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COLLEGE OF AGRICULTURE AND ENVIRONMENTAL SCIENCE
DEPARTMENT OF PLANT SCIENCE
M.Sc. in Plant Protection Graduate Program
Economic Pests of Ethiopia
BY:
Zemed Wobale Birhanie ……ID: BDU1401871
Review on sorghum anthracnose and sorghum aphids
Submitted to Asmare D. (PhD)
July, 2022
Bahir Dar, Ethiopia
INTRODUCTIONS
Sorghum [Sorghum bicolor (L.) Moench] is an important cereal crop that belongs to the grass
family Poaceae (Dahlberg, 2000). It is tolerant to harsh growing conditions with best adaptation
under drought-prone and the semi-arid tropical regions of the world (Ng'uni et al., 2011; Burrell
et al., 2015). Sorghum is called 'camel of crops' as it can withstand drought greatly. It has four
features, which make it one of the drought resistant crops, such that it has a very large root to
leaf-surface area; in time of drought, it will roll its leaves to lessen water loss; if drought
continues, it will go into dormancy rather than drying; and leaves are protected by a waxy cuticle
(Khosla et al., 1995). Sorghum is cultivated both in tropical and temperate climates for multiple
uses (Dahlberg et al., 2011). Sorghum is the most important cereal crop and is the dietary staple
of millions people in many countries in the world (Reddy et al., 2010). Further, it is a major
source of food, feed, and fuel and fiber crop across a range of environments and production
systems. Hence, sorghum is a major crop for the sustenance of human and livestock populations
in hot and dry areas of the world (Sharma et al., 2012).
Sorghum is one of the crops for which Ethiopia has been credited as being a center of origin
and/or diversity (Firew et al., 2009). Doggett (1988) suggested that sorghum originated and is
domesticated in the northeast quadrant of Africa, most likely in the Ethio-Sudan border regions.
Ethiopia is believed to be the center of origin and diversity of sorghum due to the presence of
diverse wild sorghum types. In the country, wild types of sorghum are prevalent (Doggett,
1991).Amsalu et al. (2001) reported that the presence of wild sorghum [Sorghum bicolor sp.
verticilliforum (L) Moench] in five districts of western, northwestern and southwestern Ethiopia.
Similarly, survey results of Tesso et al. (2008) indicated that wild sorghum has been found in
eastern, northeastern and northern parts of Ethiopia. In addition, sorghum genetic resources from
Ethiopia have been widely used in various breeding programs throughout the world (Smith and
Frederiksen, 2000). The greatest genetic diversity of sorghum is found in Ethiopia and adjacent
areas of northeast Africa (Poehlman and Sleper, 1995).
Sorghum is one of the most important cereal crops in Ethiopia and is produced in most parts of
the country. It is grown as a major food crop and the second most important cereal crop for
making local bread (Injera) next to tef (Teshome et al., 2007). The grain is generally used for
human food in different forms in different parts of the country. It is used as porridge,
"Injera","Kitta", "Nefro", infant food, and local beverages known as "Tella" and "Areke". The
leaves and stalks are used for animal feed and further the stalks are also used for construction of
houses and fences, and as fuel wood (MoA, 2010). Industrially, the grain is used to manufacture
alcohol, dextrose agar, edible oil, starch, syrup and wax (Rainford, 2005; Mamoudou et al.,
2006).
Ethiopia is the fifth sorghum-producing country in the world next to USA, India, Nigeria and
Argentina (Anonymous, 2010). The north central, western, northwestern and the eastern
midaltitude of Ethiopia are the areas of greater concentration of sorghum production (Wortmann
et al., 2013). Sorghum is the fifth most important cereal crop after maize, rice, wheat and barley
in the world (FAOSTAT, 2017). On the other hand, sorghum is the third most important cereal
crop next to maize and tef, which is cultivated annually on 1.88 million hectares, contributing
4.75 million tons to annual grain production in Ethiopia (CSA, 2017). Sorghum production
constituted 15.0% of land and 16.4% share of production from cereals cultivation, while it
covered 304,006.8 ha of land with a total production of 0.7 million tons in eastern Ethiopia
(CSA, 2017).
Production and productivity of sorghum is constrained by both biotic and abiotic stresses in the
world (Tari et al., 2013). The productivity of the crop in developing countries is low because of
the use of low-yielding cultivars and traditional production practices. In addition, sorghum
production by small scale farmers in arid areas is constrained by lack of inputs and seeds (Muui
et al., 2013). In Ethiopia, sorghum production and productivity is low due to abiotic and biotic
stresses. In the semi-arid areas of Ethiopia, inadequate soil water and nutrient supply pose major
threat and challenges to sorghum production (Gebreyesus, 2012).
Plant diseases, weeds and insect pests are recognized as major constraints of sorghum production
in Ethiopia. Several biotic constraints are responsible for more than 70% of annual sorghum
yield reduction (Berenji and Dahlberg, 2004). Sorghum is attacked by fungal, bacterial and viral
pathogens causing root, stalk, foliar, panicle and caryopsis diseases (Waniska et al., 2001; Prom
et al., 2005). Among these, fungal pathogens cause severe diseases, such as
anthracnose(Colletotrichum sublineolum), root and stalk rot (Fusarium moniliforme, Fusarium
thapsinum, or Colletotrichum sp.), seedling diseases (Pythium sp.), foliar diseases, such as leaf
blight (Exserohilum turcicum), zonate leaf spot (Gloeocercospora sorghi), sooty stripe
(Ramulispora sorghi), rust (Puccinia purpurea), ergot (Claviceps sorghi), aphides and head smut
(Sphacelotheca reliana) cause severe disease (Prom et al., 2005)
1.2. Objectives
 Determine the distribution and importance of sorghum anthracnose (C. sublineolum) and
aphides in major sorghum-growing areas.
