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. 2026 Aug 10;42(5):606–616. doi: 10.5423/PPJ.OA.05.2026.0071

Host-Associated Distribution and Mycological Characteristics of Fusarium sambucinum Species Complex and Other Fusarium Species in Rice, Maize, and Weeds in Korea

Eunji Jeong 1, Jeong-Ah Seo 1,*
PMCID: PMC13655601  PMID: 42572801

Abstract

Fusarium asiaticum and Fusarium graminearum, belonging to the Fusarium sambucinum species complex (FSAMSC), are primary causal agents of Fusarium head blight (FHB) and associated mycotoxin contamination in cereal crops worldwide. In this study, we performed a nationwide survey of Fusarium populations in rice, a staple crop in Korea, and examined the distribution of trichothecene genotypes and the mycological characteristics of dominant species. A total of 1,008 Fusarium isolates were obtained from rice (375) and plants surrounding rice paddy fields, including maize (133) and gramineous weeds (500). FSAMSC members accounted for the majority (59–70%) of Fusarium isolates from the three plants, with F. asiaticum predominating in both rice (57%) and weeds (44%), whereas F. graminearum was dominant in maize (41%). Trichothecene genotype analysis revealed that the nivalenol (NIV) type of F. asiaticum predominated in rice (82%) and weeds (70%), whereas the 15-acetyldeoxynivalenol (15ADON) type of F. graminearum was more common in maize (83%). A comparison of the mycological traits of two dominant species highlighted that F. asiaticum NIV type strains exhibited significantly higher pathogenicity in rice than F. graminearum 15ADON type strains (P = 0.01), despite lower growth and reproductive capacity (P < 0.05). By integrating this large-scale distribution survey with mycological analyses, we showed that F. asiaticum is a major contributor to FHB and trichothecene risks in the rice cultivation system in Korea.

Keywords: Fusarium asiaticum, nivalenol chemotype, pathogenicity, rice, trichothecene

Fusarium is a major plant pathogenic fungus responsible for economically devastating diseases of cereal crops worldwide, leading to substantial agricultural losses and posing a serious threat to food safety through mycotoxin contamination (Goswami and Kistler, 2004; Matelionienė et al., 2024). Among these, Fusarium asiaticum and Fusarium graminearum, representative species of the Fusarium sambucinum species complex (FSAMSC), are the causal agents of Fusarium head blight (FHB) in cereals such as rice and wheat (del Ponte et al., 2022; McMullen et al., 2012). FHB pathogens cause significant yield reductions and quality deterioration, largely due to the production of harmful mycotoxins, trichothecenes, in infected grains (Chen et al., 2019; Subramanian et al., 2025). These fungi persist across a range of agroecosystem reservoirs, including crop residues and alternative hosts, which serve as a constant inoculum source for the next cultivation term (Dong et al., 2021; Matelionienė et al., 2022; Trail, 2009). Therefore, understanding their ecological distribution and pathogenic potential is essential for effective management of FHB and other Fusarium-related diseases (Jeon et al., 2025; Lee et al., 2010).

Rice, the principal cereal crop in Korea, is often cultivated in association with wheat or barley as double crops in the southern regions and with maize cultivated in marginal areas adjacent to paddy fields in Gangwon Province (Choi et al., 2023a; Kim et al., 2024; Seo et al., 2021; Song et al., 2014). Within these rice-based cropping systems, previous field surveys have reported a diverse range of Fusarium species across various cereal hosts (Choi et al., 2021; Ok et al., 2014). F. asiaticum is the dominant species in rice (Jang et al., 2019; Shin et al., 2018), and other species of FSAMSC, Fusarium incarnatum–equiseti species complex (FIESC), and Fusarium fujikuroi species complex (FFSC) are also found in the previous reports (Choi et al., 2018; Kim et al., 2012; Thao et al., 2025). In wheat and barley, FSAMSC members similarly dominant and frequently co-occur with species within Fusarium tricinctum species complex (FTSC) (Choi et al., 2019a; Yang et al., 2021). In maize, both FSAMSC and FFSC, specifically F. graminearum and Fusarium verticillioides, have been commonly reported (Lee et al., 2012). Given the widespread occurrence of diverse Fusarium species across rice and co-cultivated crops in Korea, an integrated analysis of multihost-associated composition of the Fusarium community is needed (Choi et al., 2019b; Yang et al., 2021).

