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Tecnológico Superior Corporativo Edwards Deming. Quito - Ecuador Frequency April–June Vol.
1, No. 29, 2026 pp. 63–76 http://centrosuragraria.com/index.php/revista Dates of receipt Received: January 22, 2026 Approved: March 25, 2026 Corresponding author nerazo@espoch.edu.ec Creative Commons License Creative Commons License, Attribution-NonCommercial-ShareAlike
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Evaluación de la
Capacidad Antagonista de Bacillus subtilis frente a Hongos Agrícolas
Fitopatógenos y Benéficos
Norma Soledad Erazo Sandoval1
Gabriela Andrea Rosero Obando2
Pablo Israel Álvarez Romero3
Carlos Rolando Rosero Erazo4
Doctora en Ciencias Ambientales. Facultad de
Recursos Naturales, Grupo de Investigación y Desarrollo para el
Ambiente y el Cambio Climático, Escuela Superior Politécnica de Chimborazo https://orcid.org/ 0000-0001-8495-1468 nerazo@espoch.edu.ec Ingeniera en Biotecnología ambiental Grupo de Investigación y Desarrollo para el
Ambiente y el Cambio Climático, Doctor scientiae en fitopatologia Ingeniero. Magíster en Cambio climático
Escuela Superior Politécnica de Chimborazo
https://orcid.org/0000-0002-2855-6292
gabriela.rosero@espoch.edu.ec
Facultad de Recursos Naturales,
Grupo de Estudios Fitoentomológicos,
Escuela Superior Politécnica de Chimborazo
https://orcid.org/0000-0003-0743-5210
pabloi.alvarez@espoch.edu.ec
Facultad de Ciencias – Facultad de Recursos Naturales,
Escuela Superior Politécnica de Chimborazo (SEDE-ORELLANA),
Riobamba – Ecuador
https://orcid.org/0000-0003-2691-5578
carlos.rosero@espoch.edu.ec
Keywords: Bacillus subtilis, Trichoderma harzianum, secondary metabolites, nematophagous
fungi, entomopathogenic fungi, phytopathogenic fungi.
Resumen: La agricultura convencional ha generado un
desgaste en los ecosistemas y contribuye al aumento del efecto invernadero, al
desarrollo de resistencia genética y afectaciones a la salud humana. El uso de
microorganismos en el control biológico favorece una agricultura sostenible. El
objetivo de esta investigación fue evaluar la capacidad antagonista de Bacillus
subtillis F. Cohn in vitro, frente a cepas de hongos fitopatógenos y benéficos.
Se usaron los métodos de enfrentamiento dual y de compuestos volátiles en dos
etapas de crecimiento. Se realizó la medición del crecimiento micelial y se
determinó el porcentaje de inhibición y la tasa de crecimiento. Se contabilizó
las esporas para determinar el porcentaje de inhibición en la esporulación de
los hongos. El método dual mostró que el mayor porcentaje de inhibición ocurrió
en Moniliophthora roreri con un 61,85 % vertical y 27,43 % horizontal, mientras
que, en Aspergillus spp., y Trichoderma. harzianum el porcentaje fue menor al
25 %. En el método de compuestos volátiles Arthrobotrys conoides y Beauveria
bassiana presentaron un porcentaje de inhibición mayor al 50%, por su parte,
Penicillium spp. y Metarhizium anisopliae presentaron un porcentaje menor al 25
%. B. subtilis provocó antagonismo frente a la mayoría de los hongos
fitopatógenos, mientras que, causó un efecto mínimo en el desarrollo de
Aspergillus spp., y T. harzianum y, un efecto medio-alto en el desarrollo de
hongos nematófagos. Además, las colonias de B. subtilis provocaron cambios en
las características macroscópicas y microscópicas de los hongos estudiados,
debido al amplio número de compuestos que pueden producir.
Palabras clave: Bacillus subtilis, Trichoderma harzianum,
metabolitos secundários, hongos nematófagos, hongos entomopatógenos, hongos
fitopatógenos.
Introduction
Conventional agricultural practices
are becoming increasingly unsustainable, negatively impacting environmental,
economic, and social aspects in the areas where they are carried out. The
excessive use of pesticides and chemical fertilizers, along with inappropriate
farming practices and the expansion of agriculture, has damaged ecosystems and
contributed to the greenhouse effect (Baquero et al., 2007) .
It is important to recognize that
agriculture is a crucial activity for the world’s economic, social, and
environmental development, as it provides approximately 80% of the food
consumed. However, pests and diseases—including bacteria, nematodes, viruses,
insects, and especially fungi—affect approximately 20% to 30% of annual
agricultural production (FAO, IFAD, PAHO, WFP, 2020) .
