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Moisés Gilberto Yáñez-Juárez, Carlos Alfonso López-Orona, Felipe Ayala-Tafoya*, Leopoldo Partida-
Ruvalcaba, Teresa de Jesús Velázquez-Alcaraz y Raymundo Medina-López, Facultad de Agronomía, Uni-
versidad Autónoma de Sinaloa. Carretera Culiacán-Eldorado Km 17.5, Aparatado Postal 25, CP. 80000, Culia-
cán, Sinaloa, México. *Autor para correspondencia: tafoya@uas.edu.mx.
Yáñez-Juárez MG, López-Orona CA, Ayala-Tafoya F, Abstract. Phosphites are compounds derived
Partida-Ruvalcaba L, Velázquez-Alcaraz TJ, Medina- from phosphorous acid used as an alternative for
López R. 2017. Phosphites as alternative for the mana-
gement of phytopathological problems. Revista Mexica- the control of phytoparasitic organisms and their
na de Fitopatología 36(1): 79-94. effectiveness has been tested against protozoa,
DOI: 10.18781/R.MEX.FIT.1710-7 oomycetes, fungi, bacteria and nematodes;
however, compared to conventional synthesized
Primera publicación DOI: 01 de Enero, 2018.
First DOI publication: January 01, 2018. fungicides, phosphites are generally less effective
at reducing damage by phytopathogens. The
phosphite ion is easily transported in the plants
Resumen. Los fosfitos son compuestos deriva- via xylem and phloem, so it has been used in
dos del ácido fosforoso empleados como alternati- foliar application, drench of plant root and neck,
va para el control de organismos fitoparásitos y su injection trunk, through drip irrigation mixed in the
eficacia se ha probado contra protozoarios, oomy- nutrient solution in hydroponics, seed treatment,
cetes, hongos, bacterias y nematodos; sin embargo, aerial application in low volume, or as treatment
en comparación con los fungicidas convencionales in immersion of seeds and fruits. The mechanisms
sintetizados, generalmente son menos eficaces para of action involved in the prophylactic effects of
disminuir el daño por fitopatógenos. El ion fosfito phosphites are diverse and include the stimulation
es fácilmente transportado en las plantas vía xilema of biochemical and structural defense mechanisms
y floema, por lo que se ha utilizado en aplicación in plants and direct action that restricts the growth,
foliar, baño a la raíz y cuello de la planta, inyección development and reproduction of phytopathogenic
al tronco, a través de riego por goteo mezclado en organisms.
la solución nutritiva en hidroponía, tratamiento a
semilla, aplicación aérea en bajo volumen o como Key words: potassium phosphite, biostimulant,
tratamiento en inmersión de semillas y frutos. Los control of phytopathogens.
mecanismos de acción involucrados en los efectos The irrational use of synthetic pesticides in
profilácticos de los fosfitos son diversos e incluyen agriculture increases problems of environmental
la estimulación de los mecanismos de defensa bio- pollution and public health, and decreases
química y estructural en las plantas, además de la biodiversity in the agroecosystems as well as
acción directa que restringe el crecimiento, desa- the development of resistant-phytopathogenic
rrollo y reproducción de los organismos fitopató- organisms. Furthermore, the current agricultural
genos. marketing requires products that are safe for
consumers and created using low environmental
Palabras clave: fosfito de potasio, bioestimulante, impact processes. Thus, disease control based on
control de fitopatógenos. the use of inorganic salts, which aside from being
effective in managing crop diseases have minimum
adverse consequences, becomes relevant. To this
El uso irracional en la agricultura de pesticidas regard, Deliopoulos et al. (2010) reported that 34
sintetizados, incrementa los problemas de contami- different salts have been used for such purpose, and
nación ambiental, salud pública, disminución de la suggested that phosphite salts stand out because of
biodiversidad en los agroecosistemas y desarrollo their use frequency and control effectiveness.
de organismos fitopatógenos con resistencia, ade- Phosphites are oxyanions derived from
más, la comercialización agrícola actual demanda phosphorous acid (H3PO3-) regularly combined
productos inocuos para los consumidores, prove- with non-metal cations, such as potassium, sodium,
nientes de procesos con bajo impacto ambiental. calcium or ammonia. The terms “phosphite” and
Así, adquiere relevancia el control de enfermeda- “phosphonate” are commonly used in the literature
des basado en el empleo de sales inorgánicas, que to refer to salts derived from phosphorous acid, the
además de ser eficaces para el manejo de enfer- same as “hydrogen phosphates”, “orthophosphites”,
medades en los cultivos muestren mínimas conse- “phosphonic acid compounds” and “phosphorous
cuencias adversas. Al respecto, Deliopoulos et al. acid compounds” (Deliopoulos et al., 2010).
