Crop Protection 22 (2003) 247–252

Review
Research progress on microbial herbicides
Yongquan Li*, Ziling Sun, Xiaofeng Zhuang, Ling Xu, Shifei Chen, Mingzhi Li
Department of Biotechnology, College of Life Science, Yuquan Campus, Zhejiang University, Hangzhou 310027, People’s Republic of China
Received 5 August 2002; received in revised form 10 September 2002; accepted 12 September 2002

Abstract

A description is given on principles, characters, action mechanism and applications of microbial herbicides, such as bacterial,
fungal and actinomycete derived products. The present situation in this area both in China and abroad is also reviewed.
r 2003 Elsevier Science Ltd. All rights reserved.

Keywords: Microbial-herbicide; Bacteria; Fungi; Atinomycete; Phytotoxin; Weed control

Contents

1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 247
2. Status of microbial herbicides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 247
2.1. Microbial preparation herbicide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 248
2.2. Microbial-derived herbicide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 249
2.2.1. Phytotoxins from fungi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 249
2.2.2. Phytotoxins from bacteria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 249
2.3. Phytotoxins from actinomycetes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 250
3. Current international research on herbicides in the world . . . . . . . . . . . . . . . . . . . . . . . . . 250
4. Future . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 251
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 251

1. Introduction of the global environmental consciousness, bioherbi-

About 30,000 kinds of weeds are widely distributed in
the world; yield losses caused by 1800 kinds of weeds are
about 9.7% of total crop production every year. So
weed control has always been an important aspect of
environmental protection. Over the past century,
chemical herbicides have been effectively employed to
control various weeds. However, They may have caused
many serious side-effects such as herbicide-resistant
weed populations, reduction of soil and water quality,
herbicide residues and detrimental effects on non-target
organisms (TeBeest and Templeton, 1985; Beckie and
Morrison, 1993; Heap et al. 1993). With the heightening
*Corresponding author. Tel.: +86-571-88866337; fax: +86-571-
87951358.
E-mail address: lyq1962@yahoo.com (Y. Li).
cides especially microherbicides, which are highly
effective for weed control and environmentally friendly
as well, are very attractive both for research and for
application. However, the full impact of their use has
not been investigated in detail.
2. Status of microbial herbicides
It has been 200 years since control of weeds based on
biological methods was applied. However, these were all
ecological approaches through making use of natural
enemy of weeds such as herbivorous animals and
pathogenic microorganisms. Real microbial herbicides,
by definition, began to be studied in the middle period of
the last century and developed fast with the advance of

