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MECHANISMS OF DETOXIFICATION TOLERANCE TO HEAVY METALS IN WHEAT
Wu Liuliu
PhD student
Sumy National Agrarian University, Sumy, Ukraine ORCID: 0000-0001-2345-6789 Tao Ye PhD student
Sumy National Agrarian University, Sumy, Ukraine ORCID: 0000-0002-4675-1294 Zhatova H.
PhD (Agricultural Sciences), Professor Sumy National Agrarian University, Sumy, Ukraine ORCID: 0000-0002-8606-6750
ABSTRACT
As a non-essential nutrient element, cadmium in soil and water can be absorbed and accumulated by crops, affecting the normal growth and development of plants, and then causing serious phytotoxic reactions in a variety of physiological levels, thus affecting the health of animals and humans through the food chain, causing great harm to human health. This paper discusses the mechanism of plant resistance to heavy metal detoxification, studies the molecular mechanism of cadmium absorption, transport and exportation of wheat, reduces the accumulation of cadmium in crops, the treatment of cadmium pollution and the creation of wheat varieties with low accumulation of cadmium, which is of great significance to ensure food safety and food safety.
Keywords: wheat, cadmium, absorption, transport, distribution, tolerance mechanism, molecular mechanisms.
Soil contains excessive or harmful heavy metals, absorbed by plant roots to the plant, in the process of long-term heavy metal stress, plants through various resistance mechanisms and regulatory mechanism to reduce or avoid heavy metal poisoning, allow it to grow in the high concentration of heavy metals in the environment to complete its development process, this defense mechanism is called detoxification tolerance mechanism. Since it is difficult to control the accumulation of heavy metals in soil environment, it is necessary to master the detoxification mechanism and defense system of plants to heavy metals, and to understand the way that plants absorb, transport and accumulate heavy metals, so as to better avoid the heavy metal stress of plants [1-2]. Tolerance and detoxification mechanisms in plants can be divided into two categories: internal tolerance and external rejection. The internal tolerance mechanism is the complexation and chelation of some substances in plants with heavy
metals[3], Limiting heavy metals to some specific tissue parts of plants can reduce the effectiveness of heavy metals and alleviate the toxic effects of heavy metals on plants. The mechanism of external rejection is that plants prevent heavy metal ions from entering the plant cells or expel excessive heavy metals from the cells to avoid accumulation in the cells. These detoxification mechanisms are not independent, but mutually rein-forcing[3]. So the patient detoxification mechanism of heavy metals by plants, separation and accumulation of cadmium absorption or cloning of functional genes, reveal the accumulation of low accumulation or not of grain crops cultivated on the molecular mechanism of the absorption, transport and accumulation of cadmium, can clear the key process plants absorb cadmium, cadmium accumulation in the crop resistance control, reduce the consumption risks of heavy metals.
Detoxification of plant root exudates
The first barrier for cadmium to enter the plant is the root system, which is the most critical part to reduce its toxic effect. When the root system is under cadmium stress, some organic acids and sugars will be secreted by the root system[4], it can form soluble complex with cadmium and other heavy metal ions, reduce the efficiency and mobility of heavy metal ions, inhibit the transport of heavy metal ions, so as to reduce the absorption of heavy metals by plants. Studies have shown that under heavy metal stress, plant roots change the form and availability of heavy metals in rhizosphere soil by regulating the composition of organic acids with low molecular weight, so that plants can adapt to the external environment [5]. Other studies have shown that maize root tips can secrete viscous substances with strong affinity, which reduces the mobility of metal ions, and thus most metal ions are retained outside the roots[6]. Many microorganisms also play a significant role in plant detoxification[7]. kai[8] Studies have shown that plants can automatically adjust the pH gradient distribution in the rhizosphere environment when exposed to aluminum poisoning, and aluminum deposition around the root system, thus reducing aluminum entry into the plant body. In conclusion, the detoxification of plant root exudates plays a connecting role in coping with heavy metal stress.