(2010) reportan que se han utilizado 34 diferentes The chemical difference between phosphate
sales con esa finalidad, además indican que las sa- (H2PO4-) and phosphite (H2PO3-) is an oxygen atom
les de fosfito destacan por su frecuencia de utiliza- that is replaced by another of hydrogen (Figure
ción y eficacia en el control. 1). When oxygen is replaced, there is a profound
Los fosfitos son oxianiones derivados del ácido difference in the way phosphates and phosphites
fosforoso (H3PO3-), que regularmente se combinan behave in living organisms (McDonald et al.,
con cationes no metales como potasio, sodio, cal- 2001; Achary et al., 2017). Due to their structural
cio o amonio. Los términos “fosfito” y “fosfana- similarity, phosphites are thought to be akin to
to” son comúnmente utilizados en la literatura para phosphates. However, the use of phosphites as
referirse a las sales derivadas del ácido fosforoso, fungicides and biostimulants to control plant
al igual que “hidrogenofosfanatos”, “ortofosfitos”, pathogens is widely accepted, but their use as a
“compuestos del ácido fosfónico” o “compuestos phosphorus source in plant nutrition is currently
del ácido fosforoso” (Deliopoulos et al., 2010). debated (Borza et al., 2014; Gómez-Merino
La diferencia química entre fosfato (H2PO4-) y and Trejo-Téllez, 2015; Alexandersson et al.,
fosfito (H2PO3-) es un átomo de oxígeno el cual es 2016; Manna et al., 2016). Such difference gives
sustituido por otro de hidrógeno (Figura 1). La sus- phosphites a greater mobility in soil and plant
titución del oxígeno da lugar a profundas diferen- tissues, as well as a greater capacity to penetrate
cias en la manera en que los fosfatos y fosfitos se through leaves, stems and roots (Tkaczyk et al.,
comportan en los organismos vivos (McDonald et 2016).
al., 2001; Achary et al., 2017). Debido a su simili- The use of phosphites in agriculture has been
tud estructural, los fosfitos son considerados como studied mainly because of their action to control
análogos de los fosfatos, sin embargo, el uso de los phytoparasite organisms or as a nutrition source for
fosfitos como fungicidas y bioestimulantes para el cultivated plants (Gómez-Merino and Trejo-Téllez,
control de patógenos de plantas es ampliamente 2015; Alexandersson et al., 2016). The latter may
aceptado, pero su utilización como fuente de fós- occur only if phosphites are applied to soil and come
foro en la nutrición de plantas es actualmente de- into contact with bacteria that have the capacity
batida (Borza et al., 2014; Gómez-Merino y Trejo- to oxidize them to phosphates (McDonald et al.,
Téllez, 2015; Alexandersson et al., 2016; Manna et 2001; Manna et al., 2016). However, since this is a
al., 2016). Además, esa diferencia da a los fosfitos very slow process that can take up to four months,
una mayor movilidad en el suelo y en los tejidos it becomes impractical in agriculture (McDonald et
de las plantas, así como una mayor capacidad para al., 2001; Lovatt and Mikkelsen, 2006).
penetrar a través de hojas, tallos y raíces (Tkaczyk In early 1930, researchers concluded that
et al., 2016). phosphites were not an efficient nutrition source
El uso de los fosfitos en la agricultura, se ha in- for plants. However, 40 years later, phosphites
vestigado principalmente por su acción en el con- returned to the agrochemicals market as an
trol de organismos fitoparásitos o como fuente de alternative for controlling plant diseases, when the
nutrición en plantas cultivadas (Gómez-Merino y French company Rhône-Poulenc offered the active
Trejo-Téllez, 2015; Alexandersson et al., 2016). ingredient fosetyl-aluminum to control mildews and
Esto último puede ocurrir únicamente si los fosfitos diseases caused the Phytophthora genus (Tkaczyk
se aplican al suelo y entran en contacto con bacte- et al., 2016; Achary et al., 2017). Phosphites, as an
rias que tienen la capacidad de oxidarlos a fosfatos alternative for controlling phytoparasite organisms,
(McDonald et al., 2001; Manna et al., 2016). No have been extensively studied and proven to be
obstante, este proceso es muy lento, y puede tomar effective against protozoa, oomycetes, fungi,
hasta cuatro meses para completarse, lo que resul- bacteria and phytoparasite nematodes (Table 1), as
ta impráctico para la agricultura (McDonald et al., well as biostimulators (Gómez-Merino and Trejo-
2001; Lovatt y Mikkelsen, 2006). Téllez, 2015) that reduce damages caused by weeds
(Manna et al., 2016; Achary et al., 2017) and UV-B
O O radiation (Oyarburo et al., 2015).
