Received: 6 January 2020  |  Accepted: 21 December 2020
DOI: 10.1111/pbr.12896  
R E V I E W  A R T I C L E
Breeding maize (Zea mays) for Striga resistance: Past, current 
and prospects in sub-s aharan africa
Abdoul- Madjidou Yacoubou1,2,3  |   Nouhoun Zoumarou Wallis1 |   Abebe Menkir2  |   
Valerien A. Zinsou1 |   Alexis Onzo1 |   Ana Luísa Garcia- Oliveira4 |   Silvestro Meseka2 |   
Mengesha Wende2 |   Melaku Gedil2  |   Paterne Agre2
1Laboratoire de Phytotechnie, 
d’Amélioration et de Protection des Plantes Abstract
(LaPAPP), Département des Sciences et Striga hermonthica, causes up to 100% yield loss in maize production in Sub-S aharan 
Techniques de Production Végétale (STPV), 
Faculté d’Agronomie, Université de Parakou, Africa. Developing Striga- resistant maize cultivars could be a major component of 
Parakou, Bénin integrated Striga management strategies. This paper presents a comprehensive over-
2International Institute of Tropical Agriculture view of maize breeding activities related to Striga resistance and its management. 
(IITA), Oyo Road, PMB 5320, Ibadan, Nigeria
3Institut National des Recherches Agricoles Scientific surveys have revealed that conventional breeding strategies have been 
du Bénin, 01 BP 884, Cotonou, Bénin used more than molecular breeding strategies in maize improvement for Striga re-
4Excellence in Breeding (EiB), CIMMYT, sistance. Striga resistance genes are still under study in the International Institute 
ICRAF House, UN Avenue, PO Box  
1041-00621, Nairobi, Kenya for Tropical Agriculture (IITA) maize breeding programme. There is also a need to 
discover QTL and molecular markers associated with such genes to improve Striga 
Correspondence
Abdoul- Madjidou Yacoubou, Laboratoire resistance in maize. Marker Assistance Breeding is expected to increase maize breed-
de Phytotechnie, d’Amélioration et ing efficiency with complex traits such as resistance towards Striga because of the 
de Protection des Plantes (LaPAPP), 
Département des Sciences et Techniques complex nature of the host- parasite relationship and its intersection with other envi-
de Production Végétale (STPV), Faculté ronmental factors. Conventional alongside molecular tools and technical controls are 
d’Agronomie, Université de Parakou, 
Parakou, Bénin. promising methods to effectively assess Striga in Sub- Saharan Africa.
Email: abdoulmadjidou.yacoubou@yahoo.com
K E Y W O R D S
Funding information
Bill and Melinda Gates Foundation breeding strategies, maize, QTL, resistance, Striga
Communicated by: Thomas Lübberstedt
1  | INTRODUC TION most widespread stresses (Atera et al., 2013; Edmeades, 2013; Das 
et al., 2019).
Maize is one of the most important cereal crops grown worldwide. Striga, is a parasitic weed belonging to the Orobanchaceae family. 
In Sub- Saharan Africa (SSA), it is regarded as the most important sta- It infests and reduces yields of many cereal crops including maize by 
ple crop with huge potential for addressing the challenge of food in- up to 100% (Atera et al., 2013; Chemisquy et al., 2010; Parker, 2012; 
security (Abdoulaye et al., 2018). However, its productivity remains Teka, 2014). Across the globe, more than 50 species belonging to the 
relatively low across SSA countries when comparing to the global Orobanchaceae family are identified and known as crop pests. In SSA, 
average production (FAO, 2018). Amongst the major constraints that S. hermonthica (Del.) Benth. and S. asiatica (L.) Kuntze are the most 
affect maize productivity, drought, low fertility and the parasitic weeds economically important species affecting maize production (Menkir 
known as Striga hermonthica, have been recognized by farmers as the et al., 2012; Teka, 2014). According to Parker (2012), the tropical 
This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, 
provided the original work is properly cited.
© 2021 The Authors. Plant Breeding published by Wiley- VCH GmbH
Plant Breeding. 2021;140:195–210. wileyonlinelibrary.com/journal/pbr © 2021 Wiley- VCH GmbH  |  195
196  |     YACOUBOU et Al.
semi- arid climatic conditions have allowed rapid development of the 2  | ECONOMIC IMPAC T OF Str iga 
Striga and even its adaptation to context. Unfortunately, S. hermonthica INFESTATION ON MAIZE PRODUC TION 
infestation appears to be worsening due to the current intensive land AND BIOLOGY OF Str iga  spp.
use, mono- cropping practices and human demographic pressure. All 
these factors lead to a continuous decline in soil fertility, which greatly 2.1 | Economic impact of Striga infestation on maize 
favours the Striga occurrence (Rich & Ejeta, 2008). In West Africa, production
Striga is widely found across the region where maize yield losses due to 
infestation can vary from 20% to 80% ( Ejeta, 2007; Kim et al., 2002). Striga parasitism is a limiting factor to maize (Zea mays L.) cropping 
In the last few decades, efforts have been made to develop meth- in the savannah zones of Sub-S aharan Africa (SSA) which constitutes 
ods for Striga control, including agronomic cultural practice, biologi- the maize belt of the sub- region (Runo & Kuria, 2018). About 75% of 
cal control, chemical, host plant resistance and genetically modified cultivated land with maize in SSA is endemic to S. hermonthica (Akaogu 
crops. However, these strategies are only moderately effective, be- et al., 2019). Maize yield losses under severe Striga infestation can 
cause Striga are still expanding its natural range by causing more yield be as high as 100% (Figure 1) and are economically estimated to $7 
losses. From existing strategies, the most effective and sustainable billion in the SSA alone (Spallek et al., 2013). The Striga problem has 
control seems to be an integrated approach that uses resistant cul- been worsened by the increasing mono- cropping practice instead of 
tivars (Chitagu et al., 2014; Hearne, 2009; Yoder & Scholes, 2010). rotation and intercropping systems, human demographic pressure on 
Striga- resistant maize can be a major component of integrated control available land where up to 300 million farmers were exposed to the 
if resistance is incorporated into adapted and, regionally productive Striga infestation in SSA (Badu- Apraku & Fakorede, 2017). Challenges 
cultivars. Resistant maize cultivars can, indeed, reduce both new Striga in managing Striga infestation lead to agricultural land abandonment 
seed production and the Striga seed bank in infested soils. Significant in several West African countries including Benin, Burkina Faso, Niger, 
progress towards the development of Striga- resistant maize variet- Nigeria and Togo (Atera & Itoh, 2011; Badu- Apraku, 2010; Badu- 
ies have been achieved around the world, particularly in Africa. The Apraku et al., 2014). Consequently, this has threatened food security 
International Institute of Tropical Agriculture (IITA), Ibadan, Nigeria and livelihoods of millions farmers in most countries in this region 
and International Maize and Wheat Improvement Centre (CIMMYT), (Menkir et al., 2020).
Zimbabwe, have developed several maize genotypes with varied Striga 
resistance levels and adapted to different eco- climatic conditions. Yet, 
very few of these varieties are effectively resistant to Striga, because, 2.2 | Biology and Striga spp. life cycle
they are continuously tolerant to the emergence of Striga plants. Thus, 
adding each year more Striga seeds into the soil after each growing Life cycle of Striga is synchronized to that of its host and in-
season. Therefore, additional genes or sources of Striga resistance volves mechanisms that coordinate lifecycles of both the para-
need to be found for introgression into maize elite varieties in order site and the host (Bouwmeester et al., 2003). Striga life cycle 
to develop varieties that support little or no Striga emergence. This re- generally involves: germination, host attachment, formation of 
view intends to give a brief update on current work towards Striga re- haustoria, penetration and establishment of vascular connec-
sistance emphasizing breeding methods for Striga resistance in Africa tions, nutrients accumulation, flowering and seed production 
and the use of integrated Striga control mechanisms on maize. (Parker & Riches, 1993) (Figure 2). Germination of Striga seeds 
F I G U R E  1   Maize field devastated by S. hermonthica in the North of Benin Republic 
Source: Yacoubou (2018) 
YACOUBOU et Al.      |  197
F I G U R E  2   The life cycle of S. 
hermonthica on a susceptible host. 
Stages indicated: A = after-r ipening and 
conditioning of S. hermonthica seed, 
B = germination of S. hermonthica seed, 
C = haustorial initiation and attachment 
of S. hermonthica to the host followed 
by a period of growth underground, 
D = emergence of S. hermonthica plants 
from the soil, E = flowering, insect 
pollination, seed set and dispersal. 
Duration of each phase of the life cycle is 
indicated.
Source: Hearne (2001) [Colour figure can 
be viewed at wileyonlinelibrary.com]
depends on the presence of hormones known as strigolactones Cultural practices such as manual weeding, push and pull, crop 
that are produced by the host and in other cases non- host species rotation with non- host intercrops (trap crops), fertilizer application, 
(Spallek et al., 2013). With the presence of strigolactones, para- soil and water management, and transplanting have been attempted, 
site seedlings attach to the host and form vascular connections but they offered limited success in controlling Striga infestation 
depriving it of its water, carbohydrates and minerals (Yoshida & (Oswald & Ransom, 2002; Fasil & Verkleij, 2007; Udom et al., 2007; 
Shirasu, 2009). Under stressful conditions plant roots exude str- Manyong et al., 2008; Ayongwa et al., 2010; Lagoke & Isah, 2010; 
igolactone hormone to promote symbiotic relationship with soil Hailu et al., 2018). Inter- cropping cereals with legumes is another 
microbes for mineral nutrient scavenging (Steven, 2014). Parasitic low-c ost and viable strategy that has been reported to influence 
plants such as Striga hermonthica have exploited these strigolac- Striga spp. infestation (Carsky et al., 2000; Akanvou et al., 2006; 
tone hormones as signals to stimulate the germination of their Kanampiu et al., 2018). Legumes, through their roots, fix atmo-
seeds (Runo et al., 2012) (Figure 3). During early stages of seed de- spheric nitrogen, add organic matter to the soil by contributing to 
velopment, before emergence, the parasite depends totally on the soil conservation, preserving the streamline soil moisture and en-
host plant (Webb & Smith, 1996). At this stage of subterranean de- hances soil biodiversity, thereby improving soil health and fertility, 
velopment, S. hermonthica inflicts maximum damage to the maize which directly contributes to Striga control. Intercropping legumes 
plant. The adverse effect of Striga on maize is manifested as stunt- with cereals reduces S. hermonthica but does not eliminate the para-
ing, chlorotic and necrotic lesions on the leaves and reduction of site (Khan et al., 2000, 2007).
ear size and grain yield (Adetimirin et al., 2000). Striga spp. take Other methods for Striga control include biological control using 
about 4–1 0 weeks to complete its life cycle after emergence and herbicide- resistant maize variety (Imazapyr treatment), develop-
this completion usually occurs after harvest of the host (Ramaiah ment of Striga- resistant germplasm, use of fungus Fusarium isolation 
et al., 1983). by applying strigolactones (Kanampiu et al., 2002; Ejeta, 2007; Illa 
et al., 2010; Nzioki et al., 2016; Uraguchi et al., 2018; Zwanenburg, 
& Blanco- Ania, 2018; Kountche et al., 2019). All these approaches 
2.3 | Striga control methods have been used with some degree of success to minimize the effect 
of Striga in maize production. The mode of action for each approach 
Striga control is essential to ensure food security in the SSA is different. For example, in the case of fungus, when F. oxysporum 
(Ejeta, 2007; Rodenburg et al., 2005). Several methods, ranging gets in contact with maize plants, there is a production of amino 
from agricultural practices to biological control exist and significant acids (L- leucine and L- tyrosine), that disrupt plant growth and de-
progress has been made in Striga control research within Africa velopment. These amino acids are toxic to Striga plants but innoc-
(Table 1). uous to maize plants (Nzioki et al., 2016). The use of this biological 
198  |     YACOUBOU et Al.