 To review on sorghum anthracnose and aphides on sorghum crops
2. LITERATURE REVIEW
2.1. Sorghum Anthracnose Distribution and Economic Importance
Anthracnose occurs in all sorghum growing areas of the world (Patil et al., 2017). However, the
disease is commonly found in tropical and subtropical environments where there are warm and
humid climatic conditions that enhance the development and spread of the disease (Thakur and
Mathur, 2000); and it is widely prevalent and economically important in warm and humid
regions of Africa, the Americas and Asia (Chala et al., 2007). It is regarded as a constraint to
sorghum production in East African countries, including Ethiopia (Chala et al., 2007). Chala et
al. (2010b) reported that the disease had severe epidemics in Ethiopia and is highly prevalent and
very severe in sorghum-growing areas of eastern Ethiopia (Aragaw et al., 2019).
Sorghum anthracnose affects grain yield and yield components directly or indirectly. It affects
directly seed density and exacerbates early abortion of seeds. On the other hand, premature
drying of leaves is indirect cause that reduces sorghum grain yield (Mathur et al., 2002). It
invades the vascular tissue and interrupts the translocation of nutrients to the grain, reducing
grain yield and quality (Patil et al., 2017). The disease is known to affect leaf, stem, panicle and
grain. However, its effect is noticeable more on the leaves, leading to a considerable reduction of
photosynthesis and hence results in yield reduction (Crouch and Beirn, 2009).
2.2. Sorghum Production
Sorghum is one of the crops for which Ethiopia has been credited as being a center of origin
and/or diversity (Firew et al., 2009). Doggett (1988) suggested that sorghum originated and is
domesticated in the northeast quadrant of Africa, most likely in the Ethio-Sudan border regions.
Ethiopia is believed to be the center of origin and diversity of sorghum due to the presence of
diverse wild sorghum types. In the country, wild types of sorghum are prevalent (Doggett,
1991).Amsalu et al. (2001) reported that the presence of wild sorghum [Sorghum bicolor sp.
verticilliforum (L) Moench] in five districts of western, northwestern and southwestern Ethiopia.
Similarly, survey results of Tesso et al. (2008) indicated that wild sorghum has been found in
eastern, northeastern and northern parts of Ethiopia. In addition, sorghum genetic resources from
Ethiopia have been widely used in various breeding programs throughout the world (Smith and
Frederiksen, 2000). The greatest genetic diversity of sorghum is found in Ethiopia and adjacent
areas of northeast Africa (Poehlman and Sleper, 1995).
Ethiopia is the fifth sorghum-producing country in the world next to USA, India, Nigeria and
Argentina (Anonymous, 2010). The north central, western, northwestern and the eastern
midaltitude of Ethiopia are the areas of greater concentration of sorghum production (Wortmann
et al., 2013). Sorghum is the fifth most important cereal crop after maize, rice, wheat and barley
in the world (FAOSTAT, 2017). On the other hand, sorghum is the third most important cereal
crop next to maize and tef, which is cultivated annually on 1.88 million hectares, contributing
4.75 million tons to annual grain production in Ethiopia (CSA, 2017). Sorghum production
constituted 15.0% of land and 16.4% share of production from cereals cultivation, while it
covered 304,006.8 ha of land with a total production of 0.7 million tons in eastern Ethiopia
(CSA, 2017).
2.3. Constraints of Sorghum Production
Production and productivity of sorghum is constrained by both biotic and abiotic stresses in the
world (Tari et al., 2013). The productivity of the crop in developing countries is low because of
the use of low-yielding cultivars and traditional production practices. In addition, sorghum
production by small scale farmers in arid areas is constrained by lack of inputs and seeds (Muui
et al., 2013). In Ethiopia, sorghum production and productivity is low due to abiotic and biotic
stresses. In the semi-arid areas of Ethiopia, inadequate soil water and nutrient supply pose major
threat and challenges to sorghum production (Gebreyesus, 2012).
Plant diseases, weeds and insect pests are recognized as major constraints of sorghum production
in Ethiopia. Several biotic constraints are responsible for more than 70% of annual sorghum
yield reduction (Berenji and Dahlberg, 2004). Sorghum is attacked by fungal, bacterial and viral
pathogens causing root, stalk, foliar, panicle and caryopsis diseases (Waniska et al., 2001; Prom
et.al.,2005). Among these, fungal pathogens cause severe diseases, such as anthracnose
(Colletotrichum sublineolum), root and stalk rot (Fusarium moniliform, Fusarium thapsinum, or
Colletotrichum sp.), seedling diseases (Pythium sp.), foliar diseases, such as leaf blight
(Exserohilum turcicum), zonate leaf spot (Gloeocercospora sorghi), sooty stripe (Ramulispora
sorghi), rust (Puccinia purpurea), ergot (Claviceps sorghi) and head smut (Sphacelotheca reliana)
cause severe disease (Prom et al., 2005).In this connection, sorghum anthracnose
[Colletotrichum sublineolum Henn. syn. Colletotrichum graminicola (Ces.) G.C. Wils., both
anamorph; Glomerella graminicola Politis, teleomorph] is one of the most important sorghum
production-limiting factors in Ethiopia (Chala et al., 2007; Thakur, 2007). Anthracnose and rust
are considered to be of high importance in the eastern areas of Ethiopia, while grain mold and
smuts are also considered to be important sorghum diseases in the country (Wortmann et al.,
2009).
2.4. Sorghum Anthracnose
Anthracnose (C. sublineolum) is one of the most important diseases, which cause significant
biomass, quantity and quality losses in sorghum production (Sharma et al., 2012; Tesso et al.,
2012). The disease was first reported in West Africa in 1902, where it caused severe losses in
grain yield and quality (Thakur and Mathur, 2000).
2.5. Disease Symptoms
Anthracnose symptoms are observed on all parts of the sorghum but it is most conspicuous on
the leaf (Felderhoff et al., 2016). Foliar infection of sorghum anthracnose occurrs at any time of
the plant growth; however, symptoms are detected 40 days after emergence (Erpelding and
Prom, 2004). Depending on the cultivar, pathogenic variability and environmental conditions,
anthracnose causes circular spots or elongated lesions with red, tan, or blackish purple margins
and abundant fruiting bodies, the acervuli (Thakur and Mathur, 2000). The fruiting bodies
(acervuli) are observed as black spots in the center of the lesions and coalesce, which results in
leaf senescence and thereby limit the plant’s photosynthetic capacity (Crouch and Beirn, 2009).