P P
APPLICATION METHODS
HO O- H O-
OH OH
The phosphite ion is easily transported in plants
A: (H2PO4-) B: (H2PO3-)
via xylem and phloem (McDonald et al., 2001;
Figura 1. Estructura del grupo fosfato (A) y fosfito (B). Tkaczyk et al., 2016), so it has been used in foliar
Figure 1. Structure of the phosphate (A) and phosphite (B) applications (Rebollar-Alviter et al., 2010; Silva et
group.
Fue a principios de 1930 cuando se concluyó al., 2011; Pagani et al., 2014; Yáñez et al., 2014;
que los fosfitos no eran fuente eficiente de fosforo Liljeroth et al., 2016; Borza et al., 2017), drench of
para la nutrición de las plantas y 40 años después, plants root and neck (Oka et al., 2007; Akinsanmi
regresó al mercado de los agroquímicos como al- and Dreth, 2013), trunk injection (Bock et al.,
ternativa para el manejo de enfermedades, cuando 2012; Akinsanmi and Drenth, 2013: Aćimović
la compañía Francesa Rhône-Poulenc ofreció el et al., 2015; Aćimović et al., 2016) through
ingrediente activo fosetil-aluminio para el control drip irrigation mixed in a nutrient solution in
de mildius y enfermedades causadas por el género hydroponics (Förster et al., 1998), seed treatments
Phytophthora (Tkaczyk et al., 2016; Achary et al., (Abbasi and Lazarovits 2006; Lobato et al., 2008),
2017). Los fosfitos, como alternativa para el con- low-volume aerial applications (Hardy et al., 2001)
trol de organismos fitoparásitos, han sido estudia- or as a treatment in seed and fruits immersion
dos ampliamente y su eficacia se ha probado contra (Anderson et al., 2012; Cerioni et al., 2013, Borin
protozoarios, oomycetes, hongos, bacterias y ne- et al., 2017).
matodos fitoparásitos (Cuadro 1); adicionalmente,
se ha comprobado su eficacia como bioestimulador EFFECTIVENESS
(Gómez-Merino y Trejo-Téllez, 2015), para dis-
minuir los daños por malezas (Manna et al., 2016; The levels of effectiveness of phosphites to
Achary et al., 2017) y radiación UV-B (Oyarburo control phytoparasite organisms vary depending
et al., 2015). on the ion bonded to phosphite, application
method, pathogen organism and host plant (Table
MÉTODOS DE APLICACIÓN 2; Figure 2). For example, mandarin orange fruits
immersed in solutions containing calcium and
El ion fosfito es fácilmente transportado en potassium phosphites contributed to reduce 50%
las plantas vía xilema y floema (McDonald et al., of fruits infected with citrus green mold caused by
2001; Tkaczyk et al., 2016), por lo que se ha uti- Penicillium digitatum (Cerioni et al., 2013); also,
lizado en aplicación foliar (Rebollar-Alviter et soja plants treated with potassium phosphite showed
al., 2010; Silva et al., 2011; Pagani et al., 2014; up to 50% less damage by Peronospora manshurica
Yáñez et al., 2014; Liljeroth et al., 2016; Borza than non-treated plants (Silva et al., 2011). Abbasi
et al., 2017), baño a la raíz y cuello de la planta and Lazarovits (2006) reported 80% less cucumber
(Oka et al., 2007; Akinsanmi y Dreth, 2013), in- plants infected by Pythium spp. when seeds were
yección al tronco (Bock et al., 2012; Akinsanmi y treated by immersion in solutions containing
Drenth, 2013: Aćimović et al., 2015; Aćimović et copper phosphite. Ogoshi et al. (2013) obtained a
al., 2016), a través de riego por goteo mezclado en 62.5% decrease in the severity of Colletotrichum
la solución nutritiva en hidroponía (Förster et al., gloeosporioides in coffee plants treated with
1998), tratamiento a semilla (Abbasi y Lazarovits potassium phosphite. In another experiment, 93%
2006; Lobato et al., 2008), aplicación aérea en bajo of papaya plants treated with potassium phosphite
volumen (Hardy et al., 2001) o como tratamiento survived the attack of Phytophthora palmivora, but
en inmersión de semillas y frutos (Anderson et al., only 24% of non-treated plants survived (Vawdrey
2012; Cerioni et al., 2013, Borin et al., 2017). and Westerhuis, 2007).