Using microorganisms for the
biological control of crop-affecting pathogens is an ecological and effective
option that promotes sustainable agriculture, as it reduces the problems
associated with the use of pesticides and chemicals (Ruiz-Sánchez et al., 2016) . Research has focused on identifying native
microorganisms that function under the specific environmental conditions of
each region and can be used to restore soil microbiota interactions. In this
way, they could be used as biofertilizers and/or biocontrol agents, mitigating
the impact of agrochemicals and reducing production costs (Orberá et al., 2014) .
Certain microorganisms produce
nutrients that serve as food for other microorganisms (Albuquerque, Elizabeth Albuquerque, 2009) . Bacteria in the body’s normal flora control
the growth of harmful microorganisms. Among the microorganisms used for this
purpose are bacteria of the genus Bacillus, which have high potential due to
their ability to exert antagonistic activity through competition, the
production of antibiotics, and the production of lytic enzymes such as
chitinases (Tejera et al., 2012) .
The development of bioproducts for
disease control focuses on the ecological preservation of the interaction
between the plant and the microorganism, strategies for applying inoculants,
the isolation of new strains, and the identification of innovative mechanisms
of action (Cobo, 2017) . Bacillus species have great potential as
antagonists due to the large number of biocidal substances they produce, which
are capable of controlling plant pathogens (Villarreal et al., 2017) .
This research is of great importance
in the field of sustainable agriculture, as it proposes the use of
microorganisms as a biocontrol strategy against pathogens that affect crops.
The identification and application of native microorganisms allow for the
restoration of soil microbiota interactions, improved nutrient availability,
and reduced production costs, thereby contributing to the mitigation of the
adverse effects of agrochemicals.
This research offers the potential
to identify effective combinations of microorganisms in sustainable
agriculture, opening up a promising field. These organisms, primarily
beneficial bacteria and fungi, perform key functions in agricultural ecosystems
and can improve agricultural productivity while reducing the need for chemical
pesticides and fertilizers, thereby promoting sustainability in agricultural
and industrial production.
Methodology
The research was conducted at the
Biological Sciences Laboratory, located in the Faculty of Natural Resources at
ESPOCH, in the Lizarzaburu Parish, Riobamba Canton, Chimborazo Province. The
environmental conditions inside the laboratory average a temperature of 23 °C
and relative humidity of 67%.
Microorganisms Used
The Bacillus subtilis F. Cohn
isolate and the seven strains of beneficial fungi were obtained from the fungal
culture collection of the Biological Sciences Laboratory, and the seven strains
of phytopathogenic fungi were obtained from the fungal culture collection of
the Plant Pathology Laboratory at the Faculty of Natural Resources of ESPOCH.
Calculation of the percentage of
mycelial growth inhibition
To determine the effect of Bacillus
subtilis F. Cohn on fungal growth, the “ method was
used, which involves measuring the horizontal and vertical diameters in the
presence of the bacterium, while Petri dishes containing only the fungi were
used as controls. Data were collected every 24 hours. Using the measurements
obtained, the percentage of mycelial growth inhibition was calculated using
Equation 1.
To determine the degree of
inhibition, the scale described in Table 1 was used.
Table1 . Scale of
percentage of inhibition
|
Low |
0%–25% |
|
Medium |
25%–50% |
|
High |
> 50% |
Calculation of the growth rate
Mycelium growth was recorded in the
Petri dish (Corrales Ramírez et al., 2012) , using the following equation:
Calculation of the percentage of
sporulation inhibition
The percentage of sporulation
inhibition was calculated once the fungal control reached its maximum growth (Rodrigues et al., 2010; Velasquez Gurrola,
2005) .
Morphological characteristics
Among the macroscopic characteristics, the color of the colonies was evaluated( Pantone® USA | Pantone Color Systems - Introduction, n.d.) , as well as the characteristics of the mycelium, as described by various authors depending on the fungus. To determine the growth rate, the scale described in Table 2 was used (Hernandez Romero, 2018) .
Table2 . Fungal growth scale
|
Fast |
Between one and two weeks |
|
Moderate |
Between two and three weeks |
|
Slow |
Between three and four weeks |
Experimental specifications
The treatments were carried out
using phytopathogenic fungi and beneficial fungi, as shown in Table 3.
