Actinomycetes: an unexplored microorganisms for plant growth promotion and biocontrol in vegetable crops

Abstract

Actinomycetes, a Gram positive bacteria, well reported as a source of antibiotics, also possess potential to control various plant pathogens, besides acting as plant growth promoting agent. Chemicals in different forms are extensively being used in vegetable farming, adversely affecting the environment and consumer health. Microbial agent like actinomycetes can substantially replace these harmful chemicals, and have now started finding a place as an important input in to farming practices. Only selected vegetable crops belonging to 11 different families have been explored with use of actinomycetes as biocontrol and plant growth promoting agent till now. It provides ample opportunities to vegetable researchers, to further explore with use of this very important group of microorganisms, in order to achieve even higher production level of safe vegetables. Mycostop and Actinovate are two actinomycetes based formulations globally available for use in vegetable farming as a substitute for chemical formulations. Present review article has summarized the literature available on use of actinomycetes in vegetable farming. Existing wide gap in knowledge, and potential thrust areas for future research have also been projected.

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Vol.:(0123456789)
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World Journal of Microbiology and Biotechnology (2018) 34:132
https://doi.org/10.1007/s11274-018-2517-5
REVIEW
Actinomycetes: anunexplored microorganisms forplant growth
promotion andbiocontrol invegetable crops
A.Chaurasia1 · B.R.Meena1· A.N.Tripathi1· K.K.Pandey1· A.B.Rai1· B.Singh1
Received: 27 May 2018 / Accepted: 9 August 2018 / Published online: 13 August 2018
© Springer Nature B.V. 2018
Abstract
Actinomycetes, a Gram positive bacteria, well reported as a source of antibiotics, also possess potential to control various
plant pathogens, besides acting as plant growth promoting agent. Chemicals in different forms are extensively being used in
vegetable farming, adversely affecting the environment and consumer health. Microbial agent like actinomycetes can sub-
stantially replace these harmful chemicals, and have now started finding a place as an important input in to farming practices.
Only selected vegetable crops belonging to 11 different families have been explored with use of actinomycetes as biocontrol
and plant growth promoting agent till now. It provides ample opportunities to vegetable researchers, to further explore with
use of this very important group of microorganisms, in order to achieve even higher production level of safe vegetables.
Mycostop and Actinovate are two actinomycetes based formulations globally available for use in vegetable farming as a
substitute for chemical formulations. Present review article has summarized the literature available on use of actinomycetes
in vegetable farming. Existing wide gap in knowledge, and potential thrust areas for future research have also been projected.
Keywords Actinomycetes· Biocontrol· Plant growth promotion· Vegetable
Abbreviations
ACC 1-Aminocyclopropane-1-carboxylate
IAA Indole actetic acid
MPPF Mycelia preparation of pathogenic fungi
PGP Plant growth promotion
DAT Day after transplantation
CFU Colony forming unit
RHB Rhizobia helper bacteria
KI Kinetin
Introduction
Actinomycetes, a Gram positive bacteria, and a major source
of most of the antibiotics being used, are also being explored
as plant growth promoting and biocontrol agent for various
crop plants. Either the organism itself as bio-formulation
or metabolites isolated from it, and/or its derivatives are
being used for this purposes. When compared with other
groups of microorganism, actinomycetes are relatively less
explored with agricultural perspective in general and veg-
etable in particular. Though vegetable crops play a major
role in achieving nutritional security in developing country
like India, apart from enhancing the per capita income of the
farmers, usually have problems of disease, insect, and pest.
To control it, and to enhance vegetables productivity chemi-
cals in form of fertilizer, pesticide, and herbicide are being
used extensively, which have led to several concerns regard-
ing environment and human health including ever increasing
cost of production, besides affecting vegetables export busi-
ness potential in international market (PIB 2015; Sinha etal.
2012). To overcome these problems, several eco-friendly
approaches including various biological formulations have
been developed and being used to replace or used in combi-
nation of these chemicals. Among the 18 major lineages rec-
ognized in bacterial domain as per Bergey’s Manual of Sys-
tematic Bacteriology (Whitman etal. 2012), actinomycetes
also known as actinobacteria is one of the largest taxonomic
unit comprising of five subclases, six orders, and 14 subor-
ders (Ludwig etal. 2012). Among different genera Strepto-
myces is extensively researched one as it is comparatively
easy to isolate. Genera difficult to isolate were termed as non
streptomyces actinomycetes or rare actinomycetes. With the
discovery of more and more specific media, researchers have
now revealed that rare actinomycetes are not in fact rare, but
* A. Chaurasia
anurag_vns1@yahoo.co.in
1 Division ofCrop Protection, ICAR-Indian Institute
ofVegetable Research, Varanasi, UttarPradesh, India