Protozoarios
Plasmodiophora brassicae Brassica rapa Potasio Kammerich et al., 2014
Oomycetes
Peronospora destructor Allium cepa Potasio Monsalve et al., 2012
P. manshurica Glicine max Potasio Silva et al., 2011
P. parasitica Brassica oleracea Potasio Becót et al., 2000
Phytophthora cinnamomi Macadamia spp. Potasio Akinsanmi y Dreth, 2013
P. cinnamomi Ananas comosus Potasio Anderson et al., 2012
Phytophthora infestans Solanum tuberosum Aluminio Kromann et al., 2012
P. infestans S. tuberosum Calcio Lobato et al., 2008
P. infestans S. tuberosum Potasio Borza et al., 2017
P. infestans S. tuberosum Potasio Kromann et al., 2012
P. infestans S. tuberosum Potasio Liljeroth et al., 2016
P. nicotianae Nicotiana tabacum Potasio Smillie et al., 1989
P. palmivora Carica papaya Potasio Smillie et al., 1989
Plasmopara viticola Vitis vinifera Potasio Pinto et al., 2012
Pseudoperonospora cubensis Cucumis melo Potasio Méndez et al., 2010
Pythium aphanidermatum C. sativus Cobre Abbasi y Lazarovits, 2006
P. irregulare C. sativus Cobre Abbasi y Lazarovits, 2006
P. ultimum C. sativus Cobre Abbasi y Lazarovits, 2006
P. ultimum C. sativus Potasio Mofidnakhaei et al., 2016
Hongos
Alternaria alternata Malus domestica Potasio Reuveni et al., 2003
Cercospora coffeicola Coffea arabica Potasio Costa et al., 2014
Cochliobolus miyabeanus Oryza sativa Potasio Nascimento et al., 2016
Colletotrichum gloeosporioides C. arabica Potasio Ogoshi et al., 2013
C. gloeosporioides Fragaria vesca Potasio MacKenzie et al., 2009
C. gloeosporioides Malus domestica Potasio Araujo et al., 2010
Fusarium solani Solanum tuberosum Calcio Lobato et al., 2008
F. solani S. tuberosum Potasio Lobato et al., 2008
Fusicladium effusum Carya illinoinensis Potasio Bock et al., 2012
Hemileia vastatrix Coffea arabica Potasio Costa et al., 2014
Oidium sp. Cucumis sativus Potasio Yáñez et al., 2012
Oidium sp. C. sativus Potasio Yáñez et al., 2014
Penicillium digitatum Citrus limon Potasio Cerioni et al., 2013
P. italicum C. limon Potasio Cerioni et al., 2013
P. expansum Malus domestica Potasio Amiri y Bompeix, 2011
P. expansum Malus domestica Potasio Lai et al., 2017
Rhizoctonia solani Solanum tuberosum Calcio Lobato et al., 2008
R. solani S. tuberosum Potasio Lobato et al., 2008
Venturia inaequalis S. tuberosum Potasio Percival et al., 2009
V. pirina Pyrus communis Potasio Percival et al., 2009
Bacterias
Erwinia carotovora Solanum tuberosum Potasio Lobato et al., 2011
E. amylovora Pyrus malus Potasio Aćimović et al., 2015
Pseudomonas syringae pv. actinidiae Actinidia deliciosa Aluminio Monchiero et al., 2015
Nematodos
Helicotylenchus spp. Musa paradisiaca Potasio Quintero-Vargas y Castaño-Zapata, 2012
Heterodera avenae Avena sativa Potasio Oka et al., 2007
Meloidogyne marylandi Triticum aestivum Potasio Oka et al., 2007
Meloidogyne spp. M. paradisiaca Potasio Quintero-Vargas y Castaño-Zapata, 2012
Pratylenchus bracyurus Glycine max Potasio Dias-Areira et al., 2012
Pratylenchus bracyurus Zea mays Potasio Dias-Areira et al., 2012
P. bracyurus Zea mays Manganeso Puerari et al., 20015
Radopholus similis M. paradisiaca Potasio Quintero-Vargas y Castaño-Zapata, 2012
Figura 2. Incidencia y severidad de Pseudoperonospora cubensis (A y A´) y Sphaerotheca fuliginea (B y B´) en plantas de
pepino cultivadas en invernadero.
