Metabolic regulation of leaf senescence: interactions of sugar signalling with biotic and abiotic stress responses

Sugars are important signals in the regulation of plant metabolism and development. During stress and in senescing leaves, sugars often accumulate. In addition, both sugar accumulation and stress can induce leaf senescence. Infection by bacterial and fungal pathogens and attack by herbivores and gall-forming insects may influence leaf senescence via modulation of the sugar status, either by directly affecting primary carbon metabolism or by regulating steady state levels of plant hormones. Many types of biotic interactions involve the induction of extracellular invertase as the key enzyme of an apoplasmic phloem unloading pathway, resulting in a sourcesink transition and an increased hexose/sucrose ratio. Induction of the levels of the phytohormones ethylene and jasmonate in biotic interactions results in accelerated senescence, whereas an increase in plant- or pathogen-derived cytokinins delays senescence and results in the formation of green islands within senescing leaves. Interactions between sugar and hormone signalling also play a role in response to abiotic stress. For example, interactions between sugar and abscisic acid (ABA) signalling may be responsible for the induction of senescence during drought stress. Cold treatment, on the other hand, can result in delayed senescence, despite sugar and ABA accumulation. Moreover, natural variation can be found in senescence regulation by sugars and in response to stress: in response to drought stress, both drought escape and dehydration avoidance strategies have been described in different Arabidopsis accessions. The regulation of senescence by sugars may be key to these different strategies in response to stress.     

Distinct Cellular and Organismic Responses to Salt Stress

We have compared metabolic effects of high salinity betweenplants and cell suspension cultures from the facultative halophyteMesembryanthemum crystallinum (common ice plant). This plantshows developmentally-programmed inducibility for a switch fromC₃-photosynthesis to CAM (Crassulacean Acid Metabolism). Themetabolic switch is enhanced by environmental factors such asdrought, low temperature, and, most effectively, soil salinity.CAM induction is dependent on organized leaf tissue and cannotbe elicited by salt stress in suspension culture cells. In contrast,the accumulation of proline [Thomas et al. (1992) Plant Physiol.98: 626] is induced by NaCl in cultured cells as well as inplants and must be considered a cellular response to stress.We have extended our observations to include another trait ofsalt- and low-temperature-stress responses in the ice plant,the accumulation of putative osmoprotective sugars and sugaralcohols. In whole plants the cyclic sugar alcohol, pinitol,accumulates to amounts that approach 1 M during stress, whilein suspension cells no increase in sugar alcohols is observed.The distribution of carbon to different sugars is markedly differentbetween cells and plants under stress. Particularly obviousis the distinction between cell types in the different compositionof sugars and polyols, as exemplified by the epidermal bladdercells of ice plants. Ion contents and the content of sugarsand sugar alcohols of bladder cells indicate that Na⁺, Cl^–,pinitol and an unknown carbohydrate compound provide osmoticpressure in these cells, while organic anion concentrationsare low. With the ice plant, we conclude that cells in culturemimic only partly the stress response mechanisms of intact plantsand we hypothesize that communication between different tissuesis required to mount a complete environmental stress response. ⁴ Present address: Department of Botany Oklahoma State University,Stillwater, OK, 74078, U.S.A.     

Sugar transporters in higher plants--a diversity of roles and complex regulation

Sugar-transport proteins play a crucial role in the cell-to-cell and long-distance distribution of sugars throughout the plant. In the past decade, genes encoding sugar transporters (or carriers) have been identified, functionally expressed in heterologous systems, and studied with respect to their spatial and temporal expression. Higher plants possess two distinct families of sugar carriers: the disaccharide transporters that primarily catalyse sucrose transport and the monosaccharide transporters that mediate the transport of a variable range of monosaccharides. The tissue and cellular expression pattern of the respective genes indicates their specific and sometimes unique physiological tasks. Some play a purely nutritional role and supply sugars to cells for growth and development, whereas others are involved in generating osmotic gradients required to drive mass flow or movement. Intriguingly, some carriers might be involved in signalling. Various levels of control regulate these sugar transporters during plant development and when the normal environment is perturbed. This article focuses on members of the monosaccharide transporter and disaccharide transporter families, providing details about their structure, function and regulation. The tissue and cellular distribution of these sugar transporters suggests that they have interesting physiological roles.     

