Reproduction of Aphelenchoides besseyi sensu lato on fungal biological control agents

ABSTRACT

Aphelenchoides besseyi sensu lato is an emerging nematode that causes green stem and foliar retention syndrome in soybeans and cotton. As sustainable agriculture gains importance, biological control agents (BCAs) have become a key strategy for pest management. However, the influence of A. besseyi s. l. mycophagous behavior on fungal BCAs remains unclear. This study examined its feeding and reproductive abilities of eight fungal isolates commonly used in disease biological control: Trichoderma afroharzianum (T10), Pochonia chlamydosporia (PC10), Pochonia boninensis (PC17), Clonostachys pseudochroleuca (C05), Clonostachys chloroleuca (C17), Clonostachys farinosa (C35), Clonostachys rogersoniana (C101), and Clonostachys rosea (C133). The isolates were cultured on oatmeal agar (OA) and potato dextrose agar (PDA) and inoculated with A. besseyi s. l. The nematode fed and reproduced on T10, PC10, PC17, and C35 across both media. On C101 and C05, reproduction occurred only on PDA and OA, respectively, with mortality observed on C05 in OA. Feeding or reproduction was absent on C17 and C133, highlighting their potential as BCAs against A. besseyi s. l. These findings emphasize A. besseyi s. l.’s capacity to exploit fungal BCAs in vitro, which may undermine fungal disease control in infested areas.

Key words
biocontrol fungi; foliar nematode; green stem; foliar retention syndrome

INTRODUCTION

Aphelenchoides besseyi has been considered one of the emerging nematode species of significant phytosanitary and economic importance, causing green stem and foliar retention syndrome (GSFRS), formerly known as crazy soybean II (Meyer et al. 2017). GSFRS causes significant damage to soybeans, with losses reaching up to 100%, particularly in regions with hot and humid climates (Meyer et al. 2017). Aphelenchoides besseyi is a facultative plant parasitic nematode, able to survive in the host plant’s absence by feeding fungi in the soil or crop residues. Under extreme dehydration conditions, it can enter anhydrobiosis (De Jesus et al. 2016).

Recently, a reclassification of A. besseyi was proposed into Aphelenchoides besseyi sensu stricto, Aphelenchoides oryzae, and Aphelenchoides pseudobesseyi, forming a cryptic species complex (Subbotin et al. 2020). This reclassification is recent; thus, much of the available knowledge, such as geographic distribution and host ranges, predates this update and needs to be reviewed for each one of those cryptic species. Also, many studies published after 2020 preferred keeping the old classification to avoid inconsistency with the established literature. As a result, many studies still refer to the group broadly as A. besseyi. Since the new classification is not yet widely adopted and key biological and genetic traits need to be re-evaluated, we refer to the group as A. besseyi sensu lato in this manuscript.

Among the alternatives for disease control, biological control has gained prominence, driven by social pressure for sustainable agriculture (Embrapa 2023). The fungi Trichoderma afroharzianum, Pochonia chlamydosporia, Pochonia boninensis, Clonostachys chloroleuca, Clonostachys pseudochroleuca, Clonostachys rogersoniana, Clonostachys rosea, and Clonostachys farinosa are species within genera known to be biocontrol agents of nematodes and other pathogens. The fungi act through mechanisms such as hyperparasitism, resource competition, production of antagonistic metabolites, and induction of plant resistance (Iqbal et al. 2018, Pacheco et al. 2020). Thus, they can be potential biological control agents for A. besseyi s. l.

Aphelenchoides besseyi s. l. can feed and reproduce on several fungal species, including phytopathogens. Fungal species known to support the growth of A. besseyi s. l. include Neocosmospora solani (Fusarium solani) (Huang et al. 1972), Alternaria alternata, F. verticillioides, F. equiseti, Phoma medicaginis (Gokte and Mathur 1989), F. oxysporum, Curvularia spp., A. longipes, A. padwickii (Rajan and Mathur 1990), Colletotrichum gloeosporioides, Botrytis cinerea, Neopestalotiopsis rosae, and Monilinia fructicola (Oliveira et al. 2022). However, studies investigating the A. besseyi s. l. mycophagous habits on potential biological control agents (BCAs) are scarce.

Due to insufficient information about the impact of A. besseyi s. l. mycophagy on the efficiency of biological control fungi, the dietary plasticity of A. besseyi s. l. should be considered. Therefore, the host status of biological control fungi should be evaluated before introducing them as a method in integrated management for GSFRS (De Souza et al. 2023). The mycophagous behavior of A. besseyi s. l. and its dietary plasticity may influence the effectiveness of biological control fungi, potentially impacting the success of biological control strategies against nematodes (Bae and Knudsen 2001, Kim and Knudsen 2021). Thus, this study aimed to evaluate the reproduction of A. besseyi s. l. on fungal isolates of T. afroharzianum, Pochonia spp., and Clonostachys spp.