It causes premature defoliation, which may cause death of plants before seed development, in
highly susceptible cultivars.
2.6. Survival and Transmission of the Pathogen
The fungus survives as mycelium, conidia and microsclerotia up to 18 months in crop debris or
above the soil surface (Mofokeng et al., 2017). The pathogen can survive in the seed for 2.5
years at room temperatures and Johnson grass serves as an alternative host for this pathogen
(Crouch and Beirn, 2009). The pathogen spores spread throughout the field by splashing of
rainwater and/or wind (Mofokeng et al., 2017).
Transmission of C. sublineolum is known to occur through the transfer of falcate-shaped conidia,
and the process is primarily dependent on water splash and blowing rain drops (Murphy et al.,
2008). The spores adhere to the host by the aid of mucilaginous hemicelluloses (Sugui et al.,
1998). The conidia germinate and undergo mitotic divisions to produce germ tubes with
appressoria that attach to the host tissue and form penetration pegs at the base. The fungus then
penetrates the host cell through the cuticle by turgor pressure developed within individual
appressorium (Patil et al., 2017). The infection peg enlarges into a globose infection vesicle,
giving rise to hyphae that grow intracellular and colonize the adjacent cells. The secondary
hyphae develop throughout the epidermal, mesophyll, and vascular tissue to begin the necrotic
phase of infection (Crouch and Beirn, 2009). This is followed by formation of the acervuli and
further spread of infection.
3. Sorghum aphids (Melanaphis sacchari)
Aphids are tiny, soft-bodied pests with piercing mouthparts that they use to siphon plant juice.
Usually, these aphids target newly emerging sorghum leaves. Lemon-colored yellow sugarcane
aphids are a pest. A significant sorghum pest in the world and many other nations is the sorghum
aphid (Singh et al. 2004).
Both the adult and the nymph aphids harm the sorghum plant during the entire lifespan. Aphids
initially appear on the lower leaves before moving up to the higher leaves and stalks. The
sorghum aphids' sorghum piercing and sucking of the sorghum juice hinder plant growth and
even result in plant death. The maize dwarf mosaic virus, which affects the growth of sorghum,
is also spread by sorghum aphids (Wang et al. 2009).
Figure 1.The illustration of sorghum aphid life cycle.
The length of a life cycle varies depending on the surroundings (temperature and humidity).
Aphids spend the winter as their winter hosts' eggs. Up until the third generation, which is an
alate virginoparae, the eggs are of the apterous kind and give birth to nymphs before being
transferred to the summer hosts. Surprisingly quickly, they reproduce roughly five days per
generation on the summer host. The aphids gather, move, and severely harm the summer plants
during this time. The alate virginoparae migrates back to the winter hosts at the end of the fall
and gives birth to the following female generation as sexual forms. The ladies lay their eggs on
the winter hosts after mating with the alate males, who are from the summer host. Alate, a
winged aphid; apterous, an aphid without wings.
3.1. Reproduction
Aphids that feed on sorghum can produce over ten generations annually. Depending on the
weather, Liaoning Province may experience 19–20 generations. A female aphid can give birth to
50–80 young in 3–4 weeks. Depending on the temperature, the newly born aphids may finish
growing and developing in 1-2 weeks. This explains how significant losses in sorghum output
can occur when big populations of aphids develop quickly (Singh et al. 2004).
For the development of pest control technology, such as the discovery of insecticides and
biotechnology, laboratory insect rearing is necessary . For the Asian corn borer , sorghum strip
borer, cotton bollworm, and yellow peach moth, our lab has successfully established rearing
techniques. Under laboratory settings, all of these insects go through the phases of eggs, larvae,
pupae, and adult/oviposition in their life cycles, which are listed in figure 2. The insects'
maturation stage is more uniform than it would be in the field since artificial feeds and regulated
environmental factors including temperature, humidity, and light were used. The life cycles of
these pests are on average relatively shorter in the laboratory than in the field. These rearing
features facilitate insect larva supply for developing pest controlling technologies.
Figure 2. The life cycle of Asian corn borer under laboratory conditions (temperature, 27 ± 1 °C;
relative humidity, 70–80%; and light: dark period, 16:8 h)
There are six larvae instars in its cycle which takes 27–30 d (egg, 4–5 d; larvae, 12–14 d; pupae,
3–4 d; and adult and oviposition, 6–8 d).
Table 2. Life cycle information of four sorghum insect pests under laboratory conditions
Items Insects
Asian corn
borer
Sorghum strip
borer
Cotton
bollworm
Yellow peach
moth
Life cycle (d) 27–30 45–55 29–34 32–37
Egg (d) 4–5 4–5 3–4 5–7
Larva (d) 12–14 28–35 14–16 12–15
Pupa (d) 3–4 7–10 5–7 7–8
Adult/oviposition
(d)
6–8 6–8 6–8 6–8
No. of instars 6 4–9 6 5
Light: dark (h) 16:8 16:8 16:8 16:8
Temperature (°C) 27 ± 1 27 ± 1 28 ± 1 26 ± 1
Relative humidity
(%)
70–80 80–90 60–70 >70
3.3. Control and Management
3.3.1. Insect resistant varieties
Breeding aphid cultivars is a common tactic for sorghum aphides management. Numerous
quantitative trait loci (QTLs) for insect resistance have been found by sorghum geneticists and
breeders. The advancements in this field have been thoroughly evaluated and described by
(Sharma et al. 2005).
One of the most harmful pests to sorghum during the seedling stage is the shoot fly (A. soccata).
Several male-sterile and maintainer lines, restorer lines, and their F1 hybrids were tested by
Sharma et al. (2006a,2006b) against three species of sorghum shot flies: shoot flies, spotted stem
borer (Ceramby-cidae partellus), and sugarcane aphid (M. sacchari). Satish et al. 2009) reported
QTLs for sorghum shoot fly resistance. They used repeated QTL mapping to find 29 QTLs. For
the majority of the QTLs, IS18551 supplied resistance alleles, and the related QTLs were co-
localized, suggesting they may be closely linked genes. Intriguingly, insect resistance loci are
conserved between maize and sorghum as evidenced by the insect resistant QTLs' locations in
syntenic maize genomic areas.