Unraveling Physiological and Metabolomic Responses Involved in Phlox subulata L. Tolerance to Drought Stress

Metabolic responses are important for plant adaptation to abiotic stress. To investigate the responses of Phlox subulata L. to drought stress, we analyzed its physiological and metabolic changes using gas chromatography-mass spectrometer. Based on the physiological indices, P. subulata L. has tolerance to drought to some degree. Our results showed that there were a total of 30 key metabolites induced by drought stress, including amino acids, organic acids, sugars and sugar alcohols, nucleic acid and its derivatives, and other organic compounds. The glutamic acid-mediated proline biosynthesis pathway is continuously upregulated under drought stress, which could regulate osmotic pressure and maintain intracellular environmental stability. More secondary metabolites are used to increase glycolysis and tricarboxylic acid cycle, to accelerate energy production and to enhance the glutamic acid-mediated proline biosynthesis pathway, which are necessary to increase osmotic regulation. Prolonged drought stress induced progressive accumulation of compatible osmolytes, such as proline and inositol, sugars, and amino acids. Therefore, drought caused systemic alterations in metabolic networks involving transamination, TCA cycle, gluconeogenesis/glycolysis, glutamate-mediated proline biosynthesis, shikimate-mediated secondary metabolisms, and the metabolism of pyrimidine. These data suggest that plants may utilize these physiological and metabolomic adjustments as adaptive responses in the early stages of drought stress. These results deepen our understanding of the mechanisms involved in P. subulata L. drought tolerance, which will help improve the understanding of drought’s effects on plant systems.

     

Physiological response of riparian plants to watering in hyper-arid areas of Tarim River,China

The physiological responses and adaptive strategies of Populus euphratica Oliv.(arbor species),Tamarix ramosissima Ldb.(bush species),and Apocynum venetum L.(herb species)to variations in water and salinity stress were studied in the hyper-arid environment of the Tarim River in China.The groundwater table,the saline content of the groundwater,as well as the content of free proline,soluble sugars,plant endogenous hormones (abscisic acid (ABA),and cytokinins (CTK))of the leaves of the three species were monitored and analyzed at the lower reaches of the Tarim River in the study area where five transects were fixed at 100 m intervals along a vertical sampling line before and after water release.Saline stress dramatically increased soluble sugar concentration of the three species.Differences in sugar accumulation were determined among the species at different transects.The free proline concentration of the leaves of T.ramosissima and P.euphratica showed a proportional decrease with various degrees of elevation of the groundwater table after water release.There was a least correlation between the soluble sugars and proline stimulation in T.ramosissima.It was strongly suggested that T.ramosissima developed a different strategy to accumulate organic solutes to adapt to the stress environment.The soluble sugars and proline accumulation responded to the changes of groundwater table independently:the former occurred under salt stress,whereas the latter was more significant under drought stress.The concentration and the increase in concentration of ABA and CTK involved in stress resistance of the three species were also determined.This increase in the hormone concentration in P.euphratica was different from that of the other two species.Expressed as a function of increase of ABA concentration in leaves,A.venetum and T.ramosissima showed a different solute accumulation in response to groundwater table.There was a significant correlation between ABA accumulation and A [proline] in A.venetum as well as between ABA accumulation and △ [sugar] in T.ramosissima.     