World Journal of Microbiology and Biotechnology (2018) 34:132
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132 Page 2 of 16
as widely available as Streptomyces genus. This group of
microorganisms is also a source of bio-active metabolites,
and can degrade a wide range of recalcitrant compounds,
so has been recommended as a best bio-degrader (Pizzul
etal. 2006; Eshelli etal. 2015). They can degrade differ-
ent types of pesticide (Lin etal. 2011; Fuentes etal. 2004),
herbicide (Esposito etal. 1998; Arya etal. 2016), insecti-
cide (Polti etal. 2014), besides acting as biopesticide and
bioherbicide agent (Zhang etal. 1998; Danasekaran etal.
2010; De Schrijver and De Mot 1999). Actinomycetes strains
have also been developed as eukaryotic expression vector
for bulk production of eukaryotic proteins (Nakashima
etal. 2005). Though many reviews are available on different
aspects of actinomycetes like as a source of novel bio-active
compounds and natural products, as a storehouse and dis-
covery of next generation of antibiotics (Barka etal. 2016;
Matsumoto and Takahashi 2017; Jose and Jha 2016; Genil-
loud 2017; Dinesh etal. 2017; Genilloud etal. 2011), and
as biocontrol and plant growth promoting agent in general
(Vurukonda etal. 2018), but information on use of actino-
mycetes as biocontrol and plant growth promoting agent
particularly for vegetable crops is by and large not avail-
able at one place. Keeping these literature constrains in view
and for the benefit of vegetable researchers effort has been
made to review and compile information pertaining to use
of actinomycetes for plant growth promotion and biocontrol
in vegetable crops.
Actinomycetes asbiocontrol andplant
growth promoting agent forvegetable crops
Solanaceous vegetables
Researchers have mainly focused on tomato and chilli while
exploring actinomycetes applicability for solanaceous veg-
etables (Table1). Endophytic Streptomycetes strain S30 iso-
lated from surface sterilized tomato root interior promoted
growth along with enhanced resistance to R. solani in tomato
but not so in cucumber seedlings (Cao etal. 2004). Further-
more endophytic Streptomycetes spp. isolated from healthy
tomato root was also reported to act as biocontrol agent
against Ralstonia solanacearum causing wilt of tomato
under invitro condition (Tan etal. 2006), and also for both
plant growth promotion and management of tomato bacterial
wilt under in-vivo condition too (Sreeja and Gopal 2013). In
contrast endophytic Streptomycetes strain DHV3-2 isolated
from root of diseased tomato plant showed potent biocon-
trol effect against Verticillium dahliae under both in-vitro
and in-vivo condition besides enhancing plant growth (Cao
etal. 2016). Therefore it is recommended that researchers
should target both healthy and diseased plants while search-
ing for potent endophytic actinomycetes strain. Endophytic
actinomycetes belonging to both streptomycetes and non-
streptomycetes genera isolated from roots of healthy native
plants Aristida pungens, Cleome arabica, Solanum nigrum,
Panicum turgidum, Astragallus armatus, Peganum harmala,
Hammada scoparia and Euphorbia helioscopia of Algerian
Sahara region were evaluated for their biocontrol potential
against Rhizoctonia solani damping-off and plant growth
promoting traits. Two strains CA-2 (Streptomyces mutabi-
lis NBRC 12800T) and AA-2 (Streptomyces cyaneofusca-
tus JCM 4364T) reduced the disease incidence which was
comparable to thioperoxydicarbonic diamide tetramethyl-
thiram (TMTD) treatment besides increasing fresh weight,
seedling and root length of tomato (Goudjal etal. 2014).
Earlier they have also isolated endophytic actinomycetes
from five different herbaceous plants (Cleome arabica,
Solanum nigrum, Astragallus armatus, Aristida pungens
and Panicum turgidum) collected from Hassi R’mel region
of Algerian Sahara, and evaluated for indole-3-acetic acid
(IAA) production. Tomato seeds treated with culture super-
natant of Streptomycetes sp. PT2 strain [isolated from Pani-
cum turgidum] containing crude IAA [100.03 ± 0.34µg/ml]
promoted seed germination and root elongation (Goudjal
etal. 2013). Endophytic Micromonospora spp. isolated
from tomato inhibited Fusarium oxysporum f. lycopersici
(Smith 1957). Micromonospora spp. isolated from sur-
face sterilized nodules of healthy alfalfa leguminous plant
minimized pathogenic fungi Botrytis cinerea leaf infection
when inoculated in tomato root. Jasmonates led induced sys-
temic resistance played a key role in defense process as JA-
deficient tomato line def1 did not show long term induced
resistance by Micromonospora inoculation. These strains
can be used both for priming the plant immunity and also as
antifungal agent against plant pathogens (Martinez-Hidalgo
etal. 2015). Thus potent actinomycetes strain obtained from
one plant can also be applied to inhibit pathogen of another
plant. Usually potent actinomycetes strain inhibits particular
pathogen infecting many different crops, possible because
their mode of action remain same. But same strain may also
inhibits different pathogens affecting different crops due to
its broad spectrum of activity as reported by Kunova etal.
(2016), and elaborated subsequently. Streptomyces vinaceus
strain St24 isolated from root-stem junction of tomato plant
was reported with acaricidal activity, and on foliar spray
protected tomato plants against Botrytis cinerea causing gray
mold disease (Wang etal. 2012). Streptomyces sp. strain
DBT204 isolated from tomato showed plant growth promot-
ing traits in seedlings of both chilli and tomato. Genes iaaM
and acdS were amplified, phytohormones like indole acetic
acid, kinetin, and six antibiotics were detected and quanti-
fied (Passari etal. 2016). They have also reported endo-
phytic Streptomyces sp. strain BPSAC 34 and Leifsonia xyli
strain BPSAC24 isolated from ethanomedicinal plants of
Mizoram India which led to maximum increase in shoot and

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Table 1 Actinomycetes application in Solanaceous vegetables
Crops/vegetables Source of isolation Actinomycetes Trait(s) Pathogen(s) References
Chilli (Capsicum annuum L.) Medicinal plants Streptomyces sp. (BPSAC34)
Leifsonia xyli (BPSAC24)
PGP and biocontrol Plant-pathogens, i.e. Rhizoc-
tonia solani (MTCC-9666),
Fusarium graminearum
(MTCC-1893) and Fusarium
oxysporum (MTCC-284)
Passari etal. (2015)
Capsicum annuum L. Rhizospheric soil Actinomycetes strains OUA3,
OUA5, OUA18, and OUA40
Biocontrol Colletotrichum capsici and
Fusarium oxysporum Ashokvardhan etal. (2014)
Chilli Probiotic bacteria Actinomycetes sp. (ATS6) Biocontrol, Improve chilli
seed quality/prevent seed
borne disease
Colletotrichum acutatum Tefa etal. (2015)
Chilli Rhizospheric soil Actinomycetes PACCH 277,
PACCH129, PACCH225,
PACCH24 and PACCH246,
Streptomyces hygroscopicus
Biocontrol and PGP, control
stem rot diseases
Sclerotium rolfsii Pattanapipitpaisal and Kam-
landharn (2012)
Chilli Chilli rhizosphere Streptomycetes indiaensis
KJ872546
Wilt causing fungal pathogen
i.e. Fusarium oxysporum Fusarium oxysporum Jalaluldeen etal. (2014)
Chilli Culture collection of Chiang
Mai University, Thiland
Streptomyces sp. Biocontrol Fusarium oxysporum f.sp.
capsici isolate FoC4
Saengnak etal. (2013)
Red Chilli Soil samples Streptomycetes ambofaciens
S2
Biocontrol, causal agent for
anthracnose
Colletotrichum gloeospori-
oides Heng etal. (2015)
Chilli pepper Antibiotic derived from
Streptomyces sp. AG-P 1441
(AG-P 1441)
Paromomycin, antibiotic
derived from Streptomyces
sp. AG-P 1441 (AG-P 1441)
Phytophthora blight and soft
rot diseases
Phytophthora capsici and Pec-
tobacterium carotovorum Balaraju etal. (2016)
Red peppers Soil Streptomyces halstedii AJ-7 Biocontrol of phytophthora
blight in red-peppers
Phytophthora capsici Joo (2005)
Tomato Endophytic Streptomyces sp. Bacterial wilt in tomato Ralstonia solanacearum Sreeja and Gopal (2013)
Tomato Rhizospheric soil Streptomycetes sp. PGP by producing ACC
deaminase, IAA
– El-Tarabily (2008)
Tomato Roots of healthy native plants
of Algerian Sahara
Streptomyces sp. PGP and biocontrol Rhizoctonia solani damping-
off of tomato seedlings
Goudjal etal. (2014)
Tomato Endophytes from root Streptomyces sp. promoted growth and
enhanced resistance to R.
solani in tomato seedlings
Rhizoctonia solani Cao etal. (2004)
Tomato Endophytes from root Streptomyces sp. Biocontrol Ralstonia solanacearum Tan etal. (2006)
Tomato Endophytes from herbaceous
plants (Cleome arabica,
Solanum nigrum, Astragallus
armatus, Aristida pungens
and Panicum turgidum)
Streptomyces sp. PGP through IAA, promoting
seed germination and root
elongation
– Goudjal etal. (2013)
Tomato Endophytes Micromonospora sp. Wilt Fusarium oxysporum f. lyco-
persici Smith (1957)

