Table 1.
Description of the herbicide mechanisms of action (MA) and their effects on treated plants related to the production of reactive oxygen species.
| HRAC Group a | Herbicide Mechanism of Action (MA) | Biological Process Committed | Herbicide Chemical Family | Herbicide Active Ingredient b | ROS Production c |
|---|---|---|---|---|---|
| A | Inhibition of acetyl-CoA carboxylase (ACCase) | Fatty acid biosynthesis | Aryloxyphenoxy-propionate “FOPS”, Cyclohexanedione “DIMs”, Phenylpyrazoline “DEN” | benzoylprop-ethyl, diclofop-methyl, haloxyfop-methyl, cyhalofop, clethodim, setoxydim, pinoxaden | No |
| B | Inhibition of acetohydroxyacid synthase (AHAS, ALS) | Amino acid biosynthesis (Leu, Ile, Val) | Sulfonylurea, Imidazolinone, Triazolopyrimidine, Pyrimidinyl(thio)benzoate | metsulfuron-methyl, chlorimuron-ethyl, nicosulfuron, imazapyr, imazethapyr, flumetsulam, cloransulam-methyl, diclosulam, flucarbazone-sodium, pyritiobac | Yes |
| C1, C2, C3 | Inhibition of photosystem II protein D1 (psbA) | Photosynthesis (electron transfer) | Triazine, Triazinone, Triazolinone, Uracil, Pyridazinone, Phenyl-carbamate, Urea, Amide, Nitrile, Benzothiadiazinone, Phenyl-pyridazine | ametryne, atrazine, simazine, hexazinone, metribuzin, amicarbazone, bromacil, pyrazon, desmedipham, chlorotoluron, diuron, linuron, propanil, bromoxynil, ioxynil, bentazon, pyridate | Yes |
| D | Diversion of the electrons transferred by the photosystem I ferredoxin (Fd) | Photosynthesis (electron transfer) | Bipyridylium | diquat, paraquat | Yes |
| E | Inhibition of protoporphyrinogen oxidase (PPO) | Photosynthesis (heme biosyn- thesis for chlorophyll) | Diphenylether, Phenylpyrazole, N-phenylphthalimide, Thiadiazole, Oxadiazole, Triazolinone, Oxazolidinedione, Pyrimidindione | acifluorfen-Na, fomesafen, lactofen, oxyfluorfen, pyraflufen-ethyl, flumioxazin, fluthiacet-methyl, oxadiazon, azafenidin, pentoxazone, butafenacil, flufenpyr-ethyl | Yes |
| F1, F2, F3 | Inhibition of phytoene desaturase (PDS) or 4-hydroxyphenylpyruvate dioxygenase (4-HPPD) or of an unknown protein | Photosynthesis (carotenoid biosynthesis) | Pyridazinone, Pyridinecarboxamide, Triketone, Isoxazole, Pyrazole, Triazole, Isoxazolidinone, Urea, Diphenylether | norflurazon, diflufenican, fluridone, mesotrione, isoxaflutole, pyrazoxyfen, amitrole, clomazone, fluometuron, aclonifen | Yes |
| G | Inhibition of 5-enolpyruvylshikimate-3-phosphate synthase (EPSPS) | Amino acid biosynthesis (Phe, Trp, Tyr) | Glycine | glyphosate | Yes |
| H | Inhibition of glutamine synthase | Amino acid biosynthesis (Gln) | Phosphinic acid | glufosinate-ammonium | Yes |
| I | Inhibition of dihydropteroate synthase | Tetrahydrofolate biosynthesis | Carbamate | asulam | Yes |
| K1, K2 | Enhancement of tubulin depolymerization | Microtubule polymerization | Dinitroaniline, Phosphoroamidate, Pyridine, Benzamide, Benzoic acid, Carbamate | oryzalin, pendimethalin, trifluralin, amiprophos-methyl, dithiopyr, propyzamide, DCPA, carbetamide | No |
| K3 | Inhibition of fatty acid synthase (FAS) | Fatty acid biosynthesis | Chloroacetamide, Acetamide, Oxyacetamide, Tetrazolinone | acetochlor, alachlor, metolachlor, napropamide, flufenacet, fentrazamide, anilofos | No |
| L | Inhibition of cellulose-synthase | Cell wall biosynthesis | Nitrile, Benzamide, Triazolocarboxamide, Quinoline carboxylic acid | dichlobenil, isoxaben, flupoxam, quinclorac | No |
| M | Uncoupling of oxidative phosphorylation | ATP biosynthesis | Dinitrophenol | dinoseb, dinoterb | Yes |
| N | Inhibition of fatty acid elongase | Fatty acid biosynthesis | Thiocarbamate, Phosphorodithioate, Benzofuran, Chloro-Carbonic-acid | butylate, cycloate, EPTC, bensulide, ethofumesate, TCA, dalapon | No |
| O | Stimulation of transport inhibitor response protein 1 (TIR1) | Regulation of auxin-responsive genes | Phenoxy-carboxylic-acid, Benzoic acid, Pyridine carboxylic acid, Quinoline carboxylic acid | 2,4-D, MCPA, dicamba, clopyralid, fluroxypyr, picloram, triclopyr, quinclorac, quinmerac, benazolin-ethyl | Yes |
| P | Inhibition of auxin transport | Long-range hormone signaling | Phthalamate, Semicarbazone | naptalam, diflufenzopyr-Na | Yes |
a The 21 known groups of herbicides classified by mechanims of action (MA) according to the global Herbicide Resistance Action Committee (HRAC; http://www.hracglobal.com, accessed on: 28 January 2019. Also, there are other MAs, but they are not completely understood as the biochemical processes were not described. Please see the HRAC website to check all MA. b Not all active herbicide ingredients are presented, please see the HRAC website to check all of them. c Indicates the ROS production in some phase of the herbicide action [41,45].