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BY 4.0 license Open Access Published by De Gruyter Open Access August 3, 2020

Microbial antagonists against plant pathogens in Iran: A review

  • Mehrdad Alizadeh , Yalda Vasebi and Naser Safaie EMAIL logo
From the journal Open Agriculture

Abstract

The purpose of this article was to give a comprehensive review of the published research works on biological control of different fungal, bacterial, and nematode plant diseases in Iran from 1992 to 2018. Plant pathogens cause economical loss in many agricultural products in Iran. In an attempt to prevent these serious losses, chemical control measures have usually been applied to reduce diseases in farms, gardens, and greenhouses. In recent decades, using the biological control against plant diseases has been considered as a beneficial and alternative method to chemical control due to its potential in integrated plant disease management as well as the increasing yield in an eco-friendly manner. Based on the reported studies, various species of Trichoderma, Pseudomonas, and Bacillus were the most common biocontrol agents with the ability to control the wide range of plant pathogens in Iran from lab to the greenhouse and field conditions.

1 Introduction

Increasing human population in the world demands more food (70 to 100%) by 2050 to supply human needs (Godfray et al. 2010). Furthermore, different pests and diseases cause annual economic losses (20 to 40%) in agricultural products by decreasing the crop yield, destroying the quality, and pollution of products with toxic chemicals (Guo et al. 2013). Therefore, growers have generally concentrated on the intensive use of chemicals for the management of pests and diseases which induce several problems, including resistance to pesticides, hazardous effects on human health, loss of beneficial soil microorganisms, entrance of residual toxic material in the food chain, and reduction in macro–microorganism biodiversity (Sindhu et al. 2016). These problems make enhanced attempts for developing ecofriendly microbe-based pesticides or biopesticides which use biological control agents (BCAs) as active ingredients and basically act different from common chemical pesticides (Sindhu et al. 2009).

Biological control, which attracted broad considerations in the past few decades, is defined as a bioeffector strategy that uses other living organisms for controlling insects, mites, weeds, and phytopathogens (Flint et al. 1998). Biocontrol agents either with antagonistic activities, or modifying effects on plant physiology and anatomy, mostly reduce the negative effects of pathogens. The advantages of beneficial microbes for associated plants are establishment of antagonistic microorganisms, prevention of phytopathogens, overall improvement of plant health, plant growth promotion, enhanced nutrient availability and uptake, and increased resistance to both biotic and abiotic stresses in the hosts (Vinale et al. 2014).

The first published studies on biological control of plant pathogens in Iran were presented in 1992. Trichoderma spp. and Gliocladium spp. were the first biocontrol agents applied against Athelia rolfsii (Sclerotium rolfsii), Rhizoctonia solani, and Fusarium solani, the causal agents of diseases on groundnut, bean, and apple, respectively (Asghari and Myee 1992; Bazgir et al. 1992; Karampour and Okhovat 1992). In the twenty-first century, with the improvement of biological control of plant pathogens throughout Iran, different biocontrol agents have been applied against the various pathogens in vitro, in greenhouse and field conditions. A large number of fungal and bacterial biocontrol agents have been found as the most important agents for plant disease management with identification of their role in plant pathogen management (Ramadan et al. 2016). Trichoderma, Pseudomonas, and Bacillus species have mostly been used for biological control of phytopathogens in Iran (Peyghami and Nishabouri 1998; Shahiri Tabarestani et al. 2000; Mostofizadeh-Ghalamfarsa et al. 2002; Niknejad-Kazempour et al. 2004a,b; Golzary et al. 2008b; Peighami-Ashnaei et al. 2009a,b; Ojaghian et al. 2010; Khalighi and Khodakaramian 2012; Naeimi and Zare 2013; Azizpour and Rouhrazi 2016; Karimi et al. 2016; Khaledi and Taheri 2016; Abdoli et al. 2018; Hosini et al. 2018; Zeynadini-Riseh et al. 2018). Furthermore, because of increasing the stability of biological agents, the bioformulation progress has recently been evaluated in Iran (Karimi and Sadeghi 2015). The current study is a comprehensive review of applying fungal and bacterial antagonists for biological control of various plant diseases caused by fungal, bacterial, and nematodes in Iran during a period of 26 years.

2 Mechanisms of biocontrol agents for the management of phytopathogens

A key factor for attaining an effective prevention of phytopathogens in their hosts is the knowledge about their mechanism of action. Understanding the mechanisms in the biological control process can allow the establishment of favorable conditions in the interaction between phytopathogen and biocontrol agent that is important in performing a successful biological control strategy in a specific pathosystem (Handelsman and Stabb 1996). The microorganisms operating for biocontrol of phytopathogens have different modes of action (Nega 2014). In the present study, the most common mechanisms of interspecies antagonisms include direct antagonism, mixed-path antagonism, and indirect antagonism (Pal and McSpadden 2006; Parveen et al. 2016), which lead to biological control of plant pathogens, have been addressed. Microbial biocontrol agents take care of plants against pathogens via different modes. These agents could induce resistance or initial enhanced resistance against pathogens without direct confrontation with the phytopathogen. Also, competitions for nutrients and spaces are additional indirect interactions with phytopathogens (Köhl et al. 2019). These agents might directly interact with the pathogens using hyperparasitism (Ghorbanpour et al. 2018) or antibiosis (Raaijmakers and Mazzola 2012). Without these agents in soil and tissues of plants, the pathogens easily attack plants and could weaken or kill considered hosts (Figure 1). These modes will be discussed in the following sentences.

Figure 1 Left: in the absence of antagonists, different pathogens especially fungi, bacteria, and nematodes can cause losses in plants. Over time, affected plants will show the weakness in the development and symptoms of diseases. Right: in the presence of antagonists with different biocontrol mechanisms, such as competition, parasitism, and antibiosis, the pathogens will not be able to progress in the host, and thus, the plant can grow and develop well rather than the absence of antagonists in soil and tissues of hosts.
Figure 1

Left: in the absence of antagonists, different pathogens especially fungi, bacteria, and nematodes can cause losses in plants. Over time, affected plants will show the weakness in the development and symptoms of diseases. Right: in the presence of antagonists with different biocontrol mechanisms, such as competition, parasitism, and antibiosis, the pathogens will not be able to progress in the host, and thus, the plant can grow and develop well rather than the absence of antagonists in soil and tissues of hosts.

2.1 Parasitism

Mycoparasitism, direct parasitism or hyperparasitism, is the ability of fungal antagonistic agents to parasite other fungi for utilizing them as food. Mycoparasitism causes either complete death of fungal propagules or destruction and lysis of their structure (Maloy 1993). Mycoparasitism depends upon the sequential occurrence of the following events: coming into close contact with fungal pathogen, mutual recognition between antagonist and pathogen, lytic enzyme secretion by antagonist, penetration into the host, active growth of antagonist into the host, and exit (Spadaro and Gullino 2004; Talibi et al. 2014). Various chemical compounds can be implicated in these processes, such as lectins, during the initial contact and recognition and cell wall-degrading enzymes (CWDEs), such as β-1,3-glucanases, chitinases, proteinases, and lipases, during the penetration process (Vos et al. 2015). Wisniewski et al. (1991) who studied biological control of Botrytis cinerea by yeast antagonist Meyerozyma guilliermondii (Pichia guilliermondii) demonstrated that lectin-like interaction resulted in firm attachment of antagonist’s cell to B. cinerea. Lysis of fungal cell wall also occurred due to the action of extracellular β-1,3-glucanase enzyme secreted by the antagonistic yeast. Trichoderma species are specific mycoparasitic fungi with the species of T. atroviride, T. virens, and T. reesei confirming that mycoparasitism is their ancestral lifestyle (Kubicek et al. 2011).

One of the main components in mycoparasitism event is CWDEs including endochitinases, β-1,3-glucanases, and proteases that are extracellular enzymes secreted by Trichoderma (Vos et al. 2015). After initial pathogen recognition by Trichoderma, hyphae wind around the pathogen’s hyphae by forming hook, the appressorium permeates into the pathogen cell, and chitin is broken down by enzymes such as chitinase and glucanase (Ghorbanpour et al. 2018). Subsequently, mycoparasitic’s hyphae release antibiotic compounds which penetrate the affected pathogen’s hyphae and resynthesize the host cell wall inhibited by these compounds (Toghueoa et al. 2016).

