Abstract
Potato is the third important food crop for global food security. To breed new varieties with stress tolerance, tuber yield and quality are a base for sustainable development of the potato industry. With climate and environmental change, the breeding of abiotic stress tolerance potato varieties has become a research hotspot. This paper summarizes the evaluation methods, genetics, and molecular mechanism of potato tolerance to abiotic stress, including salt and alkali, drought, cold, and heavy metal. The progress of transgenic breeding of potato under abiotic stress was introduced. Finally, the prospect of potato abiotic stresses breeding was put forward.
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16.1 Research on Salt and Alkali-Tolerant Breeding of Potato
16.1.1 Overview
Soil salinization is a global issue. The increase of saline alkali land area has become the main factor restricting agricultural development. The low content of organic matter, exhausted soil fertility and poor physical and chemical properties in saline alkali soil restrict the normal growth and development of plants and seriously reduce the yield of grain crops. It is also the fourth most important crop with high grain and high adaptability in the world, second only to wheat and potato. Potato is a medium salt-tolerant crop, but the potato varieties popularized at this stage have poor salt tolerance. Salt and alkali stress affects the growth and development of potato leaves, stems, roots, and tubers and seriously inhibits the yield of potato. Therefore, it is of great significance to explore the salt and alkali resistance mechanism of potato and improve the salt and alkali tolerance of potato.
Saline alkali stress inhibits the growth of plant roots, thus limiting the growth and development of plants and then affecting the yield of crops (An et al. 2021). When potato is subjected to saline alkali stress, its morphological characteristics, such as root length, plant height, leaf length, and leaf width, are inhibited, and the degree of inhibition is more obvious with the increase of soil salinization (Liu et al. 2011; Ji et al. 2021). In addition, salt alkali stress affects the yield and quality of potato. The quality indexes of potato mainly include dry matter content, starch content, protein content, reducing sugar content, vitamin C content, etc. (Li et al. 2020a, b). Yao et al. (2020) set five NaCl concentration stress treatments for potatoes. The results showed that with the gradual increase of NaCl concentration, potato yield gradually decreased, and the contents of dry matter, starch, and protein decreased, resulting in the decline of potato quality. When potato tissue culture seedlings were treated with NaHCO3, potato leaves wilted and plants withered under alkali stress, and could not grow normally (Kang et al. 2021). For different potato varieties, there are also some differences in salt and alkali tolerance (Chen et al. 2018; Zhao and Bei 2007).
16.2 Salt and Alkali Resistance Evaluation
When the total salt content in the medium reached 0.1%, the plants with the worst salt and alkali-tolerant potato clones (144-4, 3-2, 65) began to die. However, the plants with strong salt and alkali-tolerant potato clones (I, 131, 31) could survive even if the total salt content reached 0.4%. For the plants with strong salt and alkali tolerance, the pH value of culture medium gradually decreased in the process of culture and reached the range suitable for plant growth when sampling. It is suggested that salt-tolerant potato can secrete a large amount of H+ into the soil (Wang et al. 1997). The cluster analysis by using Ward deviation square sum method is divided 130 varieties into eight groups, among which Bintje, Amisk, Onaway, BelRus, Tobique, and Sierra had the strongest salt resistance, while Mainechip had the worst salt tolerance (Khrais et al. 1998). There were significant differences in salt tolerance of callus among different potato varieties. The maximum tolerance concentrations of E 1 and Dongnong 303 were greater than those of Xiabodi and Favorita (Li et al. 2004). The stems and leaves of 12 potato varieties were used as explants for callus induction to screen salt-tolerant callus. The results showed that the callus varieties with strong tolerance were Bashu 10, Jizhangshu 5, and 1867 (Zhang et al. 2005) (Table 16.1).
16.3 Physiological Response to Saline Alkali Stress
16.3.1 Affecting Endogenous Hormones
When plants are subjected to saline alkali stress, a series of responses will occur to endogenous hormones in plants (Yu et al. 2020). Under salt stress, the content of auxin in leaves of potato tissue culture seedlings will decrease, while the content of auxin and abscisic acid in roots will increase with the extension of stress time (Wang et al. 2020). Under alkaline stress, the contents of abscisic acid and brassinolide in potato increased with the increase of alkali concentration, but the content of gibberellin decreased gradually (Kang 2021).
16.3.2 Interference Ion Steady State
Saline alkali stress will increase the content of Na+ in plants, while higher concentration of Na+ will affect the absorption of K+, resulting in the increase of Na+/K+ ratio in plants. Li et al. (2010) showed that the K+/Na+ ratio in potato virus-free seedlings decreased under different concentrations of mixed salt stress. Kang (2021) showed that under the condition of alkali stress of different concentrations of NaHCO3, the Na+ content of different potato varieties increased with the increase of alkali concentration, the K+ content decreased with the increase of alkali concentration, and the content ratio of sodium and potassium ions in roots, stems, and leaves decreased with the increase of alkali stress concentration.
16.3.3 Causing Osmotic Stress
Under salt stress, osmotic substances are very important to maintain plant salt tolerance (Singh et al. 2015). As an osmotic protective agent, proline can regulate the osmotic balance in cells. Sun et al. (2009) showed that after mixed salt stress treatment, the proline content of potato leaves increased with the increase of salt concentration, and the content of soluble sugar also increased gradually. The same results were obtained after alkali stress treatment with different concentrations of NaHCO3 (Kang 2021).