F I G U R E  3   Biological functions of 
strigolactones 
Source: Yamaguchi et al. (2010) [Colour 
figure can be viewed at wileyonlinelibrary.
com]
control tool allowed the increment of more than 45% maize yield in translocation suggests a possibility that RNA- interference (RNAi) 
Striga endemic zones in Kenya (Nzioki et al., 2016). Strigolactones could be used as a potential tool to interfere in vital processes within 
(SLs) reduce the accumulation of abscisic acid (ABA) in plant by the parasite by transforming the host with an RNAi construct that 
up- regulating the ABA catabolic enzyme gene CYP707A1 (Lechat targets gene sequences specific to the parasite (Shayanowako et al., 
et al., 2015; Toh et al., 2012). The ABA is released by maize infected 2017). This technique is constrained by the lack of genes to target 
with S. hermonthica, that subsequently trigger stomatal closure to for silencing as well as by the delivery of iRNAs into the parasite 
minimize water loss. SLs also increase the production of gibberellins (Kirigia et al., 2014). This constrain can be overcome using viral in-
(GA) hormones by up-r egulating gibberellin3β- dioxygenase 1, which duced gene silencing (VIGS). Using a Tobacco Rattle Virus (TRV) – 
is involved in GA biosynthesis (Toh et al., 2015; Yao et al., 2016). VIGS system, Kirigia et al. (2014) have shown that this system works 
Although ABA and GA represent central plant hormones and are in S. hermonthica and has been proven as a useful system for candi-
known to antagonistically regulate seed germination in non-p arasitic date gene validation either in parasite development or parasitism, for 
plants, the effects of their exogenous application vary across para- the development of resistant transgenic maize.
sitic plant species. Zehhar et al. (2002) and Toh et al. (2015), reported 
that neither GA nor ABA alone is sufficient to stimulate or inhibit 
seed germination in S. hermonthica, while Kannan and Zwanenburg 3  | GENETIC S RESISTANCE MECHANISMS 
(2014) and Zwanenburg et al. (2016) reported SLs application appears TO Str iga IN MAIZE
attractive owing to their decomposition in the soil within a short pe-
riod. Nevertheless, the use of natural SLs for decomposition in soil 3.1 | Resistance mechanism to Striga in maize
does not seem a realistic alternative because the synthesis of these 
compounds is very labourious. More recently, genetic engineering Striga resistance mechanisms act either before (preattachment) or 
has offered the promise of rapidly achieving resistance against Striga after physical contact with the host (postattachment). Preattachment 
spp. Recent findings have shown that RNAs freely translocate be- resistance (Figure 4a) occurs when a host produces low amounts of 
tween parasitic plants and their hosts (Kim & Westwood, 2015). This strigolactones or when Striga receptors that perceive germination 
YACOUBOU et Al.      |  199
TA B L E  1   Striga management methods used in African countries
Factors in favour of control 
Methods options Setbacks for control options References
Manual weeding Reduction of Striga seed Yield benefit is not immediate, Babiker (2007), Ayongwa et al. (2010)
bank, easy to implement labour intensive
Crop rotation Increase soil fertility, Benefit accruement requires time, costly as Carsky et al.(2000), Manyong 
reduction of Striga seed per family food et al. (2008)
bank
Hand pulling Reduction of Striga seed Inappropriate disposal increases seed bank Jamil et al. (2011), Oswald (2005)
bank if performed before 
flowering, increase in yield
Push and pull Provide livestock feed, Costly to implement initially, Khan et al. (2010), Hailu et al. (2018)
reduction of Striga seed benefit accruement requires time, trap crop 
bank, control of stem borer, used uneconomical
improvement of soil fertility
Fertilizer Increase in yield, Costly to implement, labour Jamil et al. (2012)
Application (N improvement of soil fertility, Intensive
and P) reduction of Striga incidence
Intercropping with Reduction of Striga seed Labour intensive, trap crop used Bilalis et al. (2010), Ibrahim et al. (2014), 
Legumes bank, increase soil fertility, uneconomical Hailu et al. (2018)
provide additional income
Seed dressing Increase in yield, easy to Purchase of seed every season is costly De Groote et al. (2008), Kanampiu et al.
(herbicide) implement, Reduction of May not be easy to implement (2003)
Striga incidence
Compost Increase in yield, easy to Increase pests, labour intensive Osman et al. (2013)
application implement, reduction of 
Striga incidence, increase 
soil fertility
Resistant Easy to implement, high crop Purchase of seed every season is costly, gene Kouakou (2014), Naitormmbaide 
Varieties yield recombination's in the parasite (mutation), et al. (2015)
limited of resistance sources
Herbicide Reduction of Striga seed Unavailable to farmers, cost Hesammi (2013); Ransom et al. (2012)
Application bank prohibitive
Biocontrol agent Reduction of Striga Labour intensive, source limited Khan et al. (2010), Nzioki et al. (2016), 
emergence, Crop uneconomical to farmers Kountche et al. (2019)
improvement of crop yield without livestock
Reduction of Striga 
incidence, increase yield, 
provide livestock fed
Integrated Suppressing emergence and Failure of the host's rhizosphere to maintain Ouédraogo et al. (2018), Shayanowako 
approach: fecundity, germination and enough pathogen levels that guarantee et al.(2018), Zarafi et al. (2015)
biocontrol agent photosynthetic inhibition control of the weed.
and resistant 
varieties
Integrated Striga Reduction Striga emergence, Low adoption of these varieties, purchase of Randrianjafizanaka et al. ( 2018), Ronald 
Management: reduction Striga infection seed every season is costly, unavailability of et al. (2019), Schut et al. (2015)
agronomic levels and seed numbers in resistant varieties to Striga species attacks, 
practices and the soil, Increase in yield Mutation or geographical changes that 
resistant varieties occur over a number of years
stimulants are insensitive to the strigolactone levels produced by the resistance mechanism produce relatively low SLs, thereby inducing 
host (Lumba et al., 2017; Mutinda, 2018). Binding causes the deg- the germination of less parasitic seeds and consequently prevent 
radation of an F- box protein, which in turn activates gene regula- the host plant from parasitism. Preattachment resistance has been 
tory processes that lead to Striga germination (Lumba et al., 2017). shown in 'KSTP’94', an open- pollinated maize variety used by farm-
It can also be due to the production of low haustorial initiation fac- ers in Eastern Africa for S. hermonthica management. This maize vari-
tors whose effect leads to a failure by Striga to develop haustorium ety was shown to produce low amounts of sorgomol, a strigolactone 
effectively (Rich et al., 2004). Crop genotypes with preattachment that does not efficiently induce S. hermonthica germination (Karaya 
200  |     YACOUBOU et Al.
F I G U R E  4   Mechanisms of resistance to S. hermonthica in maize 
Source: Amusan et al. (2008) 
et al., 2012). This resistance, qualified as phenotypic resistance, been shown in the 'KSTP’94', maize open- pollinated variety (OPV) 
has been identified in other Striga- resistant crop genotypes (Jamil (Mutinda et al., 2018). However, the molecular mechanisms under-
et al., 2011; Robert, 2011). However, resistance associated with low lying postattachment Striga resistance are unknown. Preference 
production of Striga seeds germination stimulant may not be related for OPV is most likely due to the prohibitive price of hybrids or lack 
to low production of total strigolactones, but rather to the types of of availability of hybrid seed in some SSA countries (Badu- Apraku 
strigolactones released (Yoneyama et al., 2010). & Fakorede, 2017). In addition, these OPV’s are more affordable 
In contrast, postattachment mechanisms act after Striga has and consequently easy to multiply and readily available (Midega 
attached and attempted to penetrate the host (Figure 4b). These et al., 2016).
mechanisms result in physiological or biochemical barriers, that Although hybrids are known and desirable for their high produc-
prevent Striga haustorium from connecting to the host xylem (van tivity and quality, they have shown reduced pathogen resistance 
Dam & Bouwmeester, 2016). Striga hermonthica postattachment re- compared to the OPVs which have innate defence traits (Schroeder 
sistance in maize has mainly come from its wild- grass relatives Zea et al., 2013). It is, therefore, vital to understand the genetic make- up 
diploperennis (Amusan et al., 2008; Lane et al., 1997) and Tripsacum of the parents used to develop hybrids as this would be more use-
dactyloides (Gutierrez- Marcos et al., 2003). Post attachment re- ful for further development of improved maize germplasm with en-
sistance in maize expressed by an incompatibility in ZD05 inbred hanced resistance to S. hermonthica.
line with S. hermonthica has been observed (Amusan et al., 2008, 
2011). In these cases, the parasite penetrated host cortex but was 
prevented from getting into the host endodermis. The exact mech- 3.2 | Potential sources of Striga resistance in maize
anism for this parasite's inability to penetrate the endodermis re-
mains unknown. This resistance could be attributed to biochemical Genetic improvement for Striga resistance depends on the avail-
or physiological barriers from the host (Amusan et al., 2008; Yoshida ability of germplasm sources with different levels of resistance. 
& Shirasu, 2009). Recently, postattachment Striga resistance has Therefore, resistance is prioritized in maize breeding programmes 
YACOUBOU et Al.      |  201
for regions where Striga is endemic and causes major yield losses with the desired trait and adapted to various agro-e cological regions. 
to farmers. The sources of resistance to Striga have been identi- Due to Striga proneness in Eastern Africa, maize genotype 'KSTP’94' 
fied in maize and other crops such as rice, sorghum and cowpea has been developed and deployed as Striga tolerant source especially 
(Amusan et al., 2008; Haussmann et al., 2004; Mbuvi et al., 2017; in Western Kenya (Mutinda et al., 2018). 'KSTP’94' exhibits remark-
Menkir, 2006; Yonli et al., 2006) (Table 2). able resistance to Striga under field conditions; a characteristic that 
Striga resistance in maize could be sourced from wild-g rass has made it a subject of intense research in the region as well as in 
relatives like Zea diploperennis and Tripsacum dactyloides (Amusan research to understand the mechanism of Striga resistance in maize. 
et al., 2008; Gutierrez- Marcos et al., 2003; Lane et al., 1997). Such Karaya et al. (2012) and Midega et al. (2016), have identified maize 
efforts have led to the development of Striga-r esistant inbred line landraces that are less affected by Striga hermonthica comparatively 
ZD05 suitable for integration in breeding programmes in Western to hybrids in Western Kenya. These results provide an insight into 
Africa (Kim, 1991). Integrating this breeding line into the breeding the potential role of landraces which could play an important role in 
programme, IITA in collaboration with National Agricultural Research the efforts towards an integrated management approach for Striga 
Systems (NARS) have focused on developing new maize genotypes in smallholder cropping systems. The potential genetic variability for 
TA B L E  2   Potential sources of Striga resistance
Germplasm Source Name Institution References
Wild- maize genes for inhibition of Tripsacum dactyloides, Linea IITA Gurney et al. (2018)
relatives Strigahaustorial development
Resistance Zea diploperennis, Doebley et Amusan et al. (2008)
Guzman
Landraces horizontal resistance Broad base KARI Midega et al. (2016)
Inbred lines Resistance/tolerance TZi 3 (1368 STR), IITA Kim and Akintunde 
TZi 25 (9450 STR) (1989), Konate 
et al. (2017), Menkir 
et al. (2006)
9030, 1393, TESTR151, TESTR CIMMYT Karaya et al. (2014)
156, OSU231//56/44-6 - 4-1 7-3 KARI
Resistance TZill, TZil2, TZi25 TZi30 IITA
TZEIOR 108, TZEI 10, TZEI 17
TZISTR1174, TZISTR1162, IITA, Uganda National Simon et al. (2018)
TZISTR1192 Crop Resources 
Research Institute,
OPV IITA populations TZL comp1 synw- 1 and Acr94TZE IITA Menkir and Kling (2007)
Comp s- w
Resistance/tolerance TZEE- W Pop STR, TZEE- Y Pop IITA Makumbi et al. (2015), 
STR, 2004 TZEE-Y  Pop STR Menkir, Franco, 
C4, TZEE- W Pop STR QPM C0 et al. (2012), Oyekunle 
and TZEE- W Pop STR BC2 C0; et al. (2017)
TZEE- W STR 107 BC1, TZEE- W 
Pop STR C5, 2012 TZEE- Y DT 
STR C5
Striga postattachment Resistance KSTP 94, STR- VE- 216 KALRO Mutinda et al. (2018)
CIMMYT
Hybrids Resistant and tolerant PHB3253, PHB30G19, PHB30B50 Pioneer Chitagu et al. (2014)
Resistant and tolerant MH1416, MQ623, SC643, SC527, Seed Co Nyakurwa et al. (2018)
SC535 Mukushi Seeds
Resistance/Tolerance TZISTR1162 × TZISTR1198 Uganda National Crop Simon et al. (2018)
TZISTR1199 × TZISTR1181 Resources Research 
TZISTR1192 × 1368STR Institute
Tolerance 8322- 13, 8321- 18 and 9022- IITA Kim and 
13, TZEIOR 57 × TZEIOR Akintunde (1989), 
108, TZEIOR 57 × TZEIOR Konate et al. (2017)
127, TZEIOR 13 × TZEIOR 
59, TZEIOR 57 × TZEI 10 and 
TZEIOR 127 × TZEI 10
Abbreviations: KALO, Kenya Agricultural Livestock Research Organization (Kakamega, Kenya); KARI, Kenya Agricultural Research Institute.