2.2 Antibiotic

Antibiotic is a secreting secondary metabolite with low molecular weight that is deleterious to the other microorganisms at low concentrations (Fravel 1988). The antibiotic produced by biocontrol agents decreases the disease symptoms as a main contributing mechanism particularly under soil conditions (Haas and Défago 2005). Some soilborne microorganisms, such as different strains of fluorescent Pseudomonas and Bacillus (Weller 1988) and Trichoderma species (Benítez et al. 2004), have appropriate features for biocontrol abilities. Furthermore, several strains of these species are able to promote plant growth and development as well as the disease prevention (Fernando et al. 2006; Arseneault and Filion 2017). The antibiotics at subinhibitory concentrations may inhibit the release of extracellular virulence factors and adherence mechanisms in bacteria (Kumar et al. 2008). Secondary metabolites can impress the community of soil microbial ecosystems in a variety of ways and levels (Abawi and Widmer 2000). The antibiotic production has been confirmed to be an important mechanism applied by microorganisms to manage a wide range of plant pathogens (McSpadden and Fravel 2002). Even at subinhibitory concentrations, antibiotics can create physiological changes in organisms. For instance, in Pseudomonas aeruginosa quinolone and macrolide antibiotics can block cell signaling and production of virulence factors (Ulloa-Ogaz et al. 2015). Bacillus spp. produce enzymes, exotoxins, and metabolites with nematicidal activity (Engelbrecht et al. 2018). Although several rhizobacteria such as Pasteuria, Pseudomonas, and Streptomyces have nematicidal efficacy, the largest decrease in the hatching of Meloidogyne javanica eggs was found in Bacillus (74%) and Pseudomonas (54.77%) (Turatto et al. 2017). Furthermore, Bacillus spp. with antibiotic production are applied as antifungal antagonists for controlling postharvest diseases. Pyrrolnitrin antibiotic produced by Burkholderia cepacia has been used against Penicillium digitatum, B. cinerea, and Penicillium expansum pathogens. Similarly, syringomycin produced by Pseudomonas syringae was utilized to prevent citrus green mold and apple grey mold (Dukare et al. 2019). Alongside these beneficial microorganisms, Streptomyces spp. can help plants with antibiotic production against phytopathogens (Olanrewaju and Babalola 2019).

2.3 Cell wall degradation enzymes

Microorganisms which produce enzymes are able to hydrolyze chitin, proteins, cellulose, and hemicellulose and also may play a role in the suppression of plant pathogens. Chitin and β-1,3-glucans are major constituents of many fungal cell walls (Lam and Gaffney 1993). Trichoderma strains with antagonistic potential have been mainly characterized by their ability to secrete enzymes such as chitinases, glucanases, and proteases that hydrolyze the cell walls of pathogens (López-Mondéjar et al. 2011). Geraldine et al. (2013) reported that N-β-acetylglucosaminidase and β-1,3-glucanase are the key components of Trichoderma species action in biocontrol of Sclerotinia sclerotiorum in the field. Serratia marcescens which produces chitinases was found to suppress the growth of Botrytis spp., R. solani, and Fusarium oxysporum (Ningaraju 2006).

2.4 Competition for available resources

Microorganisms’ challenge for available resources is named competition. For instance, when pine stumps were inoculated by spores Phlebiopsis gigantea (Phlebia gigantea), the spores prevent from Heterobasidion annosum infections. Considering that the pathogen is non-established on the pine, the severity of root rot disease could be decreased by the biocontrol agent (Cook and Baker 1983). Despite the possibility of existing antagonistic relationship (e.g., antibiosis) between the two fungi, the achievement of available resource sites may be the first mechanism in competition (Maloy 1993). Carbon sources such as glucose and fructose are one of the important action modes in yeasts Papiliotrema laurentii (Cryptococcus laurentii) and Sporobolomyces roseus, which can control B. cinerea in decreasing its colonization and sporulation (Ghorbanpour et al. 2018). In the biological control of P. digitatum by Debaryomyces hansenii, competition plays an important role in obtaining nutrients in occupied sites (Droby et al. 1998). Furthermore, arbuscular mycorrhiza due to the creation of physiological and anatomical modifications can limit the progression of pathogen. These changes involve root lignification, creation of a thick cell wall using pectin, chitinase activation, and transfer of pathogenesis-related protein-1a to the infected area of root (Malik et al. 2016).

2.5 Siderophore

Low-molecular weight chelators with a very high and specific affinity for Fe(iii) are called siderophores (Barbeau et al. 2002). Aerobic and facultative anaerobic microorganisms with the ability of siderophore production may have an important role in microorganism interactions (Haggag and Mohamed 2007). Siderophores have been known to play a significant role in phytopathogen prevention by several bacteria as BCAs which prevent the growth, development, and metabolic activity of phytopathogens by iron chelation (Haggag Wafaa et al. 2000). Different species of Trichoderma as biocontrol antagonists release more effective siderophores that chelate iron (Fe3+) and prevent growth and development of other fungal pathogens (Naher et al. 2014). Iron competition can be a limiting factor in alkaline soils for microbial growth and development (Leong and Expert 1989). Siderophores produced by some bacteria, such as fluorescent pseudomonads, have very high dependency for iron, as a result, sequestering these limited resources from other microflora can inhibit their growth and development (Loper and Buyer 1991). In several studies, it has been reported that Pseudomonas fluorescens with siderophore biosynthesis plays an important role in the prevention of pathogen (Costa and Loper 1994). Rahnella aquatilis with siderophore production can inhibit B. cinerea and P. expansum postharvest pathogens (Calvo et al. 2007). The siderophore pulcherrimin produced by Metschnikowia pulcherrima and Monilinia fructicola yeasts was applied for biological control of postharvest apple pathogens B. cinerea, Alternaria alternata, and P. expansum (Saravanakumar et al. 2008). In particular, several species of Streptomyces detach iron by siderophore production in a way that some pathogens, owing to a lack of siderophore production, cannot take these ions for growth (Kloepper et al. 1980).

2.6 Induction of host resistance

Plant growth promoting rhizobacteria can protect plants against pathogens using induction of systemic resistance (ISR) (Sikora 1992). P. fluorescens with stimulating ISR can prevent the early penetration of Heterodera schachtii to roots (Oostendorp and Sikora 1989). The ISR stimulation by Bacillus subtilis leads to the protection of cotton plants against Meloidogyne incognita and Meloidogyne arenaria. The ISR stimulation by Pseudomonas putida and S. marcescens inhibited cucumber Fusarium wilt caused by F. oxysporum f.sp. cucumerinum. The application of Pseudomonas sp. in plants leads to systematic protection against F. oxysporum f.sp. dianthi (David et al. 2018). Flavimonas oryzihabitans, S. marcescens, and Bacillus pumilus have developed ISR against P. syringae pv. lachrymans (David et al. 2018).

The direct promotion of plant growth by plant growth promoting bacteria through the production of phytohormones has been called phytostimulation (Bloemberg and Lugtenberg 2001). The enzyme 1-aminocyclopropane-1-carboxylate (ACC) deaminase is a phytostimulation that is the most studied one. Some bacterial endophytes producing ACC deaminase have been shown to enhance plant growth, such as Arthrobacter spp., Bacillus spp., P. putida, Rhodococcus spp. (Belimov et al. 2001; Sziderics et al. 2007), and Streptomyces spp. (Palaniyandi et al. 2014; Jaemsaeng et al. 2018). The bacterial strains producing other plant hormones, including indole-3-acetic acid (IAA), jasmonates, and abscisic acid, may also contribute to plant growth stimulation (Patten and Glick 2002; Forchetti et al. 2007). IAA is synthesized by different species of Streptomyces, such as S. violaceus, S. griseus, S. exfoliate, S. coelicolor, and S. lividans (Manulis et al. 1994). Also, IAA in S. atrovirens activates growth promoting bacteria in groundnut and several crops (Reddy et al. 2016).

3 Reduction in the population of biocontrol agents

Phytopathogens may significantly alleviate the growth of biocontrol agents by using the nutrition resources within their occupied spaces more rapidly as well as by modifying their efficacy. This was found in several fungal root pathogens which can colonize the wheat rhizosphere despite the presence of P. fluorescens biocontrol agent (Mazzola and Cook 1991). Decline in the population of P. fluorescens occurs in the existence of some Pythium species. In this instance, infection by Pythium species leads to the limitation of the root surface which is available for P. fluorescens colonization and to the reduction of population of potential antagonists. Fedi et al. (1997) reported that a plant pathogenic P. ultimum with modification of gene expression of P. fluorescens tends to decrease biocontrol agent population. The competition in the rhizosphere for nutrients released from root wounds caused by P. ultimum was limited by the reduction of population size. Because of the importance of microbial community in number and diversity, competition and microorganism–microorganism interactions may also happen in phyllosphere (Vorholt 2012). On the other hand, existence of these microbial communities may also impress the efficacy of BCAs. Understanding the rhizosphere, phyllosphere, and endosphere microbial community structure and their interactions in these niches can contribute to the betterment of biocontrol (Bardin et al. 2015).

4 Improving the biocontrol agent effects

The use of combinations of BCAs may be a better method for developing biocontrol positive effects (Duffy and Weller 1995). Combined biocontrol agents with high level of biocontrol protection have been investigated for better efficacy and prevention of several phytopathogens (Mihajlović et al. 2017). It has been confirmed that natural prevention of Fusarium wilt in France (Châteaurenard soil) was related to the different mechanisms in which multiple microorganisms singly or together restricted the pathogen activation (Alabouvette et al. 1998). However, given that the application of biological control against soilborne pathogens will not be a good replacement of methyl bromide fumigation, these two methods could act together in integrated pest management (Akrami et al. 2011).