16.3.4 Weakening Photosynthesis
Chlorophyll is an important substance for plant photosynthesis. Salt alkali stress will reduce the content of chlorophyll and affect crop yield (Fang et al. 2021). Ma (2014) showed that with the gradual increase of salt concentration, the chlorophyll content of salt-tolerant and salt-sensitive strains decreased, which affected the photosynthesis of plants. When potato plants were subjected to saline alkali stress, the contents of chlorophyll a and chlorophyll b among different varieties decreased (Hu et al. 2020).
16.3.5 Leading to oxidative stress
When potato plants are subjected to saline alkali stress, the content of malondialdehyde (MDA) increases with the increase of saline alkali concentration (Li 2016). Charfeddine et al. (2019) introduced the StERF94 gene into potato plants and obtained overexpression transgenic lines. The research showed that the activity of superoxide dismutase (SOD) in the plants was high, which could reduce the damage of reactive oxygen species (ROS)-mediated membrane system caused by salt stress. Zhang (2010) and Li et al. (2020a, b) studied several potato varieties/lines with different salt sensitivities. When the stress time gradually increased, the activities of peroxidase (POD) and SOD showed a gradual downward trend, while with the increase of salt stress time, the activity of catalase (CAT) first decreased and then increased. Zhao et al. (2014) compared the morphological and physiological characters of diploid potato with different salt tolerances under alkali stress. The results showed that whether at the concentration of 5 mmol/L NaHCO3 or 10 mmol/L NaHCO3, the relative value of SOD activity from high to low was salt-tolerant group > medium salt-tolerant group > salt-sensitive group, and the relative value of POD activity from high to low was salt-tolerant group > medium salt-tolerant group > salt-sensitive group.
16.4 Salt Tolerance Gene
ABF is a bZIP transcription factor. The results of the plantlets of transgenic potato with GhABF2 showed that the biomass of GhABF2 transgenic materials T1 and T2 can be significantly increased compared with the control. The transgenic plants had significant salt stress resistance (Pei et al. 2015). Studies have shown that ABF4 can improve the salt tolerance of Arabidopsis (Pan 2020). Noelia et al. (2018) overexpressed the ABF4 gene in Arabidopsis thaliana in potato. The results showed that under normal conditions, there was no significant differences in various physiological indexes between transgenic potato and wild type, but the tuber yield and tuber quality of transgenic potato were higher than those of wild type. After salt stress treatment, the relative water content, proline content, and chlorophyll content of transgenic potato were significantly higher than those of wild type, indicating that abf4 overexpression plants had enhanced salt tolerance. Overexpression of StDREB2 gene in potato plants can improve potato salt tolerance by participating in ABA hormone signal and proline synthesis (Bouaziz et al. 2012). In addition, the researchers also found that overexpression of StDREB1 can also significantly improve the salt tolerance of transgenic potato (Bouaziz et al. 2013).
The salt and alkali tolerance of potato can be improved by increasing the activity of antioxidant enzymes, increasing the content of proline, and reducing the content of MDA through an appropriate amount of exogenous brassinolide (Hu et al. 2016). Zhou et al. (2018) overexpressed DWF4 gene in potato to obtain transgenic potato plants. After stress treatment, the physiological indexes, such as soluble sugar content, soluble protein content, and antioxidant enzyme activity of StDWF4 transgenic potato lines, are higher than those of normal plants, indicating that overexpression of StDWF4 can improve the salt tolerance of potato.
Wu et al. (2019) studied the clonal generation plants of potato transformed with HaBADH (betaine aldehyde dehydrogenase) and showed that with the gradual increase of salt concentration, compared with normal plants, the growth, weight, number, proline, and MDA contents of HaBADH transgenic plants were significantly higher than those of non-transgenic materials. The results showed that the expression of HaBADH gene under salt stress improved salt tolerance of potato transgenic lines. HAL1 (Luo 2007) and BADH (Li et al. 2007) were transferred into potato and the salt resistance of potato plants was significantly enhanced.
Chen et al. (2013) transferred the S-adenosyl-L-methionine synthetase (SAMS) gene into potato for the first time. Comparing seven potato lines (SM13, SM33, SM4, SM22, SM30, SM40, and SM27) with non-transgenic “Atlantic” varieties, the yields of four transgenic lines SM22, SM4, SM33, and SM13 were higher than those of non-transgenic plants, indicating that the transfer of GsSAMS into potato can improve its alkaline tolerance. NHX gene encodes a Na+/H+ antiporter. The study of potato lines with AtNHX1 and non-transgenic “Gannongshu 2” showed that the introduction of AtNHX1 had a significant effect on the salt tolerance of potato plants in the field (Li et al. 2017a) (Table 16.2).
16.5 Methods of Improving Salt and Alkaline Tolerance of Potato
16.5.1 Using Plant Growth-Promoting Rhizobacteria (PGPR)
PGPR can alleviate the negative effects of saline alkali stress on plants, improve plant growth, and improve the salt tolerance of various crops. It has important application value in the field of agricultural development (Kumar et al. 2019; Etesami and Maheshwari 2018). It was found that the symbiosis of plant rhizosphere growth-promoting bacteria with potato can improve the growth ability of potato plants and improve the quality of potato (Lu 2020; Li 2020). Bacillus is a common plant rhizosphere growth-promoting bacterium. When Bacillus and potato coexist under saline alkali stress, Bacillus enhances the salt alkali tolerance of potato by promoting the production of potato rhizosphere auxin, maintaining ion homeostasis and regulating the activity of antioxidant enzymes, so as to improve the biomass, number of tubers per plant, and tuber yield of potato plant (Tahir et al. 2019).