Among the mycotoxins produced by Fusarium, trichothecenes, including nivalenol (NIV), deoxynivalenol (DON), and their acetylated derivatives, are of particular concern because they frequently contaminate grains and are toxic to humans and livestock (Foroud et al., 2019; Wang and Cheng, 2017). FHB-causing pathogens produce trichothecenes during the infection process, which are important factors for pathogen aggressiveness and host colonization (Pinton and Oswald, 2014; Zingales et al., 2021). Trichothecene-producing strains are classified into three major genotypes based on their toxin-producing ability: NIV, 15-acetyldeoxynivalenol (15ADON), and 3-acetyldeoxynivalenol (3ADON) (Dong et al., 2021; Wang and Cheng, 2017). The trichothecene genotypes of F. asiaticum and F. graminearum, two principal FHB pathogens, are predominantly NIV and DON types, respectively, and are closely associated with host adaptation and pathogenicity of two species (del Ponte et al., 2022; Foroud et al., 2019; Yulfo-Soto et al., 2024). While FHB outbreaks and mycotoxin contamination in wheat are largely associated with the F. graminearum 15ADON type, the risks posed by the F. asiaticum NIV type are of concern in rice-growing regions, particularly in East Asia (Lee et al., 2009; Lipps et al., 2025; Sun et al., 2025; van der Lee et al., 2015). Given that trichothecene genotypes substantially affect the aggressiveness and ecological distribution of pathogens, understanding the competitive interactions among genotypes is crucial for assessing pathogen dynamics and mycotoxin risks in agricultural settings.

In this study, we comprehensively analyzed the species composition, trichothecene genotype distribution, and mycological traits of major FHB pathogens based on a large number of Fusarium strains isolated from rice in Korea. A total of 1,008 Fusarium isolates were obtained from rice and plants adjacent to rice fields (maize and weeds) collected nationwide, and their species compositions were determined via molecular identification. Furthermore, we investigated the distribution of trichothecene genotypes for the two major species, F. asiaticum and F. graminearum, across three host plants. By analyzing the distribution and mycological characteristics, including pathogenicity, of numerous rice FHB pathogens distributed nationwide, this study may elucidate the ecological compatibility between the major pathogens and the crops or plants that primarily occur in rice-centered agricultural systems.

Materials and Methods

Collection of rice, maize, and gramineous weed samples

Samples of rice, maize, and gramineous weed were collected from paddy fields and surrounding areas in cultivation regions nationwide in Korea (Fig. 1). A total of 70 rice samples were collected between September and October (pre-harvest stage) in 2020 and 2022 from 57 locations across central and southern provinces (Supplementary Table 1). Central provinces include Gyeonggi-do (GG) and Gangwon-do (GW), and southern provinces include Jeollabuk-do (JB), Jeollanam-do (JN), Gyeongsangbuk-do (GB), Gyeongsangnam-do (GN), Chungcheongbuk-do (CB), and Chungcheongnam-do (CN).

Fig. 1.

Fig. 1

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Geographic distribution of sampling locations of rice, maize, and weed samples nationwide in Korea. The locations where rice, maize, and weed samples were collected are indicated. The symbols represent host types: rice (red circle), weed (green circle), rice and maize (green triangle), rice and weed (black triangle), and rice, maize, and weed (black circles).

Thirteen maize samples cultivated near the paddies were collected from five locations in GW during the same period as the rice sampling (Supplementary Table 2). 342 gramineous weed samples were collected from 39 locations adjacent to rice paddies between May and June 2018 (Supplementary Table 3) (Ahn et al., 2022).