Table3 . Treatments
|
Code |
Description |
Code |
Description |
|
T1 |
Bacillus subtilis F Cohn +
Fusarium oxysporum |
PG01 |
Bacillus subtilis F Cohn + A. oligospora |
|
T2 |
Bacillus
subtilis F Cohn + Aspergillus sp. |
CH02 |
Bacillus subtilis F Cohn + A. oligospora |
|
T3 |
Bacillus subtilis F Cohn +
Colletotrichum spp. |
CH01 |
Bacillus subtilis F Cohn + A. musiformis |
|
T4 |
Bacillus
subtilis F Cohn + Penicillium spp. |
PG02 |
Bacillus subtilis F Cohn + A. conoides |
|
T5 |
Bacillus subtilis F Cohn +
Neopestalotiopsis sp. |
A21 |
Bacillus subtilis F Cohn + B. bassiana |
|
T6 |
Bacillus
subtilis F Cohn + Alternaria grandis |
A13 |
Bacillus subtilis F Cohn + M. anisopliae |
|
T7 |
Bacillus subtilis F Cohn +
Moniliophthora roreri |
Th01 |
Bacillus subtilis F Cohn + T. harzianum |
|
T8 |
Controls |
T8 |
Controls |
Results
Dual
method
In the dual method, the variable
indicating the percentage of mycelial growth inhibition in beneficial fungi
shows that A. musiformis exhibits the highest
percentage of inhibition, with 49.95% vertical inhibition and 47.7% horizontal
inhibition, with a growth rate of 6.45 mm/day. The growth rate of the genus Arthrobotrys sp. is 5 mm/day (Castillo Ávila & Medina Medina, 2014) , while T. harzianum
showed 0% vertical inhibition and 15.07% horizontal inhibition, with a growth
rate of 11.8 mm/day.
Meanwhile, among the phytopathogenic
fungi, Moniliophthora roreri
achieved the highest percentage with 61.85% vertical inhibition and 27.43%
horizontal inhibition, with vertical reductions in its growth rate of 4.2
mm/day (Suárez Contreras & Rangel Riaño, 2013) . Bacillus subtilis, through its antibiosis
action, is effective in inhibiting phytopathogenic fungi (Caulier et al., 2019) , and the variety of metabolites generated by
Bacillus subtilis, such as antibiotics and lipopeptides (Villarreal-Delgado et al., 2018) , may have antifungal effects. The other fungi
obtained percentages above 20% (vertical) and below 15% (horizontal).
Figure1 . Percentage of inhibition of beneficial and
phytopathogenic fungi against B. subtilis using the dual method.
Volatile Compounds Method
In the volatile compounds method (0
hours), the treatment with the highest percentage of inhibition of mycelial
growth in beneficial fungi was A. conoides, with
36.6% vertical inhibition and 59.31% horizontal inhibition, with a growth rate
of 5.45 mm/day. The treatment with the lowest inhibition was treatment A13
corresponding to M. anisopliae, with 0% vertical inhibition and 22.3%
horizontal inhibition, exhibiting a growth rate of 5.85 mm/day; the growth rate
for M. anisopliae is an average of 10 to 20 mm/day (Padilla-Melo et al., 2000) .
Regarding phytopathogenic fungi, Moniliophthora roreri achieved
the highest percentage of inhibition in both vertical and horizontal
directions, exceeding 40%, with a reduction in growth rate of 3.33 to 3.37
mm/day, which is lower than in previous studies ( with inhibition rates exceeding 62.5%.
Followed by Neopestalotiopsis sp., which showed an
inhibition percentage of 20–35%, with a reduction in its growth rate from 4.10
to 4.30 mm/day, due to the emission of volatile compounds by Bacillus spp.
strains, (Tahir et al., 2017) . Some of the VOCs that have demonstrated
microbial activity include benzothiazole, benzaldehyde, phenylacetaldehyde, and
2,3-butanediol (Pedraza et al., n.d.) . The other fungi obtained inhibition
percentages below 25%.
Figure 2. Percentage of inhibition of
beneficial and phytopathogenic fungi against B. subtilis using the volatile
compound method at 0 hours post-exposure.
Volatile compound method (2nd growth
stage)
Using this method on beneficial
fungi, B. bassiana showed the highest inhibition, with 37.78% vertical
inhibition and 40.07% horizontal inhibition, exhibiting a growth rate of 2.35
mm/day, similar to the growth rate of 60 mm/day for B.
bassiana reported in previous studies ( . The treatment with the lowest
inhibition percentage was T. harzianum, with 0.99%
vertical inhibition and 1.97% horizontal inhibition, exhibiting a growth rate
of 1.9 mm/day.
Against the phytopathogenic fungi, Colletrotichum spp., it had the highest inhibition percentage at 31% and a reduction in growth rate of 0.8 mm/day. However, the other treatments obtained lower values in both vertical and horizontal directions, with inhibition percentages below 22% and reductions in growth rate of up to 1.8 mm/day. Once the fungus is fully developed, it is more difficult to control it using microorganisms such as Bacillus subtilis and . It is important to note that Aspergillus spp. showed no inhibition by any method due to its ability to produce mycotoxins, which leads to greater resistance to control by microorganisms (Martínez Padrón et al., 2013) .