16.5.2 Application of Exogenous Substances
By applying exogenous regulatory substances, the ability of plants to resist salt and alkali can be improved, so as to improve plant salt tolerance (Su et al. 2021; Gao et al. 2017; Jiang et al. 2020). Faried et al. (2017) showed that salicylic acid can significantly improve the activity of antioxidant enzymes, such as SOD, CAT, and POD, regulate the content of proline and phenol, remove ROS, improve the utilization rate of potassium, and reduce the content of sodium in potato leaves, so as to endow potato with salt tolerance. Under saline alkali stress, jasmonic acid can reduce the effect of NaCl on photosynthetic pigments, maintain the osmotic substances of cells, and enhance their salt tolerance (Efimova et al. 2019). The application of exogenous phosphatidylserine can reduce the K+ outflow caused by salt stress and regulate the H+-ATPase activity on the membrane to delay leaf senescence and induce the improvement of salt tolerance of potato (Yu et al. 2019). Ca2+ is not only an important nutrient element but also an important second messenger. When plants are stressed, Ca2+ can improve the stress resistance of plants (García Bossi et al. 2020). The application of exogenous calcium ion can increase the chlorophyll content and enhance the photosynthesis of potato. Secondly, it can also improve the activity of antioxidant enzymes and maintain the stability of membrane, so as to improve the salt tolerance of potato (Wei 2014).
16.6 Research on Drought-Tolerant Breeding of Potato
16.6.1 Overview
Potato is a typical temperate climate crop. It likes the growth conditions of low temperature and cold. It is susceptible to water shortage and lacks effective drought tolerance mechanism (Qin et al. 2019). At the same time, with the rise of global temperature, the drought in potato planting areas is becoming more and more serious, and the water shortage is serious. About potato grows, resulting in serious yield reduction (Wang et al. 2016; Xu et al. 2011).
Under drought stress, the growth of plant height, main stem and root system, leaf number, leaf area, and yield are basically inhibited (Yang et al. 2016; Yao et al. 2013). Zhang et al. (2016) believed that the number of tubers per plant, weight of tubers per plant, and biomass of potato were greatly reduced by drought. Some scholars believe that potato plants will encounter drought stress when the soil water potential drops to −25 kPa or the soil relative field water capacity is less than 50%, and are sensitive to drought in all growth stages (Schafleitner et al. 2007). Especially in the tuber expansion stage, if drought stress is encountered, the yield will be affected, and serious drought can lead to a significant decline in yield. At the same time, it will cause a series of adverse reactions and reduce the quality of potatoes, such as appearance distortion, abnormal tissues and organs, metabolic disorder, and hollow potatoes (Liu et al. 2019; Qin et al. 2018). Wang et al. (2016) showed that the plant height stress index was inversely proportional to the drought resistance index, and reached a significant level of 1%. The drought resistance index will decrease with plant height.
16.7 Drought Tolerance Evaluation
16.7.1 Morphological Yield Index
Identification of drought resistance of potato by plant morphology is a widely used research method at home and abroad. The morphological characteristics of potato, such as root tension, root length, root weight, root shoot ratio, plant height, stem diameter, biomass, fresh weight, leaf weight, stomatal subsidence, and leaf area, are related to drought resistance. The research on drought-resistant morphological characteristics of potato at home and abroad mainly focuses on the root system of the plant. Root system is the organ of potato to receive soil water signal and absorb soil water. Under drought stress, root absorption and root activity decreased, resulting in nutrient imbalance (Xie and Zheng 2015). Du et al. (2012) believed that varieties with strong drought resistance have developed roots and high yield under drought conditions. At the same time, root tension is significantly proportional to root weight, root number, and root length, which is an important index to measure the degree of root development.
Under drought stress, potato yield index is more intuitive. The drought resistance is mainly reflected in the yield, which can be used as an important index to screen the drought-resistant varieties of potato. Chionoy’s drought resistance coefficient method, Fish’s drought sensitivity index (SI) method and Hu Fushun’s drought resistance index (DI) method are its traditional methods (Xie and Zheng 2015). Gu et al. (2013) considered that the change degree of potato tuber yield under water stress is the evaluation index of drought resistance, and the drought resistance index directly shows the sensitivity of plants to drought.
16.7.2 Physiological and Biochemical Indexes
The physiological indexes of plant drought resistance evaluation include soil water content, leaf water content, photosynthetic rate, water potential, respiratory rate, dry matter stress index, osmotic regulation ability, water stress index, water holding capacity of isolated leaves, etc. Du et al. (2012) showed that the canopy coverage was directly proportional to the yield, reaching a significant level of 5%, which can be used as an important physiological index for drought resistance evaluation. The results showed that 80% soil water content was the best for potato plant growth, 60% soil water content was mild drought stress, 40% soil water content was moderate drought stress, and 20% soil water content was severe drought stress. Therefore, soil water content can be used as one of the physiological indexes to evaluate potato drought resistance (Yin and Xiao 2017; Jiao et al. 2011). When potato plants suffer from drought stress and water deficit, photosynthesis will be weakened and photosynthetic rate will be reduced, resulting in crop yield reduction (Yang et al. 2019b). Therefore, photosynthetic rate is an important index for drought resistance evaluation of potato (Li and Tong 2018). The water holding capacity of leaves is an embodiment of potato drought resistance. Wang et al. (2016) showed that the varieties with lower drought resistance have stronger water loss capacity, which is equivalent to the lower water retention capacity. The drought resistance coefficient decreased with the increase of water loss. There was a negative correlation between water loss and drought resistance coefficient, which reached a significant level of 1%, and the potato yield decreased.
