Abstract
Safeners protect crops by enhancing their ability to metabolize various compounds, including herbicides. They increase a crop’s tolerance to herbicide damage, activating herbicide-metabolizing proteins, and aiding in their detoxification. This study aimed to investigate the chemical effects of safeners in tomato cultivation and focus on injury reduction and tissue protection. The experiment followed a randomized factorial design (5 × 4) with four replications repeated twice. We evaluated the effects of herbicides dicamba, 2,4-D, metribuzin, and sulfentrazone (diluted to 1% of the recommended field rate) and safeners benoxacor, fenclorim, melatonin, 2,4,6-T, and an untreated control. Safeners were applied to the seeds before sowing, and herbicides were applied as a foliar spray 25 days after sowing. Visual injury was evaluated 7, 14, and 21 days after application (DAA). Biomass measurements were taken 21 DAA. Results showed that preconditioning tomato seeds with 2,4,6-T, melatonin, and fenclorim 7 DAA significantly decreased injury by 25, 25, and 23%, respectively. Moreover, applying melatonin, benoxacor, and 2,4,6-T 21 DAA led to significantly greater dry biomass, which increased by 1.5, 1.42, and 1.44 times, respectively, compared with the control. This research provides valuable insights into the chemical effects of benoxacor, fenclorim, 2,4,6-T, and melatonin safeners in tomato cultivation. The findings demonstrate the potential for preconditioning tomato plants with 2,4,6-T, melatonin, and fenclorim to reduce herbicide injury. Additionally, melatonin, benoxacor, and 2,4,6-T stimulated growth, increasing tomato dry biomass. Understanding plant defense mechanisms and the protective effects of safeners against herbicide damage contributes to developing effective weed management strategies.
Tomatoes (Solanum lycopersicum) are a highly valued and widely cultivated crop, ranking second only to potatoes (Solanum tuberosum L.) in production. With more than 182.3 million tons of tomato fruits grown on ∼4.85 million hectares yearly (Quinet et al. 2019), tomatoes play a crucial role in global food production and supply. However, tomato production faces numerous challenges that directly and indirectly impact yields, including low-quality seeds, adverse climatic conditions, pest infestations, and weed interference (Singh et al. 2017; Tolman et al. 2004). Among these factors, weeds pose a significant threat to tomato crops. Weeds compete with tomatoes for essential resources such as water, nutrients, sunlight, and space, reducing crop growth and productivity (Clark et al. 1998). Effective weed management is, therefore, vital to maintaining high-quality tomato yields and preventing economic losses in the agricultural industry (Yaacoby et al. 2023).
Although herbicides are used for weed control in agricultural production (Gwatidzo et al. 2023), matching an appropriate herbicide for the target weed species improves efficacy and reduces wasted time and resources. In addition, environmental factors such as temperature, wind speed, and moisture levels must be considered during herbicide application to ensure optimal effectiveness and minimize unintended off-target effects (Johnson and Young 2002; Richardson et al. 2020). Tomato plants are known to be particularly sensitive to many herbicides, especially 2,4-D and dicamba. Even small amounts of these herbicides, resulting from drift or contamination, can severely compromise tomato crop growth and yield (de Paula Medeiros et al. 2023). Implementing effective weed management strategies is crucial to achieve maximum quality and high yields in tomato cultivation. A comprehensive approach may involve both chemical and nonchemical control methods. Pre-herbicides, such as sulfentrazone, metolachlor, or fomesafen, may be recommended based on their effectiveness against specific weed species and the density of weed populations (Flint and Klonsky 1985; Jablonkai 2013).
Furthermore, nonchemical controls, including mechanical and cultural practices, can supplement herbicide use and help manage weeds effectively. Farmers and agricultural practitioners can use various weed management approaches to minimize weed-related challenges, reduce resource competition, and optimize tomato crop growth and productivity. Understanding the specific weed dynamics and implementing appropriate weed management strategies are key steps toward achieving successful tomato production while preserving the agricultural systems’ overall health and sustainability. One of the promising weed control options is to use safeners to improve herbicide tolerance in tomato, thereby enhancing weed control (Rosinger 2014).
Otto Hoffman introduced the safener concept in the late 1940s after discovering that tomato plants treated with 2,4,6-T did not suffer damage after contact with 2,4-D vapor (Hoffmann 1969; Jablonkai 2013). In 1971, these compounds were introduced as a new tool to the market, and the first commercial safener, 1,8-naphthalic anhydride, was used as a seed treatment in corn (Hoffmann 1969). Since then, safeners have generated significant interest from the chemical industry due to their physiological and biochemical features to activate important mechanisms in plants without reducing herbicide efficacy in target weed species. In addition, their use could help reduce or slow the development of herbicide-resistant weed populations (Duhoux et al. 2017). The use of fenclorim (4,6-dichloro-2-phenylpyrimidine) as a safener occurred in 1991 to protect and increase the tolerance of rice (Oryza sativa L.) against pretilachlor (Chen et al. 2013; Han and Hatzios 1991). Benoxacor was also discovered in the 1990s. Cottingham and Hatzios (1991) reported that benoxacor was used as a safener in maize to protect it from metolachlor injury.
