ABSTRACT- Striga is a major constraint affecting sorghum, maize, other cereal crops, sugar cane and legume crops
production in sub Saharan Africa. Striga may result in complete crop loss under the worst of conditions. Prodigious seed
production, prolonged viability of the seeds and the subterranean nature of the early stages of parasitism make the control
of the parasite by conventional methods difficult if not impossible. The increasing incidence of Striga has been attributed
to poor soil fertility and structure, low soil moisture, intensification of land use through continuous cultivation and an
expansion of cereal production. Many potentially successful approaches developed to control this weed include using
resistant/tolerant varieties, sowing clean seeds that are not contaminated with Striga seeds, rotating cereal hosts with trap
crops that induce abortive germination of Striga seeds, intercropping, applying organic and inorganic soil amendments
such as fertilizer or manure, fumigating soil with ethylene, applying post emergence herbicides, push-pull technology and
using biological control agents. Based on some studies, the interaction of tied-ridging with N fertilizer and resistant
varieties; cereal-legume intercropping and its interaction with N fertilizer revealed low Striga infestation. No single
management option has been found effective across locations and time. Hence, an integrated Striga management
approach, currently, offers the best possibility for reducing impact at the farm level.
Key Words- Intercropping, Integrated pest management, Fertilizer, Management options, Striga
INTRODUCTION
Agriculture remains the main source of food and provides
the primary source of livelihood for 36% of the world’s
total workforce [1]. In Asia and the Pacific, 40 to 50% of
the workforce derives its livelihood from agriculture, while
in sub-Saharan Africa (SSA) two-thirds of the working
population still makes their living from agriculture.
In Ethiopia, about 85% of the population depends on
agriculture out of which over 90% still rely on rain-fed
agriculture for their livelihood [2].
The majority of the population in the Arid and Semi-arid
areas depend on agriculture and pastoralism for
subsistence. These activities face many constraints due to
predominance of erratic rainfall patterns, torrential rainfall
which is majority lost to run-off, high rate of evapotranspiration
further reducing yields, weeds growing more
vigorously than cultivated crops and competing for scarce
reserves of moisture, low organic matter levels and high
variables responses to fertilizers [3].
Among the major pests of agricultural crops, weeds alone
caused severe yield losses ranging from as low as 10% to
as high as 98% of total crop failure in the dry land regions.
It should be emphasized that yield losses caused by weeds
could vary from crop to crop and from region to region for
the same crops, in response to many factors that include:
weed pressure, availability of weed control technology, cost
of weed control and level of management practices [4].
From the parasitic weeds, Striga spp. are fairly wide spread
in semi-arid regions crops including certain legumes,
maize, pearl millet, sorghum, other cereal crops and sugar
cane production. Small holder farmers are the most affected
by the Striga problem because they have limited ways and
means of controlling it. The increasing incidence of Striga
has been attributed to poor soil fertility and structure, moisture
stress, intensification of land use through continuous
cultivation and an expansion of cereal production
[5-6]. Most Striga infested areas are characterized by
agricultural production systems exhibiting low
productivity.
DISTRIBUTION AND HOST RANGE OF
STRIGA:
Striga has been given the common name of "witchweed"
because it attaches itself to the roots of the host plant thus
depriving it (the host) of water and nutrients. Striga spp.
(witch weeds) belongs to the family Orobanchaceae [7].
Economically important Striga species are reported from
more than 50 countries, especially from East and West
Africa and Asia [8]. S. hermonthica is common throughout
northern tropical Africa and extends from Ethiopia and
Sudan to West Africa. It also extends from the western
Arabian region southwards into Angola and Namibia [6].
S. asiatica has a wider distribution and is found throughout
semi-arid areas of tropical and subtropical Africa, Asia and
Australia [6]. Nigeria, Sudan, Ethiopia, Mali and Burkina
Faso are heavily affected counties in Africa [9].
The host range is almost wide and besides the cultivated
cereals, it attacks many of the wild grasses. The traditional
crops in the African savanna attacked by the parasite are
sorghum (Sorghum bicolor L., maize (Zea mays L.), pearl
millet (Pennisetum glaucum L.), and sugarcane (Saccharum
officinarum L.) and rice (Oriza sativa L.) [10].
STRIGA BIOLOGY:
Striga plants have green opposite leaves, bright irregular
flowers with corolla tube slightly bent at the middle. The
flowers are pink, red, white or yellow. There is a considerable
variation in flower color. The plant is characterized by
herbaceous habit, small seeds and parasitism. The seeds of
S. hermonthica are extremely small, about 0.2 X 0.3mm,
weighing about 0.7µg. They are generally dispersed by
water, wind, cattle, and man .The number of seeds per
capsule ranges from 700 – 1800 depending on the species.