5 Biological control in Iran

A complete list of all pathogens and the antagonists used against them is provided in Table 1. Results showed that most studies were conducted in vitro and in greenhouse conditions, and a few cases were carried out in the farm condition in Iran. Bacterial strains belonging to 24 genera, Achromobacter, Acinetobacter, Azotobacter, Bacillus, Beauveria, Bradyrhizobium, Brochothrix, Burkholderia, Enterobacter, Erwinia, Escherichia, Flavobacterium, Lactobacillus, Mesorhizobium, Paenibacillus, Pantoea, Pasteuria, Pseudomonas, Rhizobium, Serratia, Sphingomonas, Sporolactobacillus, Stenotrophomonas, and Streptomyces, have been used in various studies. Also, fungal strains belonging to 27 genera, Acremonium, Alternaria, Arthrinium, Arthrobotrys, Aspergillus, Chaetomium, Cladobotryum, Coniothyrium, Embellisia, Fusarium, Gliocladium, Glomus, Hypsizygus, Lecythophora, Metarhizium, Paecilomyces, Penicillium, Periconia, Piriformospora, Pleurotus, Pythium, Scopulariopsis, Sebacina, Talaromyces, Trichoderma, Trichothecium, and Verticillium, have been applied in Iranian studies against different plant pathogens. Also, strains of ten genera, Candida, Galactomyces, Hanseniaspora, Metschnikowia, Meyerozyma, Pichia, Rhodotorula, Saccharomyces, Torulaspora, and Zygoascus, which belong to the yeast have been used for controlling the phytopathogens in Iran. The bacterial strains related to different species of Pseudomonas and Bacillus and fungal strains related to Trichoderma species had the greatest efficiency in biological control of different plant pathogens in Iran. These antagonists have been mostly used for biological control of fungi, bacteria, and nematodes, respectively.

Table 1

List of pathogens, hosts, and antagonists with procedures based on published research works in Iran from 1992 to 2018