202  |     YACOUBOU et Al.
S. hermonthica resistance can be harnessed from wild- grass relatives, in S. hermonthica, namely ShCCD7 and ShCCD8 has been provided 
open pollinated, inbred as well ashybrids lines (Table 2). (Liu et al., 2014). In tobacco, the silencing of CCD7 and CCD8 genes 
Promising Striga resistance genotypes have been identified for has delayed the virus parasite formation in the host, indicating that 
further testing and experimental releases in African countries under these two genes are a key in the parasitic life cycle (Aly et al., 2014). 
projects such as Stress Tolerant Maize for Africa (STMA). Recently, some significant loci on chromosomes 9 and 10 of maize 
that are closely linked to ZmCCD1 and amt5 genes, respectively, and 
may be related to plant defence mechanisms against Striga parasit-
3.3 | Genetics resistance to Striga ism have been identified (Adewale et al., 2020).
Availability of all this information on the type of gene action 
Information on the genetic basis of resistance to Striga is critical for governing the inheritance of resistance to Striga in maize genotypes 
plant breeding and selection. Genes action for grain yield and other would, therefore, contribute to the introgression of resistance genes 
agronomic traits have been reported for maize under Striga infesta- and dissemination of resistant genotypes (Akanvou & Doku, 1998).
tion (Ejeta et al., 1997). Resistance evaluation is based on grain yield 
under Striga infestation, number of Striga plants emerged on the 
host and host damage syndrome rating. However, there have been 4  | METHODS FOR SCREENING Str iga 
contradictory reports on the gene action controlling Striga resist- RESISTANCE IN MAIZE
ance in maize. It is quantitatively inherited with additive gene effects 
being more important than non-a dditive effects. This contributes to Development of Striga- resistant cultivars has been limited by the 
regulating the host plant damage syndrome rating and grain yield lack of dependable screening techniques (Yagoub et al., 2014). 
under Striga infestation (Kim, 1994; Berner et al., 995; Akanvou Some of the screening techniques that have been used include 
et al., 1997). As reported by Kim (1994) and Berner et al. (1995) dif- field techniques, screen house and laboratory methods (Rodenburg 
ferent genes control the number of emerged Striga plants and the et al., 2015).
level of host plant damage. Moreover, there is evidence that additive Field screening is an artificial technique that consists of uniform 
gene action has a higher contribution to natural gene action with infestation with Striga using appropriate experimental design. The 
regards to grain yield and Striga traits in maize (Akaogu et al., 2013; procedure of this technique has been described in detail by Badu- 
Badu-A praku et al., 2015, 2016; Menkir et al., 2010). In contrast, Apraku and Fakorede (2017). Confounding effects of environmen-
other studies reported that the impact of non-a dditive genes is more tal conditions on the polygenic inheritance of traits associated with 
important than the effect of additive genes in the control of the in- Striga resistance make field screening indispensable despite the ad-
heritance of host plant damage, while the effect of additive genes vances made in laboratory and at pot experiments stage.
is more important in the control of the number of emerged Striga Screen house technique has been used to screen maize gen-
plants (Gethi & Smith, 2004; Badu- Apraku et al., 2007; and Yallou otypes for tolerance / resistance to Striga (Chitagu et al., 2014; 
et al., 2009). A recent study reported that the dominant effects sur- Nyakurwa et al., 2018; Yohannes et al., 2016). In screen houses, 
pass the additive effects for the number of emerged Striga plants screening for varietal resistance has been performed using pots 
and inheritance of Striga resistance in maize may be conditioned and buried seed studies (Eplee & Norris, 1987; Rao, 1985; Sand 
by non-a dditive gene action (Akaogu et al., 2019). Additionally, the et al., 1990). With regard to the pot screening techniques ‘poly bag’ 
involvement of epistatic effects in the inheritance of Striga resist- and seed pan, and the ‘Eplee bag’ are used (Eplee, 1992; Rao, 1985). 
ance aa in maize has been reported (Adetimirin et al., 2001; Akaogu The most important aspect in screen house evaluation is its compat-
et al., 2019). Unlike maize, the progress in the identification of genes ibility with experiments on the efficiency in controlling the Striga 
for marker-a ssisted selection in other crops such as sorghum and vector (Kountche et al., 2019). Several studies have also demon-
rice is substantial. The identification of lg gene mutant alleles at strated the validity of the Eplee bag technique as a good screening 
the LGS1 (Low Germination Stimulant 1) locus on chromosome 5 of method (Ahonsi et al., 2002; Yonli et al., 2006). Previously, pot ex-
sorghum has reduced significantly the S. hermonthica germination periments were used to access the level of parasite variation in the 
stimulant activity (Gobena et al., 2017). This gene was found to code attachment to the roots of diverse maize inbred lines alongside the 
for a sulfo- transferase enzyme and when silenced led to a change plant host interaction (Menkir et al., 2006).
in 5-d eoxystrigol into orobanchol compounds in the root exudates Laboratory methods employed in Striga research have proven 
(Gobena et al., 2017). In addition, other loci have been reported to be the best option so far for screening infection. The use of 
to play important roles in parasitic resistance, including the genes laboratory-b ased assays has provided interactive biological pro-
CCD1, CCD7, CCD8, DAD2, MAX1, DWARF 53 (D53) and LBO (Sun cesses between Striga and the roots of the host plants during each 
et al., 2008; Hamiaux et al., 2012; Zhou et al. 2013; Aly et al., 2014; individual stage of the parasitism process. Hess et al. (1992) devel-
Zhang et al, 2014; Brewer et al., 2016). In maize, roots with mycorrhi- oped an in vitro laboratory assay termed such as the agar gel assay 
zal formations have shown a higher ZmCCD1 expression and induced (AGA) to determine the genotypic efficacy of host root exudates 
lower germination of Striga (Sun et al., 2008). Evidence for strigo- to germinate preconditioned Striga seeds. This system gave a good 
lactones and strigolactone perception genes of the MAX-2 - type correlation with field resistance (Hess et al., 1992; Ramaiah, 1987). 
YACOUBOU et Al.      |  203
These growth systems have been used to examine the architecture of susceptible cultivars (Menkir et al., 2004). It is, therefore, relevant to 
host roots and their biochemical mechanisms of resistance (Amusan explore the applicability of many conventional breeding techniques 
et al., 2011; Mohamed et al., 2010; Mrema et al., 2017). Kountche generally used in various Striga resistance- breeding programmes.
et al. (2019) used AGA to assess the germination-i nducing activity Recurrent selection is designed to increase the frequency 
of selected strigolactones (SLs) analogues on S. hermonthica seeds. of favourable alleles in a population (Hallauer, 1992; Hallauer & 
AGA is useful for screening maize genotypes with a high degree of Carena, 2012; Badu- Apraku & Fakorode, 2017). This procedure has 
success in identifying Striga- resistant varieties especially those em- been used effectively in maize to improve quantitatively inherited 
anating from the wild-s pecies relatives such as Z. diploperennis and T. traits (Badu- Apraku, 2010; Menkir & Kling, 2007). Few studies have 
dactyloides (Amusan et al., 2011; Gurney et al., 2003, 2006; Karaya been conducted on the effectiveness of recurrent selection in im-
et al., 2012 ). More recently, AGA experiments have been used to de- proving the level of Striga resistance in maize (Menkir & Kling, 2007). 
termine the levels of resistance or tolerance of new quality protein Recurrent selection methods capitalize on additive gene action under 
maize genotypes to S. asiatica (Nyakurwa et al., 2018). an effective and reliable artificial method of Striga infestation for 
Furthermore, the rhizotron screening system has been proposed the screening of progenies. It facilitates the accumulation of Striga 
as an ideal technique to circumvent the limits of field technique and resistance genes to develop germplasm with multigenic resistance 
initiate a reliable postattachment screening (Rodenburg et al., 2015). that could be sustainable over time and effective for the control of 
Rhizotrons are transparent root observation chambers that enable the parasitic weed (Badu- Apraku et al., 2012; Menkir & Kling, 2007). 
Striga attached to the host plant to be counted. The AGA technique Recurrent selection has been used successfully to improve grain 
also allows the evaluation of resistance mechanisms phenotype and yield and other agronomic traits in maize populations under infes-
determination of the effect of Striga on host biomass over a period tation (Badu-A praku, 2010; Menkir et al., 2004). Through recurrent 
of time with minimal disturbance (Rodenburg et al., 2015; Runo selection, researchers have reported genetic gains in maize grain 
et al., 2012). Rhizotron Perspex chambers have been extensively yield cultivars under Striga infestation. Menkir et al. (2004) observed 
used to screen a variety of host species including maize (Mutinda that over 2 years selection, Striga damage symptoms were reduced 
et al., 2018). by 3% per cycle, number of emerged Striga plants by 10% per cycle 
and grain yield increased by 16% per cycle under Striga infestation 
conditions. Within two periods of selection (1988– 2000 and 2001– 
5  | BREEDING APPROACHES USED FOR 2006), recurrent selection improved the annual gain yield from 
Str iga RESISTANCE IN MAIZE 0.86% to 2.11% in early maize under Striga infestation (Badu- Apraku 
et al., 2013). This approach has led to an increase in genetic gains in 
Considerable efforts have been made in breeding for Striga resist- grain yield of 498 kgha−1 cycle−1 (16.9% cycle−1) in 3 years (2014– 
ance in cereals especially in maize and significant progress has been 2017) under Striga infestation (Badu- Apraku et al., 2019). More re-
achieved in the development of improved varieties. After the iden- cently, genetic gains in maize grain yield and other agronomic traits 
tification of a potential source of resistance, the next critical step under Striga condition for periods of selection have been reported. 
in the breeding programme depends on the breeder's ability to in- Using recurrent selection, traits associated with grain yield includ-
corporate the resistance genes into the best-a dapted varieties. This ing plant height and the number of ears per plant increased, ear as-
can be performed with several strategies, amongst which are the pect and anthesis– silking interval decreased over time under Striga 
conventional and or classical breeding and the marker- assisted se- condition (Menkir & Meseka, 2019). The authors observed that on 
lection (MAS). average, hybrids developed after the 1990s yielded 64% more and 
displayed 61% less parasite emergence and 30% less parasite dam-
age at 10 weeks after planting compared with hybrids developed 
5.1 | Conventional breeding for Striga resistance before the 1990s.