METHODS

Specimens of A. besseyi s. l. (UFV-ApbSL-S) used in the experiment were extracted from infected soybean plants collected in Santa Rosa do Tocantins, TO, Brazil (10°02’01”S 48°46’00”W). Nematodes were identified based on morphological characteristics and molecular analyses. DNA extraction was performed using one or more nematodes, following the protocol described by Buonicontro et al. (2018). Molecular identification was conducted through the amplification and sequencing of four genomic regions: the near full-length small subunit ribosomal RNA gene (SSU rRNA), the D2-D3 expansion segment of the large subunit ribosomal RNA gene (LSU rRNA), the partial mitochondrial cytochrome c oxidase subunit I gene (mtCOI), and the internal transcribed spacer regions 1 and 2 (ITS-1 and ITS-2).

The nematodes were extracted using the Coolen and D’Herde technique and propagated on F. pseudocircinatum colonies (Coolen and D’Herde 1972). The F. pseudocircinatum isolate was provided by the Otávio de Almeida Drumond Culture Collection at Universidade Federal de Viçosa (UFV), where this fungus is preserved under the COAD 3434 code. The reproduction of A. besseyi s. l. was evaluated using eight fungal isolates listed in Table 1. Among them, only the P. chlamydosporia isolate PC10 is a commercial strain included in the formulation of the biological product Rizotec (Stoller, Brazil). The other isolates belong to the fungal collection of the UFV. Clonostachys spp. isolates are preserved at the Unidade de Controle Biológico, while the Pochonia sp. and Trichoderma sp. isolates are maintained at the Laboratório de Controle Biológico, both located at UFV.

The fungal isolates were cultivated on Petri dishes containing potato dextrose agar (PDA) (Kasvi, Brazil) and oatmeal agar (OA) (30 g of oatmeal flakes + 20 g of agar per liter). The colonies were incubated for 15 days at 25°C under a 12-h photoperiod. Afterward, a suspension containing 25 specimens of A. besseyi s. l. was inoculated adjacent to the center of each fungal colony. The inoculated plates were incubated for another 20 days under the same conditions used for fungal growth. After this, the nematodes were recovered from the Petri dishes by distilled water washing (Silva et al. 2024). The multiplication of A. besseyi s. l. on the BCAs was determined by counting the number of nematodes recovered from the Petri dishes, including live and dead nematodes, using a Peters chamber and an optical microscope (Olympus IX70). For this quantification, 1 mL of the nematode suspension was used for counting, and the result was then multiplied by the total volume of the suspension obtained from washing the entire content of the Petri dishes.

The experiment was set up in a completely randomized design in a factorial scheme 9 (fungal isolates) × 2 (culture media), with five replicates, and carried out twice. The F. pseudocircinatum isolate was used as a positive control for A. besseyi s. l. multiplication, as this nematode successfully feeds and multiplies on this fungus (Silva et al. 2024).

Table 1
List of fungal isolates of Clonostachys spp., Pochonia spp., and Trichoderma sp. evaluated in this study.

Isolated	Species	Substrate	City/state
C17	Clonostachys chloroleuca	Native Cerrado soil	Montividiu (GO)
C05	Clonostachys pseudochroleuca	Lychnophora pinaster (endophytic)	Ingaí (MG)
C101	Clonostachys rogersoniana	Soil under Araucária	São Paulo (SP)
C133	Clonostachys rosea	Prunus persica (epiphytic)	Viçosa (MG)
C35	Clonostachys farinose	Fragaria ananassa	Bento Gonçalves (RS)
PC10	Pochonia chlamydosporia	Soil contaminated with Meloidogyne sp.	Viçosa (MG)
PC17	Pochonia boninensis	Soil contaminated with Meloidogyne sp.	-
T10	Trichoderma afroharzianum	Pratylenchus brachyurus	-

The data from both trials were jointly analyzed using generalized linear mixed models (GLMM) with a negative binomial distribution (nbinom2), considering trials as a random effect. Isolate C17 was excluded due to zero nematode recovery across all culture media (PDA and OA), as the nbinom2 GLMM requires variability in data. Including treatments with only zero values could distort the model and result in inaccurate interpretations. Thus, C17 was excluded to ensure an accurate model fit and interpretation. Post-hoc multiple comparisons were conducted using pairwise comparisons adjusted by Šidák (1967). Estimated marginal means (emmeans) were calculated for each fungal isolate and culture medium combination, followed by pairwise comparisons. Significance was assessed at p < 0.05. All analyses were conducted in R software version 4.4.1 (R Core Team 2024).