种子脱水耐性与糖的关系

糖类在植物种子中的累积随种子的发育阶段和种子类型不同而不同,并与种子脱水耐性的变化相联系。许多正常性植物种子的发育伴随着某些糖的累积,这些糖的累积已被认为在种子脱水耐性获得中起重要作用。但糖对种子脱水耐性的影响不是单独的,而是与ABA和蛋白质等物质协同作用。种子脱水耐性不仅与糖的种类和含量有关,而且与种子所处的生理状态和发育进程有关。本文综述了种子脱水耐性与糖关系的研究进展。     

Effect of temperature on invertase, invertase inhibitor, and sugars in potato tubers

The accumulation of reducing sugars in potato tubers exposed to low temperatures occurs with concomitant formation of the enzyme invertase. During the initial period of cold treatment when reducing sugars increase rapidly, invertase formation proceeds until the level of enzyme exceeds that of an endogenous macromolecular invertase inhibitor, resulting in a basal invertase activity. As the rate of sugar accumulation decreases and the sugar level becomes nearly constant, total invertase decreases, the basal activity disappears, and a low excess of inhibitor develops. On transfer of cold-stored tubers to warmer temperatures, sugars and invertase decrease sharply and a large excess of inhibitor develops. These changes in sugars, invertase and inhibitor occur reversibly when the tubers are subjected to alternating temperatures.     

塔里木河中游地区3种植物的抗旱机理研究

对塔里木河中游地区沙吉力克、阿其河等断面地下水位进行监测并对胡杨、柽柳、芦苇3种植物的可溶性糖、脯氨酸等生理指标进行测定分析.研究显示：（1）塔里木河中游地区植物生长与地下水位变化关系密切,随着不同断面地下水位埋深程度的增加,植物体内可溶性糖与脯氨酸含量呈增加趋势;（2）在干旱胁迫情况下,植物通过可溶性糖、脯氨酸等渗透调节物质的积累来提高自身的抗旱性;（3）植物叶片可溶性糖和脯氨酸的积累存在互相补偿的关系.研究表明在相同水分胁迫下,柽柳和芦苇对地下水位的变化更为敏感,胡杨的抗旱性较强.     

Asparagine in plants

Interest in plant asparagine has rapidly taken off over the past 5 years following the report that acrylamide, a neurotoxin and potential carcinogen, is present in cooked foods, particularly carbohydrate‐rich foods such as wheat and potatoes which are subjected to roasting, baking or frying at high temperatures. Subsequent studies showed that acrylamide could be formed in foods by the thermal degradation of free asparagine in the presence of sugars in the Maillard reaction. In this article, our current knowledge of asparagine in plants and in particular its occurrence in cereal seeds and potatoes is reviewed and discussed in relation to acrylamide formation. There is now clear evidence that soluble asparagine accumulates in most if not all plant organs during periods of low rates of protein synthesis and a plentiful supply of reduced nitrogen. The accumulation of asparagine occurs during normal physiological processes such as seed germination and nitrogen transport. However, in addition, stress‐induced asparagine accumulation can be caused by mineral deficiencies, drought, salt, toxic metals and pathogen attack. The properties and gene regulation of the enzymes involved in asparagine synthesis and breakdown in plants are discussed in detail.     

The metabolic response of cultured tomato cells to low oxygen stress

The storage of fruits and vegetables under a controlled atmosphere can induce low oxygen stress, which can lead to post‐harvest losses through the induction of disorders such as core breakdown and browning. To gain better understanding of the metabolic response of plant organs to low oxygen, cultured tomato cells (Lycopersicum esculentum) were used as a model system to study the metabolic stress response to low oxygen (0 and 1 kPa O₂). By adding ¹³C labelled glucose, changes in the levels of polar metabolites and their ¹³C label accumulation were quantified. Low oxygen stress altered the metabolite profile of tomato cells, with the accumulation of the intermediates of glycolysis in addition to increases in lactate and sugar alcohols. ¹³C label data showed reduced label accumulation in almost all metabolites except lactate and some sugar alcohols. The results showed that low oxygen stress in tomato cell culture activated fermentative metabolism and sugar alcohol synthesis while inhibiting the activity of the TCA cycle and the biosynthesis of metabolites whose precursors are derived from central metabolism, including fluxes to most organic acids, amino acids and sugars.     