The half- sibling selection scheme is also one of the easiest ways 
Conventional plant breeding aims at increasing the chances of se- in developing composite populations with at least moderate re-
lecting individuals from populations generated from genetic mating sistance to S. hermonthica (John & Sleeper, 1995). The full sib and 
designs. Selection has usually been carried out at the whole- plant selection from S1 progeny tests allows for an increased scope of 
level thereby, representing the net result of the interaction between variability in progeny from source populations and greater control 
genotype and environment (Badu-A praku et al., 2017). However, over pollen, and should translate into an increased frequency of fa-
identification of potential sources of resistance is the first step of all vourable alleles for Striga resistance in populations under selection 
Striga breeding programmes. To access the genes for resistance and (Hallauer, 1992; Menkir et al., 2004).
incorporate them into well-a dapted varieties, conventional breed- The backcross breeding procedure is straight forward if a 
ing relies on techniques such as recurrent selection, half- sib or full- source population or donor, with a high frequency of desirable 
sib selection, S1 family and F1 family (hybrid) selection schemes. alleles for Striga resistance is available. Therefore, rapid progress 
Conventional breeding techniques were predominantly used in con- can be achieved in building resistance to Striga if a donor exhibit-
ferring superior combinations of Striga resistance alleles amongst ing high dominance for Striga resistance genes is identified. Under 
204  |     YACOUBOU et Al.
such condition, ideal recurrent parents would be genotypes com- Several researchers have reported the efficiency and superi-
bining early maturity and high yield (Badu- Apraku et al., 2017). ority of MAS and its effective integration into mainstream maize 
Germplasm derived through the backcross method forms the basis breeding programmes. Efforts deployed with the use of molecular 
for cultivar advancements towards achieving polygenic resistance tools can be utilized in determining families that can be bulked or 
to S. hermonthica. Such inbred from Z. diploperennis and tropical discarded. Those families could also help in the selection of parental 
maize have been essential in the development of S. hermonthica- lines for Striga- resistant hybrids development with high yields and 
resistant open- pollinated populations like Zea diplo SYNW- 1, TZL stable across many agroecologies (Akinwale et al., 2014; Mengesha 
Comp SYNW- 1. Partial resistance to S. hermonthica was also ob- et al., 2017).
served in backcross hybrids from a resistant donor T. dactyloides Molecular marker technologies and the construction of genetic 
(Gurney et al., 2018). linkage maps have made it possible to detect genetic loci associated 
Despite the low costs and yield stability benefits associated with with complex traits (Kang et al., 1998; Sibov et al., 2003). Genetic 
the recurrent use of synthetic maize populations, the superiority linkage maps and quantitative trait loci (QTL) mapping technology 
in performance of hybrid cultivars is being acknowledged with an have enhanced the efficiency of estimating the number of loci con-
increasing trend amongst southern African farmers (Badu-A praku trolling genetic variation in a segregating population and the charac-
& Fakorede, 2017). The desire to increase maize yields under mar- terization of the map positions in the genome (Xiao et al., 1996). In 
ginal growing conditions and a rise in literacy can be the major rea- maize, QTLs identification was focused mainly on abiotic and biotic 
sons behind the increase towards the complete adoption of hybrid stresses such as drought tolerance (Semagn et al., 2015; Tuberosa 
technology in countries like Zimbabwe, Ghana, Mali and Nigeria et al., 2002), low soil nitrogen (Mandolino et al., 2018; Ribeiro 
(STMA, 2019). Heterosis of hybrid varieties can be useful in miti- et al., 2018), pests (Jiménez- Galindo et al., 2017) and foliar dis-
gating the effect of Striga on maize productivity. With the increased eases (Gowda et al., 2018). In SSA, little progress has been reported 
use of hybrid maize seed in West and Central Africa (WCA), Menkir on the detection of QTLs or genes for Striga resistance in maize. 
et al. (2004) have selected S. hermonthica- resistant hybrids by cross- However, QTLs for resistance to S. hermonthica have been identi-
ing diverse inbred lines. These hybrids are able to suppress para- fied from local populations including wild relatives and successfully 
site emergence, with some of them producing high grain yield under transferred through backcross breeding into adaptable maize pop-
high Striga infestation levels (Menkir et al., 2012b). However, multi- ulations (Rich & Ejeta, 2008). Using the linkage mapping method, 
location field screening for Striga resistance resulted in significant two putative QTLs have been discovered that govern incompatible 
genotype × environment (G × E) interactions for Striga resistance response to Striga parasitism in maize amongst F2 segregated popu-
traits in maize trials (Akinwale et al., 2014; Nyakurwa et al., 2018; lations (Amusan, 2010). Whereas some QTLs have been discovered 
Simon et al., 2018). Based on these results, there is a need to select for Striga resistance in sorghum and rice (Atera et al., 2015; Yasir & 
for specific adaptation in Striga resistance breeding, particularly in Abdalla, 2013; Yohannes et al., 2015; Ali et al., 2016). Using genomic 
the case of contrasting environment where different putative Striga association wide (GWA), 24 SNPS markers associated with grain 
ecotypes may exist. yield, Striga damage at 8 and 10 weeks after planting (WAP), ears per 
plant and ear aspect under Striga infestation were detected in early 
maturing maize inbred (Adewale et al., 2020). Therefore, there is an 
5.2 | Marker-a ssisted breeding for Striga resistance urgent need to identify QTLs for Striga resistance to facilitate the 
rapid and efficient transfer of the genes into other maize genotypes.
Marker- assisted selection (MAS) is an indirect selection process 
where a trait of interest is selected based on a marker linked to the 
trait, rather than on the trait itself (Ribaut et al., 2001). This breed- 6  | WAY FORWARD ON Str iga RESISTANCE 
ing method allows the performance of a selected phenotype to be IN MAIZE AND CONCLUSION
predicted based on the use of molecular markers at early generation.
Application of molecular markers has provided significant op- Breeding maize for Striga resistance is challenging due to the scarcity 
portunities for breeders to characterize, evaluate and select maize of resistant sources in cultivated species. In this review, we explored 
germplasm widely used by public and private sectors. Molecular the integrated approach using resistant cultivars is the most effective 
markers are also used for screening crop genotypes for tolerance option, since Striga- resistant cultivars play a major role in reducing 
to biotic or abiotic stress. Using SSRs and SNPs markers, some elite Striga pressure, both in terms of Striga count and vigour compared 
genotypes for the breeding of Striga resistance are selected and with individual control options. In general, many breeding tech-
new makers have been identified, which significantly contributed niques are used in maize breeding programmes for Striga resistance. 
to the differentiation of Striga tolerant and susceptible genotypes However, conventional breeding techniques through the screening of 
(Bawa et al., 2015; Shayanowako et al., 2018). Molecular markers resistant genotypes are the most frequently used in the maize breed-
can better help in the assessment of relatedness in isogenic lines to ing programmes in Africa. Screening of resistant genotypes under ar-
determine families that can be bulked or discarded, which in turn can tificial Striga infestation is very expensive, time- consuming and labour 
reduce maintenance costs (Dean et al., 1999). intensive. Moreover, obtained results are often not consistent due to 
YACOUBOU et Al.      |  205
genotype by environment interactions, inability to assess evenness Adewale, S. A., Badu- Apraku, B., Akinwale, R. O., Paterne, A. A., Gedil, 
of Striga distribution and ascertain contact between Striga and host M., & Garcia- Oliveira, A. L. (2020). Genome- wide association study 
of Striga resistance in early maturing white tropical maize inbred 
roots. lines. BMC Plant Biology, 20, 203.
Another possibility is to develop high yielding maize genotypes Ahonsi, M. O., Berner, D. K., Emechebe, A. M., & Lagoke, S. T. (2002). 
with resistance to Striga using genome editing of SLs genes, which Selection of rhizobacterial strains for suppression of germination 
are responsible for Striga germination and attachment. It might be a of Striga hermonthica (DEL.) Benth. seeds. Biological Control, 24(2), 
143–1 52. https://doi.org/10.1016/S1049 - 9644(02)00019-  1
direct way of increasing maize grain yield in Striga endemic locations Akanvou, L., Doku, E. V., & Kling, J. G. (1997). Estimates of genetic 
of SSA. At present, accumulation of resistance QTLs in most pro- variances and interrelationships of traits associated with Striga re-
grammes may be facilitated by conventional breeding techniques and sistance in maize. African Crop Science Journal, 5, 1–8 . https://doi.
the use of cost- effective molecular markers (Badu- Apraku & Fakorede, org/10.4314/acsj.v5i1.27864
Akanvou, L., & Doku, E. V. (1998). Heritability of traits associated with 
2017). The present challenge is to convert a large amount of avail- Striga (Striga hermonthica (Del.) Benth.) resistance in an open polli-
able genetic information into a large set of markers useful for Striga nated maize population. African Crop Science Journal, 6(2), 129– 135. 
resistance breeding in maize and to integrate such markers into a sus- https://doi.org/10.4314/acsj.v6i2.27808
tainable breeding scheme. Further exploration of closely related QTL Akaogu, I. C., Badu- Apraku, B., Adetimirin, V. O., Vroh- Bi, I., Oyekunle, 
gene markers related to Striga will help in the effective trait pyramid- M., & Akinwale, R. O. (2013). Genetic diversity assessment of extra- 
early maturing yellow maize inbreds and hybrid performance in 
ing gene actions that can contribute to maize effective production. Striga-i nfested and Striga- free environments. Journal of Agricultural 
Some effective molecular docking approaches such as CRISPR/Cas9 Science, 151(4), 519–5 37. https://doi.org/10.1017/S00218 5961 
genome editing of strigolactone genes, which are responsible for Striga 2000652
germination and attachment could also be considered for the develop- Akaogu, I. C., Badu-A praku, B., Tongoona, P., Ceballos, H., Gracen, 
V., Offei, S. K., & Dzidzienyo, D. (2019). Inheritance of Striga her-
ment of high yielding maize genotypes with resistance to Striga. monthica adaptive traits in an early- maturing white maize inbred line 
containing resistance genes from Zea diploperennis. Plant Breeding, 
ACKNOWLEDG EMENTS 138(5), 546– 552. https://doi.org/10.1111/pbr.12707
Laboratoire de Phytotechnie, d'Amélioration et de Protection Akinwale, R. O., Badu- Apraku, B., Fakorede, M. A. B., & Vroh- Bi, I. (2014). 
Heterotic grouping of tropical early- maturing maize inbred lines 
des Plantes (LaPAPP), Département des Sciences et Techniques based on combining ability in Striga- infested and Striga-f ree envi-
de Production Végétale, Faculté d'Agronomie de l'Université de ronments and the use of SSR markers for genotyping. Field Crops 
Parakou and the Stress Tolerant Maize for Africa Project are highly Research, 156, 48–6 2. https://doi.org/10.1016/j.fcr.2013.10.015
acknowledged. The authors express their appreciation to all scientist Ali, R., Hash, C. T., Damaris, O., Elhussein, A., & Mohamed, A. H. (2016). 
Introgression of striga resistance into popular Sudanese sorghum va-
involved in the improvement of this manuscript. We acknowledge rieties using marker assisted selection. World Journal of Biotechnology, 
financial support for the Bill and Melinda Gates Foundation. 1, 48–5 5. www.scipl atform.com/wjb
Aly, R., Dubey, N. K., Yahyaa, M., Abu- Nassar, J., & Ibdah, M. (2014). Gene 
CONFLIC T OF INTERE S TS silencing of CCD7 and CCD8 in Phelipanche aegyptiaca by tobacco 
rattle virus system retarded the parasite development on the host. 
The authors declare that there is no conflict of interests. Plant Signaling & Behavior, 9(8), e29376.
Amusan, I., Rich, P., Housley, T., & Ejeta, G. (2011). An in- vitro method for 
AUTHOR CONTRIBUTIONS identifying postattachment Striga resistance in maize and sorghum. 
A.M. Yacoubou, wrote the draft manuscript with the contributions Agronomy Journal, 103, 1472–1 478.
of all the co- authors. Amusan, I. O., Rich, P. J., Menkir, A., Housley, T., & Ejeta, G. (2008). 
Resistance to Striga hermonthica in a maize inbred line derived from 
Zea diploperennis. New Phytologist, 178(1), 157–1 66. https://doi.