3.3.2. Insecticides
To manage sorghum insect pests, several pesticides have been created and are now being
utilized. Numerous pesticides used to combat aphids were investigated by Tiwari and Bhamare
in 2006. Dimethoate 30 EC at 0.03 percent and Imidacloprid 17.8 SC at 0.009 percent, two of the
insecticides they tested, were the most successful in reducing the aphid population to 1.17 and
1.84 aphids per cm2 leaf, respectively. The sorghum plant was highly protected in these studies
by Dimethoate treatment, which also produced the maximum grain production (2,205.75 kg/ha).
The concentration, insect stage, sorghum developmental stage, and application technique all
affect how effective an insecticide treatment is. Costs could go up if pesticide levels are raised.
Low chemical application levels, though, might not be successful. In comparison to other phases,
applying pesticides at the early larval and eclosion stages may be more effective in controlling
insects. The most crucial time for preventing pest damage to sorghum plants is during the
blossoming stage.
3.3.3. Seed treatments
Another typical technique for controlling insects is seed treatment (Wang et al. 2009). In rabi
sorghum, Balikai and Bhagwat (2009) conducted seed treatment studies to control shoot flies,
shoot bugs (Peregrinus maidis), and aphids. Their findings showed that one of the best
approaches for controlling these three insects was treatment with thiamethoxam 70 WS at 3 g/kg
seed.
Imidacloprid is a highly efficient pesticide for aphid control by seed treatment, according to
Wang and Liu (1999). The growth of the seedlings and the germination of the seeds were
unaffected by this treatment. The leading stage may also experience the protective effects. This
seed treatment can also reduce black cutworm and boost yield.
3.3.4. Crop rotation and other managements
Since it efficiently reduces the accumulation of sorghum insects on the same field, crop rotation
is a widely used technique. In a three-year field study, Chilcutt and Matocha (2007) examined
the impact of fertilizer treatments, crop rotation, and tillage on sorghum head insects. Although
there is a higher density of Oebalus pugna x., lowering tillage may occasionally result in a drop
in O. pugnax when sorghum is rotated with cotton.
According to Spurthi et al. (2009), intercropping the sorghum hybrid CSH-14 with red gram
(Cajanus Cajan) or soybean (Glycine max) could dramatically lessen the stem borer (C.
partellus) infestation while also increasing yields.
In the soil or in crop waste, many sorghum insects overwinter. Therefore, since they limit the
sources of pests, traditional tillage and the removal of previous-plant leftovers (crop and weeds)
can lessen insect damage. Depending on the insect type, seeding either early or late can be
successful in various regions. Insect damage can be efficiently reduced by varying planting times
to create a mismatch between insect larvae stages and plant growth phases.
Light or sex pheromone mass-trapping have been used in China to control insect pests (Wang et
al. 2009) based on insect's habits. Qing et al. (1990) reported that sex pheromone mass-trapping
is an economical and effective method for managing sorghum strip borer. A total of 451 moths
were caught per pheromone mass-trapping container from April to September.
3.3.5.Biological control
Kudachi and Balikai (2009) studied the efficacy of botanicals for the management of lesser grain
borer (Rhizopertha dominica) in sorghum during storage under their laboratory conditions. Their
experiments included 15 treatments and the results showed that calamus rhizomes (1%) were
significantly superior in protecting sorghum grains from R. dominica up to 180 d after treatment.
Diarisso et al. (2005) reported that spraying extracts from the leaves of Dursban or neem seed
(Azadirachta indica) could effectively control aphids. Sorghum plants treated with 5.3 mg
Dursban/L or 200 g neem seed jelly/L had only a few aphid bugs. Deepthi et al. (2008) evaluated
biorational pesticides for the management of stem borer (Ceramby-cidae partellus) in sweet
sorghum. Their treatments include endosulfan, carbofuran, neem seed kernel extract,
Metarhizium anisopliae, nimbecidine, Bacillus thuringiensis (Bt), plant mixture, and Vitex
negundo. The Bt treatment gave the best leaf protection and fewer dead hearts. The plants treated
with neem seed extract showed the best stem protection. Kandalkar and Men (2006) reported that
three sprays of Bt (var. kurstaki) effectively controlled sorghum stem borer (C. parellus) and
gave the maximum grain yield compared to the control or other treatments. These observations
indicate that Bt technology has great potential for controlling the sorghum stem borer.
4. CONCLUSIONS
5. References
Kandalkar HG, Men UB (2006) Efficacy of Bacillus thuringiensis var.kurstaki against sorghum
stem borer, Chilo parellus (Swinhoe).J. Biol. Control 20, 101–104.
Deepthi J, Shekharappa, Patil RK (2008) Evaluation of biorational pesticides for the
management of stem borer, Chilo partellus Swinhoe in sweet sorghum. Karnataka J. Agr.
Sci. 21, 293–294.
Diarisso NY, Diourte M, Pendleton BB ´ (2005) Use of local plants to control sorghum insect
pests in the field. Int. Sorghum Millets Newsl.46, 60–62.
Spurthi GS, Shekarappa, Patil RK, Puttanavar MS, Ramegowda GK(2009) Effect of
intercropping on the incidence of stem borer and armyworm in sorghum. J. Entomol. Res.
33, 89–92.
Wang LJ, Xu XD, Jiang Y, Dong HY, Li LK, Luan SY (2009) Control technology for major
sorghum pests. Bull. Agr. Sci. Technol. Sin.12, 159–160.
Qing J, Chen C, Wang K, Huang R (1990) Study on fluctuation,harmfulness and the techniques
of sex pheromone mass-trapping for forecasting spotted sugarcane borer. J. Southwest
Agr. Univ.Sin. 12, 30–32.
Chilcutt CF, Matocha JE (2007) Effects of crop rotation, tillage, and fertilizer applications on
sorghum head insects. J. Ecol. Entomol.100, 88–94.