Fixation and dynamic response of cell wall
Plant cell wall is the necessary part and primary site for plant root to absorb cadmium[9]It plays an important role in plant tolerance and detoxification. The cell wall has negatively charged binding sites that bind positively charged metal ions[3], adsorbed or complexed heavy metal ions, and prevented cadmium and other heavy metals from entering the cell interior;Plant cell wall can improve the ability to accumulate and detoxify heavy metals by adjusting its components. Zhang Guojun et al. found that heavy metal ions can enter cell protoplasts in cell walls to protect and reduce the damage of heavy metal cadmium to plant cells[10]. Krzeslowska[11] It was found that under the stress of Cd and other heavy metals, the activity of pectin methyl ester in plant cell wall increased, and the pectin content increased and rearranged in space, which significantly improved the absorption and accumulation capacity of plant cell wall to heavy metals[12]. Other scholars have found that increasing the contents of cysteine protein and hemicellulose in root cell wall can significantly increase the adsorption and fixation ability of plant cell wall to heavy metal ions[13]. In conclusion, cell wall is involved in the transport process of heavy metal ions in plant cells, and the fixation and adsorption of cell wall can reduce the influx of heavy metal ions into the cytoplasm.
Selective permeability of cytoplasmic membrane
The plasma membrane is also a natural barrier for material exchange in plant cells and a key part of blocking heavy metal ions from entering cells. The surface of plasma membrane has strong electronegativity, which can adsorb positively charged cadmium ions on its surface, thus affecting the transmembrane transport of cadmium ions in cells. Studies have shown that plasma membrane transporters can transport metal ions
to the extracellular level and play an important role in the selective transport of metal ions[11, 14], ABC membrane transporter is the main representative of membrane transporter. It has been reported that a metal-excreting protein OsPDR9 located on the plasma membrane can transport heavy metal Cd to vacuoles, which is closely related to the detoxification of heavy metal[15]. Heavy metal transporters, such as AtPDR8, AtHMA2 and AtHMA4, are involved in excreting heavy metal Cd from the cytoplasm and transporting it to the cell wall to precipitate heavy metals outside the cell membrane[16-21].
Compartmentalization of vacuoles
Vacuole compartmentalization refers to the ability of plant cells to isolate heavy metal ions within vacuoles. When cadmium passes through the plant cell wall and enters the protoplast through the plasma membrane, it accumulates in vacuoles[22, 23]. Vacuole is rich in organic acids and can form complex with heavy metal cadmium and reduce its toxicity[24]. Shim etc[25]. The expression of vacuolar transporter gene ScYCF1 in poplar seedlings could improve their ability to repair Cd contaminated soil. Plant research invented chelating peptide (PCs) can combine with heavy metals such as cadmium form a complex, enter the vacuole and plant combining chelating peptide, formation of lower tox-icity of high molecular weight compounds, and the membrane transporters HMT1, HMT2, transferred to the vacuole in heavy metal chelate form, in the physical section of cadmium in the immobilized and isolation, Alleviate the toxicity of heavy metals to plants [26-28]. Vacuolar compartmentalization is considered to be an effective way to degrade cadmium toxicity.
chelation of metal-organic ligands
Chelation of heavy metal ions in plants is mainly through inducing the synthesis of metal ligands in plants to form metal-organic ligands, so as to change the mobility and utilization of heavy metal ions and reduce their toxicity to plants [29-31]. Metallothionein (MT) and phytochelin (PC) are mainly studied proteins that chelate heavy metal ions[32, 33]. After some heavy metals pass through the cell wall and cell membrane into the cytoplasm, they can form complex stable che-lates with organic acids, proteins and other substances, and transform into non-toxic or less toxic combination forms, alleviating the toxic effects of heavy metals on plants[34]. When plants are exposed to external cadmium stress, PCs, MT and Cd combine to form nontoxic compounds, which reduce the concentration of in-tracellular free heavy metal ions, thus alleviating cadmium toxicity of rice[34, 35].
Antioxidant effect
The antioxidant system in plants is a widely available detoxification strategy. The antioxidant system in plants can protect cells from oxidative stress by scavenging free radicals produced by heavy metal stress[36-39]. Studies have shown that scavenging free radicals produced by heavy metal stress can improve tolerance of plants to heavy metals[7]. Under heavy metal stress, rice produces a variety of antioxidant defense systems, which scavenge oxygen free radicals and prevent plant cells from being damaged[40-42]. Wang Haiou etc[43]. It was found that under copper and cadmium stress, wheat
leaves produced a large amount of GSH to effectively remove reactive oxygen species and free radicals and prevent membrane lipid peroxidation, suggesting that GSH plays a major role in leaf detoxification. Therefore, the study of antioxidant system is of great significance to understand the physiological mechanism of plant stress adaptation and heavy metal detoxification mechanism.