ORCID org/10.1111/j.1469- 8137.2007.02355.x
Abdoul- Madjidou Yacoubou  https://orcid. Atera, E. A., Ishii, T., Onyango, J. C., Itoh, K., & Azuma, T. (2013). Striga 
org/0000-0002-3496-0072 infestation in Kenya: Status, distribution and management options. 
Sustainable Agriculture Research, 2(2), 99. https://doi.org/10.5539/
Abebe Menkir  https://orcid.org/0000-0002-5907-9177 sar.v2n2p99
Melaku Gedil  https://orcid.org/0000-0002-6258-6014 Atera, E., & Itoh, K. (2011). Evaluation of ecologies and severity of 
Striga weed on rice in sub- Saharan Africa. Agriculture and Biology 
R E FE R E N C E S Journal of North America, 2(5), 752–7 60. https://doi.org/10.5251/
abjna.2011.2.5.752.760
Abdoulaye, T., Wossen, T., & Awotide, B. (2018). Impacts of improved Atera, E. A., Onyango, J. C., Thanh, P. T., Ishii, T., & Itoh, K. (2015). 
maize varieties in Nigeria: Ex-p ost assessment of productivity Identification of QTL for Striga hermonthica resistance using back-
and welfare outcomes. Food Security, 10, 369–3 79. https://doi. cross population derived from a cross between Oryza sativa (cv. 
org/10.1007/s12571 -0 18- 0772- 9 Nipponbare) and O. rufipogon. Journal of Agricultural Science, 7(2), 
Adetimirin, V. O., Aken’Ova, M. E., & Kim, S. K. (2001). Detection of 99–1 05. https://doi.org/10.5539/jas.v7n2p99
epistasis for horizontal resistance to Striga hermonthica in maize. Ayongwa, G. C., Stomph, T. J., Hoevers, R., Ngoumou, T. N., & Kuyper, 
Maydica, 46, 27–3 4. T. W. (2010). Striga infestation in northern Cameroon: Magnitude, 
Adetimirin, V. O., Kim, S. K., & Aken Ova, M. E. (2000). Responses of mid- dynamics and implications for management. NJAS - Wageningen 
altitude maize inbred lines to Striga hermonthica infestation. Cereal Journal of Life Sciences, 57, 159– 165. https://doi.org/10.1016/j.
Research Communications, 28(4), 469– 475. njas.2010.04.003
206  |     YACOUBOU et Al.
Babiker, A. G. T. (2007). Striga: The spreading scourge in Africa. Regulation Carsky, R. J., Berner, D. K., Oyewole, B. D., Dashiell, K., & Schulz, S. 
of Plant Growth & Development, 42, 74– 87. (2000). Reduction of Striga hermonthica parasitism on maize using 
Badu-A praku, B. (2010). Effects of recurrent selection for grain yield and soybean rotation. International Journal of Pest Management, 46, 115– 
Striga resistance in an extra- early maize population. Crop Science, 50, 120. https://doi.org/10.1080/09670 87002 27471
1735– 1743. Chemisquy, M. A., Giussani, L. M., Scataglini, M. A., Kellogg, E. A., & 
Badu- Apraku, B., Akinwale, R. O., Franco, J., & Oyekunle, M. (2012). Morrone, O. (2010). Phylogenetic studies favour the unification of 
Assessment of reliability of secondary traits in selecting for improved Pennisetum, Cenchrus and Odontelytrum (Poaceae): A combined 
grain yield in drought and low- nitrogen environments. Crop Science, nuclear, plastid and morphological analysis, and nomenclatural com-
52(5), 2050–2 062. https://doi.org/10.2135/crops ci2011.12.0629 binations in Cenchrus. Annals of Botany, 106, 107– 130. https://doi.
Badu-A praku, B., Akinwale, R. O., & Oyikunle, M. (2014). Efficiency of org/10.1093/aob/mcq090
secondary traits in selecting for improved grain yield in extra-e arly Chitagu, M., Rugare, J. T., & Mabasa, S. (2014). Screening maize (Zea mays) 
maize under Striga- infested and Striga-f ree environments. Plant genotypes for tolerance to witchweed (Striga asiatica L. Kuntze) in-
Breeding, 133, 373–4 38. fection. Journal of Agricultural Science, 6(2), 160– 169. https://doi.
Badu-A praku, B., Fakorede, M. A. B., & Akinwale, R. O. (2017). Key chal- org/10.5539/jas.v6n2p160
lenges in maize breeding in sub- Saharan Africa. August, 51– 86. https:// Das, B., Atlin, G. N., Olsen, M., Burgueño, J., Tarekegne, A., Babu, R., 
doi.org/10.19103/ as.2016.0001.03 Ndou, E. N., Mashingaidze, K., Moremoholo, L., Ligeyo, D., Matemba- 
Badu-A praku, B., Fakorede, M. A. B., & Fontem Lum, A. (2007). Evaluation Mutasa, R., Zaman- Allah, M., San Vicente, F., Prasanna, B. M., & 
of experimental varieties from recurrent selection for Striga resis- Cairns, J. E. (2019). Identification of donors for low- nitrogen stress 
tance in two extra-e arly maize populations in the savannas of West with maize lethal necrosis (MLN) tolerance for maize breeding in 
and Central Africa. Experimental Agriculture, 43(2), 183– 200. https:// sub- Saharan Africa. Euphytica, 1, 215–2 80. https://doi.org/10.1007/
doi.org/10.1017/S0014 47970 6004601 s1068 1-0 19-2 406-5 
Badu- Apraku, B., Fakorede, M. A. B., Gedil, M., Annor, B., Talabi, A. O., De Groote, H., Wangare, L., Kanampiu, F., Odendo, M., Diallo, A., Karaya, 
Akaogu, I. C., Oyekunle, M., Akinwale, R. O., & Fasanmade, T. Y. (2016). H., & Friesen, D. (2008). The potential of herbicide resistant maize 
Heterotic patterns of IITA and CIMMYT early- maturing yellow maize technology for Striga control in Africa. Agricultural Systems, 97, 
inbreds under contrasting environments. Agronomy Journal, 108(4), 83– 94.
1321– 1336. https://doi.org/10.2134/agron j2015.0425 Dean, R. E., Dahlberg, J. A., Hopkins, M. S., Mitchell, S. E., & Kresovich, S. 
Badu- Apraku, B., Fakorede, M. A., Oyekunle, M., Yallou, G. C., Obeng- (1999). Genetic redundancy and diversity among orange accessions 
Antwi, K., Haruna, A., Usman, I. S., & Akinwale, R. O. (2015). Gains in the U.S. national sorghum collection as assessed with simple se-
in grain yield of early maize cultivars developed during three breed- quence repeat (SSR) markers. Crop Science, 39, 1215– 1221. https://
ing eras under multiple environments. Crop Science, 55(2), 527– 539. doi.org/10.2135/crops ci1999.00111 83X00 390004 0043x
https://doi.org/10.2135/cropsc i2013.11.0783 Edmeades, G. O. (2013). Progress in achieving and delivering drought toler-
Badu- Apraku, B., Talabi, A. O., Fakorede, M. A. B., Fasanmade, Y., Gedil, ance in maize- An update. International Service for the Acquisition of 
M., Magorokosho, C., & Asiedu, R. (2019). Yield gains and associ- Agri- Biotech Applications.
ated changes in an early yellow bi- parental maize population follow- Ejeta, G. (2007). Breeding for resistance in sorghum: Exploitation of an 
ing genomic selection for Striga resistance and drought tolerance. intricate host– parasite biology. Crop Science, 47, 216– S227.
BMC Plant Biology, 19(1), 1– 17. https://doi.org/10.1186/s1287 Ejeta, G., Butler, L. G., Hess, D. E., Obilana, T., & Reddy, B. V. (1997). 
0- 019- 1740- z Breeding for Striga resistance in sorghum. International Conference on 
Badu- Aprakua, B., Yallou, C. G., & Oyekunlea, M. (2013). Genetic gains Genetic Improvement of Sorghum and Pearl Millet (pp. 504– 516).
from selection for high grain yield and Striga resistance in early Eplee, R. E. (1992). Witch weed (Striga asiatica): An overview of man-
maturing maize cultivars of three breeding periods under Striga- agement strategies in the USA. Crop Protection, 11, 3– 7. https://doi.
infested and Striga- free environments. Field Crops Research, 147, 54– org/10.1016/0261-2 194(92)90071 - C
67. https://doi.org/10.1016/j.fcr.2013.03.022 Eplee, R. E., & Norris, R. S. (1987). Field research techniques. In L. J. 
Bawa, A., Abdulai, M. S., & Addai, I. K. (2015). Evaluation of inbred lines Musselman (Ed.), Parasitic weeds in agriculture. Volume 1. Striga (pp. 
and hybrid maize (Zea mays L.) for tolerance to striga hermonthica 271–2 80). CRC Press, Inc.
(Del.) Benth in the guinea savanna agro- ecological zone of Ghana. FAOSTAT. (2018). FAOSTAT database. FAO. http://faosta t.fao.org
American Journal of Agricultural and Biological Science, 10(3), 128–1 36. Gethi, J. G., & Smith, M. E. (2004). Genetic responses of single crosses 
https://doi.org/10.3844/ajabs sp.2015.128.136 of maize to Striga hermonthica (Del.) Benth. and Striga asiatica (L.) 
Berner, D. K., Kling, J. G., & Singh, B. B. (1995). Striga research and con- Kuntze. Crop Science, 44, 2068– 2077.
trol. A perspective from Africa. Plant Disease, 97, 652–6 60. https:// Gobena, D., Shimelis, M., Rich, P. J., Ruyter- spira, C., Bouwmeester, H., 
doi.org/10.1094/PD- 79-0 652 & Kanuganti, S. (2017). Mutation in sorghum LOW GERMINATION 
Bilalis, D., Papastylianou, P., Konstantas, A., Patsiali, S., Karkanis, A., STIMULANT 1 alters strigolactones and causes Striga resistance. 
& Efthimiadou, A. (2010). Weed-s uppressive effects of maize- Proceedings of the National Academy of Sciences, 114, 4471– 4476.
legume intercropping in organic farming. International Journal of Pest Gowda, M., Beyene, Y., Makumbi, D., Semagn, K., Olsen, M. S., Bright, 
Management, 56, 173–1 81. https://doi.org/10.1080/096708 7090 J. M., Das, B., Mugo, S., Suresh, L. M., & Prasanna, B. M. (2018). 
3304471 Discovery and validation of genomic regions associated with re-
Bouwmeester, H. J., Matusova, R., Zhongkui, S., & Beale, M. H. (2003). sistance to maize lethal necrosis in four biparental populations. 
Secondary metabolite signalling in host–p arasitic plant interactions. Molecular Breeding, 38(5), 1– 16. https://doi.org/10.1007/s1103 
Current Opinion in Plant Biology, 6, 358– 364. https://doi.org/10.1016/ 2-0 18- 0829-7 
S1369 -5 266(03)00065 - 7 Gurney, A. L., Grimanelli, D., Kanampiu, F., Hoisington, D., Scholes, 
Brewer, P. B., Yoneyama, K., Filardo, F., Meyers, E., Scaffidi, A., Frickey, J. D., & Press, M. C. (2003). Novel sources of resistance to 
T., Akiyama, K., Seto, Y., Dun, E. A., Cremer, J. E., & Kerr, S. C. (2016). Striga hermonthica in Tripsacum dactyloides, a wild rela-
LATERAL BRANCHING OXIDOREDUCTASE acts in the final stages tive of maize. New Phytologist, 160(3), 557– 568. https://doi.
of strigolactone biosynthesis in Arabidopsis. Proceedings of the org/10.1046/j.1469- 8137.2003.00904.x
National Academy of Sciences of the United States of America, 113, Gurney, A., Slate, J., Press, M. C., & Scholes, J. D. (2006). 
