Global climate change manifested in average annual temperature rise and imbalance of most natural factors, such as changes in annual mean rainfall, air humidity, average temperature of cold and warm months, soil quality, etc., lead to climatic zones displacement. All these have a significant impact on agricultural production in total, including cotton growing. Cotton is one of the most important technical crops in the world. However, it is very sensitive to environmental changes. The influence of abiotic stresses (high temperature, changes in the mean rainfall and soil salinity) causes a dramatic decrease yield of this crop. Moreover, temperature anomalies and climatic zones displacement cause a change in the area of pathogens and pests distribution, which also reduces the cotton yield. One of the possible ways to increase the cotton yield under the influence of abiotic and biotic stresses is the development of new resistant varieties, using both classical breeding methods and genetic engineering achievements.
- global climate change
- abiotic and biotic stresses
- cotton breeding
- genetic engineering
Because cotton is a subtropical plant, it is well adapted to survive with dry and hot environment . Despite this, cotton nevertheless reacts to an environmental change such as temperature and rainfall in instance. Long-term exposure of negative factors such as a drought, salinity, and temperature stress causes a significant decrease of yield and fiber quality [4, 5]. Such negative effect on cotton is due to the fact that drought, salinity, and temperature stress cause osmotic imbalance, membrane disorganization, growth decrease, inhibition of cell fission and reproduction; this also leads to decline of photosynthesis level and hyperproduction of reactive oxygen species (ROS) [6, 7].
In addition to abiotic stresses, the cotton production is greatly influenced by biotic factors, such as pests and diseases that also cause a significant (up to 10–30%) reduction in yield . At the same time, global climate changes responsible for temperature factors and climatic zone displacement also affect their development, geographical distribution, pathogenicity or injuriousness .
In this regard to these threats to cotton production, breeders are facing the important task of new cotton varieties resistant to abiotic and biotic stresses. However, this problem-solving by the classical genetics methods has become complicated due to the resistance traits generally having multigenic nature with a complex type of inheritance . Additionally, the breeding oriented on resistance is further complicated by the “bottle-neck” effect such as narrow genetic basis typical for cultivated cotton . Nevertheless, these disadvantages may be successfully overcome by the use of genetic engineering methods: transgenesis, RNA interference, and genome editing approaches.
In this chapter, we would like to analyze and summarize information about increase of cotton resistance to abiotic and biotic factors using genetic engineering approaches.
2. Increasing resistance to abiotic stresses
Abiotic stresses are a direct consequence of climate change. The world increase of temperature is primarily caused by carbon dioxide effect, i.e., its content in the atmosphere. The increase in the average annual temperature is the cause of increase of water evaporation from the soil, which directly leads to osmotic (by drought) and salt stress. One of the features of abiotic stresses is their simultaneous exposure. In other words, they have usually a similar effect on plants and defense mechanisms appearance in plants .
Abiotic stresses may affect cotton upon all development stages and lead to significant decrease in both yield and quality of cotton fiber [12, 13]. Thus, an increase of temperature at 2–3°C from the optimum can decrease biomass and yield, as well as increase fiber micronaire . Drought and salinity also cause a decrease in the yield and quality of cotton fiber [6, 14].
In this regard, increase of a cotton resistance to abiotic stresses will reduce a negative effect and can raise the yield and quality of fiber. In this chapter, we consider the impact of abiotic stresses on the morphological and physiological parameters, as well as the mechanisms of resistance development and methods for increasing the adaptive potential of cotton to negative environmental factors.
2.1 Influence of abiotic stresses on morphophysiological parameters in cotton
Influence of abiotic stresses on cotton plants manifested in various forms of morphophysiological and biochemical changes, which reduce yield and fiber quality of cotton [6, 13, 14]. They negatively affect both morphological (seed germination, plant height and architecture, length and area of root system, leaf area, shoot and root biomass, boll development) and physiological parameters (chlorophyll content, photosynthetic efficiency, transpiration rate, stomatal conductance) [6, 13, 14].