The seeds can remain viable in the field for as long as
14-20 years .The minimal length of the life cycle of the
parasite, from germination to seed production comprises an
average of 4 months [10].
Since Striga is a parasitic weed the seedlings cannot sustain
themselves on their own resources for particular long after
germination. Therefore, they need to find a host root
shortly after germination and the germination needs to be
perfectly timed with the presence of a host root. Exogenous
germination stimulants called strigolactones are produced
by the host’s root and also by some non-host (usually
referred to as trap crops) roots (Gossypium sp.). They are
plant hormones which inhibit shoot branching [11] but also
signals to seeds of parasitic weeds such as Striga to start
germinate. Strigolactones are also involved in other
physiological processes such as abiotic response and the
regulation of the plants structure is also regulated by
strigolactones. Strigol, a synthetic compound belonging to
the strigolactones, was first isolated from cotton
(Gossypium sp.) and is used as a germination trigger for
Striga [12]. When the seed have been germinated the
seedling can live for 3 to 7 days without a host. After that it
will die if it is not attached to a root and there has been able
to create a parasitic link to that particular root. The seedling
finds its way to the host root by chemical signals and then
creates a xylem-to-xylem connection between the seedling
and the root. However, the seedlings cannot be at a greater
distance from the root than 2 to 3 mm to find its way there.
When the seedlings have attached to the root it grows
underground for 4-7 weeks before they emerge and are
actually seen in the field. One plant can host many Striga
plants and Striga affects the plant mostly before its
emergence. The symptoms are however hard to distinguish
from symptoms caused by drought, lack of nutrients and
other diseases [10].
Subsequent to germination, which occurs in close
proximity of the host roots, a haustorium (organ of
attachment and a physiological bridge between the host and
the parasite) is produced on perception of a host-derived
chemical signal [13]. Haustorium initiation, which
represents the switch from the vegetative to the parasitic
mode of life, occurs on or near the host. The haustorium
attaches, penetrates the host root and establishes connection
with the host xylem. Following attachment, the parasite
remains subterranean for six to eight weeks [13]. During
this period, the parasite is completely dependent on its host
and is most damaging. Generally, the below ground and
above ground development of Striga is shown in the life
cycle of Striga (Figure 1.).
Fig 1 : General life cycle of Striga species
Source: Striga Research Methods [14].
ECONOMIC IMPORTANCE OF STRIGA:
Although there are more than 35 species, only three species
are recognized as economically important [15].
S. hermonthica (Del.) Benth and
S. asiatica (L.) Kuntze are
the two most widespread and the most economically significant
species that parasitize on sorghum (
Sorghum bicolor
L. Moench), pearl millet (
Pennisetumglaucum L.), maize
(
Zea mays L.) and rice (
Oryza sativa L.), whereas
S. gesnerioides (Willd.) Vatke attacks crops such as cowpea
(
Vignaunguiculata L. Walp.) and peanut (
Arachishypogaea
L.) [16]. Of these species
Striga hermonthica is the most
serious biotic problem to cereal production [10].
S. hermonthica
is a debilitating root parasite. It causes damage in two
ways, first by competition for carbon and nutrients and
second through physiological interactions, and metabolic
processes the bulk of which is unknown [17].
The effect of
Striga damage on crops is a reduction in yield.
The extent of yield loss is related to the incidence and
severity of attack, the host’s susceptibility to
Striga,
environmental factors (edaphic and climatic) and the
management level at which the crop is produced [18]. Its
effects on crops range from stunted growth, through wilting,
yellowing, and scorching of leaves, to lowered
yields and death of many affected plants. A report by [9]
indicated that annual sorghum losses attributed to
Striga in
SSA are estimated at 22-27% and specifically at 25% in
Ethiopia, 35% in Nigeria, and 40% in Mali. In terms of
monetary value, the annual cereal losses due to
Striga are
estimated at US$7 billion in SSA. In Ethiopia, Mali and
Nigeria, the annual losses are estimated at US$75 million,
US$87million and US$1.2 billion respectively [9]. In
Sudan, more than 500,000 hectares under rain fed
cultivation are heavily infested by
Striga, which commonly
results in yield losses of 70–100% and thus severe
Striga
infestation can result in complete crop failure [10].
MANAGEMENT AND CONTROL OPTIONS OF
STRIGA:
[19] opined that
Striga is a particular problem in areas with
low moisture and where soil fertility is being eroded
through increased population pressure, decreased use of
fallow and minimal use of organic or inorganic fertilizer.