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Table 1 (continued)
Crops/vegetables Source of isolation Actinomycetes Trait(s) Pathogen(s) References
Tomato Nodules of alfalfa Micromonospora sp. Control fungal pathogen and
stimulate plant immunity
Leaf infection by Botrytis
cinerea Martínez-Hidalgo etal. (2015)
Tomato diverse Greek habitats/Rhizos-
pheric soil
Streptomyces rochei
ACTA1551
protect tomato seeds from
F. oxysporum infection
invivo, promote the growth
of tomato plants when the
pathogen was absent
Fusarium oxysporum f.sp.
lycopersici Kanini etal. (2013)
Tomato Polyene antibiotics strevertene
A strevertene B from culture
exrtact of S. psammoticus
Streptomyces psammoticus
strain KP1404
Fusarium wilt of tomato Alternaria mali, Aspergil-
lus oryzae, Cylindrocarpon
destructans, Colletotri-
chum orbiculare, Fusarium
oxysporum f.sp. lycopersici
and Sclerotinia sclerotiorum
Kim etal. (2011)
Tomato Endophytes from root of
diseased tomato plant
Streptomyces sp. strain
DHV3-2
biocontrol of fungal diseases
caused by Verticillium dahl-
iae and PGP
Verticillium dahliae Cao etal. (2016)
Tomato root-stem junction of tomato
plant
Streptomyces vinaceus St24 Biocontrol of gray mold
disease
Botrytis cinerea Wang etal. (2012)
Tomato Culture collection Bacillus thuringiensis CR-371,
Streptomyces avermectinius
NBRC14893
Bacterial wilt, root-knot
nematod
Soilborne pathogens Ralstonia
solanacearum, Meloidogyne
incognita
Elsharkawy etal. (2015)
Tomato Culture collection Saccharothrix algeriensis
NRRL B-24137
Tomato wilt Fusarium oxysporum f.sp.
lycopersici (FOLy)
Merrouche etal. (2017)
Tomato, Chilli Termites Streptomyces rubrolavendulae
S4
Biocontrol Phytophthora infestans, caus-
ing damping off disease in
nurseries
Loliam etal. (2012)
Tomato, Chilli Endophytes from tomato Streptomycessp. strain
DBT204
PGP [phytohormones IAA,
KI]
– Passari etal. (2016)
Cultivated rocket, Lamb, Let-
tuce and Tomato
Endophytic Streptomycetes
spp. available at Culture
Collection, Università degli
Studi di Milano, Italy
S. cyaneus ZEA17I, S. anula-
tus CMJ58I and S. albidofla-
vus VT111I
Biocontrol and/or PGP Soil borne fungal pathogens
Sclerotinia sclerotiorum
FW361, Rhizoctonia solani
FW408, Fusarium oxyspo-
rum f.sp. lactucae L74,
Pythium ultimum FW407,
Phytophthora sp. FW409
and Thielaviopsis basicola
FW406
Kunova etal. (2016)

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Page 5 of 16 132
root length of chilli plant when used as a mixture (Passari
etal. 2015). Endophytic Streptomycetes strains S. cyaneus
ZEA17I, S. anulatus CMJ58I and S. albidoflavus VT111I,
[from microbial collection of Department of Food, Environ-
mental and Nutritional Sciences (DeFENS), University of
Milan, Italy] showed broad-spectrum to specific inhibition
against six soil borne pathogens Sclerotinia sclerotiorum
FW361, Rhizoctonia solani FW408, Fusarium oxysporum
f.sp. lactucae L74, Pythium ultimum FW407, Phytophthora
sp. FW409 and Thielaviopsis basicola FW406. These strains
were used to colonize seeds of vegetable crops like tomato,
cultivated rocket, lamb lettuce, and lettuce. Invitro experi-
ment showed improved radical and hypocotyl growth in
tomato seedlings. Occasional negative influence on plant
growth was overcome during further development of plant
(Kunova etal. 2016).
Rhizosphere competent 1-aminocyclopropane-1-carbox-
ylic acid deaminase- producing Streptomycetes filipinensis
no.15 and S. atrovirens no.26 strains isolated from tomato
rhizospheric soil exhibited plant growth promoting traits.
Streptomycetes filipinensis no.15 was superior in PGP per-
formance on tomato compared with S. atrovirens no.26, as
it also produced IAA along with ACC deaminase activity
which enhanced plant growth by reducing endogenous eth-
ylene through in planta ACC (El-Tarabily 2008). Strepto-
myces rochei ACTA1551, isolated from rhizospheric soil
of plant Pinus brutia of Greek, protected tomato seeds from
Fusarium oxysporum f.sp. lycopersici infection invivo,
and promoted tomato plant growth in absence of pathogen
(Kanini etal. 2013). Strevertenes A and B isolated from
culture extract of Streptomyces psammoticus strain KP1404
retarded Fusarium wilt development on tomato compara-
ble to that of benomyl (Kim etal. 2011). Bacterial wilt and
root-knot nematode was suppressed when tomato root was
treated with both Bacillus thuringiensis CR-371 and Strem-
ptomyces avermectinius NBRC 14893 strains (Elsharkawy
etal. 2015). Soil pretreatment with actinobacteria Saccharo-
thrix algeriensis NRRL B-24137 (SA) significantly reduced
tomato wilt caused by F. oxysporum f.sp. lycopersici (Mer-
rouche etal. 2017). Colonization of tomato and chili seed-
lings with Streptomyces rubrolavendulae S4 strain isolated
from termite mounds considerably increased seedlings sur-
vival in Phytophthora infestans contaminated peat moss,
comparable to fungicide metalaxyl, and as such has been
recommended against damping off disease in tomato and
chili nursery (Loliam etal. 2012).
Actinomycetes strains isolated from chilli rhizospheric
soils manifested antifungal activity against Colletotrichum
capsici and Fusarium oxysporum along with production of
chitinases and other hydrolytic enzymes emphasizing their
use as biocontrol agent (Ashokvardhan etal. 2014). Chilli
seeds coated with probiotic bacteria including actinomy-
cetes exhibited antagonistic activity against Colletotrichum
acutatum to prevent seed borne diseases besides increas-
ing seed viability (Tefa etal. 2015). Actinomycetes strain
PACCH24 (Streptomyces hygroscopicus) inhibited Sclero-
tium rolfsii, and showed biocontrol activity on chilli seed-
lings, and can be used to control stem rot disease and for
better growth of chilli (Pattanapipitpaisal and Kamlandharn
2012). Streptomycetes indiaensis strain KJ872546 isolated
from chilli rhizospheric soil was reported to be antagonistic
against wilt pathogen (Fusarium oxysporum) as it produces
hydrogen cyanide, volatile and diffusible antifungal metabo-
lites, and ammonia (Jalaluldeen etal. 2014). Streptomyces
spp. strain NSP4 from culture collection of Chiang Mai
University, Thiland reduced Fusarium wilt disease (Fusar-
ium oxysporum f.sp. capsici isolate FoC4) in green house
(Saengnak etal. 2013). Streptomyces ambofaciens strain S2
was reported effective against Colletotrichum gloeospori-
oides, causing anthracnose disease in red chilli fruits, and
showed no sign of infection when its ethyl acetate extract
was sprayed on chilli fruit. (Heng etal. 2015). Chili peeper
plants treated with aminoglycoside paromomycin, derived
from Streptomyces sp. AG-P 1441 (AG-P 1441) controlled
Phytophthora blight and soft rot diseases caused by Phy-
tophthora capsici and Pectobacterium carotovorum respec-
tively (Balaraju etal. 2016). Red-pepper seeds soaked in
culture broth of soil isolate Streptomyces halstedii strain
AJ-7 reduced growth of soil inoculated Phytophthora capsici
causing Phytophthora blight in red-pepper (Joo 2005). Only
two actinomycetes genera (Streptomyces and Micromonos-
pora) have been used for solanaceous vegetables till date,
leaving much scope for researchers to explore with majority
of left out genera of this group of microorganisms. While
on other side potato and eggplant belonging to vegetable
genera Solanum of this family, Indian nightshade (Solanum
indicum), ground cherry (Physalis pubescens), and many
other cultivated solanaceous vegetables still need research-
ers attention from actinomycetes applicability point of view.
Cucurbitaceous vegetable crops
Cucumber and melon are two major cucurbitaceous veg-
etable have been explored with actinomycetes applica-
tion (Table2). Glucanase producing three cucumber root
endophytic actinomycetes i.e. Actinoplanes campanula-
tus, Micromonospora chalcea, Streptomyces spiralis when
applied in combination were found as effective as metal-
axyl a recommended fungicide for Pythium disease. These
actinomycetes strains successfully reduced damping-off, and
crown and root rot of cucumber by suppressing pathogenic
activities of P. aphanidermatum on seedling and mature
cucumber plant besides promoting plant growth (El-Tarabi-
lyet etal. 2009). Rhizosphere competent non-streptomycete
actinomycetes (Actinoplanes philippinensis Couch, Micro-
bispora rosea Nonomura and Ohara, Micromonospora