In addition, prolonged exposure to abiotic stresses leads to a decrease in yield and fiber quality. Yield reduction manifested by both a decrease in the number and weight of bolls and fiber yield [6, 13, 14]. At the same time, this negative effect on yield is explained by a decrease in the activity of catabolic processes, including photosynthesis [6, 13, 14]. The fiber quality reduction manifested in a decrease in fiber length and an increase in micronaire. Such influence of abiotic factors on one of the most important agronomic traits of cotton is caused both by reduction of carbohydrate synthesis due to reduction of photosynthesis activity and by disruption of elongation process due to changes of membrane permeability and organization of microtubules and cytoskeleton [13, 14].
Disruption of photosynthesis under abiotic stresses is associated with an increase in ion permeability of chloroplast thylakoids and a decrease in chlorophyll levels, as well as inhibition of the activity of the key enzyme of carbohydrate synthesis 1,5-bisphosphate carboxylase [13, 14, 15, 16].
2.2 Mechanisms of resistance to abiotic stresses
To reduce the negative impact of abiotic stresses in plants, including cotton, they have developed some adaptations on physiological and molecular level.
Molecular defense mechanisms against abiotic stresses include accumulation of osmolytes (proline, betaine, and soluble sugars), changes in activity of antioxidant system reducing level of ROS, regulation of cell ion balance and hormonal activity, as well as changes in activity of heat stress proteins [6, 7, 13, 14, 17]. Let us consider each mechanism separately.
The excessive accumulation of Na+ in the cytoplasm also results in the accumulation of Ca2+, which interacts with SOS3/SCaBP8, activating the serine/threonine protein kinase SOS2. Then, SOS2 phosphorylates SOS1, which increases Na+/H+-antiporter activity, restoring the ion balance in the cell and enhancing salt tolerance [7, 20, 26]. SOS3/SCaBP8-SOS2 also regulates the activity of other transporters involved in ion homeostasis: K+- and Na+-transporters, vacuolar Na+/H+-exchanger (NHX), vacuolar H+-ATPases, and pyrophosphatases (PPase) [18, 20, 26].
Thus, plants have numerous mechanisms to promote abiotic stresses resistance. The genes mediating these defense mechanisms may be potential objects of interest for enhancing the adaptive potential of plants to environmental stress conditions.
2.3 Improving the adaptive potential of cotton to abiotic stresses
A significant decrease in the yield and fiber quality under the influence of abiotic stresses assigns a task for breeders to create cotton varieties resistant to these stresses. To solve this problem, it can use the methods of classical breeding, methods of molecular selection, and genetic engineering. Let us consider the application, advantages, and disadvantages of these methods.
Presently, various strategies, including genotyping by sequencing (GBS), SNP arrays, and genome-wide association study (GWAS), as well as populations of recombinant inbred lines (RIL) and backcross inbred lines (BIL), are used to improve the efficiency of QTL mapping . Thus, 95 loci that associated with salt tolerance in
In addition, the use of meta-analysis allows improving the accuracy of QTL mapping associated with abiotic stresses. For example, this approach has identified 23 stress tolerance QTL clusters on 15 different cotton chromosomes: c3, c4, c5, c6, c7, c11, c14, c15, c16, c19, c20, c23, c24, c25, and c26 .
Summarizing the above, the use of molecular markers and associative mapping data can significantly reduce the time to breed resistant cotton varieties.
Application of transgenic approaches also allows increasing cotton resistance to drought and salinity. Many transcription factors, regulating the activity of functional genes, can influence drought and salt tolerance in cotton . Thus, overexpression of transcription factor
Increase in defense capacity of cotton due to increase level of osmoprotectants and activity of antioxidant enzymes and ion antiporters also enhance the adaptive resistance of the crop to abiotic stresses . Overexpression in cotton of
Regulation of hormonal status by overexpression of their biosynthesis genes can also increase the adaptive potential of cotton resistance to salt and osmotic stress. Thus, overexpression of isopentenyltransferase (
In this way, the application of transgenic methods makes it possible to effectively increase cotton resistance to abiotic stresses. However, those approaches are limited by the legislative regulation of GMO in many countries, according to this all transgenic crops obliged to undergo a full cycle of biosafety assessment .