Most importantly, it mostly affects the livelihoods of poor
subsistence farmers in cereal-based agricultural systems in
Africa. Prodigious seed production, prolonged viability of
the seeds and the subterranean nature of the early stages of
parasitism make the control of the parasite by conventional
methods difficult if not impossible [10]. Several measures
have been tried and adopted for control of
Striga. Many
potentially successful approaches developed to control this
weed include using resistant/tolerant varieties, sowing
clean seeds that are not contaminated with
Striga seeds,
rotating cereal hosts with trap crops that induce abortive
germination of
Striga seeds, intercropping, applying organic
and inorganic soil amendments such as fertilizer or
manure fumigating soil with ethylene, hoeing and hand
pulling of emerged
Striga, applying post emergence herbicides,
push-pull technology and using biological control
agents [10]. Generally, the approaches can be grouped in to
four independent
Striga control options, namely cultural,
chemical, genetic, and biological.
Cultural management practices:
Effective control of
Striga has been difficult to achieve
through conventional hand or mechanical weeding as the
parasite exerts its greatest damage bewitching the crop
before its emergence above ground, and providing evidence
for host plant infection. Many of the traditional control
methods, including crop rotation, soil fertility, trap and
catch cropping, intercropping, hand-pulling and fertilization
are still in vogue [10]. Still these practices are not
adopted by farmers. Because they are perceived by poor
farmers as unaffordable or uneconomical, labor intensive,
impractical, or not congruent with their other farm
operations. A lot of studies have been reported mainly on
the effect of intercropping and fertilizer against
Striga as
follows.
A. Intercropping practice on Striga management:
Weed control is an important aspect in intercropping
because chemical control is difficult once the crops have
emerged. A study by [20] showed that intercropping maize
with legumes considerably reduced weed density in the
intercrop compared with maize pure stand due to decrease
in the available light for weeds in the maize-legume intercrops,
which led to a reduction of weed density and weed
dry matter yield compared with sole crops. Similarly, [21]
demonstrated that intercropping maize or sorghum with the
fodder leguminous
Desmodium uncinatum (Jacq.) DC. and
D. intortum (Mill.) Urb, significantly reduced
S. hermonthica
infestation and increased grain yield. Similar studies
in Kenya indicate that intercropping with cowpeas between
the rows of maize significantly reduced
Striga numbers
when compared to within the maize rows [22]. Moreover,
finger millet (
Eleusinecoracana) intercropped with green
leaf desmodium (
Desmodium intortum) reduced
Striga
hermonthica counts in the intercrops than in the monocrops
[23]. [24] also reported related findings on sorghumcowpea
intercropping where
Striga emergence was lower
under intercrops than sole crops. Generally, various studies
have shown that intercropping cereals, mainly with
legumes such as cowpea (
Vignaunguiculata), peanut
(
Arachis hypogaea) and green gram (
Vigna radiate) can
reduce the number of
Striga plants [25]. Potentially, they
might be acting as traps crops, stimulating suicidal Striga
germination or the microclimate under the crop canopy
may be altered and interfere with
Striga germination and
development [26]. It is also hypothesized that nitrogen
fixed by the legumes might interact with
Striga growth, as
increasing the amount of available nitrogen can reduce
Striga densities [27].
B. Fertilizer application on Striga management:
As
Striga is more favor in less fertile soil, a system that
would improve soil fertility to increase yield as well as
reduce
Striga infestation will be also of double advantage.
Good soil management practices involving the use of crop
residues and organic manure have been effective control
measure against
Striga. [28] observed that
Striga infestation
decreased with increasing organic matter of the soil and
that organic matter content seemed to be the most important
factor which preserved the soil fertility. Since soil
microbial biomass flourishes better in a medium rich in
organic matter, organic or inorganic soil amendments may
increase soil suppressiveness to
Striga spp. and also
improve soil conditions to increase yield of subsequent
cereal. Different research findings were reported by
authors. According to [29], 55-82% reduction in number
and weight of
S. hermonthica recorded due to application
of N using urea in Niger. [30] also reported that N
fertilizers altered assimilate partitioning in favour of the ear
and increased maize grain yield and reduced
Striga count
by 64%. Similarly, the study of [31] conducted in North
east Nigeria showed a reduction in Striga infestation and
damage with the application of N fertilizer on maize varieties.