Figure 2. Incidence and severity of Pseudoperonospora cubensis (A and A´) and Sphaerotheca fuliginea (B and B´) in cucum-
ber plants cultivated in the greenhouse.
las plantas de papaya tratadas con fosfito de potasio amylovora using streptomycin sulfate, compared
sobrevivieron al ataque de Phytophthora palmivo- with the incidence of the pathogen when they used
ra, en contraste con el 24% de sobrevivencia de las potassium phosphite (Figure 4); the compounds
plantas sin tratar (Vawdrey y Westerhuis, 2007). were injected into apple tree trunks. Meyer and
En comparación con los fungicidas convencio- Hausbeck (2013) reported a significant decrease
nales sintetizados, los fosfitos generalmente son in the incidence of death squash plants infected by
menos eficaces para disminuir el daño por los pa- Phytophthora capsici when they were treated with
tógenos que atacan plantas, al respecto Méndez et fluopicolide, mandipropamid or dimethomorph
al. (2010) consiguieron que plantas de melón trata- fungicides, compared with the incidence of plants
das con clorotalonil y mancozeb, mostraran menor treated with potassium phosphite. In general, the
daño por Pseudoperonospora cubensis que aqué- reports on the effectiveness of phosphites and
llas que recibieron tratamiento con fosfito de pota- conventional fungicides in controlling phytoparasite
sio (Figura 3). También, Aćimović et al. (2016) lo- organisms suggest that phosphites are less effective
graron incidencia de Erwinia amylovora significa- and could not replace fungicides at all, but that
tivamente inferior con sulfato de estreptomicina, en if they are included as part of an integrated pest
comparación con la incidencia obtenida con fosfito management program they could help reduce the
de potasio, cuando esos compuestos se inyectaron use of fungicides as well as the probability that
al tallo de árboles de manzana (Figura 4). Meyer y the organisms develop resistance (Liljeroth et
Hausbeck (2013) reportaron disminución significa- al., 2016). Méndez et al. (2010) reported similar
tiva en la incidencia de plantas de calabaza muertas effectiveness against Pseudoperonospora cubensis
in squash plants by alternately using chlorothalonil/
600
534.9 b
500 100
90
400 80
68.5
ABCPE
306.3 a 70
300
Incidencia (%)
215.7 a 60 52.5
200 50 47.3
43.5
40
100 30
20
0
Clorotalonil/ Fosfito de Clorotalonil/ 10
Mancozeb potasio Mancozeb + 0
Fosfito de Agua Fosfito de Acibenzolar Sulfato de
potasio potasio S metil estreptomicina
Figura 3. Promedio del área bajo la curva del progreso de Figura 4. Control de Erwinia amylovora después de dos
la enfermedad (ABCPE) obtenido con el trata- aplicaciones de inductor de resistencia o antibió-
miento para el combate de Pseudoperonospora tico, al tronco de árboles de manzana (Elabora-
cubensis en plantas de pepino (Elaboración de ción de los autores con datos de Aćimović et al.,
los autores con datos de Méndez et al., 2010). 2016).
Figure 3. Average area under the progress curve (ABCPE) Figure 4. Control of Erwinia amylovora after two appli-
from a treatment against Pseudoperonospora cations of a resistance inducer or antibiotics to
cubensis in cucumber plants (Developed by the trunks of apple trees (Developed by the authors
authors with data from Méndez et al., 2010). with data from Aćimović et al., 2016).
por Phytophthora capsici, cuando fueron tratadas mancozeb and potassium phosphite compared
con los fungicidas fluopicolide, mandipropamid o with the effectiveness when using only fungicides
dimethomorph, en comparación con la incidencia (Figure 3). Liljeroth et al. (2016) also reported that
registrada en las plantas tratadas con fosfito de po- the protection against Phytophthora infestans in
tasio. En general, los reportes comparativos de la potato was similar using fluazinam, cyazofamid,
eficacia en el control entre los fosfitos y los fun- mandipropamid, metalaxyl + fluazinam or
gicidas convencionales, indican que los primeros fluopicolide + propamocarb at 100% of the
son menos eficaces y que no los podrían sustituir recommended dose, or at 50% of the dose mixed
por completo, pero su integración como parte de un with potassium phosphite.