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Table 2 Actinomycetes application in Cucurbitaceous vegetables
Crops/vegetables Source of isolation Actinomycetes Trait(s) Pathogen(s) References
Cucumber Root Actinoplanes campanulatus,
Micromonospora chalcea, Strep-
tomyces spiralis
PGP and biocontrol Pythium aphanidermatum El-Tarabily etal. (2009)
Cucumber Cucumber phyllosphere Actinomycete XN-1 Biocontrol Corynespora cassiicola Wang and Ma (2010–2011)
Cucumber Rhizospheric soil Streptomyces sp. Biocontrol Root pathogen Pythium aphani-
dermatum Postma etal. (2005)
Cucumber Cucumber rhizosphere soil Actinoplanes philippinen-
sisCouch,Microbispora
roseaNonomura and
Ohara,Micromonospora chalcea
Post emergence damping-off of
cucumber
Pythium aphanidermatum El-Tarabily (2006)
Cucumber From various habitats in China Streptomyces rimosus Fusarium wilt of cucumber, bio-
control and PGP
Fusarium oxysporum f.sp. cuc-
umerinum Lu etal. (2016)
Cucumber Soil Streptomyces bikiniensis HD-087 Cucumber Fusarium Wilt Fusarium oxysporum f.sp. cuc-
umerinum Zhao etal. (2012)
Cucumber Endophytes from medicinal plant
Alisma orientale Bioactive compound staurosporine
from strain CNS-42
PGP and biocontrol F. oxysporum f.sp. cucumerinum Li etal. (2014)
Cucumber Soil and compost Paenibacillus sp. 300 and Strepto-
myces sp. 385
Fusarium wilt of cucumber
(Cucumis sativus) caused by
Fusarium oxysporum f.sp. cuc-
umerinum
Fusarium oxysporum f.sp. cuc-
umerinum Singh etal. (1999)
Cucumber Culture collection Streptomyces albospi-
nusCT205alone or combined
with organic fertiliser (BOF-
CT205)
Cucumber Fusarium wilt Fusarium oxysporum Wang etal. (2016)
Cucumber Endophytes from cucumber
(Cucumis sativus) and pumpkin
(Cucurbita moschata)
Streptomycetes sp. MBCu-56 Cucumber anthracnose Colletotrichum orbiculare Masafumi etal. (2009)
Cucurbits Soil Streptomyces sp. Biocontrol Cucurbit plant pathogens Fusar-
ium sp./Alternaria sp.
Zhao etal. (2013)
Cucumber Soil samples Actinoplanes sp. HBDN08
Antifungal compound-
5-hydroxyl-5-methyl-2-hexenoic
acid
Antifungal activity Cucumber-B. cinerea [cucumber
gray mold (CGM)], C. cucumer-
inum [cucumber scab (CS)], C.
cassiicola [cucumber target leaf
spot (CTLS)]
Zhang etal. (2010)
Melons Cucurbit fruits Actinobacteria Biocontrol Pseudomonas syringae pv. lach-
rymans Glassner etal. (2015)
Melon Capsicum crop Streptomyces sp. Biocontrol Macrophomina phaseolina Etebarian (2006)

