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- Herausgeber: Bentham Science Publishers
- Kategorie: Wissenschaft und neue Technologien
- Sprache: Englisch
The knowledge of plant responses to various abiotic stresses is crucial to understand their underlying mechanisms as well as the methods to develop new varieties of crops, which are better suited to the environment they are grown in. Environmental Stress Physiology of Plants and Crop Productivity provides readers a timely update on the knowledge about plant responses to a variety of stresses such as salinity, temperature, drought, oxidative stress and mineral deficiencies. Chapters focus on biochemical mechanisms identified in plants crucial to adapting to specific abiotic stressors along with the methods of improving plant tolerance. The book also sheds light on plant secondary metabolites such as phenylpropanoids and plant growth regulators in ameliorating the stressful conditions in plants. Additional chapters present an overview of applications of genomics, proteomics and metabolomics (including CRISPR/CAS techniques) to develop abiotic stress tolerant crops. The editors have also provided detailed references for extended reading to support the information in the book.
Environmental Stress Physiology of Plants and Crop Productivity is an informative reference for scholars and researchers working in the field of botany, agriculture, crop science and physiology, soil science, and environmental sciences.
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Veröffentlichungsjahr: 2021
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FOREWORD
I am delighted to jot down the foreword for this book that deals with plants’ response to numerous abiotic stresses and various approaches and concepts to mitigate their adverse effects. While the demand for food is increasing continuously, agriculture productivity is threatened by various environmental factors, often related to temperature change and warming. The most typical abiotic stresses affecting the plants are high temperature and drought, resulting in unbearable crop losses. Global temperature change further increases the frequency of warmth stress, flood, and drought, which have a negative impact on crop yield and poses a heavy challenge to global food security. Presently, the best priority to satisfy the worldwide food demand is to sustain and improve crop yield.
This book “Environmental Stress Physiology of Plants and Crop Productivity” edited by Prof. Saroj Arora and Dr. Tajinder Kaur presents a broad picture of plant responses to major abiotic stress factors, plants adaptations during stress conditions like the role of antioxidative system and plant growth regulators, biotechnological approaches for abiotic stress tolerance in plants for sustainable agriculture, and the role of secondary plant compounds and signaling molecules to modulate the oxidative stress in plants under stressful conditions. These topics, written by experts in their respective fields, make this book highly useful to the scientific community. It covers a good range of subjects that provide the eader a comprehensive overview of stress tolerance to varied environmental factors affecting plant growth and development.
I congratulate the editors for compiling this publication that provides the reader the simplest way forward in abiotic stress management to enable more productive agriculture. I am sure that the readers within the field of abiotic stress management, agriculture, and biotechnology will find this book very useful.
PREFACE
The human population is increasing worldwide at a much faster rate and is expected to increase from ~7 billion to ~ 10 billion by the end of the year 2050. On the other hand, agricultural productivity is not increasing at the desired rate to feed all the people due to the negative impact of various environmental stresses. Stresses in plants include heat, cold, drought, flooding, salinity, radiations, heavy metal toxicity, and nutrient loss, limiting agricultural productivity. In the present scenario of global environmental change, increasing crop productivity and minimizing the losses in crop yield is a major concern for all nations to cope with increasing food requirements.
The present book “Environmental Stress Physiology of Plants and Crop Productivity” integrates a broad cross section of scientific knowledge and expertise about the mechanisms underlying plant responses against different environmental stressors. It is a timely contribution to a subject that is of eminent importance. Chapter one gives an overview of major abiotic stress factors affecting growth and development in plants. Chapter two and three focus on two major abiotic stressors, i.e., drought and salinity, that adversely affect crop productivity and quality. Chapter four covers factors responsible for temperature variations around the globe, plants’ response to temperature variations, and its impact on crop yield. Chapter five covers the effect of mineral deficiency on plant stress responses. Chapter six deals with the role of nitric oxide in providing tolerance to plants under salt stress. Chapter seven highlights the role of antioxidative defense system of plants in mitigating the harmful effects of excessive production of reactive oxygen species under various types of stress factors. Chapters eight and nine discuss the importance of melatonin and phenylpropanoid, respectively, in mitigating the adverse effects of abiotic stress in plants. Chapter ten comprehensively deals with the role of different plant growth regulators in abiotic stress tolerance. Chapter eleven present an overview of genomics, proteomics, and metabolic approaches, and twelve deals with the advent of new technologies like CRISPR gene technology to develop plant resistance against various environmental changes.