programa de manejo integrado puede permitir dis-
minuir el uso de fungicidas y reducir la posibilidad ACTION MODE
de generar resistencia por los organismos (Liljeroth
et al., 2016). Al respecto, Méndez et al. (2010) de- The action mechanisms involved in the
terminaron eficacia similar en el combate de Pseu- prophylactic effects of phosphites include direct
doperonospora cubensis en plantas de calabaza con and indirect action. It has been stated that when the
clorotalonil/mancozeb y fosfito de potasio aplica- phosphite ion come into contact with phytopathogen
dos de manera alternada, comparada con la eficacia organisms, it affects their growth and reproduction,
registrada cuando se utilizó únicamente los fungi- because it influences the expression of genes that
cidas (Figura 3). También, Liljeroth et al. (2016) encode the synthesis of essential compounds in
reportaron que la protección contra Phytophthora their cell structure and physiology (direct action).
infestans en papa, fue semejante cuando los fun- When the phosphite ion enters the plant tissue
gicidas fluazinam, cyazofamida, mandipropamida, cells (indirect action) activates biochemical
metalaxyl + fluazinam o fluopicolide + propamo- (production of: polysaccharides, proteins related
carb, se usaron al 100% de la dosis recomendada o to pathogenesis, phytoalexins, etc.) and structural
al 50% de la dosis en mezcla con fosfito de potasio. defense (such as callose deposition) mechanisms
that restrict pathogens penetration and survival in
MODO DE ACCIÓN the plant (Figure 5).
estructurales de defensa (como la deposición de ca- digitatum (Cerioni et al., 2013), Phytophthora
losa) que restringen la penetración y supervivencia cinnamomi (Wilkinson et al., 2001; Won et
de los patógenos en la planta (Figura 5). al., 2009; King et al., 2010), P. cinnamomi, P.
palmivora and P. nicotianae (Smillie et al., 1989),
Acción directa P. infestans (Bórza et al., 2014), P. plurivora (Dalio
et al., 2014) and Pythium aphanidermatum, P.
Los fosfitos adicionados al medio de cultivo ultimum, P. irregulare, P. myriotylum, P. torulosum,
para organismos fitopatógenos, originaron que dis- P. volutum and P. graminicola (Cook et al., 2009).
minuyera el crecimiento del micelio, el número de The decrease level in the microorganisms’ growth
estructuras generadoras de esporas, la cantidad de as a consequence of the phosphite ion is determined
esporas producidas y su germinación (Figura 6). by the organism tested, amount of phosphite added
Así, mediante esa práctica se reportó disminución to the culture medium, type of ion bonded to
en el crecimiento de Alternaria alternata (Reuve- phosphite and pH produced by the culture medium
ni et al., 2003), Colletotrichum gloeosporioides (Lobato et al., 2010).
(Araujo et al., 2010), Fusarium culmorum y F. gra- The susceptibility of the microorganisms to
minearum (Hofgaard et al., 2010), Mycosphaerella the phosphite ion was demonstrated by Wong et
Figura 5. Modo de acción de los fosfitos, representación esquemática (Elaboración de los autores con datos de: Daniel y
Guest, 2006; Jackson et al., 2000; King et al., 2010; Eshraghi et al., 2011; Olivieri et al., 2012; Cerioni et al., 2013;
Dalio et al., 2014).
Figure 5. Action mode of phosphites, schematic representation (developed by the authors using data from Daniel and
Guest, 2006; Jackson et al., 2000; King et al., 2010; Eshraghi et al., 2011; Olivieri et al., 2012; Cerioni et al., 2013;
Dalio et al., 2014).
A A’
B B’
Figura 6. Crecimiento de Pythium aphanidermatum en medio de cultivo Papa-Dextrosa-Agar (izquierda) y agua destilada
(derecha). A y A´ sin fosfito de potasio; B y B´ con fosfito de potasio.