PathogensHostAntagonistsProcedureRef.
Rhizoctonia solaniBeanGliocladium sp.In vitro and greenhouseBazgir et al. (1992)
Athelia rolfsii (Sclerotium rolfsii)GroundnutTrichoderma harzianumGreenhouseAsghari and Myee (1992)
Fusarium solaniAppleT. koningii, T. viride, T. harzianum, and T. virens (Gliocladium virens)GreenhouseKarampour and Okhovat (1992)
Colletotrichum coccodesPotatoTrichoderma spp.In vitroOkhovat et al. (1994)
R. solaniRiceT. koningii, T. viride, T. harzianum, and T. virensFieldIzadyar and Padasht (1994)
R. solaniRiceTrichoderma sp.In vitroPourabdullah and Binesh (1994)
Phytophthora erythrosepticaPotatoT. harzianum, T. viride, and T. koningiiIn vitroZafari et al. (1994)
R. solaniBeanT. viride, T. harzianum, and T. virensGreenhouseBazgir et al. (1994a)
R. solaniBeanT. viride, T. harzianum, and T. virensFieldBazgir et al. (1994b)
Scelotinia sclerotiorumEggplantT. reesei, T. hamatum, T. longibrachiatum, T. koningii, T. viride, T. virens, and Gliocladium sp.In vitroAmir-Sadeghi et al. (1994)
Macrophomina sp. and Rhizoctonia sp.SoybeanBacillus subtilisIn vitro and greenhouseSanei and Ghobadi (1995)
Heterodera schachtiiSugar beetPaecilomyces farinosusIn vitroAhmadi et al. (1995a)
H. schachtiiSugar beetF. solaniIn vitroAhmadi et al. (1995b)
H. schachtiiSugar beetAcremonium spp., Embellisia chlamydospora, Fusarium spp., P. lilacinus, Scopulariopsis brevicaulis, Verticillium chlamydosporium, and Verticillium lecaniiIn vitroHojjat Jalali and Coosemans (1995)
Tilletia laevisCucumberT. virideGreenhousePeyghami and Babadoost (1996)
T. controversaWheatT. virideGreenhousePeyghami and Babadoost (1996)
F. solaniChickpeaT. koningii, T. viride, T. harzianum, and T. virensGreenhouseOkhowat and Karampour (1996)
M. javanicaPasteuria penetransGreenhouseDamadzadeh et al. (1996)
M. javanica, M. incognita, and M. arenariaP. penetransIn vitroAmeri et al. (1996)
Macrophomina phaseolina and R. solaniSoybeanB. subtilisIn vitroSanei and Ghobadi (1996)
H. schachtiiSugar beetE. chlamydospora, Acremonium spp., S. brevicaulis, P. lilacinus, Fusarium spp., V. chlamydosporium, and V. lecaniiIn vitroHojjat-Jalali and Coosemans (1995)
Pythium ultimumChickpeaT. viride and T. virensFieldShahriary et al. (1996)
M. javanicaTomatoP. lilacinusGreenhouseFatemy (1996)
H. schachtiiSugar beetP. farinosusIn vitroAhmadi et al. (1996a)
H. schachtiiSugar beetF. solaniIn vitroAhmadi et al. (1996b)
R. solani, Colletotrichum coccodes, and Phytophthora drechsleriTrichoderma spp. and Gliocladium sp.In vitroOkhovat (1997)
H. schachtiiBeetPaecilomyces fumosoroseusGreenhouseFatemy and Ahmadian Yazdi (1997)
F. o. f.sp. cucumerinumCucumberT. harzianumGreenhousePeyghami and Nishabouri (1998)
A. rolfsii (S. rolfsii)GroundnutT. aureoviride, T. hamatum, T. longibrachiatum, T. harzianum, T. virensIn vitroMirhosaini et al. (1998)
R. solani, Bipolaris sorokiniana, and Fusarium culmorumWheatPenicillium polonicumGreenhouseMansoori (1998)
Ph. capsiciPepperTrichoderma sp. and Gliocladium sp.In vitroBehboodi et al. (1998)
P. ultimumPythium oligandrumIn vitroRahnama and Cooke (1998)
Pythium butleriAspergillus nigerIn vitroRouhani and Safari 1998
Mauginiella scaettaeDate palmT. koningii and T. virideFieldShetab-Booshehri et al. (1998)
Xanthomonas translucens pv. cerealisWheatPantoea agglomerans and Pseudomonas fluorescensGreenhouseMarefat and Rand Rahimian (1998)
F. o. f.sp. lycopersiciTomatoT. harzianum and T. virideGreenhouseNiknejad et al. (2000)
Erwinia amylovoraPearErwinia herbicola and P. fluorescensIn vitro, greenhouse, and fieldAhmadi et al. (2000)
S. sclerotiorumAubergineT. harzianum, T. virens, T. koningii, Trichoderma pseudokoningii, and Gliocladium deliquescensIn vitro and greenhouseOmrani et al. (2000)
M. phaseolinaSoybeanT. viride, T. koningii, and T. harzianumIn vitroGhaffarian et al. (2000)
R. solaniRiceT. viride, T. koningii, and T. harzianumFieldIzadyar et al. (2000a)
Verticillium dahliaeCottonTalaromyces flavusIn vitro and greenhouseNaraghi et al. (2000)
V. dahliaeCottonPseudomonas sp. and Bacillus sp.In vitroAzad Disfani et al. (2000)
R. solaniSugar beetT. harzianum, T. viride, and T. virensIn vitro and greenhouseShahiri Tabarestani et al. (2000)
R. solaniRiceT. harzianum, T. viride, T. koningii, and T. virensIn vitroIzadyar et al. (2000b)
R. solaniRiceT. harzianum, T. viride, and T. virensIn vitro and greenhouseNiknejad-Kazempour et al. (2000)
Neofusicoccum mangiferaeCitrusT. harzianum, T. virens, T. koningii, and T. longibrachiatumIn vitroTaheri et al. (2000)
S. sclerotiorumMulberryT. harzianum, T. viride, T. aureoviride, T. koningii, T. saturniporum, T. pseudokoningii, and T. longibrachiatumIn vitroMerat et al. (2000)
Fusarium spp., Sclerotium cepivorum, Pythium spp., and R. solaniOnionT. harzianum and T. virideGreenhousePeyghami (2001)
R. solaniRiceT. harzianum, T. viride, and T. virensIn vitro and greenhouseNiknejad Kazempour et al. (2002)
Gaeumannomyces graminis var. triticiWheatT. harzianum and T. virideGreenhouseForoutan et al. (2002)
F. avenaceum, F. graminearum, F. culmorum, F. moniliforme, F. oxysporum, F. solani, F. semitectum, F. sambucinum, F. proliferatum, and F. tricinctumWheatP. fluorescens, P. syringae, P. putida, P. cichorii, P. aeruginosa, P. aureofaciens, and P. viridiflavaIn vitroMostofizadeh-Ghalamfarsa et al. (2002)
G. graminis var. triticiWheatPseudomonas spp.In vitro and greenhouseSedaghatfar et al. (2002)
G. graminis var. triticiWheatT. harzianum and T. virideIn vitro and greenhouseForoutan et al. (2002)
F. graminearum, F. moniliforme, F. nygamai, F. oxysporum, F. proliferatum, F. sambucinum, F. semitectum, F. solani, and F. tricinctumWheatPseudomonas spp.In vitroMostofizadeh-Ghalamfarsa et al. (2002)
F. o. f.sp. melonisMelonStreptomyces sp., T. harzianum, T. virens, and T. virideIn vitro and greenhouseAshrafizadeh et al. (2002)
M. javanicaTomatoP. lilacinusGreenhousePakniat and Banihashemi (2002)
Ph. drechsleriCucurbitStreptomyces sp.In vitroHeidari Faroughi et al. (2002)
F. graminearumWheatStreptomyces sp., Pseudomonas sp., and Bacillus sp.In vitroNorouzian et al. (2002)
Sclerotinia minorSunflowerT. harzianum, T. viride, and T. virensIn vitroAbdollahzadeh et al. (2003)
Ph. drechsleriCantaloupeT. harzianum, T. viride, and T. virensGreenhouseHeidari Faroughi et al. (2004)
Tilletia indicaWheatT. longibrachiatum, T. harzianum, and T. virideGreenhouseBeeazar and Torabi (2004)
F. o. f.sp. ciceriChickpeaT. longibrachiatumGreenhouseKarimi et al. (2004a)
F. o. f.sp. ciceriChickpeaBacillus sp.In vitroKarimi et al. (2004b)
F. o. f.sp. dianthiCarnationP. fluorescens and Bacillus sp.GreenhouseKarimi et al. (2004c)
R. solaniChickpeaT. harzianum, T. viride, and T. virensGreenhouseMohammadi et al. (2004)
B. sorokinianaWheatB. subtilis, P. fluorescens, and Bacillus pumilusGreenhouseMohammadi et al. (2004)
M. phaseolinaSoybeanT. harzianumGreenhouseBarari et al. (2004)
Armillaria melleaCladobotryum polypore, C. varium, C. dendroides, and C. verticillatumIn vitroAsef and Mohammadi-Gholtapeh (2004)
T. laevisWheatB. subtilisGreenhouseKhodaygan et al. (2004)
B. sorokinianaWheatTrichoderma sp. and Streptomyces sp.GreenhouseSalehpour et al. (2004)
R. solaniRiceP. fluorescensField and greenhouseNiknejad-Kazempour (2004)
F. moniliformeRiceP. fluorescensIn vitroNiknejad-Kazempour et al. (2004)
R. solaniRiceBacillus cereus and P. fluorescensIn vitroSajjadi et al. (2004)
Pyricularia griseaRiceBacillus megaterium, B. subtilis, Bacillus circulans, and P. fluorescensFieldPadasht-Dehkaei et al. (2004)
F. oxysporumOnionB. cereus, B. subtilis, and P. fluorescensFieldSaberi-Riseh et al. (2004)
Ph. citrophthoraPistachioP. fluorescensFieldSaberi-Riseh et al. (2004)
P. ultimumCucumberB. subtilis, Trichoderma sp., and P. fluorescensGreenhouseTaghinasab et al. (2004)
F. oxysporumBasalB. cereus and P. fluorescensGreenhouse and fieldRamezani-Baghmishezad et al. (2004)
S. sclerotiorumRapeseedBacillus spp.In vitroAkbari-Kiarodi et al. (2004)
R. solaniCottonBacillus sp. and P. fluorescensIn vitro and greenhouseHeydari et al. (2004)
Gibberella fujikuroiRiceT. virens, T. harzianum, Bacillus sp., B. subtilis, and B. circulansIn vitro and greenhousePadasht Dehkaei et al. (2004)