6301– 6306. A novel form of resistance in rice to the angiosperm 
YACOUBOU et Al.      |  207
parasite Striga hermonthica. New Phytologist, 169, 199– 208. https:// Kanampiu, F., Makumbi, D., Mageto, E., Omanya, G., Waruingi, S., 
doi.org/10.1111/j.1469-8 137.2005.01560.x Musyoka, P., & Ransom, J. (2018). Assessment of management op-
Gutierrez-M arcos, J. F., Pennington, P. D., Costa, L. M., & Dickinson, H. tions on striga infestation and maize grain yield in Kenya. Weed 
G. (2003). Imprinting in the endosperm: A possible role in preventing Science, 66, 516– 524. https://doi.org/10.1017/wsc.2018.4
wide hybridization. Philosophical Transactions of the Royal Society B: Kanampiu, F. K., Ransom, J. K., Friesen, D., & Gressel, J. (2002). Imazapyr 
Biological Sciences, 358(1434), 1105–1 111. https://doi.org/10.1098/ and pyrithiobac movement in soil and from maize seed coats controls 
rstb.2003.1292 Striga while allowing legume intercropping. Crop Protection, 21, 611– 
Hailu, G., Niassy, S., Zeyaur, K. R., Ochatum, N., & Subramanian, S. (2018). 619. https://doi.org/10.1016/S0261 -2 194(01)00151-  X
Maize– legume intercropping and push– pull for management of fall Kang, H. J., Lee, Y. T., Cho, Y. G., Eun, M. Y., & Shim, J. U. (1998). QTL map-
armyworm, stemborers, and striga in Uganda. Agronomy Journal, ping of genes conferring days to heading, culm length and panicle 
110(6), 2513–2 522. https://doi.org/10.2134/agron j2018.02.0110 length based on molecular map of rice (Oryza sativa L.). RDA Journal 
Hallauer, A. R. (1992). Recurrent selection in maize. Plant Breeding of Crop Science, 40, 55–6 1.
Reviews, 9, 115–1 79. Kannan, C., & Zwanenburg, B. (2014). A novel concept for the control 
Hallauer, A. R., & Carena, M. J. (2012). Recurrent selection methods to of parasitic weeds by decomposing germination stimulants prior 
improve germplasm in maize. Maydica, 57, 266– 283. to action. Crop Protection, 61, 11– 15. https://doi.org/10.1016/j.
Hamiaux, C., Drummond, R. S. M., Janssen, B. J., Ledger, S. E., Cooney, cropro.2014.03.008
J. M., Newcomb, R. D., & Snowden, K. C. (2012). DAD2 is an α/β hy- Karaya, H., Kiarie, N., Mugo, S., Aringa, E., Nderitu, J., & Kanampiu, F. 
drolase likely to be involved in the perception of the plant branching (2014). Combing ability of maize (Zea Mays) inbred lines resistant to 
hormone, strigolactone. Current Biology, 22, 2032–2 036. striga hermonthica (Del.) Benth evaluated under artificial striga in-
Haussmann, B. I. G., Hess, D. E., Omanya, G. O., Folkertsma, R. T., Reddy, festation. African Journal of Agricultural Research, 9(16), 1287–1 295. 
B. V. S., Kayentao, M., Welz, H. G., & Geiger, H. H. (2004). Genomic re- https://doi.org/10.5897/AJAR12.1315
gions influencing resistance to the parasitic weed Striga hermonthica Karaya, H., Njoroge, K., Mugo, S., Ariga, E. S., Kanampiu, F., & Nderitu, J. 
in two recombinant inbred populations of sorghum. Theoretical and H. (2012). Determination of levels of Striga germination stimulants 
Applied Genetics, 109(5), 1005– 1016. https://doi.org/10.1007/s0012 for maize gene bank accessions and elite inbred lines. International 
2-0 04- 1706- 9 Journal of Plant Production, 6(2), 209– 224. https://doi.org/10.22069/ 
Hearne, S. J. (2001). Morphological, physiological and molecular interac- ijpp.2012.776
tions between maize and the parasitic angiosperm Striga hermonthica. Khan, Z. R., Midega, C. A. O., Bruce, T. J. A., Hooper, A. M., & Pickett, 
Doctor of Philosophy Thesis, University of Sheffield. 191 p. J. A. (2010). Exploiting phytochemicals for developing a “push- pull” 
Hearne, S. J. (2009). Control – The Striga conundrum. Pest Management crop protection strategy for cereal farmers in Africa. Journal of 
Science, 65, 603– 614. Experimental Botany, 61(15), 4185– 4196. https://doi.org/10.1093/
Hesammi, E. (2013). Striga and ways of control. International Journal of jxb/erq229
Food and Allied Sciences, 2, 53–5 5. Khan, Z. R., Midega, C. A. O., Hassanali, A., Pickett, J. A., & Wadhams, L. 
Hess, D. E., Ejeta, G., & Butler, L. G. (1992). Selecting sorghum geno- J. (2007). Assessment of different legumes for the control of Striga 
types expressing a quantitative biosynthetic trait that confers hermonthica in maize and sorghum. Crop Science, 47(2), 730– 736. 
resistance to Striga. Phytochemistry, 31(2), 493–4 97. https://doi. https://doi.org/10.2135/cropsc i2006.07.0487
org/10.1016/0031-9 422(92)90023- J Kim, S. K. (1991). Breeding maize for Striga tolerance and the develop-
Ibrahim, A., Ahom, R. I., Magani, I. E., & Musa, M. I. (2014). Spatial distri- ment of a field infestation technique. In Combating striga in Africa (pp. 
bution and density of Striga hermonthica (Del.) Benth infestation as- 96– 108). IITA.
sociated with cereal production in southern guinea savanna farming Kim, G., & Westwood, J. H. (2015). Macromolecule exchange in Cuscuta- 
systems. Journal of Biodiversity and Environmental Sciences, 5, 419–4 27. host plant interactions. Current Opinion in Plant Biology, 26(Table 1), 
Illa, A., Odhiambo, G., & Dida, M. (2010). Increasing imazapyr- resistant 20– 25. https://doi.org/10.1016/j.pbi.2015.05.012
maize yield by increasing plant density under natural Striga her- Kim, S. K. (1994). Genetics of maize tolerance of Striga hermonthica. Crop 
monthica infestation. Agriculture and Biology Journal of North America, Science, 34, 900–9 07.
1(5), 1061– 1068. https://doi.org/10.5251/abjna.2010.1.5.1061.1068 Kim, S. K., Adetimirin, V. O., Thé, C., & Dossou, R. (2002). Yield losses 
Jamil, M., Kanampiu, F. K., Karaya, H., Charnikhova, T., & Bouwmeestera, in maize due to Striga hermonthica in West and Central Africa. 
H. J. (2012). Striga hermonthica parasitism in maize in response International Journal of Pest Management, 48(3), 211– 217. https://doi.
to N and P fertilisers. Field Crops Research, 134, 1– 10. https://doi. org/10.1080/096708 7011 0117408
org/10.1016/j.fcr.2012.03.015 Kim, S. K., & Akintunde, A. Y. (1989). Recent progress on Striga resistance 
Jamil, M., Rodenburg, J., Charnikhova, T., & Bouwmeester, H. J. breeding in maize at IITA. In C. Fajemisin, J. M. Muleba, N. Emechebe, 
(2011). Pre- attachment Striga hermonthica resistance of New & A. M. Dabire (Eds.), Proceedings of the SAFGRAD Workshop (pp. 55– 
Rice for Africa (NERICA) cultivars based on low strigolac- 62). SAFGRAD.
tone production. New Phytologist, 192(4), 964–9 75. https://doi. Kirigia, D., Runo, S., & Alakonya, A. (2014). A virus- induced gene silencing 
org/10.1111/j.1469- 8137.2011.03850.x (VIGS) system for functional genomics in the parasitic plant Striga 
Jiménez- Galindo, J. C., Ordás, B., Butrón, A., Samayoa, L. F., & Malvar, hermonthica. Plant Methods, 10(16), 1– 11.
R. A. (2017). QTL mapping for yield and resistance against mediter- Konate, L., Baffour, B. A., & Traore, D. (2017). Combining ability and 
ranean corn borer in maize. Frontiers in Plant Science, 8(May), 2–1 1. heterotic grouping of early maturing provitamin A maize inbreds 
https://doi.org/10.3389/fpls.2017.00698 across Striga infested and optimal growing environments. Journal 
John, M., & Sleeper, D. (1995). Breeding field crops. Iowa State University of Agriculture and Environment for International Development, 111(1), 
Press. 157–1 73. https://doi.org/10.12895/ jaeid.20171 11.572
Kanampiu, F. K., Kabambe, V., Massawe, C., Jasi, L., Friesen, D., Ransom, Kouakou, C. (2014). The use of genetically tolerant maize (Zea mays 
J. K., & Gressel, J. (2003). Multi- site, multi- season field tests demon- L.) in the control of Striga hermonthica in Northern Côte d’Ivoire. 
strate that herbicide seed- coating herbicide- resistance maize con- American Journal of Experimental Agriculture, 4(5), 563– 574. https://
trols Striga spp. and increases yields in several African countries. doi.org/10.9734/ajea/2014/7269
Crop Protection, 22(5), 697– 706. https://doi.org/10.1016/s0261 Kountche, B. A., Jamil, M., Yonli, D., Nikiema, M. P., Blanco- Ania, D., 
-2 194(03)00007-  3 Asami, T., Zwanenburg, B., & Al- Babili, S. (2019). Suicidal germination 
208  |     YACOUBOU et Al.
as a control strategy for Striga hermonthica (Benth.) in smallholder Menkir, A., Kling, J. G., Badu-A praku, B., & Ibikunle, O. (2006). 
farms of sub-S aharan Africa. Plants, People, Planet, 1(2), 107–1 18. Registration of 26 tropical maize germplasm lines with resistance to 
https://doi.org/10.1002/ppp3.32 Striga hermonthica. Crop Science, 46, 1007– 1009.
Lagoke, S. T. O., & Isah, K. M. (2010). Reaction of maize varieties to Striga Menkir, A., Kling, J. G., Badu- Apraku, B., Thé, C., & Ibikunle, O. (2004). 
hermonthica as influenced by food legume intercrop, spacing and Recent advances in breeding maize for resistance to Striga her-
split application of compound fertilizer. Nigerian Journal of Weed monthica (del.) Benth. In Integrated Approaches to Higher Maize 
Science, 23, 45–5 8. Productivity in the New Millennium: Proceedings of the Seventh Eastern 
Lane, J. A., Child, D. V., Moore, T., Arnold, G. M., & Bailey, J. A. (1997). and Southern Africa Regional Maize Conference. CIMMYT.
Phenotypic characterisation of resistance in Zea diploperennis to Menkir, A., Makumbi, D., & Franco, J. (2012). Assessment of reaction 
Striga hermonthica. Maydica, 42, 45–5 1. patterns of hybrids to Striga hermonthica (del.) benth. under artifi-
Lechat, M. M., Brun, G., Montiel, G., Véronési, C., Simier, P., Thoiron, S., cial infestation in Kenya and Nigeria. Crop Science, 52(6), 2528–2 537. 
Pouvreau, J. B., & Delavault, P. (2015). Seed response to strigolactone is https://doi.org/10.2135/crops ci2012.05.0307
controlled by abscisic acid- independent DNA methylation in the obligate Menkir, A., & Meseka, S. (2019). Genetic improvement in resistance to 
root parasitic plant, Phelipanche ramosa L. Pomel. Journal of Experimental striga in tropical maize hybrids. Crop Science, 59, 2484–2 497. https://
Botany, 66(11), 3129– 3140. https://doi.org/10.1093/jxb/erv119 doi.org/10.2135/cropsc i2018.12.0749
Liu, Y., Wang, L., Sun, C., Zhang, Z., Zheng, Y., & Qiu, F. (2014). Genetic Midega, C. A. O., Pickett, J., Hooper, A., Pittchar, J., & Khan, Z. R. (2016). 
analysis and major QTL detection for maize kernel size and weight Maize landraces are less affected by Striga hermonthica relative to 
in multi-e nvironments. TAG. Theoretical and Applied Genetics., 127, hybrids in western Kenya. Weed Technology, 30(1), 21–2 8. https://doi.