We express our thanks to all the contributors. We would like to thank Prof. (Dr) K. Muthuchelian for writing the foreword. Finally, it is a profound pleasure to thank Bentham Science for taking up the publication of this book. We hope that this book will provide current knowledge on abiotic stress in plants and will lead to new discussions and efforts to deal with various environmental stress factors.
Summary
The knowledge of plant responses to various abiotic stresses is crucial to understand their underlying mechanisms as well as the methods to develop new varieties of crops, which are better adapted. Environmental Stress Physiology of Plants and Crop Productivity will provide a timely update on plants’ response to a variety of stresses such as salinity, temperature, drought, oxidative stress, and mineral deficiencies. The book will focus on how plants adapt to abiotic stress and methods of improving plants’ tolerance to abiotic stresses. The book will be beneficial to scholars and researchers working in the field of botany, agriculture botany, crop physiology, soil science, and environment sciences.
List of Contributors
Abiotic Stress in Plants: An Overview
Sharad Thakur1,2,Ravinder Singh3,Jaskaran Kaur3,Manik Sharma2,3,Kritika Pandit3,Satwinderjeet Kaur4,Sandeep Kaur3,4,*Abstract
Environmental stress is one of the major limiting factors for agricultural productivity worldwide. Plants are closely associated with the environment where they grow and adapt to the varying conditions brought about by the huge number of environmental factors resulting in abiotic stress. Abiotic factors or stressors include high or low temperature, drought, flooding, salinity, mineral nutrient deficiency, radiation, gaseous pollutants, and heavy metals. High salinity, drought, cold, and heat are the major factors influencing crop productivity and yield. The negative impact of various abiotic stress factors is the alteration in the plant metabolism, growth, and development and, in severe cases, plant death. Abiotic stress has been becoming a specific concern in agriculture leading to unbearable economic loss to the breeders. Thus, understanding these stresses will help in achieving the long-term goal of crop improvement, therefore, minimizing the loss in crop yield to cope with increasing food requirements. With this chapter, an attempt has been made to present an overview of various environmental factors that are hostile to plant growth and development, thereby leading to great loss in crop yield.
INTRODUCTION
Plants continuously face unfavorable environmental conditions that affect their growth and development by altering their metabolic activities, eventually leading to plant death. Abiotic stress is defined as environmental conditions that cause alterations in plant growth and development and limit yield below optimum levels [1-3] and can result in unacceptable economic losses. Abiotic stress factors include water stress (drought, flooding, waterlogging), extreme temperatures
(heat, cold, and freezing), too high or too low irradiation, insufficient mineral nutrients in the soil, excessive soil salinity, gaseous pollutants, and heavy metals. However, drought and salt stress pose severe threats to agriculture resulting in loss of crop yield. These threats may become more intense by global climate change and population growth [4]. The ability of crops to achieve their yield potential (maximum possible yield) if all inputs are non-limiting can be affected by a number of stress events. In most agricultural systems, crops hardly reach their yield potential due to stress events, be they abiotic or biotic. Water is often the largest restraint to limiting crop yield. Water deficiency reduces the rate of transpiration and photosynthesis in addition to the absorption of nutrients and water by the root surface, resulting in lower crop yield as compared to yield potential [5]. The average yield of major crop plants is reduced worldwide by more than 50% due to abiotic stress [6, 7]. It is predicted to become even more severe as desertification increases. The present yearly loss of arable area may double due to global warming by the end of the century [8, 9]. Crop production may be reduced to about 70%, with the majority of the crops performing at only 30% of their genetic potential with regard to yield [10]. The problem will become more intense by a simultaneous increase in population growth, creating more pressure on existing cultivated land and other resources [11]. Given the increasing human interference by humans with nature, only 2.75% of the global land area is not affected by some environmental limitations, according to its yearly report by Food and Agriculture Organization (FAO), 2018. It has been projected that more than 90% of the land in rural areas is distressed by various environmental stress factors at some point during the growing season [1].