Xanthomonas axonopodis pv. citriCitrusP. fluorescens and P. putidaIn vitro and greenhouseKhodakaramian (2004)
F. oxysporumChickpeaB. subtilis and P. fluorescensGreenhouse and fieldJamali et al. (2004)
R. solaniSugar beetB. subtilisIn vitro and greenhouseShahiri et al. (2005)
B. sorokinianaWheatB. subtilis and P. fluorescensIn vitro and greenhouseMohammadi et al. (2005)
Ph. capsiciPepperT. viride, T. koningii, T. harzianum, and T. virensIn vitro and greenhouseBehboudi et al. (2005)
S. sclerotiorumRapeseedB. cereus, B. subtilis, and P. fluorescensIn vitroAkbari et al. (2005a,b)
F. graminearumWheatP. aeruginosa, B. subtilis, and P. fluorescensGreenhouseForoutan et al. (2005)
V. dahliaeStreptomyces plicatus and Frankia sp.In vitroShahidi Bonjar and Aghighi (2005)
Pseudomonas tolaasiiAgaricus bisporusP. fluorescensIn vitro and greenhouseKhabbaz Jolfaei et al. (2005)
R. solani, F. oxysporum, F. solani, and C. coccodesPotatoT. harzianumGreenhouse and fieldSoltani et al. (2006)
S. sclerotiorumSunflowerT. harzianum, T. viride, and T. virensIn vitroAbdollahzadeh et al. (2006)
M. javanica and M. incognitaPistachioP. penetransGreenhouseKarimipourfard and Damadzadeh (2006)
R. solani, F. oxysporum, F. solani and Lasiodiplodia sp.MulberryP. fluorescens and Bacillus spp.In vitroNiknejad-Kazempour et al. (2006)
Ph. cactorumAppleP. fluorescens and B. subtilisGreenhouseFarzaneh et al. (2006)
F. o. f.sp. tuberosiPotatoP. fluorescensGreenhouseKhorasani-Aghazadeh et al. (2006)
R. solaniCommon beanBurkholderia cepaciaGreenhouseAhmadzadeh et al. (2006b)
Ascochyta rabieiChickpeaT. harzianumGreenhouseBahrami et al. (2006)
F. graminearumWheatStreptomyces sp., P. fluorescens, and B. subtilisIn vitro and greenhouseNourozian et al. (2006)
Bipolaris spiciferaWheatBacillus sp. and P. fluorescensGreenhouseBehdani et al. (2006)
R. solaniBeanP. fluorescensGreenhouseAfsharmanesh et al. (2006)
M. phaseolina, R. solani, Ph. nicotianae var. parasitica, Pythium sp., and Fusarium sp.Soybean, pistachio, bean, pepper, and cucumberPseudomonas spp.In vitroAhmadzadeh et al. (2006a)
V. dahliaeCucumberB. subtilis, P. fluorescens, and B. pumilusGreenhouseAhmadifar et al. (2006)
R. solaniRiceP. fluorescensIn vitroKazemzadeh et al. (2006)
T. laevisWheatP. putida and P. fluorescensGreenhouseKhodaygan et al. (2006)
Ph. sojaeSoybeanPseudomonas spp.GreenhouseZebarjad et al. (2006)
R. solaniRiceB. cereus, B. subtilis, and P. fluorescensGreenhouseSajadi et al. (2006)
M. phaseolinaMelonT. harzianum and T. virensIn vitro and glasshouseEtebarian (2006)
F. graminearumWheatB. cereus, B. subtilis, P. fluorescens, and E. herbicolaGreenhouse and fieldAlimi et al. (2006)
Ralstonia solanacearum and Pectobacterium carotovorum subsp. carotovorumPistachio, olive, potato, and cottonS. plicatus and Frankia sp.In vitroShahidi Bonjar et al. (2006)
V. dahliaePistachio, olive, potato, and cottonS. plicatus and Frankia sp.In vitroAghighi et al. (2006)
Sclerotinia sclerotiorumSunflowerConiothyrium minitansIn vitroPourmehdi Alamdarlou et al. (2006)
Ustilago hordeiBarleyBacillus licheniformis, B. cereus, and P. fluorescensFieldEtebarian et al. (2007)
S. sclerotiorumCanolaP. fluorescensIn vitro and greenhouseBehnam et al. (2007)
Ophiostoma novo-ulmiElmT. harzianum and T. virensIn vitroIraqi et al. (2007)
G. graminis var. triticiWheatT. virens, T. koningiopsis, T. koningii, and T. viridescensIn vitro and greenhouseMehrabi Koshki et al. (2007)
R. solaniBeanB. subtilis and P. fluorescensGreenhousePeighamy-Ashnaei et al. (2007)
R. solaniRapeB. cepaciaIn vitroSharifi-Tehrani et al. (2007)
Ph. cactorumAppleP. fluorescensIn vitro and greenhouseFarzaneh et al. (2007)
P. griseaRiceB. circulans, B. megaterium, B. subtilis, and P. fluorescensFieldPadasht and Izadyar (2007)
F. o. f.sp. dianthiCarnationB. cereus, B. subtilis, and P. fluorescensIn vitro and greenhouseKarimi et al. (2007)
R. solaniColzaP. fluorescensIn vitro and greenhouseSarani et al. (2008b)
S. sclerotiorumTobaccoT. citrinoviride, T. harzianum, T. atroviride, T. virens, T. koningii, T. ghanense, and T. longibrachiatumIn vitro and greenhouseSajadi and Asemi (2008)
M. phaseolinaEggplantT. hamatum, T. harzianum, T. polysporum, and T. virideIn vitro and fieldRamezani (2008)
M. javanicaTomatoT. harzianumIn vitroGolzary et al. (2008b)
A. melleaFruit treesT. harzianum and T. virensIn vitroAsef et al. (2008)
Penicillium digitatumOrangePseudomonas spp.In vitroZamani et al. (2008a)
P. digitatumOrangeP. agglomeransGreenhouseZamani et al. (2008b)
F. o. f.sp. tuberosePotatoBrevibacillus brevis, B. subtilis, and P. fluorescensGreenhouseKhorasani et al. (2008)
Colletotrichum gloeosporioidesCitrusB. subtilisIn vitroSalari et al. (2008a)
Penicillium expansumAppleT. virensGreenhouseTabe-Bordbar et al. (2008b)
Aspergillus flavusPistachioB. subtilis, B. licheniformis, B. cereus, and P. fluorescensIn vitroHaghdel et al. (2008a,b)
M. javanicaTomatoP. fluorescensGreenhouseMokhtari et al. (2008)
M. javanicaTomatoP. fluorescensIn vitroGolzary et al. (2008a)
Magnaporthe salviniRiceP. fluorescensIn vitroAhmaddeh et al. (2008)
F. oxysporumPotatoP. putida, P. fluorescens, and P. aeruginosaFieldOmmati et al. (2008)
P. digitatumCitrusT. viride, P. fluorescens, and B. subtilisIn vitroZamani et al. (2008c)
F. solani and P. ultimumB. subtilisIn vitroSelselehzakeri et al. (2008)
R. solaniSugar beetT. harzianum and T. virideFieldSafaee et al. (2008)
R. solaniSugar beetP. oligandrumIn vitro and greenhouseSalari et al. (2008b)
R. solaniCanolaP. fluorescens, B. cepacia, B. subtilis, and Streptomyces sp.In vitro and greenhouseSarani et al. (2008a)
Penicillium solitumAppleT. viride and T. virensGreenhouseTabe Bordbar et al. (2008a)
O. novo-ulmiElmB. subtilisIn vitroIragi et al. 2008
R. solaniRiceT. atroviride, T. harzianum, and T. virensIn vitroKhalili and Sadravi (2008)
Botrytis maliAppleCandida membranifaciensGreenhouseAlavifard et al. (2008a)
Botrytis cinereaAppleC. membranifaciens, Rhodotorula mucilaginosa, and Meyerozyma guilliermondii (Pichia guilliermondii)In vitro and greenhouseAlavifard et al. (2008b)
P. expansumAppleC. membranifaciensIn vitro and greenhouseGholamnejad et al. (2008)
S. sclerotiorumPotatoT. ceramicum , T. koningii, T. koningiopsis, T. virens, T. viridescens, T. orientalis, and T. atrovirideIn vitro and greenhouseOjaghian et al. (2008)
G. graminis and M. phaseolinaPiriformospora indica and Sebacina vermiferaIn vitroAbbaszadeh and Mohammadi Goltapeh (2008a)
M. phaseolinaSoybeanT. harzianum, T. viride, P. indica, and S. vermiferaIn vitro and greenhouse and fieldAbbaszadeh and Mohammadi Goltapeh (2008b)
Pyricularia oryzaeRiceStreptomyces spp.In vitro and greenhouseEbrahimi-Zarandi et al. (2008)
P. expansumAppleP. fluorescensIn vitro and greenhouseKhazaee et al. (2008)
Ph. nicotianaeP. fluorescensIn vitro and greenhouseNazerian et al. (2008)
S. sclerotiorumCanolaB. subtilisIn vitroNasrolah Nejad and Rahnama (2008)
P. syringae pv. tomatoTomatoP. fluorescensIn vitroMousavi et al. (2008a)
Clavibacter michiganensis subsp. michiganensisTomatoP. fluorescensIn vitroMousavi et al. (2008b)
E. amylovoraPearP. fluorescens and Pantoea sp.In vitroMirzaie et al. (2008)
M. phaseolinaSoybeanT. harzianumIn vitroMontazernia et al. (2008)
S. sclerotiorumCanolaP. fluorescens and B. subtilisIn vitro and greenhouseMansouripour et al. (2008)
G. graminis var. triticiWheatAzotobacter isolatesIn vitroMaghsodloo et al. (2008)
B. cinereaAppleB. subtilis, Pichia membraniciens, and Candida guilliermondiiIn vitro and greenhouseZangoie et al. (2008)
X. axonopodis pv. citriCitrusP. fluorescensGreenhouseKhodakaramian et al. (2008)
F. graminearum, R. solani AG4, R. solani AG5, M. phaseolina, and Ph. cactorumWheat, sugar beet, potato, soyabean, and appleT. hamatum, T. harzianum, T. virens, and Trichoderma sp.In vitroHajieghrari et al. (2008)
G. graminis var. triticiWheatT. koningiopsis, T. brevicompactum, and T. viridescensGreenhouseZafari et al. (2008)
M. javanicaTomatoT. harzianumIn vitro and greenhouseMaleki Ziyarati et al. (2009)