Biochemical indexes of plant drought resistance include soluble sugar content, proline content, betaine content, MDA content, ATPase activity, vitamin C content, SOD activity, POD activity, CAT activity, etc. Under drought stress and water shortage conditions, potato plants will accumulate a large amount of proline, and there are differences between varieties and stress time (Li and Tong 2018). Ding et al. (2013) studied that under drought stress, the proline content of different varieties of potato increased by 1.01~5.40 times, the MDA content increased by 1.10~1.91 times, and the free proline content and MDA content of potato leaves showed an upward trend, which can be used as the biochemical index of potato drought resistance evaluation. Soluble sugar is an important osmotic regulator. In many plants, soluble sugar content will almost always increase under drought conditions (Yang et al. 2016). However, Yang (2016) studied six potato varieties, such as “Atlantic,” and the result showed that there was a negative correlation between soluble sugar content and stress time. Therefore, soluble sugar as an index of drought resistance needs to be further studied.
16.7.3 Comprehensive Index (Membership Function Method)
The comprehensive index method can accurately evaluate the drought resistance of potato. The membership function method is to accumulate the membership values of all drought resistance indexes of each variety, calculate its average, and compare among varieties, so as to evaluate its drought resistance capacity (Du et al. 2012). Wang (2014) showed that MDA content is an important index for potato drought resistance evaluation by using the principal component analysis method to analyze the physiological and yield indexes of potato. The membership function method was used to evaluate the drought resistance of potato, and the evaluation results are almost the same as the practice.
16.8 Physiological Response to Drought Stress
Under drought stress, physiological and metabolic indexes of potato, such as soil water content, free proline content, soluble sugar content, chlorophyll content, root pulling force, ATP content, leaf water potential, SOD activity, and MDA content, will change, reflecting the stress status of potato (Yin et al. 2017; Fan et al. 2006). In general, the low concentration of proline indicates the low degree of stress (Ren et al. 2011). Yin and Xiao (2017) believed that drought stress leads to a large increase in proline, soluble sugar and soluble protein in potato leaves, and a decrease in osmotic pressure, which promotes the adaptability of potato plants to arid environment and their drought resistance. Zhao et al. (2018) analyzed the relative amount of photosynthetic characteristic indexes of potato plants under drought stress for 15 days and blank treatment. The results showed that drought stress affected the photosynthetic characteristic indexes of potato plants to a great extent, resulting in the decrease of net photosynthetic rate, transpiration rate, air pore conductivity, and intercellular CO2 concentration of potato plants. Gu et al. (2013) studied the partial and energy metabolism indexes of potato leaves under drought stress and found that under stress, the soluble protein content in leaves showed an increasing trend, the leaf area index showed a decreasing trend, and the soluble protein content would increase with the decrease of water supply. The water loss rate of leaves increased and the drought resistance decreased. On the contrary, the leaf water loss rate decreased and the drought resistance increased (Du et al. 2012). Ding et al. (2011) showed that the drought resistance of the strain was positively proportional to the soluble sugar content of leaves and reached a significant level of 1%, and inversely proportional to the water loss rate and water content of leaves in vitro and reached a significant level of 5%. Liang et al. (2018) believed that drought stress caused a large accumulation of MDA in potato cells. The deepening of drought stress will enhance the activity of POD, so as to eliminate hydrogen peroxide in time to resist adversity.
16.9 Drought Tolerance Gene
With the rapid development of molecular biology, the research on plant drought resistance is also deepening, and many drought resistance genes have been found. By 2014, more than 100 drought-related genes had been found in nearly 500 plants, of which 68 genes were found in Arabidopsis and major crops (Li 2018; Blum 2014), while few drought-related genes were found in potatoes, only WRKY1, DREB1, and SnRK2 (Wang et al. 2017). When potato is under drought stress, many adverse reactions will occur. This stage is an important stage of signal recognition. After being stimulated by the outside world, it can be regulated into transcription factors by signal transduction. After being regulated, the regulatory genes of transcription factors complete the drought resistance process through metabolism (Li 2018; Wang and Guo 2017). At present, many domestic and foreign scholars gradually study the molecular mechanism of potato drought resistance and focus on using modern molecular biology technology and biochemical technology to enhance the drought resistance of potato. Pei et al. (2019) analyzed the transgenic GhABF2 of potato test tube seedlings under drought stress compared with the wild type. The results showed that the transgenic potato with GhABF2 gene could significantly increase the biomass and show strong drought resistance. Li et al. (2014) believed that the SOD activity of P5CS transgenic plants was enhanced under drought stress and salt stress, thus reducing the probability of stress. Yang et al. (2019a) transferred the Haloxylon ammodendron NAC family gene hanac1 into potato and found that it improved the drought resistance of potato by participating in a variety of hormone synthesis and signal transduction and regulating the expression of genes related to downstream stress response (Table 16.3).
