Under certain environmental conditions in North America, some sweet corn hybrid cultivars and inbreds are injured by postemergence applications of cytochrome P450-metabolized corn herbicides. Some of the sweet corn cultivars that are sensitive to specific herbicides have been identified in the past two decades in numerous field trials (Burton et al., 1994; Diebold et al., 2003, 2004; Grey et al., 2000; Monks et al., 1992; Morton and Harvey, 1992; O'Sullivan and Bouw, 1998; O'Sullivan and Sikkema, 2002; O'Sullivan et al., 1995, 2000, 2002; Robinson et al., 1993, 1994; Sikkema et al., 2008; Stall and Bewick, 1992; Williams and Pataky, 2008; Williams et al., 2005). Although useful, this information is limited because a relatively small percentage of nearly 600 sweet corn cultivars sold commercially in North America were tested in any one of these studies and the commercial availability of cultivars changes continually as new cultivars are introduced and older cultivars are discontinued. A better understanding of the genetic basis of herbicide sensitivity in sweet corn would help the sweet corn industry manage this problem more effectively and it may provide an explanation for varied responses of certain cultivars under differing conditions.
In previous research, the sweet corn inbred Cr1 was found to be sensitive to multiple postemergence herbicides, including four acetolactate synthase (ALS)-inhibiting herbicides (foramsulfuron, nicosulfuron, primisulfuron, and rimsulfuron), two 4-hydroxyphenylpyruvate dioxygenase (HPPD)-inhibiting herbicides (mesotrione and tembotrione), a growth regulator herbicide combination (dicamba + diflufenzopyr), a protoporphyrinogen oxidase (PPO)-inhibiting herbicide (carfentrazone), and a photosystem II (PSII)-inhibiting herbicide (bentazon) (Nordby et al., 2008; Pataky et al., 2006; Williams and Pataky, 2008; Williams et al., 2005). Sensitivity to these herbicides appeared to be conditioned by a single gene or closely linked genes based on segregation of progeny in F2, BC1, and BC2 generations of a cross of Cr1 with Cr2 (a herbicide-tolerant inbred) and cosegregation of phenotypic responses among BC1S1, BC2S1, F2:3, F3:4, and F3:5 families, (Nordby et al., 2008; Pataky et al., 2006; Williams and Pataky, 2008). Another independent gene from Cr2 also conditioned tolerance to bentazon.
These herbicides to which Cr1 is sensitive have different modes of action, but all are metabolized by cytochrome P450 monooxygenases. Barrett et al. (1994) previously proposed the existence of a “super P450” that metabolizes multiple herbicides. Although corn has many cytochrome P450 genes (CYP genes), the number of P450s involved in metabolic inactivation of herbicides, their expression levels, and their levels of herbicide metabolism are not clearly understood (Barrett, 1995, 2000; Frey et al., 1995, 1997; Persans et al., 2001).
A growing body of evidence suggests that sensitivity of corn to multiple P450-metabolized herbicides is regulated by a single CYP gene or a group of closely linked CYP genes on the short arm of chromosome 5. Kang (1993) associated nicosulfuron sensitivity in field corn inbred lines, including the inbred W703a, with a single, recessive gene designated as nsf1. Others also reported simple inheritance of tolerant and sensitive responses to nicosulfuron and similar ALS-inhibiting herbicides (Green, 1998, Green and Ulrich, 1993; Harms et al., 1990; Widstrom and Dowler, 1995). Green (1998) cited an unpublished report that mapped sensitivity to rimsulfuron in the field corn inbred F2 to the short arm of chromosome 5. Fleming et al. (1988), Bradshaw et al. (1994), and Barrett et al. (1997) associated bentazon and nicosulfuron sensitivity in field corn inbreds B90 and GA209 with a single, recessive gene designated as ben1. An independent, dominant gene, designated as Ben2, conditioned tolerance to bentazon in the inbred B73 (Barrett et al., 1997; Bradshaw et al., 1994). Williams et al. (2006) used a mapped-based cloning approach to locate the Nsf1 gene on the short arm of chromosome 5 and to sequence the dominant, functional allele from a nicosulfuron-tolerant inbred, B73. The Nsf1 gene was one of four closely linked genes with significant sequence similarity to CYP genes. The Nsf1 gene also was very similar in sequence and function to a CYP gene in rice that conditions tolerant responses to multiple herbicides (Pan et al., 2006). Nicosulfuron-sensitive inbreds GA209 and W703a contained a 392-bp insertion in the Nsf1 gene sequence relative to B73. Thus, it appears that the nsf1 and ben1 alleles identified from GA209 and W703a, respectively, are the same 392-bp insertion mutation of this CYP allele. The 392-bp insertion in this CYP gene sequence also occurs in several nicosulfuron-sensitive field corn inbreds (e.g., A180, B90, MS1334, NC22, and R4), whereas other nicosulfuron-sensitive field and sweet corn inbreds (e.g., B94 and Ia5125) do not contain this insertion. The nsf1/ben1 CYP gene has not been sequenced from the herbicide-sensitive sweet corn inbred Cr1; however, a QTL that conditions sensitivity to multiple P450-metabolized herbicides was detected in a segregating population of Cr1 × Cr2 on the short arm chromosome 5 in tight linkage disequilibrium with the nsf1/ben1 CYP locus (Nordby et al., 2008).
The genetic condition at a single locus explained varied responses of 149 commercially adapted sweet corn hybrid cultivars to three P450-metabolized, postemergence herbicides (foramsulfuron, mesotrione, and nicosulfuron) in 12 field trials throughout the United States (Pataky et al., 2008). Seven cultivars classified as homozygous for alleles conditioning herbicide sensitivity were injured most severely and often were killed by the two ALS-inhibiting herbicides. Ninety-five cultivars classified as homozygous for an allele conditioning herbicide tolerance were uninjured or injured least. Forty-seven cultivars classified as heterozygous with one allele each conditioning herbicide tolerance and sensitivity displayed intermediate responses that were more similar to homozygous tolerant cultivars than homozygous sensitive cultivars. Injury to heterozygous and homozygous tolerant cultivars was minimal and often not significantly different in trials in dry, western climates (e.g., Caldwell, ID; Nampa, ID; and Olathe, CO), whereas injury to heterozygous cultivars was significantly greater than injury to homozygous tolerant cultivars in trials in eastern climates (e.g., Georgetown, DE and LeRoy, NY). Nevertheless, it was not evident that the seven homozygous sensitive cultivars and the 49 heterozygous cultivars carried the same allele conditioning sensitivity. The objective of this study was to determine if sensitivity to nicosulfuron and mesotrione among commercially adapted, sweet corn cultivars and inbreds is allelic with or closely linked to the recessive gene conditioning sensitivity in the sweet corn inbred Cr1.
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