Abiotic stress exhibits a huge challenge in our goal for sustainable food generation. Drought and rising temperature are two of the main abiotic stressors around the world that reduce crop productivity and influence the ability to meet the food demands of the rising global population, particularly given the current and growing impacts of climate change. Several studies have reported that increased temperature and drought can decrease crop yields by 50% [12]. Salt stress is also one of the major severe limiting factors for crop growth and production, which has been elevated mostly by agricultural practices such as irrigation [13]. More than 6% of the world’s land is affected by salinity. Of the present 230 million hectares of irrigated land, about 45 million hectares, i.e., 19.5%, are affected by salinity. Of the 1,500 million hectares under dry land agriculture, about 32 million hectares (2.1%) are salt-affected to varying degrees [14].
Mineral stress is another constraint in plant growth. Intensive agriculture practices in developed nations have lessened the availability of nutrients in the soil resulting in its low fertility and poor availability of nutrients required for plant growth. However, plants can initiate a number of cellular, molecular, and physiological changes in response to this stress. Plant reactions to abiotic stressors are complex and dynamic; they are both reversible and irreversible. Plants respond to stress in a number of ways depending on the tissue or organ affected by the stress, in addition to its intensity and duration. Some plants complete their life cycle during less stressful periods and hence escape the effects of stress. On the other hand, some plants have evolved stress tolerance, avoidance, or resistance mechanisms that isolate plant cells from stressful conditions [15]. In this chapter, we present an overview of various environmental factors that cause stress in plants and thereby affect their overall growth and yield.
ENVIRONMENTAL CONDITION THAT CAUSE STRESS
Plants are exposed to various types of abiotic stressors like drought, salinity, cold and heat, waterlogging, hypoxia, and anoxia, which have a negative influence on the survival, biomass production, and yields of staple food crops by up to 70%, hence, putting at risk food security worldwide [16-18]. Water deficiency stresses imparted by drought, salinity, and temperature severity are the most prevalent abiotic stresses that limit plant growth and productivity. The response of plants to these abiotic stressors is multigenic, with plants responding to these stressors at the molecular level by regulating the synthesis of various secondary metabolites, which act as physiological buffers to nullify the bad effects of abiotic stressors. To understand the functional dynamics of the stress signal perception and regulation of the associated molecular networks, the stress tolerance capacity and capability of plants are being un-ravelled through high throughput sequencing and functional genomics tools. Plant acclimatisation to abiotic stress is a complex and coordinated response involving hundreds of genes and various signal transduction pathways activated in response to the various environmental factors throughout the developmental period of the plant [19-21].
Hence, a thorough understanding of the molecular pathways functioning in response to the abiotic stress in plants is essential for targeting any improvement in plant biomass or yield. A better and thorough insight into the plant responsiveness to abiotic stress will help in both traditional and modern breeding applications towards improving stress tolerance in plants. Genetic regulatory mechanisms occurring at the level of transcriptional regulation, alternative splicing molecular mechanisms, and the rapid generation of signal transduction regulatory proteins via ubiquitination, sumoylation, phosphorylation, and chromatin remodeling tend to influence complex signal transduction networks that act in turn to regulate processes such as membrane transport, the ascent of sap to maintain cellular ion homeostasis and the synthesis of the secondary metabolites.
Microbiome inhabited by the plants also facilitates the reduction of environmental stress [22].
