Review Article

Chromium(VI) Toxicity in Legume Plants: Modulation Effects of Rhizobial Symbiosis

Table 1

Major components of antioxidant defense in legume plants and root nodules.

ComponentsLocationFunction

Enzymatic antioxidants
Superoxide dismutase (SOD)Dismutation of : + + 2H+ → 2H2O2 + O2
 Cu, Zn-SODCytosol, plastids
 Mn-SODMitochondria, bacteroides
 Fe-SODPlastids, some bacteroides
CatalasePeroxisomes, bacteroidesDismutation of H2O2: 2H2O2 → 2H2O + O2
Guaiacol peroxidase (GPX)Plasma membrane, vacuolesDetoxification of H2O2: H2O2 + → H2O +
PeroxiredoxinsCytosol, mitochondriaReduction of H2O2 or alkyl hydroperoxides (ROOH) to H2O or respective alcohols (ROH): ROOH + Trx-(SH)2 → ROH → Trx-S2 + H2O
Enzymes of ascorbate-glutathione cycle (AA-GSH cycle)
Cytosol (mainly), mitochondria, plastids,
peroxisomes
Detoxification of H2O2 in a series of reactions
 Ascorbate peroxidase (APX)APX: H2O2 + AA → 2H2O + MDHA (DHA)
 Monodehydroascorbate reductase (MR)MR: 2MDHA + NADH → 2AA + NAD
 Dehydroascorbate reductase (DR)DR: DHA + 2GSH → AA + GSSG
 Glutathione reductase (GR)GR: GSSG + NADPH → 2GSH + NADP+
Enzymes of GSH metabolism
 Glutathione peroxidase (GSH-PX)Cytosol, plastids, bacteroidesDetoxification of H2O2: H2O2 + GSH → 2H2O + GSSG
 Glutathione reductase (GR)Cytosol, mitochondria, plastids, bacteroidesRegeneration of oxidized glutathione: GSSG + NADPH → 2GSH + NADP+
α-Glutamylcysteine synthetase and glutathione synthetaseCytosol, plastids, bacteroidesGSH synthesis de novo
 Glutathione-S-transferaseCytosol (mainly), plastids mitochondria, bacteroidesDetoxification of xenobiotics via conjugation with GSH

Nonenzymatic antioxidants
Ascorbic acid (AA)Cytosol, mitochondria, plastids,
peroxisomes
Direct scavenging of ROS, a substrate for APX
Glutathione (GSH)Cytosol, mitochondria, plastids, bacteroidesDirect scavenging of ROS, a cosubstrate for DR, GSH-PX, and glutathione-S-transferases
α-TocopherolPlastidsDirect scavenging of ROS, quenching of lipid radicals in membranes
CarotenoidsPlastidsQuenching of electron-excited molecules, protection of chlorophylls from photodamage
Phenol compounds (phenolic acids, flavonoids, etc.)Vacuoles (mainly), cytosol, plastids, cell wallDirect scavenging of ROS and lipid radicals, metal-chelating activity
Ferritin, phytochelatins, and metallothioneinsPlastids, vacuoles, cytosolMetal-binding activity

Trx-(SH)2/Trx-S2: reduced/oxidized thioredoxin; MDHA: monodehydroascorbate; DHA: dehydroascorbate; GSH/GSSG: reduced/oxidized glutathione.