16.10 Cold-Tolerant Breeding of Potato
16.10.1 Overview
Potatoes can be planted from 69° north latitude to 50° south latitude and from sea level to 4000 meters above sea level (Hijmans 2003). Potato tubers are rich in protein, vitamins, and dietary fiber. They have high nutritional value and play an important role in solving regional food shortage and eradicating world poverty. Potato has been cultivated in China for 400 years. At present, the planting area is six million hectares and the total output is more than 120 million tons. It has become the largest producer in the world (FAOSTAT 2017). As a cool but not cold-tolerant crop, most potato varieties are not resistant to low temperature and frost. China’s potato ecological planting areas include the first cropping area in the north, the second cropping area in the Central Plains, the single and double cropping mixed cropping area in the southwest, and the winter cropping area in the south. The first cropping area in the north is vulnerable to early autumn frost and sudden drop in humidity, the second cropping area in the Central Plains is vulnerable to early spring frost and late autumn frost, the single and double cropping mixed cropping area in the southwest is vulnerable to late spring cold, and the winter cropping area in the south is vulnerable to low temperature. Low-temperature frost seriously threatens the development of China’s potato industry. For example, in 2008, snow and ice disasters in southern China affected an area of about 409,300 hm2 of potatoes, causing economic losses of up to one billion yuan (Kou et al. 2015). In 2016, the low-temperature and cold wave in Poyang Lake area of Jiangxi Province caused the freezing death rate of potato seedlings to be 75~95%, and the yield was reduced by 40%, resulting in serious economic losses (Zhang et al. 2016). In 2018, the low-temperature and cold wave in Yulin, Guangxi province led to the failure of local potato production to meet the commercial potato standard (more than 100 g), resulting in serious economic losses (Zhu et al. 2020).
When potato plants suffer from low-temperature freezing injury, they will show that the stems are paralyzed and collapsed, and the leaves are dark green and watery. The leaves lose photosynthetic capacity and stop growing, which will seriously affect the potato yield (Li and Jin 2007). Low-temperature stress is accompanied by the changes of cell membrane structure and lipid components, mainly manifested in the changes of unsaturated fatty acid content and lipid peroxidation in plant cells. The lower the content of unsaturated fatty acids, the more prone the cell membrane to the low-temperature phase transition (Theocharis et al. 2012). Lipid peroxidation induced by low-temperature stress decreased the fluidity of the cell membrane and impaired the function of the cell membrane. Freezing injury will cause freezing between or within plant cells, directly destroy the integrity of the cell membrane, deform cells, and lose cell function. Low temperature induced photoinhibition in plants, resulting in the closure of plant stomata, blocking the source of carbon dioxide, resulting in the decline of photosynthetic rate and the inhibition of photosynthetic function.
16.10.2 Cold Resistance Evaluation
16.10.2.1 Identification Method
Vega and Bamberg (1995) first used the field natural frost method to divide the cold resistance of potato into seven grades: Grade 0 represents no damage; Grades 1 to 6 represent slight injury to top leaves, freezing death of a few top leaves, freezing death of most top leaves, freezing death of all top leaves and petioles, and freezing death of all leaves and stems, respectively. The field natural frost method depends on the natural climate environment in the open field, which can directly and accurately reflect the freezing damage of plants under natural freezing conditions (Lin et al. 2020).
Electrolyte leakage method is the most reliable identification method at present by measuring plant cell membrane conductivity and calculating half lethal temperature (LT50) by the logistic equation. It has the advantage of more accurate than physiological and biochemical index determination method (Jiao et al. 2019). The physiological and biochemical index determination method is to directly reflect the cold resistance of plants through the determination of physiological and biochemical indexes, such as soluble sugar, soluble protein (SP), proline, MDA, antioxidant enzyme SOD, CAT, POD under low-temperature stress, and simple and easy to operate. This method has been widely used to identify the cold resistance of pomegranate, kiwi fruit, potato, and wheat (Soloklui et al. 2018; Wan et al. 2001; Pan 2016; Li et al. 2019).