Balikai RA, Raju A, Biradar BD, Sajjanar GM, Hundekar AR (2009) Loss estimation due to
shoot bug, Peregrinus maidis (Ashmead) in rabi sorghum under field conditions. J.
Entomol. Res. 33, 77–83.
Tiwari RB, Bhamare VK (2006) Bioefficacy of different insecticides against sorghum aphid
(Melanaphis sacchari Zehnter). Int. J. Plant Sci. (Muzaffarnagar) 1, 113–115.
Satish K, Srinivas G, Madhusudhana R, Padmaja PG, Nagaraja Reddy R, Murali Mohan S,
Seetharama N (2009) Identification of quantitative trait loci for resistance to shoot fly in
sorghum[Sorghum bicolor (L.) Moench]. Theor. Appl. Genet. 119, 1425–1439.
Sharma HC, Reddy BVS, Dhillon MK, Venkateswaran K, Singh BU, Pampapathy G, Folkertsma
RT, Hash CT, Sharma KK (2005) Host plant resistance to insects in sorghum: present
status and need for future research. J. SAT Agr. Res. 1, 1–8.
Singh BU, Padmaja PG, Seetharama N (2004) Biology and management of the sugarcane aphid,
Melanaphis sacchari (Zehntner) (Homoptera: Aphididae), in sorghum: A review. Crop
Prot. 23, 739–755.
Sharma HC, Dhillon MK, Pampapathy G (2006a) Multiple-resistance to sorghum shoot fly,
spotted stemborer and sugarcane aphid in sorghum. Int. J. Tropical Insect Sci. 26, 239–
245.
Sharma HC, Dhillon MK, Reddy BVS (2006b) Expression of resistance to Atherigona soccata in
F1 hybrids involving shoot flyresistant and susceptible cytoplasmic male-sterile and
restorer lines of sorghum. Plant Breed. 125, 473–477.

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review on sorghum anthracnose and aphids

  • 1. COLLEGE OF AGRICULTURE AND ENVIRONMENTAL SCIENCE DEPARTMENT OF PLANT SCIENCE M.Sc. in Plant Protection Graduate Program Economic Pests of Ethiopia BY: Zemed Wobale Birhanie ……ID: BDU1401871 Review on sorghum anthracnose and sorghum aphids Submitted to Asmare D. (PhD) July, 2022 Bahir Dar, Ethiopia
  • 2. INTRODUCTIONS Sorghum [Sorghum bicolor (L.) Moench] is an important cereal crop that belongs to the grass family Poaceae (Dahlberg, 2000). It is tolerant to harsh growing conditions with best adaptation under drought-prone and the semi-arid tropical regions of the world (Ng'uni et al., 2011; Burrell et al., 2015). Sorghum is called 'camel of crops' as it can withstand drought greatly. It has four features, which make it one of the drought resistant crops, such that it has a very large root to leaf-surface area; in time of drought, it will roll its leaves to lessen water loss; if drought continues, it will go into dormancy rather than drying; and leaves are protected by a waxy cuticle (Khosla et al., 1995). Sorghum is cultivated both in tropical and temperate climates for multiple uses (Dahlberg et al., 2011). Sorghum is the most important cereal crop and is the dietary staple of millions people in many countries in the world (Reddy et al., 2010). Further, it is a major source of food, feed, and fuel and fiber crop across a range of environments and production systems. Hence, sorghum is a major crop for the sustenance of human and livestock populations in hot and dry areas of the world (Sharma et al., 2012). Sorghum is one of the crops for which Ethiopia has been credited as being a center of origin and/or diversity (Firew et al., 2009). Doggett (1988) suggested that sorghum originated and is domesticated in the northeast quadrant of Africa, most likely in the Ethio-Sudan border regions. Ethiopia is believed to be the center of origin and diversity of sorghum due to the presence of diverse wild sorghum types. In the country, wild types of sorghum are prevalent (Doggett, 1991).Amsalu et al. (2001) reported that the presence of wild sorghum [Sorghum bicolor sp. verticilliforum (L) Moench] in five districts of western, northwestern and southwestern Ethiopia. Similarly, survey results of Tesso et al. (2008) indicated that wild sorghum has been found in eastern, northeastern and northern parts of Ethiopia. In addition, sorghum genetic resources from Ethiopia have been widely used in various breeding programs throughout the world (Smith and Frederiksen, 2000). The greatest genetic diversity of sorghum is found in Ethiopia and adjacent areas of northeast Africa (Poehlman and Sleper, 1995). Sorghum is one of the most important cereal crops in Ethiopia and is produced in most parts of the country. It is grown as a major food crop and the second most important cereal crop for making local bread (Injera) next to tef (Teshome et al., 2007). The grain is generally used for
  • 3. human food in different forms in different parts of the country. It is used as porridge, "Injera","Kitta", "Nefro", infant food, and local beverages known as "Tella" and "Areke". The leaves and stalks are used for animal feed and further the stalks are also used for construction of houses and fences, and as fuel wood (MoA, 2010). Industrially, the grain is used to manufacture alcohol, dextrose agar, edible oil, starch, syrup and wax (Rainford, 2005; Mamoudou et al., 2006). Ethiopia is the fifth sorghum-producing country in the world next to USA, India, Nigeria and Argentina (Anonymous, 2010). The north central, western, northwestern and the eastern midaltitude of Ethiopia are the areas of greater concentration of sorghum production (Wortmann et al., 2013). Sorghum is the fifth most important cereal crop after maize, rice, wheat and barley in the world (FAOSTAT, 2017). On the other hand, sorghum is the third most important cereal crop next to maize and tef, which is cultivated annually on 1.88 million hectares, contributing 4.75 million tons to annual grain production in Ethiopia (CSA, 2017). Sorghum production constituted 15.0% of land and 16.4% share of production from cereals cultivation, while it covered 304,006.8 ha of land with a total production of 0.7 million tons in eastern Ethiopia (CSA, 2017). Production and productivity of sorghum is constrained by both biotic and abiotic stresses in the world (Tari et al., 2013). The productivity of the crop in developing countries is low because of the use of low-yielding cultivars and traditional production practices. In addition, sorghum production by small scale farmers in arid areas is constrained by lack of inputs and seeds (Muui et al., 2013). In Ethiopia, sorghum production and productivity is low due to abiotic and biotic stresses. In the semi-arid areas of Ethiopia, inadequate soil water and nutrient supply pose major threat and challenges to sorghum production (Gebreyesus, 2012). Plant diseases, weeds and insect pests are recognized as major constraints of sorghum production in Ethiopia. Several biotic constraints are responsible for more than 70% of annual sorghum yield reduction (Berenji and Dahlberg, 2004). Sorghum is attacked by fungal, bacterial and viral pathogens causing root, stalk, foliar, panicle and caryopsis diseases (Waniska et al., 2001; Prom et al., 2005). Among these, fungal pathogens cause severe diseases, such as anthracnose(Colletotrichum sublineolum), root and stalk rot (Fusarium moniliforme, Fusarium thapsinum, or Colletotrichum sp.), seedling diseases (Pythium sp.), foliar diseases, such as leaf