1019– 1037. https://doi.org/10.1007/s00122 - 014-2 276-0 org/10.1614/wt-d -1 5-0 0055.1
Lumba, S., Holbrook-S mith, D., & McCourt, P. (2017). The perception of Mohamed, A. H., Housley, T. L., & Ejeta, G. (2010). An in vitro technique 
strigolactones in vascular plants. Nature Chemical Biology, 13(6), 599– for studying specific Striga resistance mechanisms in sorghum. 
606. https://doi.org/10.1038/nchem bio.2340 African Journal of Agricultural Research, 5, 1868–1 875.
Makumbi, D., Diallo, A., Kanampiu, F., Mugo, S., & Karaya, H. (2015). Mrema, E., Shimelis, H., Laing, M., & Mwadzingeni, L. (2017). Genetic 
Agronomic performance and genotype × environment interaction analysis of the maximum germination distance of Striga under 
of herbicide-r esistant maize varieties in Eastern Africa. Crop Science, Fusarium oxysporum f. sp. strigae biocontrol in sorghum. Journal of 
55(2), 540– 555. https://doi.org/10.2135/cropsc i2014.08.0593 Integrative Agriculture, 17(7), 1585– 1593. https://doi.org/10.1016/
Mandolino, C. I., D’Andrea, K. E., Olmos, S. E., Otegui, M. E., & Eyhérabide, S2095-  3119(17)61790- 8 
G. H. (2018). Maize nitrogen use efficiency: Qtl mapping in A U.S. Mutinda, S. M., Masanga, J., Mutuku, J. M., Runo, S., & Alakonya, A. 
dent x argentine-c aribbean flint RILS population. Maydica, 63(1), 17. (2018). KSTP 94, an open- pollinated maize variety has postattach-
Manyong, V. M., Nindi, S. J., Alene, A. D., Odhiambo, G. D., Omanya, ment resistance to purple witchweed (Striga hermonthica). Weed 
G., Mignouna, H. D., & Bokanga, M. (2008). Farmer perceptions Science, 66(4), 525– 529. https://doi.org/10.1017/wsc.2018.24
on performance attributes of maize varieties: The frequency ap- Naitormmbaide, M., Djondang, K., Mama, V. J., & Koussou, M. (2015). 
proach. Farmer Perceptions of Imazapyr-R esistant (IR) maize technol- Criblage de quelques variétés de maïs (Zea mays L.) pour la résistance 
ogy on the control of Striga in Western Kenya (pp. 43– 82). Support au Striga hermonthica (Del) Benth dans les savanes tchadiennes. 
for Development Communication, PO Box 27628- 00506, Nairobi, Journal of Animal & Plant Sciences, 24(1), 3722–3 732.
Kenya: African Agricultural Technology Foundation Nairobi. Nyakurwa, C. S., Gasura, E., Setimela, P. S., Mabasa, S., Rugare, J. T., & 
Mbuvi, D. A., Masiga, C. W., Kuria, E., Masanga, J., Wamalwa, M., Mutsvanga, S. (2018). Reaction of new quality protein maize geno-
Mohamed, A., Odeny, D. A., Hamza, N., Timko, M. P., & Runo, S. types to Striga asiatica. Crop Science, 58(3), 1201–1 218. https://doi.
(2017). Novel sources of witchweed (Striga) resistance from wild org/10.2135/cropsc i2017.10.0639
sorghum accessions. Frontiers Plant Science, 8(FEBRUARY), 1– 15. Nzioki, H. S., Oyosi, F., Morris, C. E., Kaya, E., Pilgeram, A. L., Baker, C. 
https://doi.org/10.3389/fpls.2017.00116 S., & Sands, D. C. (2016). Striga biocontrol on a toothpick: A read-
Mengesha, W. A., Menkir, A., Unakchukwu, N., Meseka, S., Farinola, A., ily deployable and inexpensive method for smallholder farmers. 
Girma, G., & Gedil, M. (2017). Genetic diversity of tropical maize in- Frontiers in Plant Science, 7(AUG2016), 1–8 . https://doi.org/10.3389/
bred lines combining resistance to Striga hermonthica with drought fpls.2016.01121
tolerance using SNP markers. Plant Breeding, 136(3), 338– 343. Osman, G., Hassan, M., Rugheim, A., Abdelgani, M., & Babiker, A. (2013). 
https://doi.org/10.1111/pbr.12479 Effects of organic and microbial fertilizers on Striga hermonthica in 
Menkir, A. (2006). Assessment of reactions of diverse maize inbred lines maize. Universal Journal of Agricultural Research, 1(2), 24–2 9. https://
to Striga hermonthica (Del.). Benth. Plant Breeding, 125(2), 131– 139. doi.org/10.13189/ ujar.2013.010202
https://doi.org/10.1111/j.1439- 0523.2006.01175.x Oswald, A. (2005). Striga control—T echnologies and their dissemina-
Menkir, A., Adetimirin, V. O., Yallou, C. G., & Gedil, M. (2010). Relationship tion. Crop Protection, 24(4), 333– 342. https://doi.org/10.1016/j.
of genetic diversity of inbred lines with different reactions to Striga cropro.2004.09.003
hermonthica (Del.) benth and the performance of their crosses. Crop Oswald, A., & Ransom, J. K. (2002). Response of maize varieties to trans-
Science, 50(2), 602–6 11. https://doi.org/10.2135/cropsc i2009.05.0247 planting in Striga-i nfested fields. Weed Science, 50(3), 392–3 96.10.16
Menkir, A., Crossa, J., Meseka, S., & Bossey, B. (2020). Staking tolerance 14/0043- 1745(2002)050[0392:romvtt]2.0.co;2
to drought and resistance to a parasitic weed in tropical hybrid maize Ouédraogo, O., Kaboré, T. D., Noba, D. R., & Traoré, S. (2018). Polygala 
for enhancing resilience to stress. Combinations, 11(February), 1–1 6. rarifolia DC., plante faux hôte du Striga hermonthica (Del.). Benth. 
https://doi.org/10.3389/fpls.2020.00166 Journal of Applied Biosciences, 123(1), 12346–1 2353.
Menkir, A., Franco, J., Adpoju, A., & Bossey, B. (2012). Evaluating consistency Oyekunle, M., Haruna, A., Badu- Apraku, B., Usman, I. S., Mani, H., Ado, 
of resistance reactions of open- pollinated maize cultivars to Striga her- S. G., Olaoye, G., Obeng- Antwi, K., Abdulmalik, R. O., & Ahmed, H. O. 
monthica (Del.) Benth under artificial infestation. Crop Science, 52(3), (2017). Assessment of early-m aturing maize hybrids and testing sites 
1051–1 060. https://doi.org/10.2135/crops ci2011.10.0543 using GGE biplot analysis. Crop Science, 57(6), 2942– 2950. https://
Menkir, A., & King, J. G. (2007). Response to recurrent selection for re- doi.org/10.2135/cropsc i2016.12.1014
sistance to Striga hermonthica (Del.) Benth in a tropical maize popu- Parker, C. (2012). Parasitic weeds: A world challenge. Weed Science, 60, 
lation. Crop Science, 47, 674– 684. 269– 276. https://doi.org/10.1614/WS-D - 11- 00068.1
YACOUBOU et Al.      |  209
Parker, C., & Riches, C. R. (1993). Parasitic weeds of the world: Biology and in the United States. Monograph series of the Weed Science Society of 
control. CAB International. America, Number 5, Weed Science Society of America.
Ramaiah, K. V. (1987). Breeding cereal grains for resistance to witch- Schroeder, C., Onyango K’Oloo, T., Nar Bahadur, R., Jick, N. A., Parzies, 
weed. In L. J. Musselman (Ed.), Parasitic weeds in agriculture (pp. 227– H. K., & Gemenet, D. C. (2013). Potentials of hybrid maize varieties 
242). Striga CRC. for smallholder farmers in Kenya: A review based on swot analysis. 
Ramaiah, K. V., Parker, C., Vasudeva Rao, M. J., & Musselman, L. J. African Journal of Food, Agriculture, Nutrition and Development, 13, 
(1983). Striga identification and control handbook. (A. P. Patancheru 7562–7 582.
(ed.)).15, (1– 51). Andhra Pradesh, India: ICRISAT. Schut, M., Rodenburg, J., Klerkx, L., Hinnou, L. C., Kayeke, J., & Bastiaans, 
Randrianjafizanaka, M. T., Autfray, P., Andrianaivo, A. P., Ramonta, I. R., & L. (2015). Participatory appraisal of institutional and political con-
Rodenburg, J. (2018). Combined effects of cover crops, mulch, zero- straints and opportunities for innovation to address parasitic weeds 
tillage and resistant varieties on Striga asiatica (L.) Kuntze in rice- in rice. Crop Protection, 74, 158– 170. https://doi.org/10.1016/j.
maize rotation systems. Agriculture, Ecosystems and Environment, 256, cropro.2015.04.011
23– 33. https://doi.org/10.1016/j.agee.2017.12.005 Shayanowako, A. T., Laing, M., Shimelis, H., & Mwadzingeni, L. (2017). 
Ransom, J., Kanampiu, F., Gressel, J., De Groote, H., Burnet, M., & Resistance breeding and biocontrol of Striga asiatica (L.) Kuntze in 
Odhiambo, G. (2012). Herbicide applied to imidazolinone resistant- maize: A review. Acta Agriculturae Scandinavica, Section B—s oil & Plant 
maize seed as a Striga control option for smallscale African farmers. Science, 1– 11.
Weed Science, 60, 283–2 89. Semagn, K., Beyene, Y., Babu, R., Nair, S., Gowda, M., Das, B., Tarekegne, 
Rao, M. V. J (1985). Techniques for screening sorghums for resistance to A., Mugo, S., Mahuku, G., Worku, M., Warburton, M. L., Olsen, M., & 
Striga, 20, ICRISAT Patancheru P.O. Andhra Pradesh 502 324(1– Prasanna, B. M. (2015). Quantitative trait loci mapping and molec-
18). India: International Crops Research Institute for the Semi- Arid ular breeding for developing stress resilient maize for Sub- Saharan 
Tropics. Africa. Crop Science, 55(4), 1449– 1459. https://doi.org/10.2135/
Reda, F., & Verkleij, J. A. C. (2007). Cultural and cropping systems approach cropsc i2014.09.0646
for striga management — A low cost alternative option in subsistence Shayanowako, A. I. T., Shimelis, H., Laing, M. D., & Mwadzingeni, I. (2018). 
farming. In G. Ejeta, & J. Gressel (Eds.), Integrating new technologies for Genetic diversity of maize genotypes with variable resistance to 
striga control. https://doi.org/10.1142/97898 127715 06_0016 striga asiatica Based on SSR markers. Cereal Research Communications, 
Ribaut, J. M., William, H. M., Khairallah, M., Worland, A. J., & Hoisington, 46(4), 668–6 78. https://doi.org/10.1556/0806.46.2018.044
D. (2001). Genetic basis of physiological traits. In Application of phys- Sibov, S. T., De Souza, C. L., Garcia, A. A. F., Silva, A. R., Garcia, A. F., 
iology in wheat breeding. CIMMYT. Mangolin, C. A., Benchimol, L. L., & De Souza, A. P. (2003). Molecular 
Ribeiro, P. F., Badu- Apraku, B., Gracen, V. E., Danquah, E. Y., Garcia- mapping in tropical maize (Zea mays L.) using microsatellite mark-
Oliveira, A. L., Asante, M. D., Afriyie- Debrah, C., & Gedil, M. (2018). ers. 2. Quantitative trait loci (QTL) for grain yield, plant height, ear 
Identification of quantitative trait loci for grain yield and other height and grain moisture. Hereditas, 139(2), 107–1 15. https://doi.
traits in tropical maize under high and low soil-n itrogen environ- org/10.1111/j.1601- 5223.2003.01667.x
ments. Crop Science, 58(1), 321– 331. https://doi.org/10.2135/crops Simon, Z., Kasozi, L. C., Patrick, R., & Abubaker, M. (2018). Gene action 
ci2017.02.0117 for grain yield and agronomic traits in selected maize inbred lines 
Rich, P. J., & Ejeta, G. (2008). Towards effective resistance to Striga in with resistance to Striga hermonthica in Uganda. Journal of Food 
African maize. Plant Signaling & Behavior, Behavior, 3, 618–6 21. Security, 6(4), 155–1 62. https://doi.org/10.12691/j fs- 6-4 - 3
Rich, P. J., Grenier, C., & Ejeta, G. (2004). Striga resistance in the wild Smith, S. M. (2014). Q&A: What are strigolactones and why they are im-
relatives of sorghum. Crop Science, 44(6), 2221– 2229. https://doi. portant to plants and soil microbes? BMC Biology, 12, 1– 7. https://doi.
org/10.2135/cropsc i2004.2221 org/10.1186/1741-7 007- 12-1 9
Robert, O. J. (2011). Genetic Analysis of Striga hermonthica Resistance in Spallek, T., Mutuku, M., & Shirasu, K. (2013). The genus Striga: A witch 
Sorghum (Sorghum bicolor) Genotypes in Eastern Uganda, (1). – 132). profile. Molecular Plant Pathology, 14(9), 861–8 69. https://doi.