The most fundamental living system symbiotically closely associated with the plants right from seed on the earth is the microbial life. As soon as a seed comes into the soil to start its life cycle, the surrounding microbial life forms an integral part of plant growth and development. Microorganisms tend to form a symbiotic association with the plant parts like the roots, leaves, and stem. Plant microbiome provides fundamental support to plant life and helps in acquiring nutrients, resistance against diseases, and tolerance of abiotic stressors [23]. Intrinsic metabolic and genetic capabilities of the microbes make them suitable microorganisms to combat extreme conditions of the environment [24, 25]. The microbial interactions with the plants improve the metabolic capability of the plants to fight against abiotic stresses by evoking various kinds of local and systemic responses that are beneficial for the survival and adaptability of the plants to a particular environment [26].
PLANT RESPONSES TO STRESS
Plants are important components of earth. The ultimate source of oxygen on this planet, which makes it habitable for human survival and life, is due to plants. It is not easy for them also to survive on Earth without facing resistance from powers of Earth’s energy. Plants encounter a variety of environmental insults in their lifetime in the form of various stresses. It may be biotic or abiotic stresses. Both biotic and abiotic stressors cause retardation in growth and fresh weight and retard food production from plants [27]. Abiotic stressors include conditions like drought, flooding, salinity, stress due to air, water, soil pollution, nutrient imbalance, radiation, stress related to heat such as high and low temperatures, heavy metal stress etc., is due to unfavorable environmental conditions which include physical, chemical or mechanical factors [28, 29]. These conditions cause severe losses to agriculture and plant growth.
Stress at the cellular and molecular levels cause alterations of different physiological processes of plants, including inactivation and denaturation of enzymes, proteins, displacement / substitution of essential metal ions from bio-molecules, blocking of functional groups of metabolically important molecules, and conformational modifications and disruption of membrane integrity, which further lead to altered plant metabolism, inhibition of respiration, photosynthesis, and alerted activities of several important enzymes [30-32]. Abiotic stress like heavy metal pollution also stimulates the generation of free radicals and reactive oxygen species (ROS) such as hydrogen peroxide, singlet oxygen, superoxide radicals, and hydroxyl radicals and perhydroxyl radical, which disturb the redox homeostasis [31-33]. ROS are extremely reactive and rapidly interrupt normal cell metabolism. Membrane destabilization is generally attributed to lipid peroxidation, due to an increased production of active oxygen species. Many studies have been performed on the enhancement of plant tolerance to oxidative stress by modifying the plant antioxidant defense system [34]. In addition, it is believed that the alterations in antioxidant enzymes may be due to the synthesis of new isozymes or enhancement of pre-existing enzyme levels for the metabolism of ROS [35] (Fig. 1).
Fig. (1)) Various response elicited plant systems during abiotic stress.On the other hand, plants may develop an anti-oxidative system through enhancing anti-oxidative enzymes and antioxidants e.g. glutathione to combat the various abiotic stressors [36, 37]. The defence mechanism of plants towards stress includes enzymatic antioxidative machinery including catalase (CAT), ascorbate peroxidase (APX), glutathione peroxidase (GPX), superoxide dismutase (SOD), glutathione reductase (GR) while also increasing malondialdehyde (MDA) and H2O2 contents in plants [38, 39]. The Asada-Halliwell pathway is documented to be responsible for sequestration of H2O2 through various enzymes. Studies have reported that the antioxidative defence system of plants plays a significant role in sequestration of ROS by reducing the imbalance that arises due to stress [37, 40].
EFFECTS OF ABIOTIC STRESS
Overview of the Effect of Abiotic Stress on Productivity and Yield
Abiotic stress is a serious global problem limiting agricultural production and food security. Plants are exposed to different types of abiotic stressors such as salinity, drought, heavy metals, heat, cold, radiation etc. which poses harmful effects on the health and yield of crops thus affecting global food security [41-44]. The investigated effects of various abiotic stressors are given in the Table 1. The detrimental effect of these stressors continue to increase due to growing urbanization, industrialization, pollution, climate change and leads to nutritional imbalances in the crops by reducing availability of water thus increasing toxicity and reducing products. Abiotic stress damages more than 50% crops worldwide due to their harmful effects [10, 45]. In plants, abiotic stresses disturb the cellular redox homeostasis resulting in the generation of reactive oxygen species (ROS) which leads to oxidative stress [46]. ROS generation can lead to cell damage in different organelles of the cell such as nucleus, mitochondria, chloroplast and peroxisome and ultimately plant cell death [47].