RESULTS AND DISCUSSION

Since this study did not focus on taxonomy, more detailed information regarding the morphological and molecular identification of the nematodes is not presented, as these data are being prepared for publication in another manuscript.

The highest reproduction rates of A. besseyi s. l. were observed on the isolates of P. chlamydosporia (PC10) and F. pseudocircinatum (FP), regardless of the culture medium used (Fig. 1, Table 2). The nematode also reproduced on C. farinosa (C35) and T. afroharzianum (T10) isolates, regardless of the culture medium, but at significantly lower rates than in PC10 and FP. However, for the C35 isolate, a significant difference was observed only on PDA when compared with PC10 and FP (Fig. 1, Table 2). The culture medium used for fungal growth affected the reproduction rate of A. besseyi s. l. on C. pseudochroleuca (C05), C. rogersoniana (C101), and P. boninensis (PC17) (Fig. 1, Table 2). The nematode could not feed and reproduce on isolate C101 or had a lower rate of reproduction on PC17 when the fungi were grown on OA medium. However, on PDA medium, the nematode population increased 28 to 167 times on C101 and PC17, respectively (Fig. 1). On the other hand, the cultivation of isolate C05 in the OA medium favored the multiplication of A. besseyi s. l., which had a population increase of 35 times, whereas this did not occur on the PDA medium (Fig. 1). In this case, the few nematodes recovered were dead, indicating a possible antibiosis effect of this fungus on A. besseyi s. l., when C05 was cultivated on PDA (data not shown). No nematode reproduction was observed on C. chloroleuca (C17) and C. rosea (C133), regardless of the culture medium (Fig. 1).

The mycophagy of an unidentified Aphelenchoides species on T. harzianum, which is used in the biocontrol of fungi and nematodes, was previously reported (Bae and Knudsen 2001, Almeida et al. 2022). Aphelenchoides sp. reduced radial growth and decreased the biocontrol ability of T. harzianum ThzID1-M3 to mycoparasitize sclerotia of Sclerotinia sclerotiorum in soil (Bae and Knudsen 2001). In addition, Kim and Knudsen (2021) demonstrated that the introduction of T. harzianum ThzID1-M3, used to control soilborne diseases, stimulated the increase of native fungivorous nematode populations in the soil and, consequently, there was a decrease in the hyphae biomass of this fungus in the soil, affecting the antagonistic effects on S. sclerotiorum (Kim and Knudsen 2021). These pieces of evidence point out that it is critical for some biocontrol fungi to keep an optimal density of fungal biomass in the soil to ensure sustained pathogen suppression and effective disease control. Also, fungivorous nematodes, such as Aphelenchoides spp., should be considered an important biotic constraint to biocontrol activity.

Figure 1
Reproduction rate of Aphelenchoides besseyi sensu lato in different species of fungi used in biological control. An exploratory bar plot was created using the ggplot2 package to visualize the nematode reproduction rate (final population divided by initial population) across both trials, which were analyzed together due to no significant interaction (p = 0.297). The plot shows standard errors as blue bars and mean reproduction rates as values above them.

Table 2
Effect of isolates of fungal biological control agents and culture media on multiplication of Aphelenchoides besseyi sensu lato*.

Isolate code	Mean nematode number (OA)	Statistical test letters	Isolate code	Mean nematode number (PDA)	Statistical test letters
C17	0	-	C17	0	-
C101	4	AB	C133	7	a
C133	8	A	C05	29	ab
PC17	91	ABCD	C35	138	bc
C35	176	BCD	T10	323	bc
T10	187	ABC	C101	689	c
C05	865	CD	PC17	4,171	d
PC10	4,006	D	PC10	5,401	d
FP	9,170	D	FP	11,301	d

• PDA: potato dextrose agar; OA: oatmeal agar;
• ^*
  significant differences (p < 0.05) were identified using Sidak-adjusted comparisons from a log-linear regression model with the negative binomial family, presented as compact letter displays. Lowercase letters indicate differences between fungal isolates on PDA, while uppercase letters represent differences on OA. Means sharing the same letter within each medium are not significantly different.