Metolachlor is among the top five herbicides applied in the midwestern corn belt of the United States. It is so widely used that it can be found in high concentrations in groundwater, among other herbicides (Fuerst et al. 1995; Simonsen et al. 2020). Melatonin application on plants results in cell enlargement and root development, delayed flowering, improved fruit yield and quality, and preserving the mineral balance in heat-stressed tomato plants (Arnao and Hernández-Ruiz 2020; Kaya et al. 2022).
Safener use may protect various crops against herbicide drift. Consequently, this investigation examined the presence and absence of four safeners to protect tomatoes against herbicide applications with distinct modes of action.
Material and methods
Tomato transplants, preconditioning, and herbicide treatments
Experiments were conducted in 2022 and 2023 at the R.R. Foil Plant Science Research Center of Mississippi State University in Starkville, MS (lat. 32.46936111°N, long. 88.78333333°E). The study used a completely randomized design with four replicates, repeated twice and arranged in a 4 × 5 factorial design. Factor A represented the herbicide treatments, including four herbicides, while factor B consisted of four safeners and a nontreated (safener control) group.
Better Boy Plus tomato seeds were immersed in the solution with safener and methanol for 1 h before planting. Each safener was dissolved in methanol at the following micromolar concentrations: 10 µM for benoxacor (TCI America™, >98.0%), 2.98 µM for fenclorim (TCI America™, >98.0%), 5,066 µM for 2,4,6-T (TCI America™, >98.0%), and 100 µM for melatonin (Alfa Aesar, 99.0+%) (Chen et al. 2013; Hartman 1959; Irzyk and Fuerst 1993; Tiwari et al. 2021). The control was treated with methanol only. Transplants were generated by sowing the seeds into 72-cell plug trays filled with a soilless potting media (Promix BX; Rivière-du-Loup, QC, Canada) and cultivated in a greenhouse. The tomato transplants were planted 21 d later into 0.815-L pots filled with the same soilless media. Herbicides were applied at 25 d after sowing using a calibrated spray chamber delivering 187 L·ha−1 and equipped with AIXR 1102 nozzles (TeeJet Technologies, Wheaton, IL, USA) at a spray pressure of 275.8 KPa. The specific herbicide applications are listed in Table 1.
List of herbicide treatments and their rates used in the greenhouse study at the R. R. Foil Plant Science Research Center at Mississippi State University, Starkville, MS, USA.
Crop injury was evaluated at 7, 14, and 21 DAA using a visual rating scale ranging from 0% to 100%, where 0% represented normal growth without herbicide symptoms and 100% indicated complete desiccation of shoot tissues. The scoring criteria included various symptoms such as leaf chlorosis, cupping, rolling, stem twisting, stunting, necrosis, and epinasty.
Shoot biomass
At 21 DAA, the shoot tissues were harvested, stored in paper bags, oven-dried at 60 °C for 72 h, and weighed to obtain the dry biomass weights.
Statistical analysis
All data were fitted using a standard least-squares (LS) model in JMP Pro 16.1 (SAS Institute Inc., Cary, NC, USA). The main effects analyzed included the safeners (benoxacor, fenclorim, melatonin, and 2,4,6-T) and herbicides (2,4-D, dicamba, metribuzin, and sulfentrazone) at 1% of the field rate (Table 1). Field rate of 1% was selected based on previous studies conducted by Fagliari et al. (2005), which they determined that under field conditions 1% rate would be equivalent to herbicide drift. Interactions between the main effects were also assessed in relation to dry biomass and injury. For data that met the assumptions of the ANOVA test, treatment means were separated using the LS Means Differences Tukey Honestly Significant Difference test, which was conducted at a significance level of α = 0.05.
Results
Crop injury
Herbicide injury was observed in tomato leaves after application (Fig. 1).
Herbicide injury in tomato plants caused by 2,4-D (Enlist one), dicamba (XtendiMax), metribuzin (Glory), and sulfentrazone (Spartan). Herbicides were applied 25 d after the seeds were sown at 1% of the recommended field rate.