World Journal of Microbiology and Biotechnology (2018) 34:132
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Page 7 of 16 132
chalcea (Foulerton) Ørskov, and Streptomyces griseoloalbus
(Kudrina) Pridham etal.) isolated from cucumber rhizos-
pheric soil, and producing cell wall degrading enzymes,
were found to suppress Pythium aphanidermatum causing
damping-off of cucumber seedlings as comparable to meta-
laxyl (El-Tarabily 2006).
Actinomycete strain XN-1 isolated from cucumber phyl-
losphere is reported to inhibit the growth of Corynespora
cassiicola causing cucumber target leaf spot in greenhouse
(Wang and Ma 2010–2011). Filamentous actinomycetes
isolated from unsterilized reused rockwool successfully
suppressed Pythium aphanidermatum causing root rot in
cucumber grown on rockwool (Postma etal. 2005). Strepto-
myces rimosus Strain M527 isolated from China, promoted
cucumber shoot growth, and prevented wilt disease caused
by F. oxysporum f.sp. cucumerinum with 72.1% control effi-
cacy in cucumber seedlings when root was irrigated with
broth of this strain (Lu etal. 2016). Similarly fermentation
broth of Streptomyces bikiniensis strain HD-087 isolated
from China grassland soil, effectively suppressed cucumber
wilt caused by F. oxysporum f.sp. cucumerinum by inhibiting
conidia germination and destroying its membrane, leading
to cytoplasmic leakage and also triggered induced resist-
ance through increase in enzymes like peroxidase, pheny-
lalanine ammonia-lyase, and beta-1,3 glucanase activity in
cucumber. (Zhao etal. 2012). While Singh PP etal. (1999)
obtained best (p < 0.05) result when mixture of two chitino-
lytic bacterial strains Paenibacillus sp. 300 and Streptomyces
sp. 385 were used in 1:1 or 4:1 ratio, and applied as zeolite
using chitosan based media at the rate of 6g/kg of potting
medium 15days before planting cucumber seeds. Hydro-
lytic enzymes like chitinase and beta-1,3-gluccanase were
reported to play significant role in biocontrol process. Endo-
phytic strain CNS-42 isolated from Chinese medicinal plant
Alisma orientale showed largest zone of inhibition against
soil borne pathogen F. oxysporum f.sp. cucumerinum, and
significantly reduced (p < 0.05) disease severity index, and
also increased fresh shoot weight and height of cucumber
plantlets when treated with this strain. Staurosporine was
reported to be responsible for its biocontrol activity, and
it was also first time reported to be responsible for plant
growth promoting activities (Li etal. 2014). Streptomyces
albospinus CT205 in combination with organic fertilizer
(BOF-CT205) significantly reduced Fusarium wilt incidence
by reducing ca. 55%, besides enhancing cucumber yield to
8.3 × 104kg/ha (Wang etal. 2016). Endophytic Streptomyces
sp. MBCu-56 isolated from cucumber effectively controlled
cucumber anthracnose caused by Colletotrichum orbicu-
lare in seedlings (Masafumi etal. 2009). Intertwining and
degradation of fungal hypha of different Fusarium sp. and
Alternaria sp. were observed when antagonistic Streptomy-
ces sp. were grown in media containing mycelia prepara-
tion of pathogenic fungi (MPPF) of cucurbit plants as soul
carbon source because of enhanced production of extracel-
lular enzymes like cellulase, chitinase, and glucanase (Zhao
etal. 2013). Antifungal metabolite 5-hydroxyl-5-methyl-
2-hexenoic acid isolated from Actinoplanes sp. HBDN08,
a Chinese soil isolate, effectively controlled cucumber
pathogens like B. cinerea [cucumber gray mold (CGM)],
C. cucumerinum [cucumber scab (CS)], and C. cassiicola
[cucumber target leaf spot (CTLS)] at 71.42%, 78.63% and
65.13% when applied at 350mg/L in greenhouse experiment
(Zhang etal. 2010).
Endophytic bacteria like Alpha, Beta, and Gammapro-
teobacteria, Firmicutes, Actinobacteria (Streptomyces spp.)
isolated from mesocarp of Cucumis melo Reticulatus Group
‘Dulce’ fruit showed antimicrobial activity against major
cucurbit pathogens like Macrophomina phaseolina (Mac),
Fusarium oxysporum f.sp. melonis races 1 and 2 (FOM 1 and
FOM 2, respectively), F. oxysporum f.sp. radicis-cucumer-
inum (Forc) and Pseudomonas syringae (P.s.). Furthermore
these endophytes not only improved growth and protected
plant in field but also showed promising post harvest effect
by increasing fruit shelf life through delayed softening and
spoilage (Glassner etal. 2015). Streptomyces strains A22,
A20, A15, and STL obtained from glasshouse crop of cap-
sicum inhibited growth of Macrophomina phaseolina caus-
ing charcoal stem rot of melon in soil and in seed treatment
experiments (Etebarian 2006). Thus it is advisable that while
searching for competent endophytic actinomycetes strain
particularly for cucurbitaceous vegetable crops, besides root
researchers should also target other plant parts like leaves
and fruits. In consortium approach, compatibility of actino-
mycetes with other groups of microorganism, organic and/
or chemical fertilizer should also be explored. So far only
four genus of actinomycetes (Streptomyces, Actinoplanes,
Micromonospora, and Microbispora) have been used just
for two cucurbitaceous vegetables, leaving much scope to
work with rest of actinomycetes genera for many left out
important cucurbitaceous vegetables like bitter gourd, bot-
tle gourd, pointed gourd, ash gourd, watermelon, pumpkin
and squashes.
Brassicaceae orCruciferous vegetables
Cabbage, radish, cauliflower, and lettuce are major
cruciferous vegetable crops researched with actinomy-
cetes application (Table3). Cabbage endophytic strain
MBCN152-1 identified as Streptomyces humidus con-
trolled damping-off disease caused by seed-borneAlter-
naria brassicicola when artificially pathogen infested
cabbage seeds were sown in 1.5 × 107 MBCN152-1 spores
per gram of soil mix in green house experiment (Has-
san etal. 2017). Lee etal. (2008) also isolated endo-
phytic actinomycetes identified as Microbispora rosea
subsp. rosea (strain A004 and A011) and Streptomyces

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Table 3 Application of actinomycetes in Brassicaceae or Cruciferous vegetables
Crops/vegetables Source of isolation Actinomycetes Trait(s) Pathogen(s) References
1. Cabbage Endophytes from cabbage Strain MBCN152-1 -identified
as a Streptomyces humidus-
related species
Biocontrol Alternaria brassicicola Hassan etal. (2017)
2. Chinese cabbage Root Microbispora, Streptomyces,
Micromonospora Biocontrol Plasmodiophora brassicae Lee etal. (2008)
3. Cabbage seeds Culture collection (culture
filtrate of S. padanus strain
PMS-702)
Streptomyces padanus strain
PMS-702
Damping-off of cabbage Rhizoctonia solani AG-4 Shih etal. (2003)
4. Radish (Raphanus sativus) Soil Streptomyces hydrogenans
DH16
Biocontrol of seed borne
pathogens, and as spray to
control black leaf spot of
crucifers
Alternaria brassicicola, causal
agent of black leaf spot and
damping off of seedlings of
crucifers
Manhas and Kaur (2016)
5. Radish seedlings Endophytes in cacao fruits and
seeds
Streptomyces violaceusniger
clade
PGP and Biocontrol P. megakaria
P.erythroseptica
F. oxysporum
B. cinerea
A. tumefaciens
S. scabiei
Tchinda etal. (2016)
6. Cauliflower, cabbage, lettuce Finnish Sphagnum peat Mycostop (Streptomycetes
griseoviridis)
Biocontrol of seedborne and
soil borne fungal pathogens
Alternaria brassicicola on
cauliflower and cabbage, and
Botrytis cinerea on lettuce
White etal. (1990)