Striga infestation was significantly reduced at 120 kg
N ha
-1 in the early variety and 60 and 120 kg N ha
-1 in late
varieties. [32] noted that, the nitrogenous compound fertilizer
which contains urea considerably suppressed germination
of
S. hermonthica when applied during conditioning.
The germination of
S. hermonthica seed is associated with
the secretion of germination stimulants by host plants. The
secretion ultimately depends upon the nutrient status of the
soil [33]. It has been demonstrated that under N and P
deficiency, host plants secrete high amounts of germination
stimulants into the rhizosphere, while supply of sufficient
N and P reduces this secretion [34, 35]. Research studies
showed that the effect of N was less pronounced than the
effect of P on strigolactones secretion. As DAP fertilizer
contains 18% N and 46% P
2O
5, high availability of P in
DAP might lead to less production of strigolactones.
However, direct suppressing effect of N on
Striga spp.
cannot be neglected [36].
The high and increasing cost of mineral fertilizers and low
purchasing power of small scale farmers have necessitated
investigating the efficacy of fertilizer application at low to
very low levels. The use of very low doses of mineral
fertilizers and their placement near the planting hole, a
technology termed ‘microdosing’, have been shown to reduce
application rates and thus cost of fertilizer per surface
area, while still improving crop yields [37]. Microdosing of
DAP may prove to be an efficient and cost effective option
to reduce
S. hermonthica damage in sorghum in SSA, particularly
in combination with other control options, such as
intercropping, use of organic fertilizer and hand pulling of
S. hermonthica at flowering to achieve integrated
S. hermonthica
management [35].
Genetic resistance:
Striga resistance is the ability of the host root to stimulate
Striga germination but at the same time prevent attachment
of the seedlings to its roots or to kill the seedlings when
attached. The use of resistant crop cultivars is the most
economically feasible and environmentally friendly means
of
Striga control. In East Africa, the most promising new
approach to
Striga control is the use of resistant cultivars
(e.g. of sorghum).
Striga resistant cultivars have been bred
in a number of crops. However, cultivars with immunity to
Striga have not been found in all host crops. The
host/parasite relationship is governed by a series of steps
involving stimulation of germination, haustorium initiation,
penetration of the host root, connection to the host xylem
and concurrent growth [38].
Many cereals are found to be naturally resistant to
Striga
e.g.; rice, sorghum and some genotypes of maize. A
resistant plant stimulates germination of
Striga but it does
not allow it to attach to the root. Study in
Striga infested
areas revealed cultivation with resistant crops results in
fewer
Striga plants and higher crop yield than a
non-resistant genotype of the cultivated plant would do
[39].
Biological control:
Biological control is generally defined as the deliberate use
of living organisms to suppress, reduce or eradicate a pest
population [40]. Means of biological control of weeds
include herbivorous insects, microorganisms specially
fungi, and smothering plants. The insects that attack Striga
can be classified according to the site damaged into
defoliators such as
Junonia spp., gall forming as
Smicronyx
spp., shoot borers as
Apanteles spp., miners as
Ophiomyia
Strigalis, inflorescence feeders as
Stenoptilodestaprobanes
and fruit feeders as
Eulocastra spp. [41]. Twenty eight
fungi and two bacteria were found to be associated with
Strigahermonthica in Sudan. Among the fungi, only
Fusarium
nygamai and
Fusarium semitectum var. majus showed
potential to be used as bio-agents for the control of Striga
[42].
Chemical control:
Various chemicals including herbicides, fumigants (e.g,
methyl bromide) and germination stimulants (e.g, ethylene)
have been reported as means of control of
Striga [43]. Herbicides
like Imazapyr and pyrithiobac applied as seed
dressing to maize were reported to give efficient control of
the parasite [44]. The excellent control capacity of the
herbicides is most likely due to their relatively long
persistence in the rhizosphere. Furthermore, multi-location
testing showed that this herbicide provided excellent early
season control of both
S. asiatica and
S. hermonthica and
could increase yield 3 to 4-fold in heavy infested fields
[44].
Emerged
Striga plants can be successfully killed with
common herbicides. However, much damage is done by the
fully parasitic young plants before emergence, so such
herbicide treatments do not necessarily reduce yield losses.
The main strategy for control is accordingly to reduce the
seed bank of
Striga in the soil by stimulating the seeds to
germinate in the absence of host plants [45]. This can be
achieved by:
- Planting a Poaceous trap crop (susceptible cereal or
grass) which is ploughed in a few weeks after sowing
before the weeds mature and set seed;
- Sowing crops which stimulate germination, but are not
parasitized, for several seasons (e.g. sunflower, groundnut,
soybean);
- Treating the soil with ethylene which simulates the
chemical substances which exude from host roots and stimulate
germination.