RNAi is one of promising approaches both for studying of resistance genes and developing new cotton varieties resistant to abiotic stresses [10, 48]. For example, the use of VIGS-mediated RNAi revealed that R2R3-type
Application of GE approaches to increase the adaptive potential of cotton in accordance to abiotic stresses is currently quite limited. However, there are successful applications of GE in cotton. For example, the target editing of
Summarizing the above, it should be noted that presently, marker-associated selection and transgenic methods have the greatest importance in breeding of cotton resistant to abiotic stresses.
3. Improving resistance to biotic factors
Biotic factors (insect pests and pathogens) are among the most important factors that reduce cotton productivity [4, 8, 51]. For example, losses of cotton yield from pests may be up to 84%  and due to pathogens up to 30% . As in the case of abiotic factors, global climate change leads to a shift of climatic zone, affecting the growth, development, and spread of insect pests and pathogens . As results, this leads to the emergence of new pests and pathogens in these areas.
In this regard, improving plant resistance to biotic factors allows effectively control of pests and pathogens to reduce yield losses. In this part, we are looking at the characteristics of the main pests and pathogens, as well as a natural defense mechanisms and methods of improving cotton resistance to them.
3.1 Characteristics of major pests and pathogens of cotton
The second group includes sap feeding insects that damage phloem: boll weevil (
In addition, soil nematodes can also cause a significant cotton yield reduction . Nematodes parasitizing on cotton include the root-knot nematode (
The viral diseases include cotton leaf curl disease (CLCuD) and (CLCrD) . CLCuD is caused by begomoviruses that lead to leaf injury (swollen veins, leaf curl, enation, and stunting). When affected in the early stages of development, there is a significant reduction in yield . The cotton leaf curl virus (CLCrD) affects the leaves resulting in leaf discoloration and vein hypertrophy, leaf curl, shortening of internodes, and growth stunting. The infestation degree depends on the stage of plant development .
Bacterial blight of cotton is one of the most serious diseases causing significant yield losses [8, 53]. Disease results from infection by
3.2 Mechanisms of resistance to biotic factors
The long coevolution of cotton and insect pests and pathogens has resulted in mechanisms to reduce the damage from biotic factors. Molecular mechanisms of pathogen resistance include the activation of resistance genes (R-genes) in response to exposure. R-gene activation triggers a large number of intracellular cascades leading to the synthesis of protective substances that reduce the damage by pathogens [54, 55]. Morphological and chemical defense mechanisms have been developed in cotton to reduce the pest influence degree [56, 57]. Let us in more detail consider mechanisms of resistance to insect pests and pathogens correspondingly.
Terpenoids, flavonoids, tannins, and anthocyanins are among the secondary metabolites providing direct protection of cotton plants from insects . Terpenoids are the most studied protectors of cotton. Terpenoids synthesized in cotton include gossypol, hemigossypol, hemigossypolone, and heliocides H1, H2, H3, and H4 contained in small subepidermal and intracellular pigment glands . Cotton terpenoids have direct toxic effect on insect pests including
It should also be noted that damage of cotton by pests and pathogens causes induction of terpenoid biosynthesis by activation of JA-, SA-, and ethylene-dependent signaling pathways [56, 58]. These pathways activation occurs due to elicitors, which, interacting with specific receptors, lead to an increase in intracellular Ca2+. This in turn activates calcium-dependent proteins, including Ca2+-dependent protein kinases (CDPKs) . CDPK, by phosphorylating proteins and changing gene expression patterns, activates mitogen-activated protein kinases (MAPK), leading to JA and SA formation, on the one hand, and the ethylene pathway, on the other .
