Antioxidants, Oxidative Damage and Oxygen Deprivation Stress: a Review

PATHWAY OF ETHYLENE BIOSyNTHESIS 156 Methionine as an Intermediate ....... 157 S-Adenosylmethionine as an Intermediate 158 I-Aminocyclopropanecarboxylic Acid as an Intermediate . ......... 158 Methionine Cycle ........ 161 Conversion of l-Aminocyclopropanecarboxylic Acid to Ethylene 164 REGULATION OF ETHYLENE BIOSYNTHESIS 167 Regulation in Ripening Fruits and Senescing Flowers ...... 167 Auxin-Induced Ethylene Production 169 Regulation of Ethylene Biosynthesis by Ethylene 169 Stress-Induced Ethylene Production . . . . . . . . 171 Regulation by Light and Carbon Dioxide 173 Inhibitors of Ethylene Biosynthesis : 174 Conjugation of l-Aminocyclopropanecarboxylic Acid to l-(Malonylamino) cyclopropanecarboxylic Acid 178 CONCLUDING REMARKS 180

Ethylene biosynthesis and its regulation in higher plants

OXIDATIVE STRESS . . . . . . . . . . . . . .. . . . . .... . . . . .. . . . . .. . . . ... .. . . . . .... . . .. .. . . . . . ... . 84 RESPONSE OF SOD TO ENVIRONMENTAL STRESS . . . . . . . . . . . . . . . .. . . . .... . . .... . . . . . . 87 Photoinhibition . . . . . . . . . . .. . ... . . .. . . . .. . . . . ..... . . . , ... ", , ... ,' , ... . ,., . . . "" . .. . ,'.' . . ,' . . . . , ., 87 Paraquat and Other Herbicides . . .. , . . .... , ..... , . . . . ... . . . .. . . . . . . . . . . . . ,.. 91 Atmospheric Pollutants . . " .... , .... , " .... " , , . . , .... , ' 94 Waterlogging and Drought .. ", . . ... , . . . . . . , .. "", . . ", . . . . . ", . . . .. ", . . .. , . . . ... """", . . " 97 The Defense Response to Pathogens . ... . . . . "" ... "" .... " ...... " ,."" .... . , 98 The Phenomenon of Cross-Tolerance . . . , , .. . . . . ...... . . , . . . . . . , . . .... , .. . . . . . . . .. .. 101 The Mechanism of SOD Regulation ........ , , ...... , " 102 GENETIC ENGINEERING OF SOD IN PLANTS .. ' ...... ' . . . . . . . . . . . . . . . . . . . . . . . .. . . . , .... ' 104 Cu/ZnSOD and MnSOD Overexpression . . . . . .. . . . . . , . . . . . . .. . . . . . .. . . . . . , .... . . . . .. . . ,.... . . 105 Prospects for Stress Tolerance through Genetic Engineering of SOD . . . . ,,, .. , . . . . . . . 106

Superoxide dismutase and stress tolerance

The ways in which plant growth promoting rhizobacteria facilitate the growth of plants are considered and discussed. Both indirect and direct mechanisms of plant growth promotion are dealt with. Th...

The enhancement of plant growth by free-living bacteria

Dioxygen activation at mononuclear nonheme iron active sites: enzymes, models, and intermediates.

