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Int. J. Life. Sci. Scienti. Res., VOL 2, ISSUE 2, pp: 110-120 (ISSN: 2455-1716) Impact Factor 2.4 MARCH-2016
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Review on Striga Weed Management
Berhane Sibhatu*
Department of Agronomy, Ethiopian Institute of Agricultural Research, Mehoni Agricultural Research Center
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
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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.
Received: 17 Jan 2016/Revised: 14 Feb 2016/Accepted: 27 Feb 2016
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 evapotranspi-
ration 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
*
Address for Correspondence:
Berhane Sibhatu*
Department of Agronomy
Ethiopian Institute of Agricultural Research
Mehoni Agricultural Research Center
Fax: +251-347770021, P.O.Box: 71, Maichew, Ethiopia
Review Article (Open access)
Int. J. Life. Sci. Scienti. Res., VoL 2, Issue 2
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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, mois-
ture 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 consider-
able 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
Int. J. Life. Sci. Scienti. Res., VoL 2, Issue 2
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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 signif-
icant 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. hermon-
thica 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 wilt-
ing, 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].
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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 organ-
ic and inorganic soil amendments such as fertilizer or
manure fumigating soil with ethylene, hoeing and hand
pulling of emerged Striga, applying post emergence herbi-
cides, 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 fertiliza-
tion 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 inter-
crops, 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. hermon-
thica 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 sorghum-
cowpea 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].
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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 impor-
tant 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 varie-
ties. 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 ferti-
lizer which contains urea considerably suppressed germina-
tion 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% P2O5, 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 re-
duce 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, par-
ticularly 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. her-
monthica 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
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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 Fusa-
rium 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]. Her-
bicides 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:
1) 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;
2) Sowing crops which stimulate germination, but are not
parasitized, for several seasons (e.g. sunflower, groundnut,
soybean);
3) Treating the soil with ethylene which simulates the
chemical substances which exude from host roots and sti-
mulate germination.
Integrated Striga management
No single management option has been found effective
across locations and time. An integrated Striga manage-
ment 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,
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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].
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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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Review on Striga Weed Management

  • 1. Int. J. Life. Sci. Scienti. Res., VOL 2, ISSUE 2, pp: 110-120 (ISSN: 2455-1716) Impact Factor 2.4 MARCH-2016 http://ijlssr.com IJLSSR © 2015 All rights are reserved Review on Striga Weed Management Berhane Sibhatu* Department of Agronomy, Ethiopian Institute of Agricultural Research, Mehoni Agricultural Research Center 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 -------------------------------------------------IJLSSR----------------------------------------------- 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. Received: 17 Jan 2016/Revised: 14 Feb 2016/Accepted: 27 Feb 2016 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 evapotranspi- ration 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 * Address for Correspondence: Berhane Sibhatu* Department of Agronomy Ethiopian Institute of Agricultural Research Mehoni Agricultural Research Center Fax: +251-347770021, P.O.Box: 71, Maichew, Ethiopia Review Article (Open access)
  • 2. Int. J. Life. Sci. Scienti. Res., VoL 2, Issue 2 http://ijlssr.com IJLSSR © 2015 All rights are reserved 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, mois- ture 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 consider- able 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
  • 3. Int. J. Life. Sci. Scienti. Res., VoL 2, Issue 2 http://ijlssr.com IJLSSR © 2015 All rights are reserved 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 signif- icant 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. hermon- thica 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 wilt- ing, 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].
  • 4. Int. J. Life. Sci. Scienti. Res., VoL 2, Issue 2 http://ijlssr.com IJLSSR © 2015 All rights are reserved 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 organ- ic and inorganic soil amendments such as fertilizer or manure fumigating soil with ethylene, hoeing and hand pulling of emerged Striga, applying post emergence herbi- cides, 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 fertiliza- tion 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 inter- crops, 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. hermon- thica 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 sorghum- cowpea 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].
  • 5. Int. J. Life. Sci. Scienti. Res., VoL 2, Issue 2 http://ijlssr.com IJLSSR © 2015 All rights are reserved 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 impor- tant 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 varie- ties. 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 ferti- lizer which contains urea considerably suppressed germina- tion 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% P2O5, 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 re- duce 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, par- ticularly 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. her- monthica 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
  • 6. Int. J. Life. Sci. Scienti. Res., VoL 2, Issue 2 http://ijlssr.com IJLSSR © 2015 All rights are reserved 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 Fusa- rium 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]. Her- bicides 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: 1) 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; 2) Sowing crops which stimulate germination, but are not parasitized, for several seasons (e.g. sunflower, groundnut, soybean); 3) Treating the soil with ethylene which simulates the chemical substances which exude from host roots and sti- mulate germination. Integrated Striga management No single management option has been found effective across locations and time. An integrated Striga manage- ment 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,
  • 7. Int. J. Life. Sci. Scienti. Res., VoL 2, Issue 2 http://ijlssr.com IJLSSR © 2015 All rights are reserved 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].