Fusarium isolation and species identification

Fusarium isolates were obtained from rice grains, maize kernels, and weed tissues using the single-spore isolation method (Ahn et al., 2022). In total, 375, 133, and 500 Fusarium isolates were obtained from rice, maize, and weed samples, respectively (Supplementary Tables 1–3). Mycelial and conidial suspensions of each isolate were stored in 20% glycerol at −80°C until further use.

Genomic DNA was extracted from approximately 0.05 g of lyophilized mycelium using the DNeasy Plant Mini Kit (Qiagen, Hilden, Germany). Species identification was performed by PCR amplification and sequence analysis of the translation elongation factor 1 α (TEF-1α) gene (O’Donnell et al., 1998) (Supplementary Table 4). The obtained TEF-1α sequences were compared with reference sequences in the NCBI database using BLAST for species-level identification. Species complex assignments were subsequently confirmed by referring to the NCBI Fusarium Taxonomy Database. For isolates identified as F. asiaticum or F. graminearum, additional PCR assays targeting the TEF-1α and TRI12 genes were performed to respectively confirm species and determine trichothecene genotypes based on the size of the amplified PCR products (Ward et al., 2002)(Supplementary Table 4).

Mycological characterization of F. asiaticum and F. graminearum.

To compare the mycological traits of the major Fusarium species isolated from rice, we randomly selected 20 strains each of F. asiaticum NIV type and the F. graminearum 15ADON type, representing the dominant genotypes in our collection (Supplementary Table 5). Additionally, nine strains of 3ADON type for each species were included to characterize the minor genotypes. Owing to the limited number of F. asiaticum 3ADON type strains isolated from rice (n = 5), four additional strains were randomly selected from weed-derived isolates to complete the sample set. The reference strain F. graminearum Z3639 (15ADON type) was used as the positive control.

To characterize the phylogenetic relationships among the 58 strains selected for mycological trait analysis, a phylogenetic analysis based on TEF-1α gene sequences was performed (Supplementary Fig. 1). The TEF-1α gene sequences of the 58 F. asiaticum and F. graminearum strains, together with those of 11 representative species of Fusarium, were used to construct a neighbor-joining phylogenetic tree in MEGA X with 1,000 bootstrap replicates. The TEF-1α sequence of F. asiaticum KCTC 16664 (GCA_025258505.1) was extracted from its genome assembly, whereas those of the remaining 10 representative Fusarium species were obtained from the GenBank database: F. graminearum PH-1 (MW233103), F. vorosii 1339/2 (MF974401), F. boothii CBS 316.73 (MW233088), F. kyushuense CBS 121807 (PQ260875), F. poae 151947 (PQ260906), F. incarnatum CB 132907 (MN170477), F. equiseti CBS 414.86 (MN170467), F. fujikuroi NRRL 5538 (MN193860), F. tricinctum NRRL 25481 (OL772833), and F. solani CBS 102429 (KM231936). To facilitate visualization of species assignments, strains identified as F. asiaticum and F. graminearum were highlighted by shading in Supplementary Fig. 1.

The mycelial growth rate, conidial production, and perithecia formation of 58 F. asiaticum and F. graminearum strains were evaluated following established procedures (Ahn et al., 2022; Jeong et al., 2024). For all mycological assays, 4-mm mycelial plugs obtained from 3-day-old cultures of each strain on potato dextrose agar (PDA) at 25°C were used as standardized inocula. Mycelial growth was determined by measuring colony diameter after 7 days of incubation of a single 4-mm mycelial plug on PDA at 25°C. Conidial production was quantified from suspensions obtained by culturing five 4-mm mycelial plugs in potato dextrose broth at 25°C with shaking at 150 rpm for 3 days. Perithecia production was assessed by culturing a single 4-mm mycelial plug on carrot agar at 25°C for 7 days under near-ultraviolet light.

Chemical analysis of trichothecene production of F. asiaticum and F. graminearum.

