The compound dichlorophenoxyacetate (2,4-D) was the first organic, systemic, selective herbicide, and for post-emergence application to be developed in the world. Along with the green revolution, this herbicide has helped to increase the cereal production on the decades after 1950. This product is a synthetic auxin that can be used as a plant growth regulator or as an herbicide for the control of broad-leaved weeds. Several broad-leaved weed species that have proved difficult to control with other herbicides are susceptible to 2,4-D. However, the use of this herbicide is restricted due to the lack of selectivity in certain crops. In recent decades, the discovery of genes related to 2,4-D tolerance in soil bacteria and its transfer to crops enabled the development of tolerance to the herbicide. The objectives of this literature review are to describe the enzymes and genes responsible for 2,4-D tolerance; to illustrate the mechanisms involved in the selectivity to 2,4-D and to other herbicides; and to analyze some of the implications for weed management. The first 2,4-D tolerance gene to be discovered was tfdA, which was found in the plasmid pJP4 from the bacterium Cupriavidus necator. This gene encodes for the 2,4 D/oxoglutarate dioxygenase enzyme, which catalyzes the conversion of 2,4-D to 2,4 dichlorophenol and glyoxylate. During the late 1980s occurred the first insertion of the tfdA gene in plants of Nicotiana tabacum via Agrobacterium tumefaciens. This conferred tolerance of tobacco plants to 2,4-D. Similar results were obtained with insertion of this gene into plants from several crops, such as Gossypium hirsutum, Vitis vinifera, and Brassica juncea. Studies on soil bacteria have identified two other genes, the rdpA gene from Sphingobium herbicidivorans MH, which encodes the enzyme ariloxyalkanoate-dioxygenase-1 (AAD-1); and the sdpA gene from Delftia acidovorans MC1, which encodes the enzyme ariloxyalkanoate-dioxygenase-12 (AAD-12). These two enzymes are similar, but have different enzyme kinetics and are able to degrade 2,4-D and other herbicides. The AAD-1 enzyme degrades 2,4-D and, surprisingly, the ariloxyphenoxypropionate (AAPP) herbicides, which inhibit acetyl-CoA carboxylase (ACCase). The AAD-12 enzyme has a high binding affinity with the auxins 2,4-D, MCPA, triclopyr, fluroxypyr. Currently, the genes encoding these enzymes are being used for the development of soybean, cotton, and corn tolerant to 2,4-D and AAPP. Soybean plants with the gene sdpA are highly tolerant to 2,4-D. Corn plants containing the gene rdpA are also tolerant to AAPP. Work carried out with the glyphosate resistant weeds Conyza bonariensis, Conyza canadensis, and Amaranthus palmeri have shown adequate control with 2,4-D. Therefore, the sdpA and rdpA genes are good candidates to develop 2,4-D tolerant crops and should expand the options to control weeds resistant to other herbicides.
2,4-D; weeds; herbicide resistance