  • 8. Int. J. Life. Sci. Scienti. Res., VoL 2, Issue 2 http://ijlssr.com IJLSSR © 2015 All rights are reserved 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]. REFERENCES [1] FAO (Food and Agricultural Organization). Investing in Sustainable Agricultural Intensification. The Role of Conser- vation Agriculture.A Framework for Action. Technical re- port: 2008. [2] Alemie A, KeesstraSD and Stroosnijder L. A new agro-climatic classification for crop suitability zoning in northern semi-arid Ethiopia.Agricultural and Forest Meteor- ology, 2010; 150: 1057–1064. [3] CASL. Arid and Semi-arid lands: characteristics and impor- tance. Community Adaptation and Sustainable Livelihoods: 2006. [4] Rezene F and Etagegnehu GM. Development of Technology for the Dryland Farming Areas of Ethiopia (Reddy, N.S. and Giorgis, Kidane Eds.). Proc. of 1st Nat. Workshop on Dryland Farming Res. in Ethiopia, 1994; 26-28 Nov. 1991. [5] Parker C. Observations on the current status of Orobacheand Striga problems worldwide. Pest ManagSci, 2008; 65: 453- 459. [6] Gethi JG and Smith ME. Genetic responses of single crosses of maize to Striga hermonthica (Del.) Benth and Striga asia- tica (L.) Kuntze. Crop Sci, 2004; 44: 2068- 2077. [7] Matusova R, Rani K, Verstappen FWA, Franssen MCR, Beale MH and Bouwmeester HJ. The strigolactone germination stimulants of the plant- parasitic Striga and Orobanche spp. are derived from the carotenoid pathway. Plant Physiology, 2005; 139: 920- 934. [8] Aly R. Conventional and biotechnological approaches for control of parasitic weeds. Invitro Cell. Dev. Biol- Plant, 2007; 43:304- 317. [9] AATF (African Agricultural Technology Foundation). Feasi- bility Study on Striga Control in Sorghum. African Agricul- tural Technology Foundation, Nairobi: 2011: ISBN 9966- 775-12-9. [10] Babiker AGT. Striga: The Spreading Scourge in Africa. Re- gul. Pl. Grow. and Devel, 2007; 42: 74-87. [11] Gomez-Roldan V, Fermas S, Brewer PB,Puech-Pagès V, Dun EA, Pillot JP, Letisse F, Matusova R, Danoun S, Portais JC, Bouwmeester H, Bécard G, Beveridge CA, Rameau C, and Rochange SF. "Strigolactone inhibition of shoot branching". Nature, 2008; 455:180-194. [12] Cardoso C, Ruyter-Spira C, Bouwmeester HJ. Strigolactones and root infestation by plant-parasitic Striga, Orobanche and Phelipanche spp. Plant Sci, 2010; 180:414-420.
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  • 10. Int. J. Life. Sci. Scienti. Res., VoL 2, Issue 2 http://ijlssr.com IJLSSR © 2015 All rights are reserved [32] Dzomeku IK and Murdoch AJ.Effects of prolonged conditioning on dormancy and germination of Striga hermonthica.Journal of Agronomy, 2007; 6:29-36. [33] Jamil M, Charnikhova T, and Cardoso C. Quantification of the relationship between strigolactones and Striga hermon- thica infection in rice under varying levels of nitrogen and phosphorus. Weed Research, 2011; 51:373–385. [34] Lopez-Raez JA, Charnikhova T, Gomez-Roldan V. Tomato strigolactones are derived from carotenoids and their biosynthesis is promoted by phosphate starvation. New Phytologist, 2008; 178:863–874. [35] Jamil M, Kanampiu FK, Karaya H, Charnikhova T and Bouwmeester HJ.Striga hermonthica parasitism in maize in response to N and P fertilizers.Field Crops Research, 2012; 134, 1–10. [36] Simier P, Constant S, and Degrande D. Impact of nitrate supply in C and N assimilation in the parasitic plant Striga hermonthica (Del.) Benth (Scrophulariaceae) and its host Sorghum bicolor L. Plant Cell and Environment, 2006; 29:673–681. [37] Tabo R, Bationo A, Gerard B. Improving cereal productivity and farmers’ income using a strategic application of fertiliz- ers in West Africa. In: A Bationo, B Waswa, JKiharaand JKimetu (eds). Advances in Integrated Soil Fertility Management in Sub-Saharan Africa: Challenges