nph.12004.

    CAS  Article  PubMed  Google Scholar 

  46. Thornton MK. Effects of heat and water stress on the physiology of potatoes. Presented at the Idaho Potato Conference 2002 January 23.

  47. Schafleitner R, Gutierrez R, Espino R, Gaudin A, Pérez J, Martínez M, Domínguez A, Tincopa L, Alvarado C, Numberto G, Bonierbale M. Field screening for variation of drought tolerance in Solanum tuberosum L. by agronomical, physiological and genetic analysis. Potato Res. 2007;50(1):71–85. https://doi.org/10.1007/s11540-007-9030-9.

    CAS  Article  Google Scholar 

  48. Lahlou O, Ouattar S, Ledent JF. The effect of drought and cultivar on growth parameters, yield and yield components of potato. Agronomie. 2003;23(3):257–68.

    Article  Google Scholar 

  49. Stalham MA, Allen EJ, Rosenfeld AB, Herry FX. Effects of soil compaction in potato (Solanum tuberosum) crops. J Agric Sci. 2007;145(4):295–312. https://doi.org/10.1017/S0021859607006867.

    Article  Google Scholar 

  50. Hirut B, Shimelis H, Fentahun M, Bonierbale M, Gastelo M, Asfaw A. Combining ability of highland tropic adapted potato for tuber yield and yield components under drought. PLoS ONE. 2017;12(7):e0181541. https://doi.org/10.1371/journal.pone.0181541.

    CAS  Article  PubMed  PubMed Central  Google Scholar 

  51. Chipilski RR, Kocheva KV, Nenova VR, Georgiev GI. Physiological responses of two wheat cultivars to soil drought. Z Naturforschung. 2012;C67:181–6. https://doi.org/10.5560/ZNC.2012.67c0181.

    Article  Google Scholar 

  52. Huseynova IM. Photosynthetic characteristics and enzymatic antioxidant capacity of leaves from wheat cultivars exposed to drought. Biochim Biophys Acta Bioenerg. 2012;1817:1516–23. https://doi.org/10.1016/j.bbabio.2012.02.037.

    CAS  Article  Google Scholar 

  53. Loutfy N, El-Tayeb MA, Hassanen AM, Moustafa MF, Sakuma Y, Inouhe M. Changes in the water status and osmotic solute contents in response to drought and salicylic acid treatments in four different cultivars of wheat (Triticum aestivum). J Plant Res. 2012;125:173–84. https://doi.org/10.1007/s10265-011-0419-9.

    CAS  Article  PubMed  Google Scholar 

  54. Sahebi M, Hanafi MM, Rafii MY, Mahmud TMM, Azizi P, Osman M, Miah G. Improvement of drought tolerance in rice (Oryza sativa L.): genetics, genomic tools, and the WRKY gene family. BioMed Res Int. 2018;2018:3158474. https://doi.org/10.1155/2018/3158474.

    CAS  Article  PubMed  PubMed Central  Google Scholar 

  55. Lum MS, Hanafi MM, Rafii YM, Akmar ASN. Effect of drought stress on growth, proline and antioxidant enzyme activities of upland rice. J Anim Plant Sci. 2014;24:1487–93.

    Google Scholar 

  56. Yokota A, Kawasaki S, Iwano M, Nakamura C, Miyake C, Akashi K. Citrulline and DRIP-1 protein (ArgE homologue) in drought tolerance of wild watermelon. Ann Bot. 2002;89(7):825–32. https://doi.org/10.1093/aob/mcf074.

    CAS  Article  PubMed  PubMed Central  Google Scholar 

  57. Bota J, Medrano H, Flexas J. Is photosynthesis limited by decreased Rubisco activity and RuBP content under progressive water stress? New Phytol. 2004;162(3):671–81. https://doi.org/10.1111/j.1469-8137.2004.01056.x.

    CAS  Article  PubMed  Google Scholar 

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Acknowledgements

Dr. Zaki is thankful to the Cultural Affairs & Missions, Ministry of Higher Education, Egypt, for the award of Post-doctoral Fellowship, and to the Laboratory of Plant Breeding, Faculty of Agriculture, Iwate University, Morioka, Iwate, Japan, for the invitation as Visiting Scientist.

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HEMZ: conceptualization, data curation, formal analysis, investigation, methodology, project administration, software, supervision, validation, visualization, writing—original draft, writing—review and editing. KSAR: data curation, formal analysis, investigation, methodology, visualization, writing—original draft, writing—review and editing. Both authors read and approved the final manuscript.

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Correspondence to Haitham E. M. Zaki.

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Supplementary Information

Additional file 1: Table S1.

Distribution of particle size and chemical properties of the experimental site soil. Table S2. List of primers of the selected genes used for RT-PCR.

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Zaki, H.E.M., Radwan, K.S.A. Response of potato (Solanum tuberosum L.) cultivars to drought stress under in vitro and field conditions. Chem. Biol. Technol. Agric. 9, 1 (2022). https://doi.org/10.1186/s40538-021-00266-z

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Keywords

  • Drought tolerance
  • Potato
  • Root traits
  • Tuber production
  • Stress tolerance index
  • Drought-related genes
  • Physiological traits