16.10.2.2 Cold-Tolerant Potato Varieties
There are great differences in cold resistance among different potato varieties. The selection of cold-resistant potato varieties is of great significance for the cultivation of potato resistance to low temperature and frost. Potato varieties with strong cold resistance should be selected as far as possible during planting. Li (2008) evaluated 26 materials by using the field natural frost method and selected 13 frost-resistant materials, such as 03079-444, 03079-435, and 03079-343, and 3 frost-sensitive materials, such as Zhongshu 3, 03079-322, and 03088-344. Zhao (2013) treated 108 wild species test tube seedling materials from Solanum chomatophilum, Solanum acaule, and Solanum paucisectum at −3.5 °C for 24 h and screened 27 cold-resistant materials, such as Solanum acaule, and 69 low-temperature-sensitive materials, such as Solanum demissum. Tu et al. (2015) identified the cold resistance of 40 potato materials by using the electrolyte leakage method and the direct evaluation system of cold resistance identification at seedling stage, and screened and obtained 21 potato materials with strong cold resistance, such as Solanum acaule, Solanum paucisectum, and Solanum albicans. Li et al. (2016) identified 65 potato materials by conductivity measurement combined with logistic equation and selected Zhengshu 6, Guinongshu 1, and 4 progeny lines (0712801, 0917-8056, D540 and 0719017) as cold-resistant materials. Pan (2016) identified the cold resistance of 54 potato materials by electrolyte leakage method and selected 5 cold-resistant materials, such as av9, gs393, and 8033 (4) and 8033 (9) and 21 × 11、41 × 25 copies of non-cold-resistant materials, such as 5. Using the method of physiological index determination, it was found that on the fifth day after low-temperature stress, the activities of SP, CAT, SOD, and POD in the leaves of Guinongshu 1 seedlings were significantly higher than those of Favorita, showing strong cold resistance (Deng et al. 2017). The cold resistance of different winter potato varieties in Guangxi was compared. The chlorophyll, relative water content, SP, MDA content, and antioxidant enzyme activity of Lishu 6, Favorita, and Xingjia 2 were measured under 4 °C low-temperature stress. The results of cold resistance were as follows: Lishu 6 > Xingjia 2 > Favorita (Li et al. 2017a, b). Wei et al. (2017) used conductivity leakage method and natural frost method to identify the cold resistance of 116 potato varieties and screened four cold-resistant materials Jinshu 2, Kexin 2, Zhengshu 5 and Zhengshu 6. Yang and Guo (2017) analyzed the changes of leaf relative conductivity, MDA, SOD, POD, and SP contents of 10 potato varieties subjected to low-temperature stress at seedling stage, and comprehensively evaluated the cold resistance by combining principal component analysis, membership function method, and cluster analysis. The order of cold resistance is Lishu 6 > Zhongshu 20 > dianshu 701 > jizhangshu 12 > Qingshu 9 > cooperative 88 > Zhongshu 18 > Diantongshu 1 > normal university 6 > Xuanshu 2. 103 potato materials were identified by conductivity method, field natural frost method, membership function method and cluster analysis method, and 16 cold-resistant materials, such as V9, bs214, gs393, Goutou yam, and Linshu 3 and 21 frost-sensitive materials, such as anti-10, Longshu 5, and sten-1 were obtained (Ding et al. 2019b) (Table 16.4).
16.10.2.3 Physiological Response to Low-Temperature Stress
The cold tolerance of potato seedlings can be improved by spraying growth substances, cross breeding, and chemical mutation. López-Delgado et al. (2018) found that salicylic acid (SA) and H2O2 can mediate the tolerance of potato to low-temperature stress, induce the enhancement of cat enzyme activity, and improve the cold tolerance of potato. Li et al. (2018a) tested osmoregulatory substances and antioxidant enzyme activities by exogenous spraying 0.5 mmol L−1sa on potato seedlings under low-temperature stress. The results showed that exogenous spraying SA could reduce cell membrane damage and resist low-temperature damage by regulating potato osmoregulation and antioxidant capacity. Spraying potato seedlings with three growth substances at concentrations of 10 mg L−1 abscisic acid (ABA), 0.5 mmol L−1 spermidine (SPD) and 0.01 mg L−1 brassinolide (BR) can improve the low-temperature tolerance of potato (Wang et al. 2018). Ding et al. (2019a) found that the cold resistance of potato seedlings was the strongest after spraying 3 mg L−1 ds twice. Huang et al. (2019) mutated potato Favorita callus with ethyl methanesulfonate (EMS), added L-hydroxyproline (L-HYP) for culture and screening, and obtained potato mutants mutated by EMS with cold resistance. Through the determination of physiological indexes of potato plants, more accurate cold resistance resources can be obtained, but most of the potatoes with strong cold resistance are diploid wild species, which cannot be directly used in potato production. Spraying growth substances or chemical mutagenesis through exogenous sources has a positive effect on improving the freezing resistance of potato seedlings. At the same time, it will also lead to a series of problems, such as slow growth of potato, short plant, environmental damage, and waste of resources, which will restrict the development of the potato industry.
16.10.2.4 Cold Tolerance Gene
With the continuous development of molecular biology technology, a series of progress has been made in the research of potato cold resistance-related genes. The damage of low-temperature stress to plants mainly includes destroying the cell membrane and affecting photosynthesis and physiological metabolism. The up-regulation of stearoyl-acyl carrier protein desaturase (SAD) gene expression after potato cold acclimation at low temperature can enhance potato cold stress tolerance (Amiri et al. 2010). Li (2013) isolated the key dehydrogenase gene sad in the pathway of unsaturated fatty acid synthesis from three wild potato species by using transcriptome sequencing technology and successfully overexpressed the sad gene in the cultivated species Zhongshu 8, which improved the cold resistance of Zhongshu 8. Studies have shown that diploid wild species Solanum commersonii has strong cold resistance. Through transcriptome sequencing, it was found that the expression of 855 genes was up-regulated after cold acclimation, mainly including cold resistance genes, such as CBF3, E3 ubiquitin ligase (HOS1), CBF-regulated transcription factor (ICE1), and sumo E3 ligase (small ubiquitin-like modifier E3 ligase, SIZ1) (Aversano et al. 2015). Arabidopsis thaliana cold responsive-element binding factor 3 (AtCBF3) was overexpressed in potato to enhance the antioxidant effect and low-temperature tolerance of potato (Dou et al. 2015). Protoplast sugar and cell wall invertase (CWI) significantly affect the cold resistance of potato mainly by inducing ABA (Deryabin et al. 2016). Transcriptome and metabolome analysis showed that potato spermidine decarboxylase gene (ADC1) was highly expressed under low-temperature treatment. Overexpression of ADC1 resulted in an increase in putrescine content and significantly improved the freezing resistance of potato (Kou et al. 2018). Li et al. (2018a, b) overexpressed Solanum tuberosum CBF1 (StCBF1) and Solanum commersonii CBF1 (ScCBF1) genes in Arabidopsis. Compared with the control, the overexpressed plants of the two genes had stronger low-temperature tolerance, and the effect of ScCBF1 was more significant than StCBF1. Overexpression of CBF1 (SpCBF1) gene of Solanum pinnatisectum increased the expression of cold-responsive genes (COR) in potato, increased SOD activity and SS content, and improved the cold resistance of potato (Zhu et al. 2018). Xie et al. (2019) found that stumirna390 isolated from potato material gs393 was induced by low-temperature stress and inhibited the expression of its target gene ScLRRK1 (lrr-rlk). Che et al. (2020) found that overexpression of potato StSOD1 gene (Solanum tuberosum superoxide dismutase, StSOD1) can regulate antioxidant enzyme activity in potato and improve cold resistance. Overexpression of amylase inhibitor gene (Solanum berthaultii amylase inhibitor, SbAI) can reduce potato starch synthesis to resist short-term low-temperature stress (Slugina et al. 2020). Although a series of potato cold resistance genes have been reported, the molecular mechanism of potato cold resistance is not completely clear and needs further research (Table 16.5).