World Journal of Microbiology and Biotechnology (2018) 34:132
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Page 9 of 16 132
olivochromogenes (strain A018) from Chinese cabbage
root which effectively suppressed clubroot disease caused
by Plasmodiophora brassicae. Control value of 58% and
33% for strain A004 and A001, and 42% for A018 were
reported when germinated Chinese cabbage seeds were
inoculated with these actinomycetes strains, and trans-
planted to P. brassicae post inoculated pots. Cabbage
seeds treatment with culture filtrate of Streptomyces
padanus strain PMS-702 significantly reduced damping-
off disease of cabbage by antagonizing the causative
organism Rhizoctonia solani AG-4. This was the first
report of polyene macrolide fungichromin acting as active
ingredient from S. padanus filtrate controlling Rhizocto-
nia damping-off disease of cabbage (Shih etal. 2003).
Endophytes belonging to Streptomyces violaceusniger
clade isolated from cacao fruits and seeds showed plant
growth promoting ability on radish seedlings inocula-
tion (Tchinda etal. 2016). Several members of this clade
are well reported as biocontrol agent, and as antibiotic
geldanamycin producer. Soil isolate Streptomyces hydro-
genans DH16 (GanBank:JX123130) and its metabolite
significantly controlled Alternaria brassicicola causing
black leaf spot seedlings damping off of crucifers. Damp-
ing off of Raphanus sativus seedlings were controlled,
seed germination (75–80%) and vigor index (1167–1538)
improved on dressing seeds by S. hydrogenans DH16 and
its culture supernatant (10%). Manhas and Kaur (2016)
concluded its application as biofungicide to control seed
borne pathogens and black leaf spot of crucifers by seed
dressing and spray respectively. Alternaria brassicicola
on cauliflower and cabbage, and Botrytis cinerea on let-
tuce were controlled by Streptomyces griseoviridis iso-
lated from Finnish Sphagnum peat (White etal. 1990).
Only two genera (Streptomyces, Microbispora) of actino-
mycetes have been used for four cruciferous vegetables,
giving plenty of opportunities to vegetable researchers
for future research.
Amaranthaceae vegetables
Sugar beet (Beta vulgaris) is only vegetable crop of the
Amarathaceae family which has been extensively researched
for actinomycetes applicability (Table4). Endophytic actino-
mycetes identified as Streptomyces griseofuscus and Strep-
tomyces globisporus along with other bacterial and fungal
strains isolated from root and leaves of healthy sugar beet
cultivars showed plant growth promoting traits (Shi etal.
2009). In addition it was recommended to be applied to
tissue culture plantlets at nursery stage before field trans-
plantation for better establishment of endophytes. Errakhi
etal. (2007) reported Streptomyces spp. strain J-2 isolated
from Moroccan soils significantly decreased disease sever-
ity of Sclerotium rolfsii led damping-off disease in sugar
beet besides significantly increasing seedlings development.
Strain J-2 was selected after in-vitro screening experiment,
and was used for sugar beet seeds treatment at 107–108CFU/
seed which led to 66% and 47% reduction in damping-off in
non-sterilized and sterilized soil respectively. Higher per-
centage of pathogen reduction in non-sterilized soil indi-
cate that other soil microbes either enhance the biocontrol
process in consortia mode or through increased activity of
Streptomces spp. J-2 strain itself which needs to be further
researched. They further reported that inhibitor(s) obtained
from culture filtrate of Streptomyces isolates J-2 and B-11
inhibited sclerotial germination 100%, and hyphal growth by
80% (Errakhi etal. 2009). Mixture of biomass and culture
filtrate of these Streptomyces strains significantly reduced
sugar beet root rot when being applied to S. rolfsii infested
soil, and was recommended for integrated control of soil
borne plant pathogens. So while looking for consortium
approach researchers should not only confined to different
microbial strains combinations, but should also take culture
filtrate into consideration. Karimi etal. (2012) reported that
native Streptomyces isolates (C and S2) on soil treatment
(especially saline soil) inhibited root rot of sugar beet caused
Table 4 Application of actinomycetes in Amaranthaceae vegetables
Crops/vegetables Source of isolation Actinomycetes Trait(s) Pathogen(s) References
Sugar beet Leaf and root Streptomyces griseo-
fuscus, Streptomyces
globisporus
PGP – Shi etal. (2009)
Sugar beet Moroccan soils Streptomycetes sp. Damping-off of sugar
beet seeds
Sclerotium rolfsii Errakhi etal. (2007)
Sugar beet Native isolates Streptomycesisolates (C
and S2)
Biocontrol of root rot of
sugar beet
Rhizoctonia solani
AG-2, Fusarium
solani and Phytoph-
thora drechsleri
Karimi etal. (2012)
Sugar beet Native isolates Streptomyces sp. Sugar beet damping off Rhizoctonia solani Sadeghi etal. (2009)
Sugar beet Sugar beet rhizosphere
soil
Streptomyces isolates
J-2 and B-11
Root rot on sugar beet Sclerotium rolfsii Errakhi etal. (2009)

World Journal of Microbiology and Biotechnology (2018) 34:132
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132 Page 10 of 16
by Phytophthora drechsleri, Rhizoctonia solani AG-2, and
Fusarium solani. When sugar beet seeds were treated with
Streptomyces C isolate it controlled Rhizoctonia solani
damping off in naturally and artificially infested soils (Sad-
eghi etal. 2009). So far only genus Streptomyces has been
used for one amaranthaceae vegetable.
Umbelliferous vegetables
Carrot is only vegetable of the Umbellifery family which
has been researched for actinomycetes application (Table5).
Seven actinomycetes strains Streptomyces janthimts, S. cin-
erochromogenes, Streptoverticilium netropsis, Actinomadura
rubra, Actinoplanes philippinensis, Muromonospora carbo-
naceae, and Streptosporangium albidum isolated from carrot
rhizosphere and screened by invitro and invivo antagonism
against Pythium coloratum Vaartaja, producing non-volatile
antifungal metabolites significantly controlled cavity-spot
disease of carrot under natural field condition. Actino-
planes philippinensis and Muromonospora carbonaceae
also showed hyperparasitism, and collapsed the pathogen
oospore by heavily colonizing it and epiphytically growing
on hyphae. This was the first report on microbial control of
cavity-spot disease of carrot (El-Tarabily etal. 1997). Root
colonizing ability of antagonistic Streptomyces griseoviridis
was tested on carrot and turnip rape by stand-tube method. It
well competed with indigenous soil microbes for root colo-
nization when sufficiently available in soil before seed emer-
gence, but could not compete when applied at later stage,
depicting the importance of time of soil spray for root colo-
nization (Kortemaa etal. 1997). Therefore besides search-
ing for competent strains, methodology of bio-formulation
development, and its timing of application should also be an
integral component of such research projects.
Asteraceae vegetables
Lettuce (Lactuca sativa), a leafy vegetable is only mem-
ber of the family Asteraceae which has been researched for
actinomycetes application (Table5). Streptomyces viridodi-
asticus, Micromonospora carbonacea, and Serratia marces-
cens isolated from lettuce growing field of Al-Ain, United
Arab Emiratus significantly reduced growth of Sclerotinia
minor, a pathogen causing basal drop disease of lettuce in
invitro test, and produced high level of beta-1,3-glucanase
and chitinase enzyme (El-Tarabily 2000). Pathogen hyphal
plasmolysis and cell wall lysis were noticed when it was
given as sole carbon source to the isolated three strains,
which were recommended either individually or in combi-
nation to control Sclerotinia minor led basal drop disease of
lettuce under field condition.
Lettuce root and rhizosphere colonization dynamics by
genetically modified five Streptomyces spp. (transformed by
plasmid plJ8641 for enhanced GFP marker and apramycin
resistance) showing strong invitro inhibition for Sclerotinia
sclerotiorum, a major soil-borne pathogen of horticultural
crops was reported. Tagged ZEA171 strain was found to
be both rhizospheric and endophytic as was evident by re-
isolation experiment (Bonaldi etal. 2015). This indicate
that some of the rhizospheric microbes enter in to plant to
become endophytes at later stage. Thus potent endophytic
actinomycetes strain(s) may also be applied as rhizospheric
application. Strptomyces exfoliatus FT05W and Streptomy-
ces cyaneus ZEA171 strains available in culture collection of
University of Milan, Italy inhibited the growth of soil borne
pathogen Sclerotinia sclerotiorum causing lettuce drop
disease by more than 75% under invitro test. Furthermore
under field condition S. exfoliatus FT05W and S. cyaneus
ZEA171 (106CFU/mL) protected lettuce from drop by 40%
and 10% respectively when inoculated a week prior to let-
tuce showing, while Streptomyces lydicus WYEC 108 iso-
lated from commercial product ‘Actinovate’ could not pro-
tect the plant (Chen etal. 2016). When applied timely some
of these strains may perform better compared with available
commercial products. These strains were viable and per-
sistent in the rhizosphere and endorhiza up to 3 weeks as
was evident by GFP tagging and SEM analysis. Strain S.
exfoliatus FT05W has been recommended for controlling
this soil borne disease which could not be done by available
plant protection products. Only two actinomycetes genera
have been applied for one Asteraceae vegetable.
Fabaceae vegetables
Pea (Pisum sativum) is only vegetable crop of the Fabaceae
family which has been investigated for actinomycetes appli-
cation (Table5). Yuan and Crawford (1995) reported that
Streptomyces lydicus WYEC108 antagonized various plant
pathogens in plate assays, and inhibited growth of Pythium
ultimum and Rhizoctonia solani in liquid medium. Pea seeds
coated with WYEC108 was 70% protected by P. ultimum
invasion when planted 24h prior to pathogen infection.
WYEC108 when applied as spore-peat moss-sand formu-
lation (108 CFU/g) also showed plant growth promoting
traits. Tokala etal. (2002) further investigated the interaction
between WYEC108 and pea, and reported increase in root
nodulation frequency by acting on Rhizobium spp. infec-
tion level. Average nodule size was increased as S. lydicus
colonized and sporulated in it, leading to improved bacte-
roids vigor by enhanced nodular assimilation of iron and
nutrients, and reduced poly-beta-hydroxybutyrate accumu-
lation. It was hypothesized that Streptomyces act as plant
growth promoting bacteria in pea and other leguminous
plants by root and nodules colonization, which has been
proved now, and such actinomycetes strains have also been
termed as “Rhizobia helper bacteria” [RHB]. Several species