The production of type-B trichothecenes by the 58 strains of F. asiaticum and F. graminearum was examined as previously described with minor modifications (Ahn et al., 2022; Ok et al., 2014). Five 4-mm-diameter mycelial plugs from each strain were incubated in white rice medium at 25°C for 3 weeks. The cultures were air-dried for 3 days and subsequently ground into a fine powder. Trichothecenes were extracted from 5 g of the powdered culture using Q-Sep QuEChERS extraction salts and dSPE tubes (Restek, Bellefonte, PA, USA), according to the manufacturer’s protocol. The extracts were filtered through a 0.45 μm PTFE syringe filter prior to analysis.

Quantification of trichothecenes was performed using high-performance liquid chromatography equipped with a UV detector and a C18 column (4.6 × 250 mm, 5 μm, Restek). The mobile phase consisted of acetonitrile (ACN) and ultrapure water (UPW), and elution was carried out at a flow rate of 0.4 mL/min using the following gradient: 0–10 min, 10% ACN, 90% UPW; 10–22 min, 30% ACN, 70% UPW; 22–24 min, 60% ACN, 40% UPW; 24–34 min, 90% ACN, 10% UPW; 34–36 min, 60% ACN, 50% UPW; and 36–60 min, 10% ACN, 90% UPW. Detection was performed at 218 nm. A trichothecene standard mixture (Romer Labs, Union, MO, USA) containing NIV, 4-acetylnivalenol (4ANIV), DON, 15ADON, and 3ADON was used for standard.

Disease assessment of major rice-derived Fusarium genotypes on rice and wheat

The FHB-inducing ability and trichothecene occurrence of 58 strains of F. asiaticum and F. graminearum were evaluated on rice and wheat (Ahn et al., 2022; Jeong et al., 2024). Spore suspensions of each strain were spray-inoculated onto rice panicles and spot-inoculated onto wheat spikes, and the inoculated plants were incubated in a greenhouse for 3 weeks. Disease progression was monitored weekly and quantified by calculating the area under the disease progress curve (AUDPC) based on the number of grains exhibiting FHB symptoms. After 3 weeks, infected plants were harvested, and the amount of trichothecene accumulated in the plant tissues was quantified using the extraction and quantitative methods described above.

Statistical analysis

Differences in the distributions of Fusarium species and trichothecene genotypes in rice, maize, and weeds were assessed using a chi-square test (χ2, P < 0.01). Differences in mycological characteristics, trichothecene production, and pathogenicity among trichothecene genotype groups were evaluated using pairwise two-tailed Welch’s t-tests. Differences were considered statistically significant at P < 0.05 and highly significant at P < 0.01.

Results

Species composition of Fusarium populations across rice, maize, and weed

A total of 375 Fusarium isolates were obtained from rice and identified as 16 species belonging to four species complexes (Figs. 2 and 3). The FSAMSC was the most prevalent, accounting for approximately 70% of the total isolates, followed by FIESC (23%) and FFSC (7%) (Fig. 2). F. asiaticum (215 isolates, 57%) was the most frequently isolated species, followed by F. incarnatum (59 isolates, 16%) and F. graminearum (41 isolates, 11%) (Fig. 3A).

Fig. 2.

Fig. 2

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Isolation frequency of Fusarium species complexes isolated from rice, maize, and weeds. The stacked bar charts show the isolation frequencies (%) of Fusarium isolated from rice, maize, and weeds classified at the species complex level. FSAMSC, Fusarium sambucinum species complex; FIESC, Fusarium incarnatum-equiseti species complex; FFSC, Fusarium fujikuroi species complex; FTSC, Fusarium tricinctum species complex; Others, minor complexes with low frequency. The numbers inside the bars represent the percentage of each complex.

Fig. 3.

Fig. 3

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Species composition of Fusarium isolated from rice, maize, and weeds. Isolation frequencies (%) of Fusarium species isolated from (A) rice, (B) maize, and (C) weeds. Species were grouped by species complex and listed according to isolation frequency. The numbers above the bars indicate the isolation frequency of species exceeding 10% of the total isolates from each host. FSAMSC, Fusarium sambucinum species complex; FIESC, Fusarium incarnatum-equiseti species complex; FFSC, Fusarium fujikuroi species complex; FNSC, Fusarium nisikadoi species complex (Sanna et al., 2023); FTSC, Fusarium tricinctum species complex; FSSC, Fusarium solani species complex.