16.11 Heavy Metal Tolerance Breeding of Potato
16.11.1 Overview
As the fourth food crop, potatoes (Solanum tuberosum L.) are grown worldwide from sea level to over 4000 m elevation (Tabaldi et al. 2009a). Plant species differ in their aluminum (Al) tolerance and potato is inherently tolerant to Al (Little 1988). Potato crops grow well at pH 5.0–6.5, but yield decreases in the soils with a pH value below 5.0 (Castro 1983). In Southern China, most potatoes are planted in red and yellow soil or winter postharvest rice field. The pH value of winter rice field prolonged exposure to the sun quickly drops below 5.0. At that moment, Al ion mainly exists in the form of highly toxic trivalent cation. Al toxicity has become a vital factor affecting potato production. Potato showed a strong preference for cadmium absorption. The accumulation of heavy metals in potatoes was detected by planting potatoes in composted soil with municipal solid waste (Topcuoglu and Onal 2012), and potatoes were more enriched than wheat and barley. The ability of the genus is stronger (Casova et al. 2009). The accumulation of heavy metal cadmium in potato tubers is as high as 0.04–0.20 mg/kg (Fan et al. 2009), and the accumulation law in plants is stem > leaf > fruit > root (Fan et al. 2011). According to the standard, except for fruits, the cadmium content in roots, stems, and leaves of potatoes almost all exceeds the national food safety limit, which has a high ecological risk (Fu et al. 2014).
With the increase of Al concentration (1–10 mg/kg), Mg concentration in the root and stem apex of potato was declined. Low concentration of Al could improve Mg contents in potato varieties Acer and Gleditsia, but was the opposite at high Al concentration. When Al concentration in the solution was increased to 10 mg/kg, Fe accumulated in potato root (Xiao and Wang 2006). The absorption and distribution of Zn, Fe, Mn, and Cu were affected by excessive Al accumulation in roots and shoot of potato clones. With the increase of Al levels, the concentrations of Zn, Mn, and Fe in root decreased linearly in Macaca and SMIC148-A, but increased linearly in Solanum microdontum (Tabaldi et al. 2009b). Meanwhile, the competition of combining site on the root surface between Al and Cu resulted in Cu accumulation in potato root (Xiao and Wang 2006). There was a quadratic relationship in root Cu content among three potato clones (Tabaldi et al. 2009b). Al over 100 mg L−1 decreased the concentrations of chlorophyll and carotenoid in the Al-sensitive potato clones (Tabaldi et al. 2007). Characterized by the production of ROS and RNS, oxidative stress is a toxic mechanism in the Al-sensitive Macaca clone (Tabaldi et al. 2009a). Al toxicity not only depends on Al availability but growth condition and the clone used. The increasing H2O2 concentration of root and shoot was dependent on the concentration and distribution of Al (Tabaldi et al. 2009a). A13+ accelerated the death of potato cells and produced much H2O2，but Al(OH)3 caused higher H2O2 accumulation and did not cause significant death of potato cells. There was dual influence of dissolution rate of Al in soil and its form change on the resistance of potato tuber during soft rot bacteria infection (Shi et al. 2008). Lipid peroxidation was an early symptom induced by Al in the Al-sensitive Macaca clone. Protein oxidation was also observed in roots. Under Al treatment, CAT and APX activity in Macaca roots firstly were decreased and then increased. 200 mg L−1 Al lowered NPSH and AsA concentration in Macaca roots. In potato plants, the cellular redox status was sensitive to Al injury (Tabaldi et al. 2009a). Al interferes with root acid phosphatase (APase) activity in potato clones. The Apase activity in vivo cannot be used to screen Al adaptation of potato clones, because it depends on Al availability, plant organ, growth conditions, and genetic background (Tabaldi et al. 2011).