The role of amino acids and sugars in supporting of osmotic homeostasis in maize seedlings under salinization conditions and treatment with synthetic growth regulators

Stress state in plants caused by salinization conditions is characterized by the disturbance of ionic and osmotic homeostasis. The maintenance of the latter is reached by accumulation of osmolytes including free amino acids and soluble sugars in cells. The free amino acid level in the 8-day-old control seedling leaves was higher, than in the roots, whereas the contrary picture was observed in 17-day-old plant tissues. At the same time 8-day-old seedling roots contained more total sugars, than leaves, although the reduced sugar content was nearly a half of the total sugar content. A decrease of both total and reduced sugar levels was observed in 17-day-old seedling tissues. One-day exposure of 7-day-old seedlings to 0.1 M NaCl increased the free amino acid content especially in roots, than in leaves, and the total sugar content in maize leaves, whereas in roots this level remained without changes. The prolongation of salt exposure to 10 days leads to osmolyte content decrease. The seed treatment with Methyure and Ivine intensified accumulation of free amino acids and soluble sugars in the root and leaf tissues under salinization conditions.     

Assessment of proline function in higher plants under extreme temperatures

Climate change and abiotic stress factors are key players in crop losses worldwide. Among which, extreme temperatures (heat and cold) disturb plant growth and development, reduce productivity and, in severe cases, lead to plant death. Plants have developed numerous strategies to mitigate the detrimental impact of temperature stress. Exposure to stress leads to the accumulation of various metabolites, e.g. sugars, sugar alcohols, organic acids and amino acids. Plants accumulate the amino acid ‘proline’ in response to several abiotic stresses, including temperature stress. Proline abundance may result from de novo synthesis, hydrolysis of proteins, reduced utilization or degradation. Proline also leads to stress tolerance by maintaining the osmotic balance (still controversial), cell turgidity and indirectly modulating metabolism of reactive oxygen species. Furthermore, the crosstalk of proline with other osmoprotectants and signalling molecules, e.g. glycine betaine, abscisic acid, nitric oxide, hydrogen sulfide, soluble sugars, helps to strengthen protective mechanisms in stressful environments. Development of less temperature-responsive cultivars can be achieved by manipulating the biosynthesis of proline through genetic engineering. This review presents an overview of plant responses to extreme temperatures and an outline of proline metabolism under such temperatures. The exogenous application of proline as a protective molecule under extreme temperatures is also presented. Proline crosstalk and interaction with other molecules is also discussed. Finally, the potential of genetic engineering of proline-related genes is explained to develop ‘temperature-smart’ plants. In short, exogenous application of proline and genetic engineering of proline genes promise ways forward for developing ‘temperature-smart’ future crop plants.     

Sugar signalling and gene expression in relation to carbohydrate metabolism under abiotic stresses in plants

Sucrose is required for plant growth and development. The sugar status of plant cells is sensed by sensor proteins. The signal generated by signal transduction cascades, which could involve mitogen-activated protein kinases, protein phosphatases, Ca²⁺ and calmodulins, results in appropriate gene expression. A variety of genes are either induced or repressed depending upon the status of soluble sugars. Abiotic stresses to plants result in major alterations in sugar status and hence affect the expression of various genes by down- and up-regulating their expression. Hexokinase-dependent and hexokinase-independent pathways are involved in sugar sensing. Sucrose also acts as a signal molecule as it affects the activity of a proton-sucrose symporter. The sucrose transporter acts as a sucrose sensor and is involved in phloem loading. Fructokinase may represent an additional sensor that bypasses hexokinase phosphorylation especially when sucrose synthase is dominant. Mutants isolated on the basis of response of germination and seedling growth to sugars and reporter-based screening protocols are being used to study the response of altered sugar status on gene expression. Commoncis-acting elements in sugar signalling pathways have been identified. Transgenic plants with elevated levels of sugars/sugar alcohols like fructans, raffinose series oligosaccharides, trehalose and mannitol are tolerant to different stresses but have usually impaired growth. Efforts need to be made to have transgenic plants in which abiotic stress responsive genes are expressed only at the time of adverse environmental conditions instead of being constitutively synthesized.     