M. phaseolinaMelonP. fluorescens and P. putidaIn vitro and greenhouseKheiri et al. (2009)
H. schachtiiSugar beetT. harzianum and T. virensIn vitro and greenhouseMahdikhani Moghadam et al. (2009)
S. sclerotiorumCanolaT. harzianum and T. virensIn vitroNasrolah Nejad et al. (2009)
M. javanicaTomatoT. harzianumGreenhouseZiarati et al. (2009)
T. laevisWheatT. koningii, T. brevicompactum, T. harzianum, and T. virensFieldMehrabi Koshki et al. (2009)
R. solaniCommon beanP. fluorescensGreenhouseAhmadzadeh and Tehrani (2009)
B. cinereaAppleP. fluorescens and B. subtilisGreenhousePeighami-Ashnaei et al. (2009a)
S. sclerotiorumSunflowerP. fluorescensGreenhouseAshofteh et al. (2009)
Xanthomonas campestris pv. malvacearumCottonP. aeruginosaGreenhouseFallahzadeh-Mamaghani et al. (2009)
R. solaniBeanB. subtilis and P. fluorescensIn vitroPeighami-Ashnaei et al. (2009b)
R. solaniCommon beanB. cepaciaIn vitro and greenhouseAhmadzadeh et al. (2009)
P. expansumAppleSaccharomyces cerevisiaeIn vitroGholamnejad et al. (2009)
H. schachtiiSugar beetPleurotus ostreatus, P. sajor-caju, P. florida, P. flabellatus, P. eryngii, and Hypsizygus ulmariusIn vitro and greenhousePalizi et al. (2009)
P. expansumAppleR. mucilaginosa and M. guilliermondiiIn vitroGholamnejad et al. (2009)
V. dahliaeCottonGlomus etunicatum, G. intraradices, and G. versiformeGreenhouseNorouzi et al. (2009)
Meloidogyne spp.Paecilomyces lilacinusGreenhouseBoromand et al. (2010)
F. oxysporum, R. solani, M. phaseolina, and Pythium sp.Faba beanPseudomonas sp.In vitro and greenhouseGolpayegani et al. (2010)
R. solaniRiceT. harzianum, T. atroviride, and T. virensIn vitro, greenhouse, and fieldNaeimi et al. (2010)
Phytophthora sojaeT. virens, T. orientalis, T. brevicompactum, T. atroviride, T. ceramicum and T. asperellumIn vitroAyoubi et al. (2010)
B. sorokinianaWheatP. fluorescensIn vitro and greenhouseRanjbar Sistani et al. (2010)
G. fujikuroiRiceT. harzianum and T. virensIn vitro and greenhouseRoodgar et al. (2010)
Pythium aphanidermatumCucumberB. subtilis and B. licheniformisIn vitro and greenhouseSafari Asl et al. (2010)
B. cinereaTomatoT. harzianum, T. arundinaceum, T. viridescens, T. atroviride, and T. koningiiIn vitro and greenhouseEivazi et al. (2010)
P. expansumAppleB. subtilisIn vitro and greenhouseEmadi et al. (2010)
Phytophthora drechsleriCantaloupePseudomonas fluorescens, P. putida and P. aeruginosaIn vitro and greenhouseTabarraie et al. (2010)
P. expansumAppleR. mucilaginosaIn vitro and greenhouseGolamnejad et al. (2010)
Penicillium italicumOrangeM. guilliermondiiIn vitro and greenhouseGhasemi Sardareh et al. (2010)
Verticillium albo-atrumTomatoT. flavusIn vitro and greenhouseNaraghi et al. (2010)
F. o. f.sp. ciceriChickpeaB. subtilis, P. aeruginosa, and P. putidaIn vitroKarimik Amini et al. (2010)
B. sorokinianaWheatGlomus fasciculatum and B. subtilisGreenhouseHashemi Alizade et al. (2010)
F. o. f.sp. radicis-cucumerinumCucumberB. subtilisGreenhouseYousefi et al. (2010)
S. sclerotiorumPotatoT. ceramicum, T. koningii, T. koningiopsis, T. viridescens, T. virens, and Coniothyrium minitansIn vitroOjaghian et al. (2010)
Sclerotinia sclerotiorumSunflowerPseudomonas fluorescensIn vitro and greenhouseKhezri et al. (2010)
Sclerotium cepivorumGarlicBacillus spp.In vitroBabaei Nasir et al. (2010)
Fusarium oxysporum f.sp. gladioli GladiolusGarlicTrichoderma spp.In vitroBagheri et al. (2010)
F. oxysporum and F. solaniChickpeaT. harzianum and T. asprellumIn vitro and greenhouseAkrami and Ibrahime (2010)
M. phaseolinaSunflowerB. subtilisIn vitro and greenhouseIraqi and Rahnama (2011)
R. solaniCanolaB. cepaciaIn vitro and greenhouseSarani et al. (2010)
P. carotovorumPotatoP. putida, P. aeruginosa, and P. fluorescensFieldKhodakaramian and Zafari (2010)
X. axonopodis pv. citriCitrusP. fluorescens, P. viridiflava, and P. syringaeIn vitroMontakhabi et al. (2010)
G. graminis var. triticiWheatB. subtilis, B. pumilus, P. fluorescens, P. putida, P. aeruginosa, and Chromobacteria sp.In vitro and greenhouseBabaeipoor et al. (2011)
Phoma lingamRapeseedB. subtilis and T. koningiiIn vitro and greenhousePanjehkeh et al. (2011)
H. schachtiiSugar beetT. harzianum, T. virens, and B. subtilisFieldMahdikhani Moghadam and Rouhani (2011)
F. oxysporumLentilP. fluorescensGreenhouseAkrami et al. (2011)
F. culmorumB. subtilisGreenhouseKhezri et al. (2011)
S. sclerotiorumSunflowerP. fluorescensGreenhouseHeidari-Tajabadi et al. (2011)
P. italicum and P. digitatumCitrusP. syringae and Candida famataGreenhouseNasrollahi Omran et al. (2011)
G. graminis var. triticiWheatP. fluorescensGreenhouseBagheri et al. (2011)
P. expansumAppleR. mucilaginosaIn vitroGholamnejad et al. (2011)
Phytophthora parasitica and Ph. citrophthoraPistachioStreptomyces sp.In vitro and greenhouseSalari et al. (2011)
Ph. drechsleriSugar beetT. asperellum, T. atroviride, T. harzianum, and T. virensGreenhouseMoayedi and Mostowfizadeh-Ghalamfarsa (2011)
M. javanicaOliveP. fluorescens and P. putidaGreenhouseKhalighi and Khodakaramian (2012)
Fusarium solaniPotatoT. brevicompactum, T. longibrachiatum and T. asperellumIn vitro and greenhouseOmmati and Zaker (2012)
M. javanicaTomatoT. harzianumGreenhouseNaserinasab et al. (2012)
P. carotovorumPotatoPseudomonas spp.In vitro and greenhouseGhods-Alavi et al. (2012)
P. griseaRiceT. harzianumGreenhouseRaeesi et al. (2012a)
B. cinereaT. harzianumIn vitro and greenhouseRaeesi et al. (2012b)
R. solaniRiceP. fluorescens and P. aeruginosaIn vitro and greenhouseKazemzadeh et al. (2012)
Ph. sojaeSoybeanBradyrhizobium japonicum, T. spirale, T. orientale and T. brevicompactumIn vitro and greenhouseAyoubi et al. (2012)
P. aphanidermatumCucumberT. longibrachiatum and T. atrovirideGreenhouseAle Aghaee et al. (2012)
Ph. parasiticaCitrusStreptomyces sp.In vitro and greenhouseSadeghi (2012)
F. o. f.sp. lycopersiciTomatoStreptomyces sp.In vitroFadaei et al. (2012)
F. solani f.sp. pisiChickpeaT. harzianum and T. virideIn vitro and greenhouseAfrousheh et al. (2012a)
Fusarium subglutinansCucumberStreptomyces spp.In vitro Sadeghi and Hatami (2012)
F. solani f.sp. pisiPeaT. harzianum and T. virideIn vitro and greenhouseAfrousheh et al. (2012b)
Phytophthora sojaeSoybeanT. orientals, T. brevicompactum and T. spirale and Bradyrhizobium japonicumIn vitro and greenhouseNajmeh et al. (2012)
Rosellinia necatrixT. flavusIn vitroMasudi and Shahidi (2012)
P. aphanidermatumCucumberT. virens, T. harzianum, and T. atrovirideIn vitro and greenhouseHosseyni et al. (2012a)
R. solani, M. phaseolina, F. graminearum, and S. sclerotiorumT. virideIn vitroSoofi et al. (2012)
Bipolaris australiensis and B. cinereaSaffronT. virens, T. harzianum, and T. koningiiIn vitroRoohabadi et al. (2012)
F. graminearumWheatT. harzianum and T. virensFieldBaghani et al. (2012)
F. o. f.sp. radicis-cucumerinumCucumberT. harzianumIn vitro and greenhouseJavanshir Javid et al. (2012)
Monosporascus cannonballusMuskmelonT. atroviride, T. harzianum, and T. virensIn vitro and greenhouseKeshavarzi et al. (2012a)
F. graminearumWheatT. harzianum and T. virensFieldBaghani et al. (2012)
R. solani and F. solani f.sp. tuberoseSugar beetP. putidaIn vitroNazari et al. (2012)
Alternaria alternataPotatoT. viride, T. orientalis, T. arundinaceum, and T. harzianumIn vitro and greenhouseNasiri et al. (2012)
P. aphanidermatumCucumberBacillus sp. and B. subtilisGreenhouseHosseyni et al. (2012b)
F. solani and R. solaniStreptomyces sp.In vitroVasebi and Dehnad (2012)
B. cinereaAppleHanseniaspora occidentalisIn vitroAzadrooh et al. (2012)
A. flavusPistachioT. harzianum and T. longibrachiatumIn vitroChegini et al. (2012)
M. cannonballusCucumis melonT. harzianum, T. virens, T. atroviride, and Chaetomium globosumIn vitroKeshavarzi et al. (2012b)
M. javanicaPenicillium griseofulvum, Penicillium chrysogenum, and Penicillium coprophilumIn vitroKarkhaneh et al. (2012)
V. albo-atrum, F. oxysporum, and R. solaniPotatoT. flavusIn vitroNaraghi et al. (2012a)
R. solaniCommon beanStreptomyces microflavusIn vitro and greenhouseMoazenian et al. (2012b)
A. flavusPistachioT. harzianum and T. koningiiIn vitroKahnooji et al. (2012)