16.11.2 Evaluation of Heavy Metal Resistance
Lee (1971b) founded that Al tolerance of four potato varieties decreased in the order: Netted Gem > Sedago > Katahdin > GreenMountain. Tabaldi et al. (2007) evaluated Al tolerance of four potato clones. Based on relative root growth and Al content in roots, the screening results arranged from strong to weak as SMIC148-A > Solanum microdontum > Dakota Rose>Macaca. 200 mg L−1 Al significantly lowered relative root growth in Macaca clone, which did not depend on exposure time. Al content in Macaca roots increased linearly in a dose-dependent manner. Therefore, Macaca was considered Al-sensitive potato clones, whereas SMIC148-A was considered to be Al tolerant. Al depressed plant growth and number and yield of tubers for both potato varieties Netted Gem and Sebago. Al treatment lowered the yield of knobby tubers. 20 ppm Al decreased small tuber yield and increased the dose-dependent yield of larger tubers and the specific gravity of the tubers for both potato varieties (Lee 1971a). Therefore, Al accumulation in potato tissues affects the growth and plant yield. The yield per plant, leaf dry weight, stem dry weight, tuber dry weight, and root dry weight were declined under Cd stress. For both varieties, Cd toxicity to different organs ordered as follows: root > tuber > leaf > stem. The resistance of Kexin 1 to Cd was better than that of Favorita (Bai et al. 2012) (Table 16.6).
16.11.3 Physiological Response to Heavy Metal Stress
Through the proliferation of deep roots, the root of SMIC148-A potato clone was growth into the higher Al level soil zones (Tabaldi et al. 2009a). Al avoidance responses differed because of the distinct Al sensitivity in potato clones. Al avoidance was more obvious for the Al-sensitive clone, whereas SMIC148-A had a stronger antioxidant response to Al stress (Tabaldi 2008). The alteration of nutrient element distribution in potato root can improve Al tolerance of potato. Al tolerance among certain potato varieties may be related to the absorptive ability of plant roots to Mg and K (Lee 1971b), which are in accordance with the results of Liu and Jiang (1995). The alteration of shoot Cu concentration mainly occurred in Al-sensitive clones. The higher concentrations of Zn, Fe, and Mn in the roots were associated with Al tolerance in Solanum microdontum (Tabaldi et al. 2009b). Potato roots accumulate more Al and more micronutrients could not be transported to the shoot. So antioxidant systems in roots efficiently scavenged the side effect of Al stress (Tabaldi et al. 2007).
In terms of potato, the enhancement of antioxidant capacity has been identified as an Al-tolerant mechanism. CAT activity in SMIC148-A roots was increased, but APX activity was decreased with the increase of treatment time. Higher level non-protein thiol groups (NPSH) and ascorbic acid (AsA) in root occurred at 120 h. Al supplies could not alter H2O2 concentration in root and shoot in Al-tolerant potato clones (Tabaldi et al. 2007). The transgenic potato expressing tomato Cu-Zn SOD enhanced oxidative stress defense (Perl et al. 1993). With the cadmium concentration increased，the activities of SOD and CAT in the two cultivars of potato leaves increased firstly and then decreased. The content of MDA and proline in the two cultivars of potato leaves increased with the cadmium concentration increased (Zhou et al. 2012).
16.11.4 Heavy Metal Tolerance Gene
Compared with the wild type, the transgenic potato mutant overexpressing PME was more sensitive to Al (Schmohl et al. 2000). Zimmermann et al. (2004) reported that Solanum tuberosum purple acid phosphatase (StPAP1) was expressed more lowly in young leaves, stolons, and flowers. In contrast, P starvation induced the higher expression of StPAP2 and StPAP3 in roots and stem. Due to precipitation of Al phosphates, the relative Al avoidance was associated with the response of local P sources to Al-induced internal P shortage. The gene could be induced by heavy metals Pb2+, Ni2+, Zn2+, Cu2+, and Cd2+, and the StERF10 expression level varied with different times and different tissues. Among the five heavy metal stresses, StERF10 gene was the most sensitive to Ni2+ stress and it was speculated that StERF10 might be involved in the regulation of abiotic stress response in potatoes (Meng et al. 2020a). NAC is one of the unique transcription factor families in plants. The expression of StNAC2 gene was inhibited under high concentration stress and StNAC2 gene could involve in cadmium stress (Meng et al. 2020b). Potatoes had certain tolerance under low concentration of Pb or Cd，and their injuries of active oxygen were alleviated through increasing the activity of antioxidant enzymes and decreasing the membrane lipid peroxidation, but the resistance gradually reduced along with increase of the stress (Li and Liu 2014). As an important antioxidant in plants, glutathione (GSH) can effectively remove the oxidative damage. Glutathione synthetase (GS) is the key enzyme of GSH biosynthesis (Hasanuzzaman et al. 2017). StGS is a cadmium-responsive gene, because it is differently expressed in various organs of potato plants in response to cadmium stress (Tian et al. 2020). Genes containing a heavy metal associated (HMA) domain are required for the spatiotemporal transportation of metal ions that bind with various enzymes and co-factors within the cell. 36 gene members in the StHMA family were identified and divided into six subfamilies by phylogenetic analysis (He et al. 2020). ATP-binding cassette (ABC) transporter proteins without transmembrane domains may play important roles in heavy metal detoxification. A new approach was established to evaluate ABC transporter family functions using microRNA targeted inhibition (He et al. 2021) (Table 16.7).
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He, H., He, LF. (2023). Advances in Potato Breeding for Abiotic Stress Tolerance. In: Singh, S., Sharma, D., Sharma, S.K., Singh, R. (eds) Smart Plant Breeding for Vegetable Crops in Post-genomics Era . Springer, Singapore. https://doi.org/10.1007/978-981-19-5367-5_16
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