Genetic variation in kernel traits under lead and tin stresses in spring wheat diverse collection
Introduction
Pollution with heavy metals is a common problem worldwide due to the increased population and industrialization, excessive use of pesticides and fertilizer, and rapid progress in chemical and petrochemical industries (Chen et al., 2013; Murtaza et al., 2008). In Egypt, a high concentration of harmful heavy metals was reported in the soil of Delta, Asyut, and El-Minya (El-Bady and Metwally, 2019; Mekky et al., 2019; Salman et al., 2019; Shokr et al., 2016). Lead (Pb) is one of these harmful heavy metals that exists with a concentration exceeding the allowed level for human health in soils and irrigation water including wastewater and Nile river water (Elgharably and Mohamed, 2016; Hassan and Elhassan, 2016; Naggar et al., 2014). Furthermore, tin (Sn) contamination, an important component of plastic products and canned food, has been reported widely in agricultural soils and water irrigation (Noubissié et al., 2017, 2016). However, there is not enough quantification and identification of tin in soils to provide a full risk assessment. Generally, the uptake of heavy metals usually occurs by different plant species which alter the plant growth, could cause cell death, and reduce crop yield (Ashfaque et al., 2016; FL et al., 2015). The effect of heavy metals on plant growth could be illustrated by the physiological changes that occur in the plant cell such as increasing peroxidation and membrane lipids and decreasing the activity of antioxidant enzymes (Rizvi et al., 2020). Consumption of such contaminated plant tissues with heavy metals affects human health and causes severe diseases like kidney dysfunction, diabetes, osteoporosis, high blood pressure, cancer, and infertility (Debnath et al., 2019; Morias et al., 2012). For non-smoking people, the main source of heavy metals is contaminated food; mainly cereals, potatoes, and vegetables (WHO, 2011; Wu et al., 2016).
Wheat (Triticum aestivum L.) is the third most widely grown cereal globally. It is the most important cereal crop in Egypt and is the main energy source in the Egyptian diet. Bread is the main staple food in the typical Egyptian diet especially for families living under the poverty level. The consumption of this cereal per Egyptian capita is among the highest in the world. Due to this high consumption of wheat per capita, Egyptian human health is threatened with a high concentration of heavy metals (Bermudez et al., 2011). Pb was found to be among the highest concentrated heavy metal in the kernels of wheat plants irrigated with sewage water (Hassan et al., 2013; Khan et al., 2018; Yu et al., 2016). So, it is very important to reduce these heavy metals content in wheat parts, especially grains to reduce its effect on human health. Many approaches are possible to reduce the concentration of heavy metals in wheat plants such as; using polyelectrolyte-coated fly ash and sophorolipids produced by Starmerella bombicola (Olabemiwo et al., 2017; Qi et al., 2018), reducing plant uptake by increasing the PH of the soil (Naidu et al., 1994) and removing accumulated heavy metals from grains during food processing. However, these approaches are slow, costly and affect the concentration of other useful metals in wheat grains like Calcium (Ca) and Magnesium (Mg). On the other hand, producing wheat varieties that have reduced concentrations of heavy metals combined with high yield and high quality by using breeding programs is a more effective approach (Zaid et al., 2018]; Mondal et al., 2021). Therefore, selection criteria should be effective in improving both targets simultaneously.
Wheat grain yield is a complex trait that is controlled by multiple genes/QTLs interacting with each other and with the environment. Therefore, in recent decades there is an indirect trend to increase grain yield by increasing its components. Kernel length (KL), kernel width (KW), and thousand kernel weight (TKW) have been reported as important components associated with kernel weight in durum and bread wheat (Arriagada et al., 2020; Xin et al., 2020). Previous studies reported that TKW is mainly affected by KL and KW hence both are very important components in increasing grain yield (Campbell et al., 1999). In addition, TKW, KL, and KW have been reported as easy and cheap predictors of milling quality as larger kernels usually have higher flour yield than those smaller kernels (Berman et al., 1996; Marshall et al., 1986; Wiersma et al., 2001). A positive correlation was reported between KL, KW, and kernel area with flour-water dough quality and endosperm separation index and friability (Baker et al., 1999; Breseghello and Sorrells, 2006; Morgan et al., 2000). As a result, selection for higher values of kernel traits under Pb and Sn conditions will help in breeding for high yield, high end-use quality, and tolerant genotypes.
