Abstract Trehalose is a nonreducing disaccharide composed of glucose molecules connected by αglycosidic bond. This soluble substance plays an important role of protecting green algae and other lower plants from stress. It can help plants cope with extreme environments such as severe cold， drought and high salinity， regulate the stomatal conductance and water utilization rate of plants， and participate in the growth and metabolism regulation of plants as a signal molecule. As an impermeable cryoprotectant， trehalose is widely used in the refrigeration protection of various animal cells and tissues due to its nontoxicity and high efficiency. According to the research results at home and abroad in recent years， the protection， regulation and mechanism of trehalose on plant tissues and animal cells were summarized， so as to provide a theoretical basis for the further development and utilization of trehalose.
　　Key words Trehalose; Abiotic stress; Signal molecule; Cryoprotectant; Action mechanism
　　 The full name of trehalose is αDglucopyranosylαDglucopyranoside， which is a nonreducing disaccharide formed by condensation of two glucose molecules through a hemiacetal hydroxyl group. The molecular formula is C12H22O11with the molecular weight of 342.3， and the molecular structure is shown in Fig. 1[1]. The molecular structure of this symmetrical disaccharide is far more stable than other small sugar molecules such as maltose， sucrose， glucose， etc.， making it an important component in moisturizing and cell activitymaintaining cosmetics， as well as a unique food ingredient that prevents food degradation， maintains fresh flavor and enhances food quality.
　　Trehalose is present in all living organisms， serving as an emergencymetabolite and signal molecule. It is synthesized in a large quantity in cells in high temperature， freezing， radiation， drying， high osmotic pressure and other adverse environments， and is rapidly used as an energy substance after the crisis is relieved[2]. Trehalose can help bacteria， fungi， lower plants and invertebrates to resist environmental stress and ensure longterm survival in extreme environments such as high salinity and acidity[3-4]， thus protecting the original morphological structure of the living body[5]. Therefore， the protective property is one of the most important functions of trehalose in addition to its use as a sweetener， which can longtermly maintain the stability of various tissues and biomacromolecules such as biological membranes and proteins in extreme environments including abiotic stresses[6-7]， to protect the structure of the two intact， without degradation or denaturation. This kind of protection has attracted much attention， and the research on its action mechanism is controversial and endless. In this paper， we summarized the regulation of trehalose in plant tissues and its mechanism of enhancing plant stress resistance， and the mechanism of trehalose on animal cell cryoprotection， so as to update theoretical research progress and application scope of trehalose in planting， breeding and refrigerated transport. The paper will provide a theoretical basis and reference for the indepth development of trehalose. 　　Regulation and Stress Protection Mechanism of Trehalose in Plant Tissues
　　Trehalose as a metabolite regulates the growth and development of plants
　　In microorganisms and invertebrates， trehalose is mainly involved in physiological processes such as storage and transportation of carbohydrates and stress protection; trehalose in plants is not only directly involved in metabolism， but also participates in the metabolic regulation and gene expression regulation of plants as an important metabolite of plant growth and development[8]. Its metabolic precursor， trehalose6phosphate （T6P）， can maintain the balance of carbon content in plants， delay leaf senescence， and affect fruit setting rate. The SnRK1 signaling pathway involving T6P regulates the metabolism of plant respiration， starch synthesis， starch and sucrose in two ways， and responds to the growth environment and growth stage of different plants correspondingly to ensure the growth and development of plants and simultaneously improve the adaptability of plants to adverse environmental conditions such as drought and cold damage[9-10].
　　Trehalose as a stress protectant enhances the ability of plants to resist abiotic stress
　　Abiotic stress refers to the adverse effects of any abiotic factors on plants under specific circumstance such as salt， alkali， drought， high temperature， severe cold， flooding， lack of minerals and unfavorable pH. Among them， salinealkali and drought are the two main stress factors that restrict plant growth. Under abiotic stress conditions， trehalose can act as a carbon source and a stress protectant to improve the stress resistance of plants， reduce the damage of plant tissues caused by stress and maintain the normal growth of plants[11].