Heavy metal transporters
Heavy metal transporters are involved in heavy metal uptake and transport in plants and play a key role in tolerance mechanisms such as organic ligand chela-tion and vacuolar compartmentation. Heavy metal transporters can transport heavy metal ions out of the cytoplasm or localize them to specific organelles, or transport heavy metals from non-plastids or organelles to the cytoplasm for detoxification of heavy metals[3]. Atp-binding cassette transporters (ABC transporters) are widely distributed in prokaryotes and eukaryotes. They mainly bind and hydrolyze ATP to release energy to achieve substrate transport across membranes. The ABCB transporter gene mediates the transport of heavy metal ions such as cadmium, lead and aluminum, and improves the tolerance of plants to heavy metals[44]. Many authors have found that heavy metal excretion proteins associated with vacuolar compartmentaliza-tion in plants can transport heavy metals to vacuoles. Amtp1 and Ahmtp1 transporters can transfer excess Zn2 +To take the excess zinc out of the cell, or from the cell2 +Compartmentalization to reduce Zn2 +Damage to cells[45-47]. Other studies have shown that the heavy metal absorbing proteins AtNRAMP1, AtNRAMP3 and AtNRAMP4 can transport heavy metal ions such as Cd from vacuoles to the cytoplasm[48-49]. Moons et al. demonstrated that OsPDR9 transported heavy metal cadmium to vacuoles[50]. Other studies have shown that proteins Athma1 and Athma6 can transport excess Zn and Cu from cytoplasm to chloroplast[51-52]. Proteins Athma7 and Athma8 can transport excessive Cu from cytoplasm and chloroplast matrix to golgi apparatus and thylakoid[53, 54]. The Zinc-iron transporter (ZIP) family (such as IRT1, Osirt1, Osirt2) is involved in the absorption and chelation of Cd and other heavy metals in plants[55-57]. Thus, for different kinds of heavy metal transport protein, under the stress of different heavy metals, transportation way and the way of tolerance is differ, the combined transport way how, in the same kind of plants of different heavy metal detoxification of heavy metal ion transporters exists between joint coordination effect, are subject to further study.
To sum up, it is an effective way to improve heavy metal pollution to further study the mechanism of heavy metal detoxification tolerance, use advanced molecular research methods to isolate heavy metal resistance genes and analyze the causes of key plants' heavy metal resistance[58]. To summarize and analyze the detoxification mechanism of heavy metal transporters, antioxidant enzyme system, cell wall and vacuole in plants, in order to provide reference for future detoxification mechanism and tolerance mechanism of plants.
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ПРОДУКТИВНОСТЬ СЕЛЬСКОХОЗЯЙСТВЕННЫХ КУЛЬТУР ЗАВИСИМО ОТ СПОСОБОВ ОБРАБОТКИ ПОЧВЫ В УСЛОВИЯХ ПОЛЕСЬЯ УКРАИНЫ
Земляной Е.А.
магистр Бобовский Б.Л.
магистр Панченко А.Н.
магистр Галицкий Л. С.
магистр Васильчук В.А. магистр
Полесский национальный университет, Украина
PRODUCTIVITY OF AGRICULTURAL CROPS DEPENDING ON THE METHODS OF SOIL TILLAGE IN THE CONDITIONS OF POLISSYA OF UKRAINE
Vasylchuk V.
master Halitskyi L. master Bobovskyi B.
master Zemlianyi Ye. Master
Panchenko A.
Master
Polissya National University, Ukraine
АННОТАЦИЯ
Одна из ключевых задач современного сельского хозяйства - эффективное управление почвенной экосистемой, предотвращение распространения процессов деградации почвы и сохранение его энергетического потенциала. В этом смысле целью исследования было изучение возможности улучшения основных агрофизических и гидрофизических показателей пахотного слоя светло-серой лесной почвы как компонента управления плодородием легких почв Полесья и формирование продуктивных агроценозов.