Different selection indices have been used in selecting tolerance genotypes to the different stresses such as mean productivity (MP) (Rosielle and Hamblin, 1981), geometric mean productivity (GMP) (Fernandez, 1992), tolerance (TOL) (Rosielle and Hamblin, 1981), stress susceptibility index (SSI) (Fischer and Maurer, 1978), harmonic mean (HM) (Bidinger et al., 1987), stress tolerance index (STI) (Fernandez, 1992), yield index (YI) (Gavuzzi et al., 1997), yield stability index (YSI) (Bouslama and Schapaugh, 1984), and relative stress index (RSI) (Fischere and Wood, 1979). These indices differ in their ability to distinguish tolerant and susceptible genotypes (Belay et al., 2021; Moursi et al., 2020; Negarestani et al., 2019; Pour-Aboughadareh et al., 2020; Sallam et al., 2018, 2015; Thabet et al., 2021). Previous studies reported that select resistant genotypes based on a single index might be problematic. Thus, it is better to select resistant genotypes based on all indices together. The iPASTIC orders the genotypes based on all indices and provides an average rank for each one of them. Using this rank is very helpful in selecting the resistant/tolerant genotypes (Pour-Aboughadareh et al., 2019).
The objectives of this study are (1) study the genetic variation in kernel traits in spring wheat core collection under controlled and heavy metals conditions (Pb and Sn), (2) identifying the effect of Pb and Sn on kernel traits and some physiological parameters in the flag leaf, and (3) Select resistant genotypes to each metal based on kernel traits.
Section snippets
Plant materials and experimental design
In this study, a set of 103 spring bread wheat genotypes were evaluated for lead and tin stress tolerance. The seeds of the tested genotypes were obtained from the USDA-ARS, United States, and representing fourteen different countries (Supplementary Table 1). Out of these 103- genotypes, fifteen are Egyptian varieties that are usually planted in the Egyptian fields. The remaining 88 worldwide genotypes were evaluated in the Egyptian fields and found to be well adapted to the Egyptian
Genetic variation of kernel traits in the tested genotypes under controlled conditions
The analysis of variance revealed highly significant differences among the tested genotypes for all the studied kernel traits (Supplementary Table 2). The phenotypic variation in KL, KD, KW, and TKW is presented in Fig. 1a–d, respectively. Under controlled conditions, KL ranged from 0.5 mm to 0.9 mm with an average of 6.79 mm. KD ranged from 2.3 to 4.11 mm with an average of 3.13. KW ranged from 2.8 mm to 4 mm with an average of 3.23 mm, while TKW ranged from 25.87 to 63.27 gm with an average
Genetic variation in kernel traits in the tested genotypes
The recent study discussed the genetic variation in kernel traits under Pb and Sn contaminated conditions. Pb and Sn heavy metals are widely accumulated in soils. Unfortunately, not enough studies investigate the effect of these two metals on wheat grain. Therefore, it is very important to provide intensive information about their effect on wheat and identify the tolerant genotypes to these heavy metals. Due to the high cost of measuring heavy metals content in wheat flour, it is not
Conclusion
In conclusion, the effect of Pb and Sn, for the first time, were investigated on kernel traits in a highly diverse wheat core collection. The high significant correlation between TKW and the studied kernel traits confirmed their efficiency to be used as indirect selection criteria for high yielding genotypes under controlled, Pb, and Sn conditions. However, to select tolerant genotypes to both metals together, KL is the best selection criteria that should be used. Our study concluded that there
Author contributions
AM conducted the experiment, performed all the genetic and phenotyping analyses and wrote all the manuscript. A.A.A. helped in soil contamination. M.F.A.D. conducted the physiological analyses. All the authors agreed to be accountable for the content of the work.
Funding
This work was financially partially supported by the cultural affairs and mission sector, the Egyptian government.
Declaration of Competing Interest
The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.
Acknowledgments
The authors would like to thank the staff of the Agronomy research farm, Faculty of Agriculture, Assuit University for their help in conducting the experiment.
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