　　Trehalose enhances the adaptability to high salinity stress in plants
　　Salt stress can affect the growth and physiological metabolism of plants. During seed germination， a higher salt concentration in the soil leads to the reduction of the germination rate， germination index and vigor index of the seeds， and finally inhibits the germination and seedling growth. During the growth and development of the plants， salt stress will cause the plants to develop slowly， inhibit the growth and differentiation of plant tissues and organs， advance the development of plants， and ultimately inhibit plant growth. In addition， salt stress can disrupt the normal respiratory metabolism of plants and destroy the synthesis of proteins in plants[12]. Therefore， improving the salt tolerance of plants is of great significance for promoting high yield and stable yield of crops. 　　Studies have confirmed that the transformation of trehalose synthaserelated genes into rice， wheat and other plants that do not synthesize trehalose can improve the adaptability of these plants to salt stress， and immersing plants with exogenous trehalose can also alleviate the inhibition of salt stress on seedling growth. Yan et al.[13]studied the effects of low concentration trehalose on the growth and physiological characteristics of Yangmai 19 seedlings under salt stress. The results showed that the soaking treatment with 2 mmol/L trehalose could improve the synthesis and accumulation of dry matter in Yangmai 19 seedlings， promote the synthesis of carbohydrates in Yangmai 19 plants， alleviate the damage of chlorophyll in seedlings caused by salt stress， and reduce the inhibition of salt damage on leaf photosynthesis， and as a result， plants can adapt to salt stress more quickly. It can be seen that trehalose can be used as an inducer of plant stress adaptability， inducing plants to adapt to later salt stress through gene expression and physiological changes; and at the same time， as a signal molecule， it can induce Yangmai 19 plants to synthesize certain amino acids and absorb potassium ions to balance the damage caused by the accumulation of sodium ions， to thereby improve the adaptability of plants to salt stress.
　　Trehalose enhances drought resistance of plants
　　Extended growth is the result of interactions between the environment and metabolic activities within plants， and water availability is dominant among various plantrelated environmental factors. Whether for cell division and differentiation or volume expansion， it is related to water absorption， solute accumulation and cell wall loosening. Any factor that directly or indirectly affects one of the three can affect plant growth. In the shortwater environment， plants reduce leaf area by reducing the growth rate of leaves and detaching old leaves in a short period of time to reduce transpiration and water loss， and longterm water shortage will cause the decrease in growth rate of roots， the inhibition of activities in stems， the sluggishness of elongation， the reduction of plant height and the thinning of stalk. Drought can also cause different degrees of disturbance and damage to the plants carbon cycle， nitrogen metabolism and information transfer and other metabolic processes[14]. Therefore， improving the viability of plants after drought stress can help to reduce crop yield losses. 　　In the experiment of detecting the expression of trehalose synthetase gene （MeTPS13） in cassava under drought stress by realtime quantitative PCR[15]， when cassava was subjected to drought stress， the upregulated expression of trehalose synthetase gene in roots was the most obvious， indicating that trehalose plays an important regulatory role in cassava when subjected to drought stress. The gene was transcribed into the leaves of Nicotiana benthamiana， the trehalose content in the leaves of the plants was measured. It was found that the MeTPS13 gene was transcribed in the receptor plants and began to express and synthesize trehalose. The drought resistance test confirmed that the waterretaining capacity of the transgenic N. benthamiana strains under drought stress and the recovery capacity after rehydration were enhanced， the electrolyte leakage rate was lower than that of the control， the seed germination power was stronger than the control， and the natural drought resistance ability was enhanced[16]. It can be seen that trehalose is of great significance to the improvement of drought resistance in plants.
　　Trehalose increases the resistance of plants to high temperature stress
　　High temperature stress destroys photosynthetic apparatus and reduces photosynthetic capacity easily， thereby inhibiting photosynthesis and causing crop yield reduction. During the growing season of crops， the crop yield will drop by 17% for every 1 ℃ increase in the average temperature. Organisms will generate a heat shock reaction under heat stress conditions to respond to the increase in temperature， and have certain heat resistance characteristics. However， with the degree of stress increasing， high temperature will destroy the chloroplast structure， degrade chlorophyll， reduce the solubility of CO2， and ultimately affect the photosynthetic rate of plants. The continuous high temperature directly affects the vitality of respiratory enzymes， causing irreversible inactivation， which inhibits the respiration of plants. Furthermore， when the temperature rises and stresses the plants， the stomatal opening on leaves decreases， the transpiration decreases， and the transport of mineral ions by plants and the demand for water are reduced. In addition， high temperature can cause varying degrees of damage to the cell membrane system， osmotic pressure and antioxidant systems in plants[17]. Therefore， high temperature stress is a major environmental stress factor that limits plant distribution， growth and productivity. 　　Under high temperature stress， plant cells produce excessive amounts of superoxide anion （O2-）， hydrogen peroxide （H2O2） and other reactive oxygen species （ROS）， which can damage cellular DNA， proteins， lipid membranes， etc.， causing lipid peroxidation[18-21]. Studies have shown that moderate levels of reactive oxygen species act as signaling molecules in cells to protect against diseases or regulate physiological processes such as pathogen defense and programmed cell death. However， when the concentration reaches the toxic level， if it is not removed in time， it will cause oxidative damage to the body. During the resuscitation of Selaginella tamariscina， the antioxidant system content increases with dehydration and decreases with rehydration[22]， which is exactly the timely response to reactive oxygen species. Trehalose can effectively defense against the damage by these hydroxyl radicals. It has been found from experiments that when the trehalose phosphate synthase 1 gene （TPS1） is overexpressed in tobacco and tomato， the plants significantly improve their tolerance under oxygen stress. The level of trehalose at the milligram numerator level can protect the superoxide dismutase from heat shock inactivation[23]. This protection mechanism can be well promoted in the agricultural production of crops such as wheat to improve the resistance against high temperature and oxidative stress in crops.