  • 4. blight (Exserohilum turcicum), zonate leaf spot (Gloeocercospora sorghi), sooty stripe (Ramulispora sorghi), rust (Puccinia purpurea), ergot (Claviceps sorghi), aphides and head smut (Sphacelotheca reliana) cause severe disease (Prom et al., 2005) 1.2. Objectives  Determine the distribution and importance of sorghum anthracnose (C. sublineolum) and aphides in major sorghum-growing areas.  To review on sorghum anthracnose and aphides on sorghum crops
  • 5. 2. LITERATURE REVIEW 2.1. Sorghum Anthracnose Distribution and Economic Importance Anthracnose occurs in all sorghum growing areas of the world (Patil et al., 2017). However, the disease is commonly found in tropical and subtropical environments where there are warm and humid climatic conditions that enhance the development and spread of the disease (Thakur and Mathur, 2000); and it is widely prevalent and economically important in warm and humid regions of Africa, the Americas and Asia (Chala et al., 2007). It is regarded as a constraint to sorghum production in East African countries, including Ethiopia (Chala et al., 2007). Chala et al. (2010b) reported that the disease had severe epidemics in Ethiopia and is highly prevalent and very severe in sorghum-growing areas of eastern Ethiopia (Aragaw et al., 2019). Sorghum anthracnose affects grain yield and yield components directly or indirectly. It affects directly seed density and exacerbates early abortion of seeds. On the other hand, premature drying of leaves is indirect cause that reduces sorghum grain yield (Mathur et al., 2002). It invades the vascular tissue and interrupts the translocation of nutrients to the grain, reducing grain yield and quality (Patil et al., 2017). The disease is known to affect leaf, stem, panicle and grain. However, its effect is noticeable more on the leaves, leading to a considerable reduction of photosynthesis and hence results in yield reduction (Crouch and Beirn, 2009). 2.2. Sorghum Production Sorghum is one of the crops for which Ethiopia has been credited as being a center of origin and/or diversity (Firew et al., 2009). Doggett (1988) suggested that sorghum originated and is domesticated in the northeast quadrant of Africa, most likely in the Ethio-Sudan border regions. Ethiopia is believed to be the center of origin and diversity of sorghum due to the presence of diverse wild sorghum types. In the country, wild types of sorghum are prevalent (Doggett, 1991).Amsalu et al. (2001) reported that the presence of wild sorghum [Sorghum bicolor sp. verticilliforum (L) Moench] in five districts of western, northwestern and southwestern Ethiopia. Similarly, survey results of Tesso et al. (2008) indicated that wild sorghum has been found in eastern, northeastern and northern parts of Ethiopia. In addition, sorghum genetic resources from Ethiopia have been widely used in various breeding programs throughout the world (Smith and
  • 6. Frederiksen, 2000). The greatest genetic diversity of sorghum is found in Ethiopia and adjacent areas of northeast Africa (Poehlman and Sleper, 1995). Ethiopia is the fifth sorghum-producing country in the world next to USA, India, Nigeria and Argentina (Anonymous, 2010). The north central, western, northwestern and the eastern midaltitude of Ethiopia are the areas of greater concentration of sorghum production (Wortmann et al., 2013). Sorghum is the fifth most important cereal crop after maize, rice, wheat and barley in the world (FAOSTAT, 2017). On the other hand, sorghum is the third most important cereal crop next to maize and tef, which is cultivated annually on 1.88 million hectares, contributing 4.75 million tons to annual grain production in Ethiopia (CSA, 2017). Sorghum production constituted 15.0% of land and 16.4% share of production from cereals cultivation, while it covered 304,006.8 ha of land with a total production of 0.7 million tons in eastern Ethiopia (CSA, 2017). 2.3. Constraints of Sorghum Production Production and productivity of sorghum is constrained by both biotic and abiotic stresses in the world (Tari et al., 2013). The productivity of the crop in developing countries is low because of the use of low-yielding cultivars and traditional production practices. In addition, sorghum production by small scale farmers in arid areas is constrained by lack of inputs and seeds (Muui et al., 2013). In Ethiopia, sorghum production and productivity is low due to abiotic and biotic stresses. In the semi-arid areas of Ethiopia, inadequate soil water and nutrient supply pose major threat and challenges to sorghum production (Gebreyesus, 2012). Plant diseases, weeds and insect pests are recognized as major constraints of sorghum production in Ethiopia. Several biotic constraints are responsible for more than 70% of annual sorghum yield reduction (Berenji and Dahlberg, 2004). Sorghum is attacked by fungal, bacterial and viral pathogens causing root, stalk, foliar, panicle and caryopsis diseases (Waniska et al., 2001; Prom et.al.,2005). Among these, fungal pathogens cause severe diseases, such as anthracnose (Colletotrichum sublineolum), root and stalk rot (Fusarium moniliform, Fusarium thapsinum, or Colletotrichum sp.), seedling diseases (Pythium sp.), foliar diseases, such as leaf blight (Exserohilum turcicum), zonate leaf spot (Gloeocercospora sorghi), sooty stripe (Ramulispora sorghi), rust (Puccinia purpurea), ergot (Claviceps sorghi) and head smut (Sphacelotheca reliana)