0261-2194/03/$ - see front matter r 2003 Elsevier Science Ltd. All rights reserved.
PII: S 0 2 6 1 - 2 1 9 4 ( 0 2 ) 0 0 1 8 9 - 8
248 Y. Li et al. / Crop Protection 22 (2003) 247–252
phytotoxin science. In last 10 years, some phytotoxins
screened from pathogenic weeds showed potential
herbicidal activity. Some experts suggest these phyto-
toxins be developed as a new kind of bio-herbicide as an
alternative chemical herbicides. It will require further
work based on discovery, study and the separation of
phytotoxins.
Phytotoxins used for microbial herbicides can be
divided into three types: bacterial, fungal, actinomycete
derived product. The pathogens which produce phyto-
toxins as a microbial herbicide must fit certain require-
ments: (1) be reproduced by biological technique,
(2) grow fast after spraying or be capable of killing
weeds within definite time, (3) suit industrial pro-
duction, and (4) be suitable for packaging, transport
and use.
The first studied microbial herbicides in China was a
culture suspension of C. gloeos poroides f. sp. cuscutae
with the name Lubao No. 1 to control dodder (Cuscutae
australis) in the 1960s (Zhaoyuan Gao, 1992). De Vine
was registered first in America. Other bioherbicides such
as COLLEGO, Dr. BioSEDGE, BioMal, Stumpout
followed them (Yongqang Chen, 1998; Charudattan
and Dinoor, 2000). All of these microbial herbicides use
fungal spores preparations. However, they are difficult
to make in mass production, and because of their
specific requirement for action conditions, these agents
have not been accepted widely and did not bring
significant economic profits (Makowski, 1993; TeBeest
et al., 1992).
In the 1990s, scientists turned their interests to
bacteria. They isolated some strains possessing herbici-
dal activity from rhizobacteria of weeds and acquired
exogenous toxins through cultures for controlling plants
(Boyetchko and Holmstrom-Ruddick, 1996). While the
structure and property of fungal herbicides have been
studied extensively, the bacterial-herbicides are based on
report that some crude toxins from bacterial fermenta-
tion can control and kill weeds (Begonia et al., 1990;
Dorworth, 1992; Harris and Stahlman, 1992; Kennedy
et al., 1995; Kremer et al., 1990; Norman and Kim,
1994). In Western Canada, more than 1000 strains of
rhizobacteria from prairie soils have been screened for
their inhibitory effects against annual grassy weeds
(Boyetchko and Holmstrom-Ruddick, 1996), which
indicated that these pathogens from rhizobacteria were
possible as potential biocontrol agents. Host-range tests
of several strains that could suppress downy brome
(Bromus tectorum) demonstrated that the effects of the
rhizobacteria and their secondary metabolites were host
specific and the amount of reduction in root growth was
found to be concentration dependent. Tests on non-
target organisms showed little or no detrimental effects
on root and shoot growth and improvements to plant
growth in some cases (Kennedy et al., 1991). Comparing
with fungi, bacteria hold some advantageous character-
istics such as short-growth period, simple fermentation
technique, and easily controlled production process. In
addition, bacteria can produce secondary metabolites
unlike fungal spores, which need strict conditions for
action as herbicides, and their residues are easily
degraded. Bacterial herbicides have a good prospect in
application and exploitation, and may become a focus in
this research field.
Microbial herbicides can be divided into microbial
preparation herbicide and Microbe-derived herbicide by
virtue of the effective components from the pathogen
itself or its phytotoxin.
2.1. Microbial preparation herbicide
Microbial preparation herbicide is defined as micro-
oganism that can control the weeds. Up to now, most
research reports are relevant to plant fungal pathogens,
for bacterial pathogens with weed control activity are
scarce. The earliest reported mycoherbicide is a culture
suspension of C. gloeos poroides f. sp. cuscutae for
controlling dodders with the name Lubao No. 1, which
was studied by Shandong Academy of Agriculture
Science in China in the 1960s. However, its commercial
development was not successful. In the middle of 1990s,
three mycoherbicides have been developed and regis-
tered in the United States, one was registered in Canada
and one was in South Africa (Charudattan and Dinoor,
2000). The first registered mycoherbicide under the
commercial name De Vine is a chlamydospores suspen-
sion of Phytophthora palmivora for controlling strange-
lervine (Morrenia odorata). It kills up to 96% weeds in
fruit garden, and its effects can last at least 2 years
(McRae, 1988). Abbott Laboratory introduced this
mycoherbicide into the American market and began
sales in 1995.
Colletotrichum gloeosporioides (Penz) Penz. & Sacc. f.
sp. aeschynomene can produce anthracnose symptoms
on the host plant to control northern jointvetch