Salt Stress
Salinity stress being one of the major environmental threats causes both ionic toxicity and osmotic stress in plants [57, 58]. Salt stress severely affects the crop yield because of oxidative damage, nutritional disorders, and negative impact on ion imbalance and water relations and thus limits plant growth. Disturbance in cellular ion homeostasis negatively affects the uptake of soil nutrient resulting in nutrition deficiencies in plants under salt stress. It reduces the growth and development of crops by inhibiting several vital biochemical and physiological processes [59-61].
Siddiqui et al. [62] found that salt stress decreases the activities of enzymes of sulfate and nitrate assimilation pathway in plants, and thus increases the demand for sulfur and nitrogen. The accumulation of reactive oxygen species (ROS) in the plants under salt stress results in oxidative stress that leads to cellular damage, such as lipid peroxidation, DNA damage and enzyme inactivation [63, 64]. Moreover, salinity stress also causes membrane disintegration, ion imbalance, metabolic function loss, breakdown of DNA and following cell injury and cell necrosis [65].
Temperature Stress
Like other stresses, high (heat) and low (cold) temperature stress have been a major limiting factor to the growth and development of plants leading to reduced yield of crops. Heat stress leads to oxidative stress, enzyme inactivation, lipid peroxidation, membrane damage, degradation of proteins and ultimately DNA defragmentation in plants [66]. According to various reports, there is an increase in the expressions of various genes in plants under temperature stress [58]. Plants have evolved various molecular and physiological mechanisms to overcome heat stress. High temperature reduced germination potential of the seeds, photosynthetic rate and crop yield. Increased temperature have various harmful effects in the plants during the reproduction period such as loss of functional tapetal cells, formation of dysplastic anther, inhibition of the swelling pollen grains during flowering period, thus leads to the poor shedding of pollen grains as well as the indehiscence of anthers [67, 68]. Heat stress decreased the number of spikes and florets in rice plants as well as seed-set in sorghum [69].
High temperatures cause significant decrease in the growth and net assimilation rate in maize and sugarcane plants [70, 71] as well as reduction in the length of internodes, early leaf senescence and accumulation of biomass in sugarcane [72]. In plants, cold stress affects various biochemical and physiological processes and ROS-homeostasis [66, 73, 74]. High temperature reported to decrease productivity in peanut [75], tomato and rice [76] as well as reduce starch, oil and protein contents in cereals and oilseed crops [77]. High temperature was shown to damage various PSII components in barley and wheat [78] as well as reduce photosynthetic rate in two varities of rice i.e. IR64 and Huanghuazhan [76].
Heavy Metal Stress
Heavy metal stress poses harmful effects on the growth of the plants and induces many physiological changes in plants which results in the reduction of productivity, thus threaten horticultural cultivars. Cadmium toxicity causes changes in metabolic processes, mineral deficiency, osmotic imbalance, inhibited essential elements uptake as well as reduction in growth and photosynthetic efficiency in plants, which ultimately results in reduced productivity [79].
In pea plant, cadmium has been reported to decrease chlorophyll content, plant growth, photosynthetic and transpiration rates as well as damage the antioxidant defense system and cause nutrient imbalance [80]. The roots of alfalfa showed oxidative and glutathione depletion under cadmium and mercury stress [81]. Nickel stress results in the enhancement of O22- and H2O2 contents in the roots of wheat seedlings [74]. Arsenic toxicity leads to the generation of reactive oxygen species (ROS) which induce oxidative stress and harmfully affected primary metabolism in Pistia stratiotes [82].