  • 7. cause severe disease (Prom et al., 2005).In this connection, sorghum anthracnose [Colletotrichum sublineolum Henn. syn. Colletotrichum graminicola (Ces.) G.C. Wils., both anamorph; Glomerella graminicola Politis, teleomorph] is one of the most important sorghum production-limiting factors in Ethiopia (Chala et al., 2007; Thakur, 2007). Anthracnose and rust are considered to be of high importance in the eastern areas of Ethiopia, while grain mold and smuts are also considered to be important sorghum diseases in the country (Wortmann et al., 2009). 2.4. Sorghum Anthracnose Anthracnose (C. sublineolum) is one of the most important diseases, which cause significant biomass, quantity and quality losses in sorghum production (Sharma et al., 2012; Tesso et al., 2012). The disease was first reported in West Africa in 1902, where it caused severe losses in grain yield and quality (Thakur and Mathur, 2000). 2.5. Disease Symptoms Anthracnose symptoms are observed on all parts of the sorghum but it is most conspicuous on the leaf (Felderhoff et al., 2016). Foliar infection of sorghum anthracnose occurrs at any time of the plant growth; however, symptoms are detected 40 days after emergence (Erpelding and Prom, 2004). Depending on the cultivar, pathogenic variability and environmental conditions, anthracnose causes circular spots or elongated lesions with red, tan, or blackish purple margins and abundant fruiting bodies, the acervuli (Thakur and Mathur, 2000). The fruiting bodies (acervuli) are observed as black spots in the center of the lesions and coalesce, which results in leaf senescence and thereby limit the plant’s photosynthetic capacity (Crouch and Beirn, 2009). It causes premature defoliation, which may cause death of plants before seed development, in highly susceptible cultivars. 2.6. Survival and Transmission of the Pathogen The fungus survives as mycelium, conidia and microsclerotia up to 18 months in crop debris or above the soil surface (Mofokeng et al., 2017). The pathogen can survive in the seed for 2.5 years at room temperatures and Johnson grass serves as an alternative host for this pathogen
  • 8. (Crouch and Beirn, 2009). The pathogen spores spread throughout the field by splashing of rainwater and/or wind (Mofokeng et al., 2017). Transmission of C. sublineolum is known to occur through the transfer of falcate-shaped conidia, and the process is primarily dependent on water splash and blowing rain drops (Murphy et al., 2008). The spores adhere to the host by the aid of mucilaginous hemicelluloses (Sugui et al., 1998). The conidia germinate and undergo mitotic divisions to produce germ tubes with appressoria that attach to the host tissue and form penetration pegs at the base. The fungus then penetrates the host cell through the cuticle by turgor pressure developed within individual appressorium (Patil et al., 2017). The infection peg enlarges into a globose infection vesicle, giving rise to hyphae that grow intracellular and colonize the adjacent cells. The secondary hyphae develop throughout the epidermal, mesophyll, and vascular tissue to begin the necrotic phase of infection (Crouch and Beirn, 2009). This is followed by formation of the acervuli and further spread of infection.
  • 9. 3. Sorghum aphids (Melanaphis sacchari) Aphids are tiny, soft-bodied pests with piercing mouthparts that they use to siphon plant juice. Usually, these aphids target newly emerging sorghum leaves. Lemon-colored yellow sugarcane aphids are a pest. A significant sorghum pest in the world and many other nations is the sorghum aphid (Singh et al. 2004). Both the adult and the nymph aphids harm the sorghum plant during the entire lifespan. Aphids initially appear on the lower leaves before moving up to the higher leaves and stalks. The sorghum aphids' sorghum piercing and sucking of the sorghum juice hinder plant growth and even result in plant death. The maize dwarf mosaic virus, which affects the growth of sorghum, is also spread by sorghum aphids (Wang et al. 2009). Figure 1.The illustration of sorghum aphid life cycle. The length of a life cycle varies depending on the surroundings (temperature and humidity). Aphids spend the winter as their winter hosts' eggs. Up until the third generation, which is an alate virginoparae, the eggs are of the apterous kind and give birth to nymphs before being transferred to the summer hosts. Surprisingly quickly, they reproduce roughly five days per generation on the summer host. The aphids gather, move, and severely harm the summer plants during this time. The alate virginoparae migrates back to the winter hosts at the end of the fall and gives birth to the following female generation as sexual forms. The ladies lay their eggs on
  • 10. the winter hosts after mating with the alate males, who are from the summer host. Alate, a winged aphid; apterous, an aphid without wings. 3.1. Reproduction Aphids that feed on sorghum can produce over ten generations annually. Depending on the weather, Liaoning Province may experience 19–20 generations. A female aphid can give birth to 50–80 young in 3–4 weeks. Depending on the temperature, the newly born aphids may finish growing and developing in 1-2 weeks. This explains how significant losses in sorghum output can occur when big populations of aphids develop quickly (Singh et al. 2004). For the development of pest control technology, such as the discovery of insecticides and biotechnology, laboratory insect rearing is necessary . For the Asian corn borer , sorghum strip borer, cotton bollworm, and yellow peach moth, our lab has successfully established rearing techniques. Under laboratory settings, all of these insects go through the phases of eggs, larvae, pupae, and adult/oviposition in their life cycles, which are listed in figure 2. The insects' maturation stage is more uniform than it would be in the field since artificial feeds and regulated environmental factors including temperature, humidity, and light were used. The life cycles of these pests are on average relatively shorter in the laboratory than in the field. These rearing features facilitate insect larva supply for developing pest controlling technologies. Figure 2. The life cycle of Asian corn borer under laboratory conditions (temperature, 27 ± 1 °C; relative humidity, 70–80%; and light: dark period, 16:8 h)