Figure 6. Pythium aphanidermatum in Potato Dextrose Agar (PDA) culture medium (left) and distilled water (right). A and
A´ without potassium phosphite; B and B´ with potassium phosphite.
fijiensis (Mogollón y Castaño, 2012), Penicillium al. (2009), who determined the negative effect of
digitatum (Cerioni et al., 2013), Phytophthora cin- phosphite and the positive effect of phosphate on
namomi (Wilkinson et al., 2001; Won et al., 2009; Phythopthora cinnamomic growth, when grown in
King et al., 2010), P. cinnamomi, P. palmivora y P. a culture medium enriched with salts containing
nicotianae (Smillie et al., 1989), P. infestans (Bór- each ion individually.
za et al., 2014), P. plurivora (Dalio et al., 2014) As for interspecific susceptibility, Hofgaard et
y Pythium aphanidermatum, P. ultimum, P. irregu- al. (2010) were able to reduce Fusarium culmorum,
lare, P. myriotylum, P. torulosum, P. volutum y P. F. graminearum and Microdochium majus
graminicola (Cook et al., 2009). El grado de dis- mycelium growth by 60, 80 and 90%, respectively,
minución en el crecimiento de los microorganis- using 10 μl ml-1 of potassium phosphite. To inhibit
mos como consecuencia del ion fosfito, está deter- Phytophthora infestans in vitro growth, Lobato et
minado por el organismo ensayado, la cantidad de al. (2010) used a lower dose, followed by the dose
fosfito agregado al medio de cultivo, el tipo de ion used to inhibit Streptomyces scabies, Rhizoctonia
unido al fosfito y el pH que se origine en el medio solani and Fusarium solani growth.
de cultivo (Lobato et al., 2010). The intraspecific susceptibility was studied by
La susceptibilidad de los microorganismos al Wilkinson et al. (2001), who determined that from
ion fosfito fue comprobada por Wong et al. (2009), 21 Phytophthora cinnamomi isolates collected
quienes determinaron el efecto negativo de fosfi- in western Australia, 9% were susceptible, 82%
to y el positivo de fosfato sobre el crecimiento de intermediate susceptible and 9% tolerant. Smillie
Phythopthora cinnamomi, cuando creció en me- et al. (1989) showed Phytophthora cinnamomi,
dio de cultivo enriquecido con sales que contenían P. palmivora and P. nicotiana susceptibility to
cada ion de manera individual. potassium phosphite, and explained that as the
En relación a la susceptibilidad interespecífica, concentration of phosphite increased in the culture
Hofgaard et al. (2010) obtuvieron disminuciones medium, the weight of the biomass produced
de 60, 80 y 90% en el crecimiento del micelio de by Phytophthora species decreased. Cook et
Fusarium culmorum, F. graminearum y Microdo- al. (2009) also evaluated the concentration of
chium majus, respectivamente, con 10 μl ml-1 de phosphite and mefenoxan fungicide needed to
fosfito de potasio. Lobato et al. (2010) determina- inhibit 50% mycelium growth of eight Pythium
ron que para inhibir el crecimiento in vitro, la dosis aphanidermatum isolates, and reported that all the
menor se empleó para Phytophthora infestans, se- isolates were susceptible to both compounds, but
guida de la usada para inhibir a Streptomyces sca- that the concentrations of phosphite were higher
bies, Rhizoctonia solani y Fusarium solani. than those of the fungicide.
La susceptibilidad intraespecífica fue estudiada On the other hand, the phosphate ion reduced
por Wilkinson et al. (2001), quienes determinaron pH in the culture medium at acidity levels that
que de 21 aislamientos de Phytophthora cinnamomi were associated with Phytophthora infestans,
colectados en el oeste de Australia, 9% resultaron Rhizoctonia solani, Fusarium solani and
susceptibles a fosfito de potasio, 82% mostraron Streptomyces scabies low growth (Lobato et al.,
susceptibilidad intermedia y 9% fueron tolerantes. 2010). Araujo et al. (2010) reported a larger decrease
Smillie et al. (1989) determinaron la susceptibili- in the diameter of Colletotrichum gloeosporioides
dad de Phytophthora cinnamomi, P. palmivora y colonies as well as in the growth speed index using
P. nicotiana a fosfito de potasio, explicando ade- formulations containing potassium phosphite,
más que a medida que la concentración de fosfito which reduced pH in the medium culture.
se incrementó en el medio de cultivo, disminuyó Also, potassium phosphite affected Alternaria
el peso de la biomasa producida por las tres espe- alternata (Reuveni et al., 2003), Colletotrichum
cies de Phytophthora. También Cook et al. (2009) gloeosporioides (Ogoshi et al., 2013), Penicillium
evaluaron la concentración necesaria de fosfito y digitatum (Cerioni et al., 2013), P. expansum (Amiri
el fungicida mefenoxan, para inhibir el 50% del and Bompeix, 2011) and Peronospora parasitica
crecimiento del micelio de ocho aislamientos de (Becót et al., 2000) spore germination, as well as
Pythium aphanidermatum, y reportaron que todos Fusarium oxysporum f sp. cubense (Davis and
los aislamientos fueron susceptibles a los dos com- Grant, 1996), Mycosphaerella fijiensis (Mogollón
puestos, pero que las concentraciones de fosfito uti- y Castaño, 2012), Peronospora sparsa (Hukkanen
lizadas fueron mayores que las del fungicida. et al., 2008), Phytophthora plurivora (Dalio et al.,
Por otro lado, el ion fosfito disminuyó el pH en 2014) and P. cinnamomi (Wong et al., 2009) spore
el medio de cultivo a niveles de acidez que se rela- production.