Thus, plants have a various mechanisms that provide resistance to pests and pathogens. Genes mediating these defense mechanisms may be potential genes for improving cotton resistance to biotic factors. In addition, the control of genes of the causative agents themselves, playing an important role in their life activity, may also be of potential interest.
3.3 Improving cotton resistance to biotic factors
Biotic factors (pathogens and pests) are one of the main reasons for significant yield losses (up to 84% due to insects and up to 30% for pathogens) in agriculture [4, 8, 51]. At the same time, strategies to control infestations are an increase in the internal defense mechanisms of plants or introduction of pathogen-targeted constructs into the genome [62, 63]. Methods of classical breeding, molecular breeding, and genetic engineering are used to develop new varieties that are resistant to the impact of biotic factors. Let us consider the application, as well as advantages and disadvantages of each of these methods.
Such interspecific crossing for the purpose of transfer wilt resistance genes is complicated by different type of these traits inheritance in
Furthermore, a meta-analysis of the consensus map of Cotton Marker Database (CMD) based on
The obtained data of a QTL mapping can be successfully used in further MAS and genomic breeding programs.
Thus, vegetative insecticidal proteins (
In order to create varieties resistant to fungal pathogens (
In accordance with above, the application of transgenic technology is currently the most used and commercially successful for creating pest and pathogen resistant crops. However, the most serious disadvantage of this technology is the need for long-term biosafety assessment of transgenic cotton to minimize risks of human health and the environment .
The use of HIGS to the genes encoding proteins that play an important role in the life maintenance of insect allows the development of cotton IR lines. Thus, silencing of cytochrome P450 gene of insect monooxygenase (
Another approach to improve cotton resistance to biotic stresses is virus-induced gene silencing (VIGS) of the host genome . Thus, VIGS-mediated suppression of
Summarizing these, RNAi is a promising approach to develop cotton varieties resistant to biotic stresses. However, the application of this approach is limited by high probability of effect on nontarget organisms and complexity of cotton genome, due to tetraploidy .
Thus, summarizing the data above, transgenic methods are currently the most used and commercially successful strategy for developing of new insect pest and pathogen-resistant varieties.
4. Conclusion and future perspectives
Global climate change has a significant impact on cotton production through the complex impact of abiotic and biotic factors, reducing yields and fiber quality [4, 8, 13, 14, 51]. This poses a task to breeders of developing new cotton varieties that are resistant to abiotic and biotic stresses. To challenge it, breeders use both classical and molecular breeding methods and genetic engineering.
By developing cotton varieties resistant to abiotic stresses, molecular breeding methods are more often used, while genomic transgenomic methods improve resistance to insect pests and pathogens [8, 14]. However, the use of modern genetic engineering approaches, including cis- and intragenesis methods, is limited by the complexity of the genome of cultivated tetraploid cotton species. Therefore, the application of RNAi and GE methods to obtain cotton varieties resistant to abiotic and biotic stresses is currently insignificant [8, 14].
In addition, the insignificance of using molecular breeding methods to create pest and pathogen-resistant cotton varieties should be noted. This is due to the insignificant number of mapped insect and pathogen resistance loci in the cotton genome [59, 60, 61, 62, 63, 64, 65, 66, 67, 68, 69, 70].
Fundamental understanding of molecular and genetic mechanisms underlying cotton resistance to abiotic and biotic stresses will allow application of cis- and intragenesis methods as well as RNAi and GE technologies in new resistant varieties development. Thus, the genes encoding DRE-binding protein 1 (
Studying the mechanisms of interaction between the host plant and insect pests or pathogens, as well as the molecular and genetic basis of life support functions of causative agents, will allow more successful use of the HIGS, RNAi, and GE technologies to suppress key genes and cisgenesis technologies to enhance the host plant defense mechanisms. Genes encoding Vacuolar-type ATPase (
Thus, modern molecular biology technologies have great potential to reduce the negative effect of global climate changes on cotton yield and fiber quality.
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