Figure 3. Percentage of inhibition of
beneficial and phytopathogenic fungi against B. subtilis using the volatile
compounds method 96 hours after exposure.
Percentage of inhibition of fungal
sporulation
Bacillus subtilis possesses
antimicrobial activity capable of combating spores from various types of
microorganisms, such as bacteria and fungi (Ruiz-Sánchez et al., 2016) . Likewise, it has been demonstrated to
possess an exceptional ability to inhibit the growth and sporulation of various
phytopathogenic fungi, both in vitro and in vivo.
Figure 4 shows the percentage of
spore formation inhibition in beneficial fungi; the highest inhibition was
found for CH02, corresponding to A. oligospora, with
a percentage of 89.32%, and PG02, corresponding to A. conoides,
with a percentage of 87.42%. On the other hand, the lowest inhibition
percentages were observed for M. anisopliae, at 12.78%, and T. harzianum, at 7.34%.
Figure 4. Percentage of spore formation
inhibition by beneficial and phytopathogenic fungi.
Regarding phytopathogenic fungi,
Alternaria grandis showed the highest percentage at 86.14%, due to the
difficulty of sporulation for this fungus (Woudenberg et al., 2014) ; followed by Moniliophthora
roreri, which obtained 63.05%, consistent with
results showing sporulation inhibition values exceeding 60.85% (Vera Loor et al., 2021) .
Furthermore, the production of
hydrolytic enzymes degrades the cytoplasmic membrane of filamentous fungi,
thereby reducing fungal sporulation. Similarly, research indicates that against
M. roreri and Fusarium oxysporum
f. sp. Lycopersici, the presence of Bacillus subtilis
was observed to inhibit spores with an effectiveness of over 90% (Guato-Molina et al., 2019) .
In the case of phytopathogenic
microorganisms, changes in morphological characteristics occurred in most
treatments with Bacillus subtilis, with the exception of Aspergillus spp. In
the other treatments, the changes were noticeable, whether in color, growth
pattern, or similarly, in the mycelium; this is due to the production of lytic
enzymes that aid in the degradation of the main polysaccharides of the fungal
cell wall through the hydrolysis of their glycosidic bonds (Caulier et al., 2019) . Similarly, the production of lipopeptides
causes the formation of pores and, consequently, an osmotic imbalance in the
cytoplasmic membrane (Villarreal-Delgado et al., 2018) .
Iturine and fengicin
are compounds with strong biocontrol properties, as they are capable of
inhibiting the action of a wide variety of plant pathogens (Ragazzo-Sánchez et al., 2011) . On the other hand, surfactin
alone is not capable of inhibiting fungal growth, but it has been shown to have
a synergistic effect with the antifungal activity of iturine
A.
It has been demonstrated that
Trichoderma spp. is capable of synthesizing secondary metabolites involved in
the production of volatile compounds with antimicrobial properties (Hernández-Melchor et al., 2019) . Among these compounds, tetracyclic
diterpenes such as harziandione, sesquiterpenes such
as trichothecenes, trichodermin, and harzianum A, and
the triterpene viridine stand out.
Conclusions
Bacillus subtilis F. Cohn causes a decrease in growth in the fungi A. oligospora (CH02), A. musiformis
(CH01), A. conoides (PG02), and B.
bassiana (A21), with mycelial growth inhibition exceeding 50%, affecting
the growth rate. Moniliophthora roreri had the highest inhibition percentage, with 61.85%
vertical and 27.43% horizontal inhibition in the dual method, and over 40% in
volatile tests. The Aspergillus spp.
strain, A. oligospora (PG01), M.
anisopliae (A13), and T. harzianum (Th01) showed the lowest inhibition
percentage in all methods used.
Alternaria grandis, Moniliophthora roreri, Penicillium spp., A. oligospora
(PG01), A. oligospora (CH02), A. musiformis
(CH01), A. conoides (PG02), and B.
bassiana (A21) had the
highest spore formation inhibition percentages, exceeding 50%. In
contrast, the fungi Neopestalotiopsis sp., Aspergillus spp., M.
anisopliae (A13), and T. harzianum (Th01)
showed percentages below 25%.
Through this study, the
morphological characteristics of each of the phytopathogenic fungi were
determined, concluding that Bacillus
subtilis F. Cohn causes changes in both macroscopic and microscopic
characteristics due to the wide range of substances and compounds it is capable
of producing.
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