In our study, the feeding and reproductive abilities of A. besseyi s. l. on fungi used or potentially useful for disease biocontrol were evident, especially on PC17 and PC10. Under the tested conditions, those fungi supported a nematode population increase of 167 to 216 times, in only 20 days (Fig. 1). At an intermediary rate, the isolates C101 and C05 can also support a considerable increase in the population of this nematode, raising the population by 28 to 35 times (Fig. 1). Thus, although our results demonstrate that A. besseyi s. l. was able to reproduce on all evaluated fungal isolates under laboratory conditions, including on strains considered potential biological control agents, caution is needed when extrapolating these findings to field conditions. The controlled and simplified environment of Petri dish assays does not fully represent the complexity of the soil and rhizosphere, in which multiple biotic and abiotic factors could limit nematode development and fungal interactions. Even the isolates C35 and T10, which supported the lowest reproductive rates in vitro, may not behave similarly in the field, as environmental conditions, microbial competition, plant root exudates, and other stressors could alter the dynamics of nematode-fungus interactions. Therefore, while the findings raise an important hypothesis about the potential risk of mycophagy by A. besseyi s. l. on beneficial fungi, further studies under greenhouse and field conditions are needed to validate these results and to assess the real impact on GSFRS incidence in infested areas.

The nutritional status of the biocontrol fungus can affect the ability of A. besseyi s. l. to feed and reproduce in its colonies, as observed in the case of isolates PC17, C05, and C101 (Fig. 1, Table 2). However, it is still unclear whether the fungal hostability to the nematode would be favored under restrictive or enriched nutritional conditions. OA medium generally promoted greater pigmentation, particularly in Clonostachys spp. All nematode inoculations were performed after complete fungal colonization, ensuring a food source was present (Honório 2025). However, variations in mycelial structure or density, influenced by the culture medium, may have impacted the nematode’s feeding and reproduction.

The hypothesis that would explain this variation in the hostability of the biocontrol fungus to A. besseyi s. l., influenced by the culture medium, would be due to changes in the palatability of these fungi or the effect of antibiosis on the nematode (Iqbal et al. 2018). This control is likely due to antibiosis, as these fungi are known to produce nematicidal toxins. Therefore, it is necessary to consider the possible antibiosis effect of biocontrol fungal isolates on A. besseyi s. l. or another type of antagonism, such as induced systemic resistance, that could happen in field conditions but could not be observed in vitro.

We presented evidence of the possible contribution of fungal BCAs such as P. chlamydosporia, P. boninensis, and T. afroharzianum to the survival and multiplication of A. besseyi s. l. This could increase the severity of GSFRS in cases in which these species are employed in the management of soil-borne diseases. Currently, information on how mycophagy affects the efficiency of fungi used in biological control is scarce. In this context, reproduction of A. besseyi s. l. on BCAs should be considered, as it may interfere with fungi used as biocontrol agents against various pathogens, including A. besseyi s. l. (Santamaria et al. 2023).

CONCLUSION

The results showed that A. besseyi s. l. reproduced efficiently on P. chlamydosporia (PC10) and P. boninensis (PC17), indicating that these fungi are suitable hosts in the tested conditions. Lower reproduction rates were observed on C. farinosa (C35) and T. afroharzianum (T10). In contrast, C. chloroleuca (C17) and C. rosea (C133) did not support nematode reproduction, suggesting potential as safe biological control agents. These findings highlight the importance of evaluating the host status of fungal isolates before field application.

ACKNOWLEDGMENTS

We thank Professor Emerson Medeiros Del Ponte for his support with the statistical analysis and to the PIBIC CNPq - UFV scholarship awarded to the first author of the manuscript.

• How to cite:
  Matos, E. A. N., Honório, A. P., Silva, M. F., Abreu, L. M., Freitas, L. G. and Buonicontro, D. S. (2025). Reproduction of Aphelenchoides besseyi sensu lato on fungal biological control agents. Bragantia, 84, e20250057. https://doi.org/10.1590/1678-4499.20250057
• FUNDING
  Coordenação de Aperfeiçoamento de Pessoal de Nível Superior
  Finance Code 001
  Conselho Nacional de Desenvolvimento Científico e Tecnológico
  Grant No.: 139960/2022-8 and 141347/2021-0

DATA AVAILABILITY STATEMENT

The image plates documenting the biocontrol fungal agents and nematodes are available in the OSF repository at https://osf.io/z43b5. All other data generated or analyzed during this study are contained within the article.

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Edited by

• Section Editor:
  Luis Garrigós Leite https://orcid.org/0000-0001-7947-5698

Publication Dates

• Publication in this collection
  17 Oct 2025
• Date of issue
  2025

History

• Received
  12 Mar 2024
• Accepted
  28 July 2025