Citation: HortTechnology 34, 6; 10.21273/HORTTECH05480-24
Significant crop injury was observed in tomato leaves 7, 14, and 21 days after herbicide application (DAA) (P < 0.05) (Fig. 2). At 7 DAA, fenclorim, 2,4,6-T, and melatonin treatments resulted in lower injury than the control, while the highest injury levels were observed in the benoxacor and control treatments (33% and 32%, respectively). The fenclorim treatment showed the lowest injury (23%), followed by melatonin and 2,4,6-T (25% each) (Fig. 2) at 7 DAA. At 14 DAA, the benoxacor and fenclorim treatments had injury comparable to the control, while the 2,4,6-T and melatonin treatments had reduced injury (Fig. 2). At 21 DAA, the 2,4,6-T and melatonin treatments effectively reduced herbicide-induced injury, showing significant differences from the fenclorim, benoxacor, and control treatments (Fig. 2).
Injury across herbicide 2,4-D, dicamba (auxin), sulfentrazone (PPO), and metribuzin (PSII) treatments 7, 14, and 21 d after application (DAA) in safer-conditioned tomato plants. Means were separated within assessments using a least-squares Means Tukey honestly significant difference separation at P < 0.05. Bars sharing the same letters are not significantly different.
Citation: HortTechnology 34, 6; 10.21273/HORTTECH05480-24
All herbicides caused more than 50% injury in tomato leaves and stems 21 DAA, except for fenclorim, which reduced injury from sulfentrazone by more than 50%, and 2,4,6-T, which reduced injury from dicamba, 2,4-D, and metribuzin. Melatonin also reduced 2,4-D injury by more than 50%; however, benoxacor did not reduce herbicide injury (Fig. 3).
Visible estimates of injury at 21 d after application (DAA) of herbicides 2,4-D, dicamba (auxin), sulfentrazone (PPO), and metribuzin (PSII) in tomato plants treated with safeners before sowing. Means were separated within assessments using a least-squares Means Tukey honestly significant difference separation at P < 0.05. Means followed by the same letter do not differ.
Citation: HortTechnology 34, 6; 10.21273/HORTTECH05480-24
Shoot dry biomass
The shoot dry biomass of tomato plants 21 DAA showed significant differences (P < 0.05) among the treatments. The melatonin, 2,4,6-T, and benoxacor treatments had more dry biomass after herbicide application than the control. In contrast, the fenclorim treatment did not differ in accumulation from the treatments or the control (Fig. 4). The preconditioning with melatonin resulted in 1.50 times more dry biomass accumulation than the control, whereas 2,4,6-T and benoxacor accumulated 1.44 and 1.42 times more, respectively. The fenclorim treatment dry biomass was not statistically different from the control. Still, the seed treated with this safener had 1.36-fold more tomato biomass than the control (Fig. 4).
Dry biomass of safener-conditioned tomato plants at 21 d after application of herbicide 2,4-D, dicamba (auxin), sulfentrazone (PPO), and metribuzin (PSII). Means were separated within assessments using a least-squares Means Tukey honestly significant difference separation at P < 0.05. Means followed by the same letter do not differ.
Citation: HortTechnology 34, 6; 10.21273/HORTTECH05480-24
The dry biomass of tomatoes preconditioned with safeners was greater among the herbicide treatments than the non-preconditioned control. The combination of herbicides and safeners showed a significant difference compared with the control plants (Figs. 5 and 6). Although dicamba was not significantly different compared with metribuzin and sulfentrazone, it resulted in the lowest increase in tomato dry biomass (1.10 times), while metribuzin and sulfentrazone treatments (1.21 and 1.33, respectively) allowed for more dry biomass accumulation (Fig. 5). Conversely, 2,4-D treatment resulted in the greatest increase (1.76 times) in tomato biomass compared with the control.
Shoot dry biomass values of tomato plants with and without safeners compared under different herbicide applications, 21 d after herbicide treatment. Means were separated using the least-squares Means Tukey honestly significant difference at P < 0.05. Means followed by the same letter do not differ.
Citation: HortTechnology 34, 6; 10.21273/HORTTECH05480-24
Safener-conditioned tomato plants 7 d after herbicide application. Safeners were applied at the following rates: 2.98 µM fenclorim, 10 µM benoxacor, 100 µM melatonin, 5066 µM 2,4,6-T, and a non–safener-conditioned control. Herbicides were applied at 1% of the field rate. Visible estimates of injury were recorded for each treatment.