Detection of sugar accumulation and expression levels of correlative key enzymes in winter wheat (Triticum aestivum) at low temperatures

Carbohydrate accumulation is common in frost-resistant plants, and many enzymes participate in this process. The sugar content and expression levels of metabolic enzymes related to sugar biosynthesis in response to drops in temperature were measured in two cultivars of winter wheat (Triticum aestivum) with different cold tolerances. The results indicate that the two cultivars examined, Dongnongdongmai 1 and Jimai 22, accumulated high levels of carbohydrate before November 4 (above 0°C), and that accumulation decreased as temperatures fell. However, this decrease was more modest in Dongnongdongmai 1, which had a higher sugar content. Sucrose and fructose were the main soluble sugars, indicating an important role in freezing tolerance. Gene expression studies revealed that expression of the genes encoding chloroplastic enzymes was significantly upregulated in the tillering nodes. Expression upregulation of TaSS and TaTPT may be helpful for sugar accumulation before November 4.     

The Implication of Manganese Surplus on Plant Cell Homeostasis: A Review

Manganese management in plant cells is important for providing cells with appropriate development conditions, as both manganese deficiency and excess interfere with redox homeostasis and activate oxidative stress in cells. Excessive amounts of Mn affect morphological and anatomical parameters but the precise mechanism of manganese toxicity has not been fully understood yet. This review presents the impact of Mn on plant metabolism. We discuss the role of this element in the reduction and oxidation, as well as the alterations appearing in the cells as a result of excessive accumulation of manganese. The closest attention is paid to the transport of Mn into the cells and its interaction with other essential elements. We also summarize the observed alterations in physiological and biochemical properties of plant cells, with regard to the influence of Mn on defense mechanism initiated in its presence. Finally, this review recapitulates possible cell defense strategies under excessive manganese accumulation.

     

Impact of mild heat treatments on induction of thermotolerance in the biocontrol yeast Candida sake CPA-1 and viability after spray-drying

Aims: The objective of this study was to examine the induction of thermotolerance in the biocontrol agent Candida sake CPA‐1 cells by mild heat treatments to enhanced survival of formulations using spray‐drying. The possible role of heat‐shock proteins (HSPs) biosynthesis in induced thermotolerance and the role of sugars and sugar alcohols were also determined. Methods and Results: Studies were conducted on C. sake cells grown in molasses medium and exposed to mild temperatures of 30 and 33°C during mid‐ (16 h), late‐exponential (24 h), early‐ (30 h) and mid‐stationary (36 h) growth phases. The effect on viability was determined both before and after spray‐drying. Cycloheximide and chloramphenicol were used to examine the role of HSPs and HPLC was used to analyse the accumulation of sugar and sugar alcohols. The results indicate that both temperatures induced thermotolerance in cells of C. sake. Mild heat‐adapted cells at 33°C in the early‐ or mid‐stationary phases had survival values after spray‐drying significantly higher (P ≤ 0·05) than nonadapted cells. However, viabilities were not high enough to be considered for commercial use with values up to 17%. HSPs were not implicated in thermotolerance acquired by mild heat‐adapted cells as similar viabilities were obtained in the presence of protein inhibitors. Little change was observed in sugar and sugar alcohols with an increase in glucose and arabitol in some treatments. Conclusions: This study suggests that it is possible to induce thermotolerance in biocontrol yeasts such as C. sake. However, this does not improve survival of cells exposed to spray‐drying sufficiently to consider this a suitable formulation method for this biocontrol agent. HSPs, sugars and sugar polyols were not directly responsible for induced thermotolerance in yeast cells. Significance and Impact of the Study: This type of information can be effectively applied to improve the viability of cells in the process of formulation.