(Aeschynomene virginica), a leguminous weed in rice
(Oryza sativa L) and soybeans ( Glycine max L). Its
product can be applied into the foliage with conven-
tional herbicide sprayers and was registered as ‘‘Colle-
go’’, a wettable powder comprised of dried spores
(Mortensen, 1988; Boyetchko, 1997).
DR. BioSEOGE, produced by Puccinia canaliculata,
can control yellow nutsedge (Cyperus esculentus L)
in all cropping areas and completely inhibit weed
flowering in early spring and reduce yellow nutsedge
stand density and new tuber formation by 46% and
66%, respectively (Boyetchko, 1997). In addition,
urediniospores of Puccinia canaliculata. applied in
mid-June did not reduce the population of yellow
nutsedge, but did decrease its tuber biomass by
32% (TeBeest and Templeton, 1985). Although DR.
BioSEOGE has been exploited, it is not available in
 Y. Li et al. / Crop Protection 22 (2003) 247–252
the marketplace because it is a parasitic microoganism
in specific plants and its mass production is only
achieved by inoculating on living plants (TeBeest and
Templeton, 1985).
BioMal, the first bioherbicide registered in Canada,
is a product comprised of fungal spores of
Colletotrichum gloeosporioides f :sp. malvae and cause
round-leaved mallow (Malva pusilla Sm.) sunken lesions
on the stems and leaf petioles. A suspension of 2  106
spores/ml can give excellent weed control under a dew
period of at least 20 h along with temperature about 301
(Makowski, 1993). In addition, the fungus Colletotri-
chum truncatum (Schw) Andrus and Moore has been
identified as a promising mycoherbicide for controlling
hemps sesbania (Sesbania exaltata). Alternaria cassiae
Jurair and Khan was recognized as a possible biocontrol
agent for sicklepod (Cassia obtusifolia L), a weed found
in soybean and peanut land (ShengQiang, 1997).
Velvetleaf (Abutilon theophrasti) has been controlled
with Colletotrichun coccodes (Gotlieb et al., 1987;
Boyetchko, 1997).
2.2. Microbial-derived herbicide
Microbial-derived herbicides, especially microorgan-
ism secondary metabolites, are a new kind of microbial
herbicide to control weeds, which are always phyto-
toxins. Microorganism can produce a lot of metabolites
whose characteristics are diversity in structure and
biological activity, and easily degraded. These bioactive
components invade into the host plant, cause patho-
genicity, destroy their structure and lead them to
produce necrotic lesions or chlorotic halo. Now,
biologists and agriculturalists are paying more atten-
tion to natural products that have herbicidal activity,
for these natural phytotoxins have a specific target
and new different chemical structures which are
difficult to synthesize by common pesticide synthesis
method. They are less poisonous to most of mammalian
systems, easily degraded and so far result in no
biological disaster compared to chemical herbicides
(Charudattan, 1991).
These phytotoxins are very different in chemical
structure and size. Some of them are polypeptides,
terpenes, macrocylic and bakelite (Strobel et al., 1991).
They are also different in their host plant speciality,
they either apply to a single species or to one kind
of plant. Others are not only specific to one kind of
host plant but also to a lot of plants. Although the
latter are non-specific, they have a certain selectivity,
for example, Anisomycin from Actinomycetes is not
specific to barnyardness (Echinochloa crusgalli)
and common crabgrass (Digitaria sanguinalis), but
it shows no harm to crops such as rice paddy.
They are introduced according to different sources as
follows.
249
2.2.1. Phytotoxins from fungi
Fungi such as Alternaria, Fusarium, Coletotrichum
et al. can produce phytotoxin. Junko Ohra and Kenji
Morita (1995) found that phytotoxins from Colletotri-
chum gloeosporioides have weed control activity. The
experiment of controlling 7 different kinds of weeds
indicated that Jointvetch (Aeschynomene virginica) was
extremely damaged, Pigweed (Amaranthus retroflexus)
and Florida beggarweed (Desmodium tortuosum) were
severely burned and would not grow out of the damage,
and Johnsongrass (Sorghum halepense) was stunted but
not killed by its extract. The phytotoxin was purified by
HP-20, silica gel column chromatography and HPLC.
When analyzed by NMR and IR, it was found that this
phytotxin is ferricrocin, a kind of siderophore, whose
action mechanism has some relation with chelating
activity (Junko Ohra and Kenji Morita, 1995).
Phytotoxins from other fungi have weed control
activity such as AAL-toxin, cornexistin and tentoxin.
AAL-toxin and its analog in structure can suppress
cearmide synthetase and results in sphingol accumula-
tion that makes membrane break. Cornexitin is meta-
bolin inhibitor and action mechanism of this is similar to
aminoacetic salt. It inhibits one isoenzyme of aspar-
agines aminotransferases, but once acid from tricar-
boxylic acid cycle such as aspartic acid and glutamic