and Oppor- tunities. Springer, Dordrecht, the Netherlands: 2007, 201– 208. [38] Butler LG. Chemical Communication between the Parasitic Weed Striga and its host crop, a new dimension in allelo-chemistry, In: Ml Dakshini and FA Einhelling (eds). Allelopathy: Organisms, Processes, and Application. ACS Symposium Series 582, American Chemical Society. Wash- ington DC., USA: 1993, .158-168. [39] Rodenburg J, Bastiaans L, and Kropff MJ. Characterization of host tolerance to Striga hermonthica.Euphytica. In: Lars- son M. Soil fertility status and Striga hermonthica infestation relationship due to management practices in Western Kenya. (MSc Thesis Online publication: http://stud.epsilon.slu.se), SLU, Swedish University of Agricultural Sciences, 2006; (http://stud.epsilon.slu.se/4488/1/larssonm120704.pdf).Acces sed on April 21/2014 [40] BoyetchkoSM. Innovative application of microbial agents for biological weed control. In: KG Mukerji, BP Chamola and RKUpdahyay (eds.). Biotechnological Approaches in Bio control of Plant Pathogens. Kluwer Academic Plenum, New York, USA: 1999: pp. 73-97. [41] Kroschel J. Analysis of the Striga problem, towards joint action. In: J Kroschel, H Mercer-Quarshie, and J Sauerborn (eds). Advances in Parasitic Weed Control at On-farm Level, 1:13–26. Joint Action to Control Striga in Africa. Marggraf- Verlag, Weikersheim, Germany: 1999. [42] Zahran EB. Biological Control of Striga hermonthica (Del.) Benth. Using Formulated Mycoherbicides Under Sudan Field Conditions. Ph.D Thesis, University of Hohenheim, Germany: 2008, pp:143. [43]Egley GH, Eplee RE and Norris RS. Discovery and testing of ethylene as a witchweed germination stimulant. In: PF Sand, RE Eplee, and RG Westbrooks (eds). Witch weeds Research and Control in the United States. Weed Science Society of America Campaign, 1990; 37-45. [44] Kanampiu FK, Kabamble V, Massawe C, Jasia L, Friesen D, Ransom JK and Gressel J. Multi-site, multi-season field tests demonstrate that herbicide seed-coating, herbicide-resistance maize controls Striga spp. and increases yield in several African countries. Crop Prot, 2003; 22: 697 - 706. [45] Hesammi E. Striga and Ways of Control.Intl.J.Farm and Alli.Sci, 2013; 2(3):53-55. (http://ijfas.com/wp- content/uploads/2013/02/53-55.pdf). Accessed on April 21, 2014. [46] Marley PS, Aba DA, Sheboyan JAY, Musa R and Sanni A. Integrated management in Striga hermonthica in sorghum using a mycoherbicide and host plant resistance in the Nige- rian Sudano-Sahelian Savanna. Weed Res, 2004; 44: 157 – 162. [47] Franke AC, Ellis-Jones J,Tarawali G, Schulz S, Hussaini MA, Kureh I, White R, Chikoye D, Douthwaite B, Oyewole BD and Olanrewaju AS. Evaluating and scaling-up inte- grated Strigahermonthica control technologies among far- mers in northern Nigeria. Crop Protection,2006; 25:868– 878.
  • 11. Int. J. Life. Sci. Scienti. Res., VoL 2, Issue 2 http://ijlssr.com IJLSSR © 2015 All rights are reserved [48] Kamara AY, Chikoye D, Ekeleme F, Omoiguil O and Dugje IY.Field performance of improved cowpea cultivars under natural infestation with Striga gesneroides.International Journal of Pest Management,2008; 54(3):189–195. [49] Mamudu AY. Intercrops With Trap Crops, Nitrogen Fertili- zation For Striga hermonthica (Del.) Benth Control At Niger State. International Journal of Humanities and Social Science Invention, 2013; 2(5):64-69. [50] Temam T. Distribution of two Striga species and their rela- tive impact on local and resistant sorghum cultivars in East Ethiopia. Wiley Inter Science, Trop. Sci, 2006; 46(3), 147– 150. DOI: 10.1002/ts.70. Accessed on April 21, 2014. [51] Gebisa E and Gressel J (Eds).Integrating New Technologies For Strigacontrol: Towards Ending the Witch-Hunt.World Scientific Publishing Co. Pte. Ltd., Singapore: 2007, p:345.