cionaron con menor crecimiento de Phytophthora The effect of phosphites added to a medium
infestans, Rhizoctonia solani, Fusarium solani y culture was explained by King et al. (2010), who
Streptomyces scabies (Lobato et al., 2010). Araujo et reported changes induced by potassium phosphite
al. (2010) reportaron mayor diminución del diáme- in the expression of the genes that encode protein
tro de las colonias y el índice de velocidad de cre- synthesis and builds Phytophthora cinnamomi cells
cimiento de Colletotrichum gloeosporioides, con and cytoskeleton, which caused hyphae distortion
formulaciones de fosfito de potasio que originaron and cell wall lysis.
mayor disminución del pH en el medio de cultivo.
Además, fosfito de potasio afectó la germinación Indirect action
de esporas de Alternaria alternata (Reuveni et al.,
2003), Colletotrichum gloeosporioides (Ogoshi et The indirect action of the phosphite ion includes
al., 2013), Penicillium digitatum (Cerioni et al., stimulation of structural and biochemical defense
2013), P. expansum (Amiri y Bompeix, 2011) y Pe- mechanisms in plants. Pilbeam et al. (2011)
ronospora parasitica (Becót et al., 2000); así como described lignin and suberine deposition around
la producción de esporas de Fusarium oxysporum f tissue damaged by Phytophthora cinnamomi in
sp. cubense (Davis y Grant, 1996), Mycosphaerella eucalyptus plants treated with potassium phosphite,
fijiensis (Mogollón y Castaño, 2012), Peronospora which limited the pathogen’s development. Olivieri
sparsa (Hukkanen et al., 2008), Phytophthora plu- et al. (2012) reported an increased pectin content
rivora (Dalio et al., 2014) y P. cinnamomi (Wong in periderm and cortex tissue and tubers from
et al., 2009). potato plants treated with potassium phosphite,
El efecto de los fosfitos agregados al medio de a feature that improves resistance to different
cultivo fue explicado por King et al. (2010), quie- pathogens. Jackson et al. (2000) reported that the
nes reportaron modificación inducida por fosfito de development of lesions caused by P. cinnamomi
potasio en la expresión de genes que codifican la was highly restricted when they applied a
síntesis de proteínas que constituyen la pared y el high concentration of phosphite in Eucalyptus
citoesqueleto de las células de Phytophthora cinna- marginata tissue, and that the reduced number of
momi, lo que originó distorsión de hifas y lisis de lesions was associated with a significant increase of
pared celular. defense enzymes (4-coumarate coenzyme A ligase
(4CL) and cinnamyl-alcohol dehydrogenase)
Acción indirecta and soluble phenols. Daniel and Guest (2006)
demonstrated that when Arabidopsis thaliana
La acción indirecta del ión fosfito incluye la es- plantlets were treated with potassium phosphite
timulación de los mecanismos de defensa estructu- and inoculated with Phytophthora palmivora
ral y bioquímica en las plantas; así, Pilbeam et al. zoospores, the infected cells showed increased
(2011) describieron deposición de lignina y sube- cytoplasmic activity, development of cytoplasmic
rina alrededor del tejido dañado por Phytophtho- aggregates, release of hydrogen peroxide, located
ra cinnamomi en plantas de eucalipto tratadas con cell death, and enhanced phenolic compounds
fosfito de potasio, efecto que limitó el desarrollo accumulation. Eshraghi et al. (2011) also reported
del patógeno. También Olivieri et al. (2012) re- that when Arabidopsis thaliana plantlets were
firieron aumento en el contenido de pectina en el treated with potassium phosphite and inoculated
tejido de la peridermis y la corteza, en tubérculos with Phytophthora cinnamomi, the infected cells
procedentes de plantas de papa tratadas con fosfito increased their production of calose and hydrogen
de potasio, condición que mejora la resistencia a peroxide.
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