Citation: HortTechnology 34, 6; 10.21273/HORTTECH05480-24
Discussion
Preconditioning tomatoes with safeners melatonin, 2,4,6-T, and fenclorim reduced crop injury compared with the benoxacor and control treatments 7 DAA. These findings align with previous studies, such as Castro et al. (2020), which reported the protective effects of fenclorim on tomato seeds treated with different herbicides. The exact mechanism by which safeners protect plants against herbicides remains unclear, and further research is needed to understand their modes of action in crops. For instance, 2,4,6-T has been shown to suppress epinasty induced by 2,4-D vapor and stimulate tomato ripening (Hoffmann 1953). The advantageous effects of melatonin in plants, including its influence on cell enlargement, root development, and stress mitigation, have been highlighted in numerous studies (Arnao and Hernández-Ruiz 2020). Melatonin’s role as a potent scavenger of free radicals and its ability to detoxify various chemical contaminants make it a promising candidate for mitigating herbicide drift (Hacışevki and Baba 2018). These observations are consistent with the results obtained in this study.
At 14 DAA, plants preconditioned with melatonin and 2,4,6-T exhibited the lowest injury, while the control, fenclorim, and benoxacor treatments showed higher injuries. Melatonin’s antioxidant capacity allows it to scavenge reactive oxygen and nitrogen species, thereby protecting plants from environmental stress (Arnao and Hernández-Ruiz 2014). The protective effects of safeners have been documented in other plant species. For example, peas immersed in a solution with 2,4,6-T exhibit less stem curvature when exposed to 2,4-D (Hartman 1959).
Data from 21 DAA indicated that seeds treated with fenclorim, melatonin, and 2,4,6-T resulted in less tomato tissue injury compared with benoxacor and the control group. Furthermore, all treatments showed a greater dry biomass accumulation than the control. Fenclorim has been shown to promote antioxidant effects and protect rice plants from the effects of pretilachlor (Hu et al. 2021). Similarly, pretreatment with melatonin reduced membrane damage and lipid oxidation and stimulated antioxidant enzyme activity in poplar leaves (Hacışevki and Baba 2018). Although benoxacor did not result in less injury than the control treatment at 21 DAA (Fig. 2), seeds pretreated with benoxacor had higher dry biomass accumulation post-herbicide application than the control (Fig. 4). The biochemical mechanism of benoxacor involves the detoxification of herbicides through conjugation with the tripeptide glutathione (GSH), an antioxidant enzyme found in various organisms (Narayanankutty et al. 2019).
Despite the protective effects observed with melatonin, fenclorim, and 2,4,6-T, the toxicity of herbicides on tomato leaves increased as the number of DAAs increased.
Safeners are a diverse group of agrochemicals that selectively protect crops from harm without affecting weed control. Pesticide detoxification is a complex process that involves various signaling pathways and mechanisms. Their crop-specific modes of action have yet to be determined and will require further research (Giannakopoulos et al. 2020).
Most plants undergo a three-phase process to facilitate the detoxification of xenobiotic compounds like herbicides and insecticides and convert them into intermediate substances with reduced phytotoxicity (Sandermann 1992). Using herbicide safeners to protect plants against xenobiotic effects is known to go through glutathione S-transferase (GSTs) conjugation during phase II, promoting detoxification. Acting as the primary phase II detoxification enzymes, GSTs safeguard living cells by facilitating the conjugation of reduced glutathione (GSH) to a diverse range of electrophilic molecules originating from internal and external sources (Edwards et al. 2011).
Tomatoes are highly sensitive to various herbicides, especially 2,4-D and dicamba, where even minimal, unintended drift poses a risk to the entire crop (de Paula Medeiros et al. 2023). The recommendation for pre-herbicide application in tomato production depends on herbicide efficacy and weed density (Flint and Klonsky 1985). Therefore, effective weed management is crucial for ensuring optimum quality and high yield, which may involve nonchemical control methods such as mechanical and cultural practices.
Conclusion
This study demonstrated that preconditioning tomatoes with safeners significantly increased the tolerance of tomato plants to herbicide exposure. The results showed that herbicides of different modes of action, such as protoporphyrinogen oxidase inhibitor herbicide (PPO), Photosynthetic Inhibitors the photosystem II (PSII), and synthetic auxin, are harmful even in small doses to tomatoes, damaging their leaves and stems. However, safener fenclorim reduced sulfentrazone herbicide (PPO) damage by more than 50%; 2,4,6-T reduced PPO, PSII, and auxin damage by more than 50%; and melatonin reduced sulfentrazone (PSII) damage by more than 50% after 2,4-D (auxin) exposure. Moreover, 2,4,6-T, melatonin, and benoxacor promoted dry biomass accumulation.
Overall, this study highlights the efficacy of safeners in protecting tomato crops from herbicide injury. Implementing safener preconditioning in tomato cultivation practices can contribute to developing more effective herbicide formulations, help safeguard the productivity and quality of tomato yields, and promote sustainable agriculture practices.
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