　　Trehalose enhances cold resistance of plants
　　When plants suffer from the damage of low temperature higher than 0 ℃， physiological dehydration causes the biological membrane lipid to change from liquid crystal phase to gel phase， resulting in pores and cracks in the membrane. Then， the permeability of the membrane increases， the electrolyte and other soluble substances in the membrane are extravasated， and the intracellular enzyme reaction system and substances are out of balance， causing a series of metabolic disorders， which cause serious damage or death of the tissue. Deng et al.[24]found that after 2 d of treatment with 0.1% trehalose， rice seedlings showed the intracellular electrolyte leakage rate much lower than that of the control at low temperature and the amylase activity in the cytoplasm much higher than that of the control， and their recovery was also faster than the control[24]. Therefore， when rice is dehydrated at low temperature， trehalose may form a similar waterreplaced layer structure on the surface of cell membrane， thereby maintaining the integrity of the cell membrane structure during lowtemperature dehydration， protecting the intracellular enzyme reaction system and material balance， and endowing the plants with stronger cold resistance. 　　The protective effect of exogenous trehalose on plants in the stressresistant environment should be related to their higher glass transition temperature （Tg） and lipid membrane interaction[25]. When plant individuals and tissues surfaces respond to high temperature or cold environment， the inherent characteristics of trehaloses small free volume， restricted molecular mobility and resistance to mutual separation and crystallization protect the sample surrounded by it from various irreversible damages such as structural damage and cell dehydration. When the individuals respond to harsh environments such as salinealkali land， floods and mineral deficiency， the inherently symmetrical stable structure of trehalose helps the sample to stabilize the osmotic pressure inside and outside the tissue， avoiding disturbances in the internal environment. When the plant encounters a waterdeficient environment such as drought， the hydrophilic end of the trehalose and the lipid membrane cooperate to protect the cell membrane structure intact and avoid mechanical damage such as deformation.
　　Yiwen WANG et al. A Review on the Protection Mechanism of Trehalose on Plant Tissues and Animal Cells
　　Trehalose as a Cryogen Enhances the Survival Rate of Cryopreservation of Animal Cells
　　The cryopreservation of sperm is an important technical means to ensure the reproductive capacity of male animals. A large amount of active oxygen is produced during cryopreservation， which would destroy cell membrane structure and motility and is fatal to sperm[26]. Trehalose has the characteristics of stabilizing lipid membrane， protein and biofilm system. During the cryopreservation of sperm cells， trehalose acts as a protective agent to reduce the formation of intracellular ice crystals and maintain protein structure and significantly improve the mobility of sperm cells and the stability of acrosome and cell membrane， and thus can reduce sperm cell aberrations or lesions[27]. Therefore， the addition of antioxidants to the cryoprotectant helps protect sperm cells from reactive oxygen species and improve the resuscitation survival rate of sperm cells. At present， trehalose has been added to the cryopreservation of sperm cells of various mammals such as rabbits[26]， rams[28]， wild boars[29]， goats[30]and bulls[31]， for the improvement of the resuscitation survival rate of cells.