Reactions involving free radicals are an inherent feature of plant senescence and appear to contribute to a process of oxidative deterioration that leads ultimately to cell death. Radical species derived from molecular oxygen are the primary mediators of this oxidative damage, but non-radical excited states of oxygen, specifically singlet oxygen, may also be involved. Several lines of evidence suggest that degradation of lipids in senescing membranes and the ensuing release of free fatty acids initiate oxidative deterioration by providing substrate for lipoxygenase. In some tissues, lipoxygenase activity increases with advancing senescence in a pattern that is consistent with its putative role in promoting oxidative damage. However, there are important exceptions to this which may be explained by the fact that the timing and extent of peroxidative reactions initiated by lipoxygenase are likely to be determined more by the availability of substrate for the enzyme than by changes in its activity. There are both membranous and cytosolic forms of lipoxygenase in senescing tissues, and peroxidation of membrane lipids appears to be initiated by the membranous enzyme once the appropriate fatty acid substrates, linoleic acid and linolenic acid, become available. Since lipid peroxidation is known to form alkoxy and peroxy radicals as well as singlet oxygen, these reactions in membrane bilayers are probably a major source of activated oxygen species in senescing tissues. Further-more, there are indications that activated oxygen from the lipoxygenase reaction can become substrate for the cytosolic form of the enzyme which, in turn, may raise the titre of activated oxygen during senescence. Additional possible sources of increased free radical production in senescing tissues include peroxidase, which shows greatly increased activity with advancing age, leakage of electrons from electron transport systems to oxygen, in particular from the photosynthetic electron transport system, and decompartmentalization of iron, which would facilitate formation of the highly reactive hydroxyl radical from the less reactive superoxide anion. A variety of macromolecules can be damaged by activated oxygen. Unsaturated fatty acids are especially prone to attack, and this implies that membranes are primary targets of free radical damage. The manifestations of this damage in senescing tissues range from altered membrane fluidity and phase properties to leakiness that can be attributed to a destabilized and highly perturbed membrane bilayer. There is also a progressive breakdown of cellular protein with advancing senescence. Free radicals can inactivate proteins by reacting with specific amino acid residues, and a number of in zitro studies have indicated that such alteration renders the proteins more prone to hydrolysis by proteases. Thus, although there is no direct evidence linking enhanced proteolysis during senescence to free radical damage, there is reason to believe that this may be a contributing factor. Wounding of certain plant tissues also initiates a series of reactions that revolve around the breakdown of membrane lipids and their peroxidation. Indeed, as in the case of senescence, membrane deterioration follokving wounding appears to be facilitated by a self-perpetuating wave of free radical production emanating from peroxidation within the lipid bilayer. There is also recent evidence for activation of an O2- -producing NADPH oxidase in plant tissues following fungal infection that may be analogous to the well-characterized O2- -generating NADPH oxidase associated with the plasma membrane of polymorphonuclear leukocytes. This raises the interesting possibility that plants and animals share a common defence response to invading organisms. Contents Summary 317 I. Introduction 318 II. Species of activated oxygen 319 III. Sites of activated oxygen production 319 IV. Free radical production during senescence 323 V. Targets of free radical damage in senescing tissues 330 VI. The role of free radicals in seed ageing 336 VII. The role of free radicals in wounding 337 VIII. Concluding remarks 338 Acknowledgement 338 References 338.

The role of free radicals in senescence and wounding.

Radical scavenging properties of polyamines

Lipid metabolism during plant senescence.

Lipid analysis of rosette leaves from Arabidopsis has revealed an accumulation of triacylglycerol (TAG) with advancing leaf senescence coincident with an increase in the abundance and size of plastoglobuli. The terminal step in the biosynthesis of TAG in Arabidopsis is catalyzed by diacylglycerol acyltransferase 1 (DGAT1; EC 2.3.1.20). When gel blots of RNA isolated from rosette leaves at various stages of development were probed with the Arabidopsis expressed sequence tag clone, E6B2T7, which has been annotated as DGAT1, a steep increase in DGAT1 transcript levels was evident in the senescing leaves coincident with the accumulation of TAG. The increase in DGAT1 transcript correlated temporally with enhanced levels of DGAT1 protein detected immunologically. Two lines of evidence indicated that the TAG of senescing leaves is synthesized in chloroplasts and sequesters fatty acids released from the catabolism of thylakoid galactolipids. First, TAG isolated from senescing leaves proved to be enriched in hexadecatrienoic acid (16:3) and linolenic acid (18:3), which are normally present in thylakoid galactolipids. Second, DGAT1 protein in senescing leaves was found to be associated with chloroplast membranes. These findings collectively indicate that diacylglycerol acyltransferase plays a role in senescence by sequestering fatty acids de-esterified from galactolipids into TAG. This would appear to be an intermediate step in the conversion of thylakoid fatty acids to phloem-mobile sucrose during leaf senescence.

A Role for Diacylglycerol Acyltransferase during Leaf Senescence

https://www.cell.com/trends/plant-science/pdf/S1360-1385(04)00054-8.pdf

John E. Thompson

Papers

The role of free radicals in senescence and wounding.

Radical scavenging properties of polyamines

Lipid metabolism during plant senescence.

A Role for Diacylglycerol Acyltransferase during Leaf Senescence

Regulation of senescence by eukaryotic translation initiation factor 5A: implications for plant growth and development