Droughts and Salinity
There are various morphological, physiological, biochemical and molecular changes that arise due to abiotic stress that adversely affect the growth and productivity of plants [83]. These gross changes in morphology are underpinned by improvements in leaf anatomy. Water deficient leaves usually have smaller cells and higher stomatal density [84]. In another study, comparable effects of high temperature and water deficiency on cell density were reported, but there is limited data available regarding changes in the leaf anatomy in response to high temperatures [85]. Due to hydraulic insufficiency and chemical signalling from the roots to the shoots through the xylem, shoot growth may be limited in drying soil [72]. Growth of plant is impaired by reducing water uptake into expanding cells under water deficit pressure and enzymatically altering the cell wall's rheological properties; for example, ROS activity on cell wall enzymes [86]. High temperature, salinity or drought stress, results in oxidative stress which can cause functional and structural proteins to be denaturated [2]. In Medicago sativa (alfalfa), it was reported that length of hypocotyl, germination capacity and fresh and dry shoot and root weights were reduced by a deficiency of water induced by polyethylene glycol, while the length of root was increased [87].
Drought stress decreased grain yields in barley (Hordeum vulgare), by reducing the number of tillers, spikes and grains per plant and grain weight. Drought stress after anthesis was detrimental to grain yield regardless of the extent of the stress [88]. In maize, water stress decreased yield by delaying silking, thereby increasing the period between anthesis and silking. This trait was strongly associated with grain yield, in particular the number of ears and kernels per plant [89]. Turgor potential, relative water quality, stomatal conductance, transpiration, and water-use efficiency were reduced under drought stress in Hibiscus rosa-sinensis [90]. The degree of stomatal opening of K+-treated plants initially suggested an instant decline in drought-treated sunflower. However, diffusive resistance in the leaves of K+-treated plants remained lower than those that received no K+ at similarly low soil water capacity [85]. A major effect of drought is decrease in photosynthesis resulting from decreased leaf expansion, impaired photosynthetic machinery, premature leaf senescence, and related decrease in food production [91].
Under drought conditions, a rapid decline in photosynthesis was followed by reduced maximum velocity of ribulose-1, Rubisco 5-bisphosphate carboxylation, ribulose-1, 5-bisphosphate regeneration, Rubisco and stromal fructose bis-phosphatase activities, and higher plant quantum efficiency of Photosystem II [92]. The shoot and root biomass, photosynthesis and root respiration was decreased by severe drought. Restricted root respiration and root biomass under extreme soil drying situations can enhance growth and physiological activity of drought-tolerant wheat which is advantageous over a dry cultivar in arid regions [93]. In another study, it was found that cotton crops are very susceptible to the stress of temperature, soil salinity, heat and drought [94]. Mild drought stress during the initial stage can increase root elongation but under long-term water stress conditions root morphological and physiological activities are seriously hampered [95]. Under conditions which are water challenging, plants perceive stress through different sensors involved in signaling response. These are transduced by different pathways where several signaling and transcriptional factors play essential and specific role [96].
The transport of water within a plant occurs under strain, as determined by the availability of soil water and the deficit in atmospheric vapor pressure, causing turgor pressure within cells. Under changing environmental conditions, physiological adjustments which maintain turgor pressure are crucial. Various elements, such as root anatomy, water quality, and soil salts, influence water transport in roots [97]. Specifically, vulnerable to heat stress are reproductive processes involving pollen and stigma viability, pollen tube growth, pollination anthesis and early embryo development [98].