　　In addition to germ cells， trehalose is also used in the low temperature protection of protists and other cells[32]. The latest research results show that trehalose is present in all living organisms[33-34]and is an emergency metabolite. It will be synthesized in the cell when the cells are in high temperature， freezing， radiation， drying， high osmotic pressure and other adverse environments， and can be quickly decomposed as an energy substance after the crisis is relieved. Therefore， the protective function becomes the most important function of trehalose in addition to its use as a sweetener， and it can maintain the stability of various biological tissues and biological macromolecules such as biological membranes and proteins for a long time[35]. The research on the mechanism of trehalose on animal cell protection is mainly reflected in the following two categories： first， vitrified trehalose inhibits the production of lipid membrane kinetic energy， prevents lipid membrane molecular movement， and maintains the integrity of biofilm system due to its viscous characteristics[36-37]; and second， a certain concentration of trehalose specifically binds to lipids， protects liposomes from harmful phase transitions in freezedried environments， and maintains biological membrane permeability and homeostasis[38-39]. After years of repeated research， scientists have found that glassy trehalose does not change the phase transition temperature of lipid membrane dehydration and a high concentration of protective agent produces cytotoxicity easily[40]， and it is firmly believed that only inhibiting the membrane lipid phase transition and ensuring the selective permeability of the cell membrane are the main factors for the protective function of trehalose. 　　As for how small sugar molecules play a protective role of avoiding phase change in the biofilm system， scientists have successively proposed the water replacement hypothesis， the priority exclusion hypothesis and the hydration force action[41]. In the water replacement hypothesis， it is assumed that the living body is in a dry and waterdeficient environment， and the cell membrane is completely dehydrated. The water molecules originally bound to the polar head of the phospholipid are replaced by sugar molecules， allowing the hydrophilic head group to interact with the sugar molecules to maintain the original membrane lipid morphology[42-44]. For example， when there is lateral pressure （0.1-25 MPa） and the trehalose concentration is 2 M[45]， or under the conditions such as increased monomolecular membrane surface tension[46]， some trehalose molecules can displace water molecules that are connected to the hydrophilic group of the lipid membrane through hydrogen bonding， thereby specifically binding itself to the polar head of the lipid membrane and maintaining the hydration state of the lipid membrane to prevent the lipid membrane from phase transition. However， none of the tests described in the above papers were completed under complete water loss conditions and the monomolecular membrane type was insufficient to represent the multilayer membrane structure in the real biological membrane system. At the same time， the experimental conclusions are limited to some inferences or possible reasons， and these speculative conclusions are hardly convincing. The priority exclusion hypothesis is usually used to explain the interaction between trehalose and protein macromolecules. This theory confirms that proteins are more liable to bind to water molecules through molecular dynamics simulation experiments. Trehalose is used to stabilize the protein structure in the natural state， so as to avoid the degeneration of its chemical potential[47]. However， the tripeptide molecules constructed by such simulation experiments differ greatly from the protein structure in actual solutions， and the main component of biological membranes is the phospholipid bilayer rather than the protein molecules， so the theory cannot strongly support the membrane protection mechanism of trehalose. Another explanation， hydration force action， is the most convincing action mechanism research theory： when the water molecules on the hydrophilic side of the lipid membrane fluctuate， the amphiphilic polymer will naturally produce tension to debond the water molecules， and thousands of water molecules are embedded in the polar surface of the bilayer. Such tension is called hydration force by scientists. The hydration force between multilayered lipid membranes has an action range as long as 20 angstroms or more， and is mainly used to prevent crossrecognition of different lipid membranes due to excessive proximity to each other. In the dehydrated state， increasing hydration force increases the lateral pressure inside the lipid bilayer， causing adjacent lipid molecules to approach each other， causing the liquid phospholipid bilayer to become a gel state. At this time， the sugar molecules （sugar ring ∶lipid=1∶1） located between the membrane systems， by virtue of their nonspecific osmotic effect and volume structure， can effectively reduce the hydration force between membrane lipids and the compressive stress generated due to the pressure on membrane lipids， which in turn reduce the transition temperature required for the phase transition of the membrane and maintain the liquid lipid membrane spacing as much as possible， thereby achieving the purpose of protecting biological membranes[48]. Therefore， the hydration force effect suggests that trehalose can effectively protect frozen organisms at a certain concentration. This protection is mainly derived from the nonspecific osmotic pressure and volume effect of trehalose， while the relationship with its binding to the hydrophilic end of lipid membranes in place of water molecules is not large. 　　Studies on the cryoprotective mechanism of trehalose on biomacromolecules such as proteins or interaction with lipid membranes usually require electron microscopy， Xray diffraction techniques， and electron paramagnetic resonance for the observation of the molecular movement locus of lipid membranes and small molecular sugars at low temperatures and the evaluation and calculation of corresponding kinetic parameters， so as to analyze their movement locus and changes[49]and finally explain the possible protection mechanisms. Recent kinetic studies have found that in the aqueous solution of trehalose， the number of water molecules around the protein molecules is significantly reduced， and the trehalose surrounds water molecules， forming a cage structure， which reduces the movement rate of the water molecules trapped therein and avoids crystallization of the protein structure during cooling[50]， thus producing a cryoprotective effect. This new protection mechanism also denies the priority exclusion hypothesis.