In addition, 133 and 500 Fusarium isolates were obtained from maize and weeds, respectively (Figs. 2 and 3). Maize-derived isolates were identified as 12 species belonging to four species complexes. Similar to rice, FSAMSC (59% of total isolates) was the most dominant in maize, but at the species level, F. graminearum (59 isolates, 44%) was the most frequently isolated. Members of FFSC (29%) were also commonly appeared, including F. verticillioides (23 isolates, 17%) and F. fujikuroi (15 isolates, 11%) (Fig. 3B). Weed-originated isolates were identified as 13 species belonging to four species complexes, with FSAMSC accounting for approximately 60% of the total isolates. F. asiaticum (206 isolates, 41%) was the most often obtained species, and FTSC members were also abundant, with F. avenaceum (90 isolates, 18%) and F. acuminatum (88 isolates, 18%) (Fig. 3C).

Distribution of trichothecene genotypes in F. asiaticum and F. graminearum across rice, maize, and weed

The trichothecene genotypes of 256 isolates of F. asiaticum and F. graminearum, the two major species of FSAMSC isolated from rice, were examined, revealing four species–trichothecene genotype combinations (Fig. 4A). Overall, the F. asiaticum NIV type (210 isolates, 82%) was the most common genotype, followed by the F. graminearum 15ADON type (30 isolates, 12%). The 3ADON type was found at a low frequency in both species.

Fig. 4.

Fig. 4

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Distribution of trichothecene genotypes of F. asiaticum and F. graminearum according to the host. Trichothecene genotypes of F. asiaticum and F. graminearum isolated (A) from rice and (B) from rice, maize, and weeds. Rice- and weed-derived isolates were further grouped according to the sampling region (central and southern provinces). The genotypes included F. asiaticum NIV and 3ADON types and F. graminearum 15ADON and 3ADON types. The numbers within the bars indicate the percentages.

The distribution of trichothecene genotypes of the two species isolated from maize and weeds was also examined (Fig. 4B). The F. asiaticum NIV type predominated in both rice and weeds (195 isolates, 70%), whereas the F. graminearum 15ADON type was the major genotype in maize (58 isolates, 82%). The trichothecene genotype distribution was compared between the central and southern provinces based on the sampling locations of rice and weed samples, and the F. graminearum 15ADON type was detected more frequently in host plants from the central regions (Supplementary Tables 1 and 3). The isolation frequency of the F. graminearum 15ADON type strain increased from 7% to 29% in rice and from 6% to 53% in weeds, indicating a higher occurrence in the central region than in the southern region.

Mycological characteristics of F. asiaticum and F. graminearum.

Among the 375 Fusarium isolates isolated from rice, 29 strains each of F. asiaticum (NIV and 3ADON types) and F. graminearum (15ADON and 3ADON types) were mycologically characterized (Supplementary Table 5). The 58 selected strains were significantly differed in mycelial growth and conidial production among the trichothecene genotype groups (Table 1). In particular, F. graminearum 15ADON type strains showed significantly higher mycelial growth rates and conidial production than F. asiaticum NIV type strains (P = 0.032 and 0.033, respectively) and F. asiaticum 3ADON type strains (P < 0.001 and 0.022, respectively). F. graminearum 3ADON type strains were not different from the other genotype groups in either mycelial growth or conidial production. For perithecia formation, most F. asiaticum strains (25/29 isolates) were classified as level I, whereas most F. graminearum strains (18/29 isolates) as level IV (Table 1).

Table 1.