It has also been shown that plant hormones are involved in root and shoot long distance signaling and hydraulic conductivity control. Abscisic acid (ABA) is the most crucial hormone involved in the regulation of abiotic stress tolerance, such as drought, salinity, cold, heat and wounding [99]. Plant production is strongly affected by water deficit. The shoot and root are the most affected on a morphological level, and both are main components of plant adaptation to drought. In response to drought stress, plants typically restrict the number and area of leaves merely to raising the water budget at the expense of yield loss. Since roots are the only source of soil water, root growth, density, proliferation, and size are key plant responses to drought stress [100]. Various environmental conditions that improve the rate of transpiration often increase the pH of the leaf sap, which can encourage abscisic acid accumulation and reduce the stomatal conductance. Also, increased concentration of cytokinin in the xylem sap directly promotes stomatal opening and affects stomata's sensitivity towards abscisic acid [101].
Root growth rates are widely used in cotton crop for estimations crop yield losses. Insufficient soil moisture restricts root growth and development and thus impairs the functioning of the aerial components. Water deficiency in the upper soil profile results in deeper root penetration for further exploration of moisture and nutrients, while excess water in the upper layer results in reduced root penetration [101]. Accelerated abscission of fruits and leaves from drought-stressed cotton crops may be associated with reduction of final yields [102]. Under severe drought conditions, decreased stomatal conductance and metabolic (non-stomatal) damage, such as limited carboxylation, become major photosynthesis limitations [103]. Water and heat stress are reported to degrade proteins, decrease electron transport, and release calcium and magnesium ions from their protein-binding partners [104]. Extended exposure to high temperatures often causes a reduction in chlorophyll content, disintegration of thylakoid grana, increased amylolytic activity, and disruption of transport of assimilates [105].
Floods
Most crops are vulnerable to short-lived flood events, resulting in growth and yield reductions. The leaf structure is influenced by higher temperatures that often cause thinner leaves with a higher leaf area to grow [106]. Another mechanism which enhances plant survival under flooded conditions is to prevent the build-up of potential phytotoxins. A particular type of haemoglobin, known as phytoglobin may play such a role through the detoxification of the nitric oxide produced during root tissue hypoxia. Conversely, phytoglobin can also regenerate NAD+ and thus function as an alternative route to fermentation [107]. Oryza sativa commonly known as rice is highly tolerant to flooded conditions. It is not only capable of germinating without any oxygen, but green leaves and stems capable of elongation mediated by ethylene response [108]. Flooding may have a detrimental impact on physical and chemical properties of soil (structure, porosity, and pH) and microbial functional communities thus impacting soil production levels [109]. The loss of nitrogen (N) from flooded soil can be significant; that along with other deleterious effects on plants can result in lower crop productivity [110]. Soybean growth and grain yield are affected by the flooding from excessive rainfall or irrigation [111].
Flooding primarily prevents the diffusion of gas between the plant and its surrounding area due to physical consequences. Simple exchange of oxygen as well as CO2 cannot easily take place through stomata and underwater cell walls. This results in a lack of oxygen within flooded parts of the plants, which primarily limits heterotrophic, mitochondrial energy production. In addition, poor availability of CO2 in flooded leaves limits photosynthesis. The flooding therefore triggers an energy shortage within plant cells [112-114]. Plants growing in a flooded area with no ongoing photosynthesis will see the concentration of oxygen rapidly decrease, leading to hypoxic condition [115].
CONCLUSION
Plants and crops are exposed to various abiotic stressors like drought, salinity, cold and heat, water logging, hypoxia and anoxia which have a negative influence on its survival, biomass production and yields. However, plants and crops have antioxidative defence systems that enhance their tolerance to oxidative stress and ameliorate the harmful effects of these abiotic factors. Due to the severity of these abiotic factors, stress at the cellular and molecular levels, causes alterations of different physiological processes of plants, including inactivation and denaturation of defensive enzymes. However, to overcome the deterimental effects of these environmental factors certain effective strategies have been deve-
loped that helps to achieve the long-term goal of plant and crop improvement and also minimize the economic loss to the breeders.
CONSENT FOR PUBLICATION
Not applicable.
CONFLICT OF INTEREST
The author declares no conflict of interest, financial or otherwise.
ACKNOWLEDGEMENTS
We are grateful to DST-PURSE, Programme, Department of Science and Technology (DST), New Delhi (India) for supporting this work.