acid is added, the activity of toxin will disappear.
Tentoxin have two different action mechanisms under
different conditions. One is interrupting the formation
of chloroplast by blocking synthesis of coding nucleo-
cytoplasmic protein and the other is energytranferase
inhibitor of ATPase’s coupling factor for controlling
photophosphorylation (Duke et al. 1996).
2.2.2. Phytotoxins from bacteria
Most of the bacteria with an ability to produce toxins
are Gram-postive bacteria such as Pseudomonas, Erwi-
nia, Xanthomonas but there are a few Gram-negative
bacteria such as Streptomyces, Corynebacterium fascio-
monads and some are non-fluorescent pseudomonads
(Kremer et al., 1990).
Pseudomonas syringe pv. phaseolicola is a bacterial
plant pathogen which causes halo blight disease and
localized death on common bean (Phaseolus vulgaris L)
and kudzu (Pueraria lobata), whose toxin was called
phaseolotoxin. Once it infects plant root, it will spreads
to shoot terminus, then causes stunting, chlorosis and
even causes foliage necrotic lesions (Ninak Zidack and
Backman, 1996), Gurusiddaiah and Gealy (1994) have
purified the phytotoxin from D7, which was inhibitory
to downy brome (Bromus tectorum) and was a complex
consisting of at least two polypeptides, a chromophore,
fatty acid esters, and a lipopolysaccharide matrix.
Further purification of this compound resulted in near
complete loss of phytotoxin because purification proce-
dures may damage the phytotoxin (Gurusiddaiah and
250 Y. Li et al. / Crop Protection 22 (2003) 247–252
Gealy, 1994). For further exploring possible mechanism
of D7, Patrick et al. (1993) evaluated effects of a crude
preparation of D7 on various physical process in roots
of downy brome seedlings. It was found that cell
division, respiration, and synthesis of protein, RNA,
and DNA were not inhibited or only slightly inhibited,
whilst disruption of lipid synthesis and membrane
integrity were significant, which might account for
inhibition of root elongation. Tranel repeated this
experiment with the further purified toxin and found
the same result (Patrick et al., 1993).
Other phytotoxins from pseudomonas have different
physiological mechanisms. Phaseolotoxin, a tripeptide
from Pseudomonas syringe pv. phaseolicola, inhibits
arginine synthesis by competing with carbamoyl phos-
phate for the binding site on ornithine carbamoyl
transferase (Daly, 1981). Tabtoxin from Psyringep pv.
tabaci can be hydrolyzed to tabtoxinine-b-lactam, which
inhibits glutamine synthetase. Syringomycin, a peptide-
based phytotoxin from P. syringae pv. syringae, was
originally believed to cause cell membranes hydrolysis.
Further studies showed that this toxin could specifically
increase K+ efflux and H+-ATPase activity and was
revealed to have a possible role in Ca2+ transport
(Patrick et al., 1993).
2.3. Phytotoxins from actinomycetes
Herbicidines and herbimycins are higher-plant toxins
both produced by Streptomyces saganonensis. The
former is used to control grassy weeds in paddyfield as
selective herbicides, the latter controls monocotyledo-
nous and dicotylous weeds (Stephen and Lydon, 1987).
Anismycins from Streptomyces actinomycetes is a kind
of growth inhibitor for annual grassy weeds such as
barnyardness and common crabgrass and broad-leaved
weeds. Its mechanism is to destroy synthesis of plant
chloropyl (Yufen Zhang, 1987).
Bialaphos is a metabolite of Streptomyces viridochro-
mogenes and is widely used to control annual or
perennial grassy weeds and broad-leaved weeds. The
mechanism here is to be metabolized to phosphinothri-
cin in plant, and then phosphinothricin inhibits gluta-
mine synthesis. Anismycin can make small seedlings of
barnyardness and common crabgrass die above 50 ppm,
and inhibit radicle growth under 12.5 ppm. Its synthe-
tase may accumulate ammonia, and control photosyn-
thetic phosphorylation causing plant death (Changlin
Liu, 1999; Xiliang Jiang, 1994). Carbocyclic coformycin
and Hydantocodin is produced from Streptomyces
hygrosoopics, which can decrease synthetase of acleny-
losuccinate by increasing content of ATP and hold back
synthesis of protein (Pillmoor, 1998). In addition,
phthoxazolin, hytantocidin and homoalanosin from
Streptomyces can control several weeds (YanChu Shen,
1993).
3. Current international research on herbicides in the
world
Johnson is the first scientist who studied Xanthomonas
as bioherbicide and screened many Xanthomonas
Campestris pv. poannua for controlling annual bluegrass
(Poa annua L) in burmudagrass (Cynodon dactylon L.
Pers) (Johnson, 1994). Imaizumi used mixture of
Xanthomonas campestris pv. poannua (JT-P482) at
103-8 cfu/ml with sulfonylurea herbicides at 0.01–0.1%
to control annual bluegrass and applied for a patent in
1996 (Imaizumi and Seiko Yokohama, 1996). He found
that only this pathogen suspension of 108 cfu/ml would
give excellent control. Xanthomonas campestris pv.