Ph. drechsleriPistachioT. longibrachiatum and T. harzianumIn vitroMirkhani et al. (2012)
Ph. drechsleriPistachioT. harzianumGreenhouseAlipoor Moghadam et al. (2012)
Paecilomyces variotiiPistachioStreptomyces spp.In vitroAnsari et al. (2012)
P. tolaasiiA. bisporusPseudomonas reactants, Bacillus sp., and P. fluorescensIn vitroTajalipour et al. (2012)
V. albo-atrumPotatoT. flavusGreenhouseNaraghi et al. (2012b)
B. oryzaeRiceT. harzianum, T. atroviride, and T. virensGreenhouseKhalili et al. (2012)
F. solaniBeanT. harzianum and T. virideIn vitro and greenhouseKhodaei et al. (2012)
F. solani, R. solani, F. oxysporum, Pestalotiopsis spp., C. gloeosporioides, and P. digitatumCitrusStreptomyces sp.In vitroNoorizadeh et al. (2012)
P. italicumOrangePichia kluyveriIn vitroGhasemi Sardareh et al. (2012)
R. solaniSugar beetT. harzianumIn vitroGhanbari et al. (2012)
P. expansumAppleTorulaspora delbrueckiiIn vitroEbrahimi et al. (2012a,b)
Magnaporthe oryzaeRiceT. harzianum, T. atroviride, and T. virensIn vitroJavadi et al. (2012)
F. graminearumWheatP. fluorescens, E. herbicola, B. subtilis, and B. cereusIn vitro and greenhouseAlimi et al. (2012)
S. sclerotiorumBeanB. subtilis subsp. spizizenii and Streptomyces acrimyciniIn vitro and greenhouseGholami et al. (2012)
S. sclerotiorumCucumberBacillus sp.In vitroRostami et al. (2012)
G. graminis var. triticiWheatP. fluorescensIn vitro and greenhouseLagzian et al. (2012)
F. oxysporumCucurbitT. harzianum and T. longibrachiatumIn vitroAbdolahy and Parsaiyan (2012)
F. solaniCucurbitT. koningiiIn vitroAbdolahy and Parsaiyan (2012)
V. dahliaePistachioT. harzianum and T. koningiiIn vitroJamdar et al. (2012)
Pratylenchus loosiTeaB. subtilisIn vitroRahanandeh et al. (2012)
M. javanicaCucumberP. fluorescens, B. subtilis, and Pantoea sp.GreenhouseMajzoob et al. (2012)
Meloidogyne javanicaTomatoArthrobotrys oligospora and Paecilomyces lilacinusGreenhouseJamshidnejad et al. (2012)
B. cinereaStrawberryTrichoderma spp.In vitro and greenhouseNaeimi and Zare (2013)
P. aphanidermatumSugar beetTrichoderma erinaceum, T. koningii, T. longibrachiatum, and T. harzianumGreenhouseAbdollahi et al. (2013)
Fusarium oxysporum f. sp. tuberosiPotatoTrichoderma virens and Trichoderma asperellumIn vitro and greenhouseOmmati et al. (2013)
B. sorokinianaWheatG. fasciculatum and P. fluorescensGreenhouseHashemi Alizadeh et al. (2013)
Tylenchulus semipenetransCitrusF. solani, F. oxysporum, P. lilacinus, Cladosporium cladosporioides, and Acremonium strictumGreenhouseChavoshisani et al. (2013)
Aspergillus flavusPistachioTrichoderma harzianum and Trichoderma longibrachiatumIn vitroChegini et al. (2013)
M. javanicaTomatoGlomus mosseae and G. intraradicesGreenhouseGolzari et al. (2013)
A. flavusPistachioB. subtilisIn vitroAfsharmanesh et al. (2013)
Colletotrichum lindemuthianumBeanB. subtilis subsp. subtilis, B. atrophaeus, B. tequilensis, B. subtilis subsp. spizizenii, Streptomyces cyaneofuscatus, S. flavofuscus, S. parvus, and S. acrimyciniIn vitro and greenhouseGholami et al. (2013)
P. aphanidermatumTarragonT. asperelloidesIn vitroPakdaman et al. (2013)
E. amylovoraApple, pear, and quinceP. fluorescens, P. agglomerans, P. putida, and Serratia marcescensFieldGerami et al. (2013)
F. culmorumWheatPseudomonas spp.GreenhouseMadloo et al. (2013)
P. loosiTeaP. fluorescensIn vitroRahanandeh et al. (2013)
V. dahliaeCottonB. subtilis, Bacillus coagulans, Bacillus polymyxa, and P. fluorescensGreenhouseMansoori et al. (2013)
F. graminearumWheatT. harzianum and T. virideFieldForoutan (2013)
M. phaseolinaSoybeanP. agglomerans, Bacillus sp., and T. harzianumIn vitro and greenhouseVasebi et al. (2013)
Ph. drechsleriCucumberP. fluorescensIn vitro and greenhouseGhafelebashi et al. (2014)
B. cinereaAppleB. subtilis, C. membranifaciens and M. guilliermondiiIn vitro and greenhouseZanguei et al. (2014)
M. oryzaeRiceT. harzianum, T. atroviride, and T. virensIn vitro and greenhouseJavadi et al. (2014)
F. solani f.sp. pisiPeaG. mosseae and G. intraradicesGreenhouseSoharabi et al. (2014)
F. o. f.sp. lycopersiciTomatoT. harzianum and T. virensGreenhouseJalali (2014)
P. tolaasiiMushroomP. reactants, P. putida, P. fluorescens, and B. subtilisGreenhouseTajalipour et al. (2014)
M. javanicaTomatoP. fluorescensIn vitroBagheri et al. (2014)
A. flavusPistachioB. subtilisIn vitroAfsharmanesh et al. (2014)
P. digitatumCitrusB. subtilis, Rhizobium rubi, and P. digitatumIn vitroMohammadi et al. (2014)
Cercospora beticolaSugar beetBacillus sp., Enterobacter sp., and Enterobacter sp.In vitro and greenhouseMousavi Mirak (2014)
M. phaseolinaSoybeanT. harzianumIn vitro and greenhouseKhaledi and Taheri (2014)
R. solanacearumPotatoPaenibacillus polymyxaIn vitro and greenhouseDadjoo et al. (2014)
F. o. f.sp. melonisCantaloupeB. subtilisIn vitroHosseini Haji Abdal et al. (2014b)
F. o. f.sp. melonisCantaloupeT. atroviride and T. harzianumIn vitro and greenhouseHosseini Haji Abdal et al. (2014a)
M. incognitaTeaP. aeruginosa and P. fluorescensIn vitroRahanandeh and Moshaiedy (2014)
P. chrysogenum and B. cinereaStreptomyces griseusIn vitroDanaei et al. (2014)
R. solaniCottonP. aureofaciens and P. fluorescensGreenhouseSamavat et al. (2014)
M. phaseolinaSoybeanT. harzianumFieldBarari et al. (2014)
F. sambucinum, Fusarium subglutinans, Phoma glomerata, and N. mangiferaeCucumberStreptomyces sp.In vitroSadeghi and Hatami (2014)
M. phaseolinaSoybeanP. agglomeransIn vitro and greenhouseVasebi et al. (2015)
M. javanicaTomatoTrichoderma spp.In vitro and greenhouseKavari et al. (2015)
Ph. drechsleriCucumberT. harzianumGreenhouseDelkhah and Behboudi (2015)
M. javanicaTomatoT. harzianum and P. fluorescensGreenhouseMokhtari et al. (2015)
M. javanicaTomatoMetarhizium anisopliae and T. harzianumGreenhouseKhosravi et al. (2015)
F. graminearumWheatP. fluorescensGreenhouseShahbazi et al. (2015)
P. aphanidermatumCucumberP. fluorescensGreenhouseAkbari-Moghadam et al. (2015)
G. graminis var. triticiWheatTrichoderma spp. and T. flavusIn vitroMohammadi and Ghanbari (2015)
S. cepivorumGarlicT. asperellum, T. harzianum, and T. flavusGreenhouseMahdizadehnaraghi et al. (2015)
F. solani, R. solani, and F. oxysporumBeanPseudomonas sp. and Bacillus sp.In vitro and greenhouseFaraji et al. (2015)
R. solaniCottonP. fluorescensGreenhouseAbdollahipuor et al. (2015)
F. o. f.sp. lycopersiciTomatoB. pumilusGreenhouseHeidarzadeh and Baghaee-Ravari (2015)
Meloidogyne spp.KiwifruitPseudomonas chlororaphis subsp. aureofaciens and P. fluorescensGreenhouseBashiri et al. (2015)
Fusarium solani, Rhizoctonia solani, and Fusarium oxysporumBeanPseudomonas sp. and Bacillus sp.In vitro and greenhouseFaraji et al. (2015)
V. dahliaePistachioT. harzianumGreenhouseFotoohiyan et al. (2015)
P. aphanidermatumCucumberG. mosseaeGreenhouseHosseini et al. (2016)
F. oxysporumTomatoP. fluorescensGreenhouseJamali et al. (2016)
B. cinerea, C. cladosporioides, and Aspergillus tubingensisGrapeT. harzianum and T. hamatumIn vitroDavari and Ezazi (2016)
B. cinereaStrawberryB. subtilis and B. licheniformisGreenhouseAmini et al. (2016)
Mycosphaerella rabieiChickpeaT. atroviride, T. virens, and T. atrovirideGreenhouseNaghed et al. (2016)
A. alternata, Alternaria dumosa, Alternaria tenuissima, Alternaria mimicula, Alternaria tomaticola, and C. cladosporioidesTomatoT. harzianum and T. virensGreenhouseBeydaghi et al. (2016)
M. javanicaTomatoG. mosseaeGreenhouseAzami-Sardooie et al. (2016)
F. o. f.sp. lycopersiciTomatoT. harzianumIn vitro and greenhouseBarari (2016)
M. phaseolina, R. solani, S. sclerotiorum, and F. graminearumMelon, melon, canola, and wheatT. harzianumIn vitroAbbasi et al. (2016)
M. phaseolinaSoybeanT. harzianumGreenhouseKhalili et al. (2016)
F. solani f.sp. pisiChickpeaStreptomyces antibioticusGreenhouseSoltanzadeh et al. (2016)
A. rabieiChickpeaP. putida, P. fluorescens, Mesorhizobium ciceri, and Burkholderia multivoransIn vitroAzizpour and Rouhrazi (2016)
R. solaniSugar beetBacillus amyloliquefaciens, B. pumilus, and Bacillus siamensisGreenhouseKarimi et al. (2016)
F. o. f.sp. lycopersiciTomatoB. subtilis and P. fluorescensIn vitro and greenhouseJalali et al. (2016)