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Page 11 of 16 132
Table 5 Application of actinomycetes in vegetables belonging to Umbelliferae, Asteraceae, Fabaceae, Amaryllidaceae, Asparagaceae, Amaranthaceae, Zinzgiberaceae families
Crops/vegetables Source of isolation Actinomycetes Trait(s) Pathogen(s) References
Carrot Carrot rhizosphere Streptomyces janthinus, S. cin-
erochromogenes, Streptover-
ticillium netropsis, Actino-
madura ruhra, Actinoplanes
philippinensis, Micromono-
spora carbonaceae, and
Streptosporangium albidum
Biocontrol Pythium coloratum Vaartaja,
a causal agent of cavity-spot
disease of carrots (Daucus
carota)
El-Tarabily etal. (1997)
Turnip rape and carrot Culture collection Streptomyces griseoviridis Root colonization ability – Kortemaa etal. (1997)
Lettuce Culture collection, University
of Milan
genetically modified [GFP]
Streptomyces spp.
PGP and Biocontrol Sclerotinia sclerotiorum Bonaldi etal. (2015)
Lettuce Culture Collection, University
of Milan
S. exfoliatus FT05W, S. cyaneus
ZEA17I
Biocontrol of soil borne dis-
ease, Lettuce drop
Sclerotinia sclerotiorum Chen etal. (2016)
Lettuce Lettuce rhizospheric soil Micromonospora carbonacea Basal drop disease of lettuce Sclerotinia minor El-Tarabily etal. (2000)
Pea plant (Pisum sativum) Rhizosphere soil of linseed Streptomyces lydicus
WYEC108
Enhanced root nodulation,
at the level of infection by
Rhizobium spp.
– Tokala etal. (2002)
Sweet pea Endophytic actinomycetes of
sweet pea
Streptomycetes sp. P4-isolate Biocontrol Oidium sp. powdery mildew
pathogenic fungus
Sangmanee etal. (2009)
Pea seeds Soil samples Streptomyces lydicus
WYEC108
Pythium seed rot and root rot Pythium ultimum or Rhizocto-
nia solani Yuan and Crawford (1995)
Pea Rhizospheric soils Streptomyces St7c5 Foot rotting and blight Mycosphaerella pinodes Mohamed and Benali (2010)
Transgenic pea Culture collection chitinase (Chit30) from Strepto-
myces olivaceoviridis ATCC
11238
Inhibition or delay of hyphal
extension of T. harzanium Trichoderma harzanium Hassan etal. (2009)
Onion Egyptian soils Streptomyces coelicolorHHFA2 Onion bacterial rot disease Erwinia caroto-
vorasubsp.caroto-
voraandBurkholderia
cepacia
Abdallah etal. (2013)
Asparagus Asparagus field soil Streptomyces spp. (ME2-27-
19A)
Asparagus decline syndrome Fusarium oxysporum f.sp.
asparagi (FOA), F. monili-
forme (FM)
Elson etal. (1994)
Table beet, bush beans Clay granules Actinoplanes spp. ReducingPythiumdamping-off
of plants
Pythium ultimum Khan etal. (1997)
Ginger Soil Streptomyces species (SSC-
MB-01 to SSC-MB-06)
Biocontrol Fusarium oxysporum f.sp.
zingiberi Manasa etal. (2013)