Comparison of mycological characteristics of F. asiaticum and F. graminearum

Species Trichothecene genotype (No. of strains) Mycelial growth (cm) Conidiation (104 spores/mL) No. of strains that produce perithecia*
I II III IV
F. asiaticum NIV (20) 9.6 ± 0.7a 17.6 ± 32.0a 17 1 1 1
3ADON (9) 9.1 ± 0.5b 0.4 ± 0.2b 8 1 0 0
F. graminearum 15ADON (20) 10.1 ± 0.7c 230.9 ± 413.1c 2 1 2 15
3ADON (9) 9.7 ± 1.2a,b,c 160.6 ± 264.2a,b,c 5 1 0 3
F. graminearum Z3639 15ADON 8.3 ± 1.8 539.3 ± 146.5 - - 1 -
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Values are presented as mean ± standard deviation.

Different letters within a row indicate significant differences at P < 0.05 based on pairwise two-tailed Welch’s t-tests.

*

The level of perithecia production of each strain was assessed by dividing into four groups based on the proportion of the agar surface covered with perithecia: I, under 25%; II, 25–50%; III, 50–75%; and IV, over 75%.

Trichothecene production of F. asiaticum and F. graminearum.

Trichothecene production differed markedly among the trichothecene genotype groups (Table 2). Among the strains analyzed, F. asiaticum NIV type strains produced NIV and 4ANIV, with 4ANIV levels about threefold higher than NIV. In contrast, F. graminearum 15ADON type strains produced DON and 15ADON, with DON levels approximately 35-fold higher than 15ADON. The 3ADON type strains of both species produced DON and 3ADON, with 3ADON levels approximately threefold higher than DON. Overall, the average amount of the total trichothecenes produced by F. asiaticum strains was approximately threefold lower than that produced by F. graminearum strains (Table 2).

Table 2.

B-type trichothecene production of F. asiaticum and F. graminearum

Species Trichothecene genotype (No. of strains) B-type trichothecene production (μg/g)
NIV 4ANIV DON 15ADON 3ADON
F. asiaticum NIV (20) 157.3 ± 131.4 346.0 ± 286.3 nd nd nd
3ADON (9) nd nd 143.9 ± 110.5 nd 561.0 ± 353.4
F. graminearum 15ADON (20) nd nd 1,574.9 ± 1,177.4 44.6 ± 25.9 nd
3ADON (9) nd nd 571.7 ± 435.7 nd 1,612.0 ± 978.3
F. graminearum Z3639 15ADON nd nd 2,434.4 ± 313.9 277.9 ± 71.3 nd
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Values are presented as mean ± standard deviation.

nd, not detected.

Pathogenicity of F. asiaticum and F. graminearum on rice and wheat

All 58 strains of F. asiaticum and F. graminearum tested were confirmed as pathogenic on both rice and wheat (Supplementary Table 6), but aggressiveness differed by species (Table 3). F. asiaticum strains showed significantly higher AUDPC in rice (P < 0.01), whereas F. graminearum strains had a higher AUDPC in wheat (P < 0.01) (Table 3). F. asiaticum NIV (P = 0.018) and 3ADON type strains (P = 0.011) exhibited significantly higher AUDPC values on rice than on wheat. In contrast, F. graminearum 15ADON type strains showed significantly higher AUDPC values on wheat than rice (P < 0.01).

Table 3.

Degree of pathogenicity and B-type trichothecene occurrence of F. asiaticum and F. graminearum on different hosts

Host Species Trichothecene genotype (No. of strains) AUDPC B-type trichothecene occurrence (μg/g)
NIV 4ANIV DON 15ADON 3ADON
Rice F. asiaticum NIV (20) 148 ± 56a nd 2.8 ± 0.7 nd nd nd
3ADON (9) 162 ± 24a nd nd 1.4 ± 0.1 nd 1.5 ± 0.1
F. graminearum 15ADON (20) 105 ± 43b nd nd 1.3 ± 0.1 1.3 ± 0.0 nd
3ADON (9) 119 ± 18b nd nd 1.2 ± 0.1 nd 1.4 ± 0.2
F. graminearum Z3639 15ADON 68 nd nd 1.2 1.4 nd
Wheat F. asiaticum NIV (20) 113 ± 27a nd 2.3 ± 0.5 nd nd nd
3ADON (9) 131 ± 22a,c nd nd 1.1 ± 0.7 nd 1.8 ± 0.1
F. graminearum 15ADON (20) 151 ± 17b nd nd 2.3 ± 0.6 2.6 ± 0.8 nd
3ADON (9) 144 ± 36b,c nd nd 2.9 ± 3.7 nd 3.4 ± 1.2
F. graminearum Z3639 15ADON 70 nd nd nd 1.7 nd
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Values are presented as mean ± standard deviation.

nd, not detected.