A. flavusPistachioB. subtilisIn vitroFarzaneh et al. (2016)
Penicillium crustosum and A. tubingensisGrapePichia membranaefaciensIn vitro and greenhouseRanjbar Chaharborj et al. (2016)
S. sclerotiorum and P. aphanidermatumCucumberPseudomonas spp., Stenotrophomonas spp., and Flavobacterium spp.In vitro and greenhouseBagheri et al. (2016)
X. translucens pv. cerealisWheatPseudomonas spp.Greenhouse and in vitroFallahzadeh-Mamaghani et al. (2016)
F. o. f.sp. cucumerinumCucumberT. flavusGreenhouseShahriari et al. (2016)
M. javanicaTomatoT. harzianumGreenhouseHeidari and Olia (2016)
Agrobacterium tumefaciensTobaccoB. subtilisGreenhouseNazari et al. (2016)
R. solanacearumPotatoP. fluorescensIn vitro and greenhouseHasani and Khodakaramian (2016)
G. graminis var. triticiWheatB. subtilisIn vitro and greenhouseKhezri and Manafi Shabestari (2016)
F. solani f.sp. phaseoliBeanT. hamatum and P. fluorescensIn vitro and greenhouseKhosro-Anjom et al. (2016)
V. dahliaeTomatoB. subtilis, B. pumilus, B. atrophaeus, and Bacillus thuringiensisIn vitro and greenhouseSafdarpour and Khodakaramian (2016)
E. amylovoraAppleP. agglomeransIn vitroFirouzian Bandpey and Rahimian (2016)
F. oxysporumT. harzianum, T. koningii, and T. virensIn vitroHabibi and Rahnama (2016)
F. oxysporum f.sp. lycopersici and M. javanicaTomatoG. mosseaeGreenhouseAmirafzali et al. (2016)
Diplodia bulgaricaAppleArthrinium arundinis, Arthrinium saccharicola, Periconia sp., Penicillium sp., Aspergillus persii, C. globosum, Chaetomium sp., Trichothecium roseum, A. tenuissima, and Alternaria infectoriaIn vitroAlijani et al. (2016)
R. necatrixAppleB. siamensis and B. pumilusIn vitroBinandeh et al. (2016)
F. oxysporumCucumberT. harzianumIn vitroAkhlaghi et al. (2016)
P. aphanidermatumCucumberB. cereus, B. licheniformis, and Bacillus endophyticusIn vitroRezaei et al. (2016)
R. solaniWheatT. citrinovirideIn vitro and greenhouseSharify et al. (2016a)
R. solaniWheatT. asperellumIn vitro and greenhouseSharify et al. (2016b)
P. tolaasiiA. bisporusBrochothrix thermosphacta and Bacillus mycoidesIn vitroAslani et al. (2016)
B. cinereaGrapeT. harzianum and C. membranifaciensIn vitroKasfi et al. (2016a)
A. nigerGrapeT. harzianum and C. membranifaciensIn vitroKasfi et al. (2016b)
H. schachtiiSugar beetT. harzianum, Thalaromyces flavus, F. solani, and V. chlamydosporiumGreenhouseShirazi et al. (2016)
M. phaseolinaMungbeanT. harzianum, T. reesei, A. niger, and B. subtilisGreenhouseShahid and Khan (2016)
Curvularia lunata and Bipolaris sorokinianaWheatAchromobacter xylosoxidansIn vitroBagheri and Ahmadzadeh (2016)
R. solaniTomatoBeauveria bassianaGreenhouseAzadi et al. (2016)
Bipolaris victoriaeRiceT. harzianum, A. tenuissima, Fusarium verticillioides, Alternaria citri, A. infectoria, and Preussia sp.GreenhouseSafari Motlagh and Mohammadian (2016)
M. javanicaTomatoP. fluorescensGreenhouseTanha et al. (2016)
M. javanicaTomatoTalaromyces flavus and T. harzianumGreenhouseAbootorabi and Naraghi (2016)
F. solani and F. oxysporumChickpeaT. harzianum, T. asperellum, and T. virensFieldKhomeini et al. (2016)
M. phaseolinaSoybeanT. harzianumGreenhouseKhaledi and Taheri (2016)
C. beticolaSugar beetBacillus sp., Paenibacillus sp., Pseudomonas sp., and Enterobacter sp.In vitro and greenhouseArzanlou et al. (2016)
M. phaseolinaSoybeanT. harzianum, T. asperellum, and Trichoderma virensGreenhouseBarari and Foroutan (2016)
M. phaseolinaSoybeanT. harzianum and T. atrovirideIn vitro and greenhouseKia and Rahnama (2016)
F. o. f.sp. dianthiCarnationT. harzianumIn vitroKermajany et al. (2017)
V. dahliaePistachioT. harzianumGreenhouseFotoohiyan et al. (2017)
F. verticillioides, P. aphanidermatum, Penicillium sp., and V. dahliaeLactobacillus fermentum, Lactobacillus plantarum, Lactobacillus paralimentaris, Lactobacillus pentosus, Lactobacillus buchneri, and Sporolactobacillus terraeIn vitro and fieldKharazian et al. (2017)
Rhizoctonia solaniSuger beetBacillus subtilisIn vitro and greenhouseAhmadzadeh et al. (2017)
Macrophomina phaseolinaSoybeanTrichoderma harzianum, Tricchoderma reesei and Trichoderma atrovirideIn vitro and greenhouseAlamdarlou et al. (2017)
F. oxysporum, X. campestris, and E. amylovoraStreptomyces sp.In vitroShams and Shahnavaz (2017)
M. javanicaCucumberPseudomonas rhodesiae and Acinetobacter sp.In vitro and greenhouseAmini et al. (2017)
C. michiganensis subsp. insidiosusAlfalfaB. subtilis, Pseudomonas sp., Escherichia coli, Sphingomonas paucimobilis, and Paenibacillus glycanilyticusIn vitro and greenhouseOmidi Nasab and Khodakaramian (2017)
P. loosiTeaP. lilacinusGreenhouseYahyavi Azad et al. (2017)
P. syringae pv. syringaeOakPseudomonas protegens, Stenotrophomonas maltophilia, and Bacillus firmusIn vitroTashi‐Oshnoei et al. (2017)
F. o. f.sp. radicis lycopersiciTomatoStreptomyces carpaticusIn vitro and greenhouseZahed and Behbudi (2017)
A. niger and A. flavusB. cereusIn vitroMotamedi et al. (2017)
Xanthomonas oryzae pv. oryzaeRiceBacillus sp., B. subtilis, P. putida, and Enterobacter sp.GreenhouseYousefi et al. (2018)
R. solaniCommon beanB. pumilus, T. harzianum, and Rhizophagus intraradicesGreenhouseNasir Hussein et al. (2018)
P. syringae pv. syringae and P. tolaasiiPistachioPantoea brenneri, P. protegens, S. maltophilia, Bacillus anthracis, B. pumilus, and Serratia plymuthicaGreenhouseEtminani and Harighi (2018)
M. incognitaPistachioP. fluorescens, B. cereus, and B. subtilisGreenhouseZeynadini-Riseh et al. (2018)
Fusarium pseudograminearumWheatP. fluorescens, B. subtilis, and Streptomyces sp.In vitro and greenhouseNorouzian et al. (2018)
V. dahliaeTomatoBacillus spp., Pseudomonas spp., and Enterobacter spp.In vitroZendehdel et al. (2018)
H. schachtiiSugar beetT. harzianum, Talaromyces flavus, F. solani, and Pochonia chlamydosporiumGreenhouseHosini et al. (2018)
F. oxysporumCucumberT. harzianum and T. atrovirideIn vitroNosrati (2018)
P. carotovorum subsp. carotovorumPotatoBacillus spp.In vitroAbdoli et al. (2018)
S. sclerotiorumCucumberStreptomyces albidoflavusIn vitro and greenhouseEini et al. (2018)
P. loosiTeaP. lilacinus and Clonostachys roseaGreenhouseYahyavi Azad et al. (2018)
M. phaseolinaMelonB. amyloliquefaciensGreenhouseBakhshi et al. (2018)
S. sclerotiorumCucumberS. albidoflavusIn vitro and greenhouseZahed and Behbudi (2018)
F. o. f.sp. radices lycopersiciTomatoStreptomyces spp.In vitro and greenhouseVafaie et al. (2018)
F. oxysporumCucumberP. fluorescensGreenhouseSaberi-Riseh and Fathi (2018)
B. cinereaPichia galeiformis, Galactomyces candidum, M. guilliermondii, S. cerevisiae, Zygoascus meyerae, Pichia sp., Candida parapsilosis, Metschnikowia sp., Candida boidinii, Lecythophora sp., and Candida catenulataIn vitroGharaei et al. (2018)

6 Future perspectives

The climate changes and the growing human population with enhancing food demands have been considered as risks to food security in Iran. Different beneficial population of microbes can positively affect agricultural farms, such as pollutant degradation, productivity, and decrease in disease and pest. Maintaining and improving the performance of these microbes can increase agricultural products in different geographical area of Iran with various climates. More studies are needed to reach these aims in Iran. We need to engineer microbes in different parts and various population of plants. We can use beneficial microbes as bioformulation or other methods of treatments on plants. Since the roots are in contact with soil and rhizosphere has high biodiversity, these are the most important parts in plants for engineering the microbes. Therefore, soil improvement with beneficial microorganisms will be able to help farmers and agricultural ecosystems in the future years.

  1. Conflict of interest: Authors declare no conflict of interest.

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Received: 2019-10-27
Revised: 2020-04-19
Accepted: 2020-05-18
Published Online: 2020-08-03

© 2020 Mehrdad Alizadeh et al., published by De Gruyter

This work is licensed under the Creative Commons Attribution 4.0 International License.

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