University of KwaZulu-N atal, Pietermaritzburg, South Africa: org/10.1111/mpp.12058
ResearchSpach.. KwaZulu- Natal, Pietermaritzburg, South Africa. Sun, Z., Hans, J., Walter, M. H., Matusova, R., Beekwilder, J., Verstappen, 
http://hdl.handle.net/10413/ 9981. F. W., Ming, Z., van Echtelt, E., Strack, D., Bisseling, T., & Bouwmeester, 
Rodenburg, J., Bastiaans, L., Weltzien, E., & Hess, D. E. (2005). How can H. J. (2008). Cloning and characterisation of a maize carotenoid 
field selection for Striga resistance and tolerance in sorghum be im- cleavage dioxygenase (ZmCCD1) and its involvement in the biosyn-
proved? Field Crops Research, 93, 34–5 0. thesis of apocarotenoids with various roles in mutualistic and para-
Rodenburg, J., Cissoko, M., Kayeke, J., Dieng, I., Khan, Z. R., Midega, C. sitic interactions. Planta, 228, 789–8 01.
A. O., Onyuka, E. A., & Scholes, J. D. (2015). Do NERICA rice culti- Teka, H. B. (2014). Advance research on Striga control: A review. African 
vars express resistance to Striga hermonthica (Del.) Benth. and Striga Journal of Plant Science, 8(11), 492– 506. https://doi.org/10.5897/
asiatica (L.) Kuntze under field conditions? Field Crops Research, 170, AJPS2 014.1186
83– 94. https://doi.org/10.1016/j.fcr.2014.10.010 Toh, S., Holbrook- Smith, D., Stogios, P. J., Onopriyenko, O., Lumba, 
Ronald, M., Charles, M., Stanford, M., & Eddie, M. (2019). Mulching offers S., Tsuchiya, Y., Savchenko, A., & McCourt, P. (2015). Structure- 
protection from Striga asiatica L. Kutnze parasitism in Sorghum geno- function analysis identifies highly sensitive strigolactone receptors 
types. Acta Agriculturae Scandinavica Section B: Soil and Plant Science, in Striga. Science, 350(6257), 203– 207. https://doi.org/10.1126/scien 
69(2), 167– 173. https://doi.org/10.1080/09064 710.2018.1520916 ce.aac9476
Runo, S., & Kuria, E. K. (2018). Habits of a highly successful cereal killer, Toh, S., Kamiya, Y., Kawakami, N., Nambara, E., McCourt, P., & Tsuchiya, 
Striga. PLoS Path, 14(1), e1006731. https://doi.org/10.1371/journ Y. (2012). Thermoinhibition uncovers a role for strigolactones in ara-
al.ppat.1006731 bidopsis seed germination. Plant and Cell Physiology, 53(1), 107– 117. 
Runo, S., Macharia, S., Alakonya, A., Machuka, J., Sinha, N., & Scholes, https://doi.org/10.1093/pcp/pcr176
J. (2012). Striga parasitizes transgenic hairy roots of Zea mays and Tuberosa, R., Sanguineti, M. C., Landi, P., Giuliani, M. M., Salvi, S., & Conti, 
provides a tool for studying plant- plant interactions. Plant Methods, S. (2002). Identification of QTLs for root characteristics in maize 
8(1), 1– 11. https://doi.org/10.1186/1746- 4811- 8- 20 grown in hydroponics and analysis of their overlap with QTLs for 
Sand, P. F., Eplee, R. E., & Westbrooks, R. G. (1990). In R. G. S and, P. grain yield in the field at two water regimes. Plant Molecular Biology, 
F., Eplee, R. E., & Westbrooks (Eds.), Witchweed research and control 48(5– 6), 697– 712. https://doi.org/10.1023/A:10148 97607670
210  |     YACOUBOU et Al.
Udom, G. N., Babatunde, F. E., & Tenebe, V. A. (2007). Suppression of Yoneyama, K., Awad, A. A., Xie, X., Yoneyama, K., & Takeuchi, Y. (2010). 
witch-w eed (Striga hermonthica) in sorghum: Cowpea mixture as af- Strigolactones as germination stimulants for root parasitic plants. 
fected by cowpea varieties and planting patterns. International Journal Plant and Cell Physiology, 51(7), 1095–1 103. https://doi.org/10.1093/
of Agricultural Research, 2(3), 268–2 74. https://doi.org/10.3923/ pcp/pcq055
ijar.2007.268.274 Yonli, D., Traoré, H., Hess, D., Sankara, P., & Sereme, P. (2006). 
Uraguchi, D., Kuwata, K., Hijikata, Y., Yamaguchi, R., Imaizumi, H., Effect of growth medium, Striga seed burial distance and 
Sathiyanarayanan, A. M., Rakers, C., Mori, N., Akiyama, K., Irle, depth on efficacy of Fusarium isolates to control Striga her-
S., McCourt, P., Kinoshita, T., Ooi, T., & Tsuchiya, Y. (2018). A monthica in Burkina Faso. Weed Research, 46, 73–8 1. https://doi.
femtomolar-r ange suicide germination stimulant for the parasitic org/10.1111/j.1365- 3180.2006.00486.x
plant striga hermonthica. Science, 362(6420), 1301–1 305. https:// Yoshida, S., & Shirasu, K. (2009). Multiple layers of incompatibility to 
doi.org/10.1126/scien ce.aau5445 the parasitic witchweed, Striga hermonthica. New Phytologist, 183, 
van Dam, N. M., & Bouwmeester, H. J. (2016). Metabolomics in the rhizo- 180– 189.
sphere: Tapping into belowground chemical communication. Trends Yoshida, S., & Shirasu, K. (2009). Multiple layers of incompatibility to 
in Plant Science, 21(3), 256– 265. https://doi.org/10.1016/j.tplan the parasitic witchweed. Striga Hermonthica. New Phytologist, 183(1), 
ts.2016.01.008 180– 189. https://doi.org/10.1111/j.1469-8 137.2009.02840.x
Webb, M., & Smith, M. C. (1996). Biology of Striga hermonthica (scrophu- Zarafi, A. B., Elzein, A., Abdulkadir, D. I., Beed, F., & Akinola, O. M. (2015). 
lariaceae) in Sahelian Mali: Effects on pearl millet yield and prospects Host range studies of Fusarium oxysporum f.sp. strigae meant for 
of control. Weed Research, 36(3), 203– 211. https://doi.org/10.1111/ the biological control of Striga hermonthica on maize and sorghum. 
j.1365- 3180.1996.tb0165 0.x Archives of Phytopathology and Plant Protection, 48(1), 1–9 . https://
Yagoub, S. O., Hassan, M. M., Gani, M. E. A., El, A., & Babiker, G. E. (2014). doi.org/10.1080/032354 08.2014.880580
Screening Sorghum for resistance to Striga hermonthica (Del.) Benth. Zehhar, N., Ingouff, M., Bouya, D., & Fer, A. (2002). Possible in-
Nova Journal of Medical and Biological Sciences, 3, 1– 5. https://doi. volvement of gibberellins and ethylene in Orobanche ra-
org/10.20286/n ova- jmbs- 03034 mosa germination. Weed Research, 42(6), 464– 469. https://doi.
Yallou, C. G., Menkir, A., Adetimirin, V. O., & Kling, J. G. (2009). Combining org/10.1046/j.1365- 3180.2002.00306.x
ability of maize inbred lines containing genes from Zea diploperennis Zhang, Y., van Dijk, A. D. J., Scaffidi, A., Flematti, G. R., Hofmann, M., 
for resistance to Striga hermonthica (Del.) Benth. Plant Breeding, 128(2), Charnikhova, T., Verstappen, F., Hepworth, J. O., van der Krol, S., 
143– 148. https://doi.org/10.1111/j.1439- 0523.2008.01583.x Leyser, O., Smith, S. M., Zwanenburg, B., Al- Babili, S., Ruyter- Spira, 
Yamaguchi, I., Cohen, J. D., Culler, A. H., Quint, M., Slovin, J. P., Nakajima, C., & Bouwmeester, H. J. (2014). Rice cytochrome P450 MAX1 ho-
M., Yamaguchi, S., Sakakibara, H., Kuroha, T., Hirai, N., Yokota, T., Ohta, mologs catalyze distinct steps in strigolactone biosynthesis. Nature 
H., Kobayashi, Y., Mori, H., & Sakagami, Y. (2010). Plant hormones. Chemical Biology, 10, 1028–1 033.
Comprehensive Natural Products II Chemistry and Biology, 4, 9–1 25). Zhou, F., Lin, Q., Zhu, L., Ren, Y., Zhou, K., Shabek, N., Wu, F., Mao, H., 
Elsevier. https://doi.org/10.1016/B978-0 0804 5382- 8.00092-  7 Dong, W., Gan, L. U., Ma, W., Gao, H. E., Chen, J., Yang, C., Wang, D., 
Yao, Z., Tian, F., Cao, X., Xu, Y., Chen, M., Xiang, B., & Zhao, S. (2016). Tan, J., Zhang, X., Guo, X., Wang, J., … Wan, J. (2013). D14- SCFD3- 
Global transcriptomic analysis reveals the mechanism of Phelipanche dependent degradation of D53 regulates strigolactone signalling. 
aegyptiaca seed germination. International Journal of Molecular Nature, 504, 406–4 10.
Sciences, 17(7), 1139. https://doi.org/10.3390/ijms1 7071139 Zwanenburg, B., Mwakaboko, A. S., & Kannan, C. (2016). Suicidal ger-
Yasir, A. G., & Abdalla, H. M. (2013). Introgression of Striga resistance mination for parasitic weed control. Pest Management Science, 72, 
genes into a Sudanese sorghum cultivar, Tabat, using marker assisted 2016–2 025. https://doi.org/10.1002/ps.4222
selection (MAS). Greener Journal of Agricultural Sciences, 3(7), 550– 
556. https://doi.org/10.15580/ GJAS.2013.3.061013 654
Yoder, J. I., & Scholes, J. D. (2010). Host plant resistance to parasitic How to cite this article: Yacoubou A- M, Zoumarou Wallis N, 
weeds; Recent progress and bottlenecks. Current Opinion in Plant 
Biology, 13(4), 478– 484. https://doi.org/10.1016/j.pbi.2010.04.011 Menkir A, et al. Breeding maize (zea mays, l.) for Striga 
Yohannes, T., Ngugi, K., Ariga, E., Abraha, T., Yao, N., Asami, P., & Ahonsi, resistance: Past, current and prospects in sub- saharan africa. 
M. (2016). Genotypic variation for low Striga germination stimula- Plant Breed. 2021;140:195–2 10. https://doi.org/10.1111/
tion in Sorghum Sorghum bicolor (L.) Moench” landraces from eritrea. pbr.12896
American Journal of Plant Sciences, 07(17), 2470– 2482. https://doi.
org/10.4236/ajps.2016.717215