Different letters within a row indicate significant differences at P < 0.05 based on pairwise two-tailed Welch’s t-tests.

Consistent with the pathogenicity scores, trichothecene accumulation was higher in rice infected with F. asiaticum and wheat infected with F. graminearum (Table 3). In both hosts inoculated with F. asiaticum NIV type strains, only 4ANIV was detected, with higher levels observed in rice than wheat (P < 0.01). F. asiaticum 3ADON type produced similar levels of DON and 3ADON in both hosts. In contrast, both rice and wheat inoculated with F. graminearum 15ADON or 3ADON types contained DON along with its acetylated derivatives, with trichothecene levels approximately two- to three-fold higher in wheat (P < 0.05).

Discussion

In this nationwide survey of Fusarium populations across rice-growing regions in Korea, we determined that host plants surrounding rice paddies significantly influenced the shape of Fusarium populations. Members of FSAMSC were the most frequently isolated among Fusarium from rice and plants adjacent to paddy fields (maize and weeds); however, the frequency of major FSAMSC species and other Fusarium species differed significantly among host plants (χ2 test, P < 0.01) (Fig. 2). F. asiaticum accounted for the largest proportion of the Fusarium community in rice and was also isolated in substantial numbers in maize and weeds (Fig. 3), confirming that F. asiaticum is widely distributed across paddy fields (Cao et al., 2025; Choi et al., 2021; Choi et al., 2023b; Qiu et al., 2024). In contrast to other plants, maize had a higher prevalence of F. graminearum than F. asiaticum, along with a wide distribution of FFSC species, including F. verticillioides (Fig. 3B). This pattern suggests that maize cultivated near rice paddies serves as an alternative host contributing to the dispersal of both FHB pathogens and FFSC species responsible for bakanae disease (Basler, 2016; Sun et al., 2025; Qiu et al., 2025). Furthermore, gramineous weeds collected from paddy margins were dominated by FSAMSC species, including F. asiaticum, alongside FTSC members pathogenic to wheat and barley (Fig. 3C) (Choi et al., 2019a; Yang et al., 2021). These findings highlight those non-cultivated plants within agricultural ecosystems can act as critical reservoirs for Fusarium pathogen inoculum (Ahn et al., 2022; Dong et al., 2021; Suproniene et al., 2018). The diverse Fusarium species distributed according to host in rice paddies suggests that Fusarium populations are not randomly distributed but are fundamentally linked to plant reservoirs that contribute to the persistence and spread of these pathogens within agroecosystems.

F. asiaticum and F. graminearum, the dominant species in rice paddies identified in this study, are well-known FHB-causing pathogens that contaminate a wide range of cereal crops with trichothecenes (Aoki et al., 2012; Goswami and Kistler, 2004; Laraba et al., 2021). Previous research has shown that trichothecene synthetic profiles of these two species are associated not only with pathogenicity but also with differences in distribution of the two species depending on the host crops (Liu et al., 2026; Sun et al., 2025; Suproniene et al., 2018). Consistent with this, we observed distinct distribution patterns of trichothecene genotypes of F. asiaticum and F. graminearum across rice, maize, and weeds (Fig. 4). The F. asiaticum NIV type was predominantly isolated from rice and weeds, whereas the F. graminearum 15ADON type was more frequent in maize (Qiu and Shi, 2014; Xu et al., 2021; Yang et al., 2018). When further comparing the trichothecenes genotype of the two species from rice and weeds between the central and southern provinces where samples were collected, the F. graminearum 15ADON type was markedly more prevalent in the central region (Fig. 4B). This regional difference is likely attributable to local practices of crop cultivation in some central regions, such as the cultivation of maize in marginal areas near rice paddy fields (Lee et al., 2010; Lee et al., 2012). Thus, the proximity of maize likely promotes the accumulation and dispersal of DON-producing F. graminearum to surrounding farming areas (Jang et al., 2019; van der Lee et al., 2015; Sun et al., 2025). These findings indicate that the trichothecene genotypes of F. asiaticum and F. graminearum vary depending on the host type, indicating that nearby plants may affect the spread of NIV and DON-producing pathogens into rice paddies and the resulting mycotoxin risks.