poannua began to grow largely in inner plant above
17* (Imaizumi et al., 1998). When environment is up to
30–35*, the concentration of suspension is 1010cfu/g of
fw after a week, which can provide 75% downy brome
control after one month (Imaizumi and Fujimori, 1999).
It is reported in Business Paper that Japan Tobacco Ltd.
Company has sold a kind of bioherbicide, whose main
component is Xanthomonas campestris pv. poannua and
effect is seen after 2 h sprays. This bioherbicide from
nature has not shown any harm to environment, human
and animal, and would be suitable for use in soybean,
golf course, roadside and lawns.
It is now possible to improve efficacy of plant
pathogens by recombinant DNA methods. The use of
adjuvant in bioherbicide and supplement some compo-
nents in medium improves the performance. Charudat-
tan and Dinoor (2000) has made an attempt to modify
the host range and improve the virulence of Xanthomo-
nas campestris pv. campestris by using genes encoding
bialaphos production to control weed. Owen and Zdor
(2001) reported Pseudomonas putida ATH-1RI/9 and
Acidovarax delafieldii ATH2-2RS/1 with supplement
can produce more HCN to control velvetleaf (Abutilon
theophrasti). Gronwald et al (2002) have studied the
effect of Pseudomonas syringae pv. tagetis as a biocon-
trol agent on Canada thistle in growth chamber, and
found that foliar application of Pst109 cfu/ml plus
Silwet L-77 (0.3%, v/v) on 4–5-week-old Canada thistle
reduced shoot dry weight by 52%.
In China, study of mycoherbicide is taken up very late
However some papers about screening strain from
actinomycetes and fungi for herbicide are reported,
and studies on bacterial herbicides have not been
reported. Yongquan Li’s research group from Zhejiang
University screened some bacterial strains. They showed
strong activity of weed control from the rhizosphere of
natural dead or diseased weed in non-irrigated farm-
land, using on the model of glutamine synthetase
inhibitor and the high-throughput screening program
of chlorella pyrenoidosa combined with common isola-
tion method of microorganism. Among them, the strain
Xanthomonas L4 had more effective herbicidal activity
 Y. Li et al. / Crop Protection 22 (2003) 247–252
than others, especially for controlling broad-leaved
weeds such as shepherdspurse (Capsella bursa-pastoris)
and redroot amarant (Amaranthus retroflex). A fermen-
ted broth diluted at 1:30 is up to 85% effective in the
control of redroot amarant, 76% effective in the control
of bluegrass. However it is safe to most turf such as
burmudagrass (Cynodon dactylon L. Pers), tall fesece
(Festruca arurdinacea), and annual ryegrass (Lolium
perenne) (Yongquan Li et al., 2001). When the
concentration is higher at 20-25*, the control effect is
better. Further experiments showed that this extracel-
lular toxin is a small heat-resisting molecule and water-
soluble, strong-polarity compound. So it will be possible
to develop it as a bioherbicide for lawn.
4. Future
With the development of sustainable agriculture and
consciousness of human environmental protection,
government and enterprises will pay more attention to
the study and exploitation of microbial pesticide because
of their potential benefits for the environment. With
further study of weed control mechanism and establish-
ment of many models of screening weed pathogen,
microbial herbicide will have a high chance to develop
successfully.
At present, the exploitation of chemical pesticide
becomes more and more difficult. The chance of
screening new pesticides is lower and lower, but their
requirements become higher and higher. So screening
new pesticides becomes more and more difficult, whilst
cost is larger and larger. In 1950s the ratio of success
was one in 800 compared to only 1 in 20,000 now
Moreover, the expense for exploiting one chemical
pesticide is nearly USA$100 million over 8 years. On
the contrary, microbial pesticides on exploitation cost
USA$2 million, which is only 1 per 40 of chemical
pesticide. However their development cycle is shortened
by one–third of that of chemical herbicides. In 1990
global pesticide sales were USA$23 billion, of this
herbicide sale is USA$10 billion and microbial pesticide
sale is less than $100 million. In 1998 sale is nearly
USA$30 billion. In this herbicide is USA$15 billion,
microbial pesticide is USA$300 million (Weijing Chen,
et al, 2002; Shangcheng Xu, 2002; Yibin Zhan, 1996).
It can be expected that the growth rate of herbicide
use will continue to increase, and microbial pesticide will
also increase at the rate of 20% every year, for biology
and biotechnology are developing quickly. In the
meantime, they also give rise a new way to explore
new pesticide based on biosynthesis and molecular
modification by gene technology. Therefore, the study
and exploitation of microbial herbicide will become a
focus area of new pesticide discovery.
251
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