　　Summary and Prospect
　　The plant trehalose signaling pathway is thought to be the centrum linking metabolism to the signaling network of growth and development. It regulates the growth and development of plants by mediating internal and external signals. Its precursor T6P is directly involved in various plant physiological activities. Gradually clarifying the T6P signal conditioning mode， molecular mechanism and plant signal response can allow more clear understanding of the multiple adaptability of plants to stress.
　　The increase of soil salinity has become a major threat in agricultural planting in recent years. Trehalose plays a very good role in protecting plants from salt stress. Low concentrations of trehalose can reduce the accumulation of sodium ions in cells， and high concentrations prevent chlorophyll deficiency caused by sodium chloride and protect the integrity of plant roots[51-53]. When plants are subjected to salt stress， the expression level of trehalase is reduced， so that the trehalose accumulates in the body and exerts the protective property of the disaccharide to resist high salt corrosion.
　　Droughttolerant plants usually store large amounts of trehalose. Such plants can suspend metabolism for several years until they encounter water. During metabolic stagnation， trehalose is used to protect cell membranes， proteins， and other cellular components[54]. However， in crops， the content of trehalose is very small， and almost no change or difference in concentration is detected. Therefore， the exogenous addition of trehalose by transgenic technology[55]or the biosynthesis of trehalose by trehalose synthase[56]for the improvement of plant stress resistance is currently a more effective means of plant protection. The genetically modified rice cultivated by the researchers can effectively resist drought， cold and salt， and has high yield and high output value[57]. Wang et al.[58]introduced Agrobacterium carrying the Arabidopsis thaliana trehalase gene interference vector （iTre1285） into N. alata by the leaf disk infection method applying the RNA interference technology. They successfully interfered with the expression of trehalase gene， and inhibited the activity of trehalase in N. alata， thereby achieving the effects of reducing the degradation of trehalose， increasing endogenous trehalose content， improving the ability of transgenic plants to resist stress， effectively protecting crops against environmental disasters， and increasing crop yield. These applications have significant economic benefits for planting， especially in response to the effects of cold and dry climates on crop cultivation in the north. 　　When plants respond to temperature and pressure， their endogenous trehalose will undergo different forms of changes during the metabolism process， and complex physiological and biochemical reactions will be triggered in plants， such as the increase and decrease of transcription factors and protein molecules， changes in the contents of carbohydrates and lipids and secondary metabolites and changes in membrane structure and composition. At this time， trehalose will combine with other solute molecules to resist severe cold or high temperature environment. The proteomic data analysis showed that trehalose phosphate synthase 5 （TPS5） interacts with the transcriptional coactivator MBF1c protein at high temperature. MBF1c protein is a key regulator of plant resistance to high temperature， and TPS5 deletion may cause plants to be sensitive to high temperature. In the low temperature environment， the high expression of AtTPPA gene will increase the contents of trehalose and trehalose6phosphate[59-60]. It can be seen that trehalose participates in plant metabolism and various physical and chemical reactions when plants respond to extreme cold or high temperature weather， helping plants to cope with extreme temperatures.
　　Trehalose is widely known as a protective agent in the cryopreservation process of animal and plant cells or tissues， but because it is a disaccharide molecular structure that cannot rapidly diffuse and arbitrarily enter and leave cells， it fails to replace dimethyl sulfoxide or glycerol as an independent protective agent. At present， mediating trehalose to get through the cell membrane[61-63]by means of a carrier such as a polymer material or the electroporation technology  to allow it to evenly distribute inside and outside the cell has become a protective agent application field which is of great concern in the current research. Overcoming this problem can not only reduce the osmotic pressure caused by trehalose cryopreservation solution on cells and improve the resuscitation survival rate of cryopreserved cells， but also effectively solve the transportation and preservation problems in medicine， ecological breeding， food processing and other industries. It is expected to realize the replacement of dimethyl sulfoxide with trehalose as an independent new nontoxic protective agent that is applied to cryopreservation industries such as cell， tissue and enzyme preparation protection[64-65]. Related discussions on properties and functions of trehalose on higher plants， vertebrates and industrial enzymes in metabolic regulation， stress resistance， pathogen resistance， toxicity， and cryopreservation protection are still continuing[66-67]. Continuously exploring the action mechanism of trehalose in the process of animal and plant protection will have great significance in the development， application and promotion of the cryoprotectant industry. 　　References
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　　Editor： Yingzhi GUANG Proofreader： Xinxiu ZHU