To further understand the biological characteristics of the major species in rice-growing regions, F. asiaticum and F. graminearum, we compared the mycological characteristics, trichothecene production, and pathogenicity to rice and wheat of two species. F. graminearum strains, particularly the 15ADON type, exhibited faster mycelial growth and produce greater numbers of conidia and perithecia than F. asiaticum (P < 0.05) (Table 1) (Ahn et al., 2022; Xi et al., 2021). In addition, F. graminearum 15ADON type strains produced approximately threefold higher levels of trichothecenes than F. asiaticum (Table 2) and exhibited higher pathogenicity and DON production in wheat (P < 0.05) (Table 3). Consequently, F. graminearum, especially the 15ADON type, possesses mycological attributes that facilitate its survival and colonization in diverse environments and contribute to disease development and toxin production in cereal cultivation systems (Lee et al., 2014; Singh et al., 2024; van der Lee et al., 2015; Xu et al., 2021). In contrast, although the growth, reproductive capacity, and toxin production were significantly lower than those of F. graminearum (Tables 1 and 2), F. asiaticum NIV type exhibited high pathogenicity and 4ANIV production in rice (Table 3). Also, our study confirmed that the F. asiaticum NIV type is the dominant species in rice throughout Korea (Table 1), consistent with widespread trends in rice-growing regions of East Asia (Pu et al., 2025; Sun et al., 2025; Yuan et al., 2025). This suggests that F. asiaticum possesses rice-specific advantageous mechanisms that enable its dominance and disease development in rice, despite its limited growth and reproductive capacity (Qiu and Shi, 2014; Ramezanpour et al., 2026; Yang et al., 2018). Such host-associated traits likely provide a selective advantage to F. asiaticum in rice-growing environments by supporting efficient colonization and sustained accumulation of NIV-type trichothecenes (del Ponte et al., 2022; Dong et al., 2020; Liu et al., 2026; Xinhao et al., 2026). Overall, the differences in growth, toxin production, and host-dependent pathogenicity between F. asiaticum and F. graminearum can serve as biological basis for their contrasting host preferences and distributions in cereal-based agricultural environments (Karlsson et al., 2021; Xu et al., 2021).

Collectively, our findings indicate that, although diverse host plants in rice-based agroecosystems in Korea serve as reservoirs of Fusarium inoculum, F. asiaticum, particularly the NIV genotype, remains the predominant pathogen. Notably, F. asiaticum showed rice-specific ecological fitness, specifically its significantly higher pathogenicity and trichothecene production on rice, despite exhibiting relatively lower mycelial growth and reproductive capacity than F. graminearum. These results suggest that the dominance of F. asiaticum NIV type in rice-based agricultural systems may be associated with its higher host-specific aggressiveness and adaptation to rice. This study is expected to contribute to establishing strategic approaches for disease control by understanding the genetic and mycological characteristics of major pathogens in rice-centered agricultural ecosystems and by identifying host affinity and surrounding plants that serve as reservoirs for the pathogens.

Footnotes

Conflicts of Interest

No potential conflict of interest relevant to this article was reported.

Acknowledgments

This work was supported by the Rural Development Administration (RDA), Republic of Korea, under the grant “Cooperative Research Program for Agriculture Science and Technology Development” [RS-2025-00512693].

Electronic Supplementary Material

Supplementary materials are available at The Plant Pathology Journal website (http://www.ppjonline.org/).

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