Integrated Striga management:
No single management option has been found effective
across locations and time. An integrated
management
approach, currently, offers the best possibility for
reducing impact at the farm level. Many reports on Striga
management suggested the combined use of cultural
agronomic practices, herbicides, host plant resistance,
fertilization, trap cropping, germination stimulants and
biological control [46]. Control is most effective if a range
of practices are combined in to a program of integrated
Striga control (ISC) that can provide sustainable control
over a wide range of biophysical and socio-economic
environments [47,31]. [47] found that ISC that combined
rotation of Striga resistant maize, trap crops and fertilizer
application reduced the Striga soil seed bank by 46% and
increased crop productivity by 88% while [48] showed that
these practices reduced Striga infestation and damage on
farmers’ fields and increased productivity by more than
20%. Likewise, a report by [31] showed that applying N
fertilizer may not be feasible as a stand-alone solution to
managing purple witch weed in cereals because of the high
cost of fertilizer, but the combined use of N fertilizer and
Striga tolerant / resistant maize and sorghum varieties has
shown promise in the west African Savanas. Furthermore,
an experiment conducted in Niger State on two varieties of
maize (Jo-98 and local) intercropped with soya bean and groundnut at three level of N application (0, 50 and 100kg N
ha-1) showed highly significant (P = 0.01) effect on the severity of Striga infestation [49]. Accordingly, resistant variety
Jo-98 showed less severity of Striga and its interaction with N fertilizer at 100kg ha-1 as well as its intercropping
decreased Striga infestation as compared to local (susceptible) (Table 1).
An integrated management approach, if properly designed, using a combination of suitable control measures, has the
potential to provide a lasting solution to Striga problems. [10] reported that soil fertility and soil moisture management
should be an integral part of any Striga control strategy. A similar study by [50] pointed out that species of Striga were
controlled by using the resistant variety, fertilizer and tied ridges on farms of eastern Ethiopia which had long been
abandoned due to Striga infestation. According to Table 2, species of Striga were controlled by using the resistant variety,
fertilizer and tied ridges on farms; whereas, the local cultivar had severe infestations where the average yield of the
resistant variety was 1718 kg ha-1 as against only 216 kg ha-1 from the local variety. The Striga-infested local varieties
died, failed to produce a head or had very small heads. [51] also added that treatment combination that included resistant
variety, fertilizer and tied ridge gave significantly higher yield followed by one that combined local variety with fertilizer
and tied ridging in North wolloat Sirinka and Kobo sites.
Table 1: Effect of variety, intercropping and nitrogen rate interaction on the severity of Striga infestation
(%)
Intercropping/ Maize Variety | Nitrogen rate (kgNha-1) |
0 | 50 | 100 | Mean |
Maize only (J0-98) – Resistant | 62.2 | 39.2 | 9.4 | 36.9 |
Maize only (Local) –Susceptible | 82.2 | 40.4 | 30.4 | 51.0 |
J0-98 + Soyabean | 19.4 | 14.2 | 6.8 | 13.5 |
Local + Soyabean | 50.1 | 20.8 | 10.8 | 27.2 |
J0-98 + groundnut | 13.2 | 6.2 | 4.3 | 7.9 |
Local + Groundnut | 45.2 | 19.3 | 7.6 | 24.0 |
N-rate mean | 45.4 | 23.4 | 11.6 | - |
F-LSD (0.01) for comparing variety (V) means = 20.6; F-LSD (0.01) for comparing intercropping (I) means = 14.9; F-LSD (0.01) for comparing V x I interaction = 42.4
Source: Intercrops With Trap Crops, Nitrogen Fertilization for Striga hermonthica (Del.) Benth Control at Niger State
[49].
Table 2: Striga count and sorghum yield as influenced by variety, fertilizer and tied ridge
Treatment | Striga count/m2 |
Yield (kg/ha) |
| Babile | Fedis | Gursum | Babile | Fedis | Gursum |
Improved variety with fertilizer and tied
ridge |
1 | 2 | 4 | 1467 | 1740 | 1947 |
Improved variety without fertilizer and
tied ridge |
2 | 3 | 5 | 1200 | 980 | 1244 |
Local variety with fertilizer and tied ridge | 140 | 151 | 170 | 122 | 235 | 290 |
Local variety without fertilizer and tied
ridge |
266 | 181 | 288 | 98 | 148 | 130 |
Striga count against treatment and yield against treatment were significant at p = 0.01. Striga count against location and
yield against location were not significant
Source: Distribution of two Striga species and their relative impact on local and resistant sorghum cultivars in East
Ethiopia [50].
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