World Journal of Microbiology and Biotechnology (2018) 34:132
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132 Page 12 of 16
of genus Micromonospora have recently been reported to
play a significant role in nodular tissues of both legumi-
nous and actinorhizal plants (Trujillo etal. 2015; Martínez-
Hidalgo etal. 2014). Actinomycetes rhizobia consortia will
soon be replacing currently used rhizobia based nodulating
agents. Endophytic Streptomyces spp. isolated from sweet
pea when inoculated by seed and three foliar spray at 15, 34,
and 49days after transplantation significantly reduced the
leaf damage by powdery mildew when infected by patho-
genic fungus Oidium sp. 67 DAT (Sangmanee etal. 2009).
Therefore in addition to timing of application, plant parts to
be treated should also be worked out. When Chit30 (a fam-
ily 19 chitinase) from Streptomyces olivaceoviridis ATCC
11238 was expressed in transgenic pea it inhibited and
delayed Trichoderma harzanium hyphal extension because
of enhanced antifungal activity when compared with non-
transgenic control plant (Hassan etal. 2009). These actino-
mycetes strains may also work as a source of useful genes
for generating transgenic crops. Pea seeds treated with talc
formulation of Streptomyces St7c5, a rhizospheric isolate
enhanced seed germination, showed plant growth promo-
tion, and significantly reduced root rotting and blight caused
by Mycosphaerella pinodes (Mohamed and Benali 2010).
Actinoplane strain 25,844 (108CFU/g of granules) at 1%
(w/w) increased bush beam emergence on being planted
in Pythium ultimum infested plots after 28days (Khan
etal. 1997). Three genera of actinomycetes (Streptomyces,
Micromonospora, Actinoplanes) have been used for single
Fabaceae vegetable.
Amaryllidaceae family vegetable
There is only one report on management of onion bacte-
rial rot disease using actinomycetes as biocontrol agent
(Table5). Egyptian soil isolates Streptomyces lavendulae
HHFA1 and Streptomyces coelicolor HHFA2 controlled
onion bacterial rot in pot and field experiments, and also
showed plant growth promoting effect by enhanced photo-
synthetic pigments and foliar growth. S. coelicolor HHFA2
significantly reduced onion bacterial rot disease incidence
throughout the post harvest storage period compared with
untreated control, and has been recommended as biological
control agent for onion bacterial rot disease (Abdallah etal.
2013).
Asparagaceae family
There is only one report by Elson etal. (1994) on Strepto-
myces spp. (ME2-27-19A) isolated from asparagus field soil
inhibiting Fusarium oxysporum f.sp. asparagi (FOA) and
F. moniliforme (FM) causing asparagus decline syndrome
of asparagus root and crown respectively (Table5). Chro-
matographically purified compound(s) obtained by solvent
extraction of Streptomyces strain ME2-27-19A inhibited
FAO and FM at 40mg/ml, and at 100mg/ml when applied
in soil but asparagus shoot length was also reduced. There-
fore it is desirable that researchers should optimize the
strains for both biocontrol and PGP traits.
Amaranthaceae family
Khan etal. (1997) has reported augmentation of field soil
with Actinoplanes spp. sporangia for biological control
of Pythium damping-off (Table5). Actinoplanes strain
25,844 increased growth and reduced root rot of table beet
compared with control when applied at 5% (w/w) as gran-
ules (4 × 107 –4 × 108CFU/g of granules) to P. ultimum
(750–1000oospores/g) infested soil. Hyper-parasitic activity
of strain 25,844 for P. ultimum oospores was reported even
on six month old granules.
Zinzgiberaceae family
Manasa etal. (2013) screened Streptomyces sp. (SSC-MB-01
to SSC-MB-06) for biocontrol potential against Fusarium
oxysporum f.sp. zingiberi causing rhizome rot of ginger by
dual plate method, and also by using ethyl acetate extract of
broth by agar diffusion test. Streptomyces sp. SSC-MB-02
showed best inhibitory activity while strain SSC-MB-05 did
not inhibit pathogen in both the test. Streptomyces strain
SSC-MB-02 has been recommended for controlling soft rot
symptoms of ginger rhizome (Table5).
Commercial strains available
There are nine actinomycetes based products available in
the market worldwide and ten active substances registered
as commercial products, and all of them have been prepared
using different species of the genus Streptomyces (Kabaluk
etal. 2010; Aggarwal etal. 2016; Vurukonda etal. 2018),
besides few other strains are at various stages of develop-
ment in different laboratories in order to get commercial sta-
tus. Mycostop and Actinovate are two of these actinomycetes
based commercial products which have been in use globally
for a long time. Mycostop, registered as microbial pesticide
in Canada, European Union and USA, inhibits soil and seed
borne fungal pathogens, and has been prepared using Strep-
tomyces griseovirids isolated from Finnish Sphagnum peat
(White etal. 1990). Seed and soil treated with ‘Mycostop’
inhibits seed and soil borne fungal pathogens because of
antifungal aromatic heptaene polyene antibiotic secretion
by S. griseovirids K61 strain. Like Alternaria brassicicola
on cauliflower and cabbage, and Botrytis cinerea on let-
tuce were controlled by ‘Mycostop’ application, besides it
is also being used to control Fusarium, Phytophthora, and
Pythium led wilt and root diseases in different crops. While

World Journal of Microbiology and Biotechnology (2018) 34:132
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Page 13 of 16 132
Actinovate, registered in Canada and USA, inhibits the
growth of Pythium ultimum and Rhizoctonia solani besides
showing plant growth promoting traits, and has been pre-
pared using Streptomyces lydicus WYEC 108 strain (Yuan
and Crawford 1995). Soil borne diseases like those being
caused by Pythium, Fusarium, Phytophthora, Rhizoctonia,
Verticillium, foliar diseases such as powdery and downy
mildew, and those being caused by Botrytis and Alternaria,
Postia, Geotrichum, and Sclerotinia can be controlled using
Actinovate (Vurukonda etal. 2018). Mykocide KIBC con-
trolling powdery mildews, grey mold, brown patch and Safe-
grow KIBC being used for sheath blight and large patch
are registered in South Korea. Actofit targeting Colorado
potato beetle, web mites and other phytophags, while Bac-
tophil controlling seed germination diseases are registered
in Ukraine. Incide SP and Actin being used as insecticides
and fungicide respectively are registered in India, while well
known Bialaphos being used as herbicide is registered in
USA (Kabaluk etal. 2010).
Among the list of active substances derived from actino-
mycetes, Abamectin (Avermectins) registered globally tar-
gets beetles, mites, leaf miners, suckers and other insects
of vegetables and other crops. Streptomycin, Phytomycin,
Validamycin, and Polyoxorim registered in Canada, China,
India, New Zealand, Ukrain, and USA control various bacte-
rial and fungal led diseases of vegetables and other crops.
Polynactin controlling different types of mites, and Milbe-
mycine controlling mites and leaf miners in eggplant are
registered in Japan. While Blasticidin-S controlling rice
blast and Kasugamycin controlling Phytophthora led root
rot and leaf spot in different crops are registered in USA and
Ukraine respectively (Aggarwal etal. 2016).
Conclusions andfuture perspectives
In general potential actinomycetes strain has been isolated
either from rhizospheric soil sample, and/or as endophyte
of same or different crop. Usually researchers have focused
on roots of healthy plants while searching for potent endo-
phytes, but the diseased plants and also other parts of plant
like stem and leaves should not be ignored. Focus should
also be on isolation of rare actinomycetes and still unex-
plored actinomycetes genera using different newly reported
media. Actinomycetes strain inhibiting specific pathogen
affecting different vegetable crops, and also strain inhibiting
different pathogens because of its broad spectrum of activity
should be the target of research. Such potent strain should
be further optimized for its PGP and other useful traits.
Consortium approach should be open not only to different
species and genera of actinomycetes, but also to different
groups of microorganisms and its culture filtrates, besides
looking for combined application with different chemicals
and/or organic substitutes. Bioformulation development, and
its application on particular part of plant at specific time
should be well established. Strain specificity of both part-
ners should be workout before using actinomycetes rhizobia
consortia. Biotechnological expertise need to be develop to
convert any actinomycetes strain into competent potential
strain either by activating silent genes, and/or by transfer of
genes from other strains using modern genetic engineering
and synthetic biology tools.
It is quite evident that a limited group of vegetables
belonging just to 11 families have only been researched for
actinomycetes application. Many vegetable families are not
even touched, and single publication on application of actin-
omycetes to only one vegetable crop belonging to family like
Amaryllidaceae, Asparagaceae, Amaranthaceae, and Zinz-
giberaceae itself speak the scope for future research. Fur-
thermore even among the 12 orders with 162 genera of actin-
omycetes only few have got researchers attention. Future
research programme should not only target still untouched
vegetables, but should also focus on still unexplored genera
of this very promising group of microorganisms isolated
from different sources including endophytes. Exploration
of multifunctional actinomycetes strains having optimized
traits combination of biocontrol, PGP, herbicidal, and pesti-
cidal activities should be the target of future research.
Acknowledgements This review article has been prepared under the
institute project “Bioprospecting of microorganisms associated with
vegetables against plant pathogens-Actinomycetes component” [Project
Code IXX08678]. Facilities provided by Director, ICAR-IIVR, Vara-
nasi, and Head, Division of Crop Protection, ICAR-IIVR, Varanasi is
duly acknowledged.
Funding Funding was provided by Indian Council of Agricultural
Research.
Compliance with ethical standards
Conflict of interest The authors declare that they have no conflict of
interest.
Disclaimer The mention of specific products, trade names, or manu-
facturers is intended only for accuracy, and should not be considered
as an endorsement, or otherwise, of that product.
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