  • 11. There are six larvae instars in its cycle which takes 27–30 d (egg, 4–5 d; larvae, 12–14 d; pupae, 3–4 d; and adult and oviposition, 6–8 d). Table 2. Life cycle information of four sorghum insect pests under laboratory conditions Items Insects Asian corn borer Sorghum strip borer Cotton bollworm Yellow peach moth Life cycle (d) 27–30 45–55 29–34 32–37 Egg (d) 4–5 4–5 3–4 5–7 Larva (d) 12–14 28–35 14–16 12–15 Pupa (d) 3–4 7–10 5–7 7–8 Adult/oviposition (d) 6–8 6–8 6–8 6–8 No. of instars 6 4–9 6 5 Light: dark (h) 16:8 16:8 16:8 16:8 Temperature (°C) 27 ± 1 27 ± 1 28 ± 1 26 ± 1 Relative humidity (%) 70–80 80–90 60–70 >70 3.3. Control and Management 3.3.1. Insect resistant varieties Breeding aphid cultivars is a common tactic for sorghum aphides management. Numerous quantitative trait loci (QTLs) for insect resistance have been found by sorghum geneticists and breeders. The advancements in this field have been thoroughly evaluated and described by (Sharma et al. 2005). One of the most harmful pests to sorghum during the seedling stage is the shoot fly (A. soccata). Several male-sterile and maintainer lines, restorer lines, and their F1 hybrids were tested by Sharma et al. (2006a,2006b) against three species of sorghum shot flies: shoot flies, spotted stem borer (Ceramby-cidae partellus), and sugarcane aphid (M. sacchari). Satish et al. 2009) reported QTLs for sorghum shoot fly resistance. They used repeated QTL mapping to find 29 QTLs. For
  • 12. the majority of the QTLs, IS18551 supplied resistance alleles, and the related QTLs were co- localized, suggesting they may be closely linked genes. Intriguingly, insect resistance loci are conserved between maize and sorghum as evidenced by the insect resistant QTLs' locations in syntenic maize genomic areas. 3.3.2. Insecticides To manage sorghum insect pests, several pesticides have been created and are now being utilized. Numerous pesticides used to combat aphids were investigated by Tiwari and Bhamare in 2006. Dimethoate 30 EC at 0.03 percent and Imidacloprid 17.8 SC at 0.009 percent, two of the insecticides they tested, were the most successful in reducing the aphid population to 1.17 and 1.84 aphids per cm2 leaf, respectively. The sorghum plant was highly protected in these studies by Dimethoate treatment, which also produced the maximum grain production (2,205.75 kg/ha). The concentration, insect stage, sorghum developmental stage, and application technique all affect how effective an insecticide treatment is. Costs could go up if pesticide levels are raised. Low chemical application levels, though, might not be successful. In comparison to other phases, applying pesticides at the early larval and eclosion stages may be more effective in controlling insects. The most crucial time for preventing pest damage to sorghum plants is during the blossoming stage. 3.3.3. Seed treatments Another typical technique for controlling insects is seed treatment (Wang et al. 2009). In rabi sorghum, Balikai and Bhagwat (2009) conducted seed treatment studies to control shoot flies, shoot bugs (Peregrinus maidis), and aphids. Their findings showed that one of the best approaches for controlling these three insects was treatment with thiamethoxam 70 WS at 3 g/kg seed. Imidacloprid is a highly efficient pesticide for aphid control by seed treatment, according to Wang and Liu (1999). The growth of the seedlings and the germination of the seeds were unaffected by this treatment. The leading stage may also experience the protective effects. This seed treatment can also reduce black cutworm and boost yield.
  • 13. 3.3.4. Crop rotation and other managements Since it efficiently reduces the accumulation of sorghum insects on the same field, crop rotation is a widely used technique. In a three-year field study, Chilcutt and Matocha (2007) examined the impact of fertilizer treatments, crop rotation, and tillage on sorghum head insects. Although there is a higher density of Oebalus pugna x., lowering tillage may occasionally result in a drop in O. pugnax when sorghum is rotated with cotton. According to Spurthi et al. (2009), intercropping the sorghum hybrid CSH-14 with red gram (Cajanus Cajan) or soybean (Glycine max) could dramatically lessen the stem borer (C. partellus) infestation while also increasing yields. In the soil or in crop waste, many sorghum insects overwinter. Therefore, since they limit the sources of pests, traditional tillage and the removal of previous-plant leftovers (crop and weeds) can lessen insect damage. Depending on the insect type, seeding either early or late can be successful in various regions. Insect damage can be efficiently reduced by varying planting times to create a mismatch between insect larvae stages and plant growth phases. Light or sex pheromone mass-trapping have been used in China to control insect pests (Wang et al. 2009) based on insect's habits. Qing et al. (1990) reported that sex pheromone mass-trapping is an economical and effective method for managing sorghum strip borer. A total of 451 moths were caught per pheromone mass-trapping container from April to September. 3.3.5.Biological control Kudachi and Balikai (2009) studied the efficacy of botanicals for the management of lesser grain borer (Rhizopertha dominica) in sorghum during storage under their laboratory conditions. Their experiments included 15 treatments and the results showed that calamus rhizomes (1%) were significantly superior in protecting sorghum grains from R. dominica up to 180 d after treatment. Diarisso et al. (2005) reported that spraying extracts from the leaves of Dursban or neem seed (Azadirachta indica) could effectively control aphids. Sorghum plants treated with 5.3 mg Dursban/L or 200 g neem seed jelly/L had only a few aphid bugs. Deepthi et al. (2008) evaluated biorational pesticides for the management of stem borer (Ceramby-cidae partellus) in sweet
  • 14. sorghum. Their treatments include endosulfan, carbofuran, neem seed kernel extract, Metarhizium anisopliae, nimbecidine, Bacillus thuringiensis (Bt), plant mixture, and Vitex negundo. The Bt treatment gave the best leaf protection and fewer dead hearts. The plants treated with neem seed extract showed the best stem protection. Kandalkar and Men (2006) reported that three sprays of Bt (var. kurstaki) effectively controlled sorghum stem borer (C. parellus) and gave the maximum grain yield compared to the control or other treatments. These observations indicate that Bt technology has great potential for controlling the sorghum stem borer.
  • 16. 5. References Kandalkar HG, Men UB (2006) Efficacy of Bacillus thuringiensis var.kurstaki against sorghum stem borer, Chilo parellus (Swinhoe).J. Biol. Control 20, 101–104. Deepthi J, Shekharappa, Patil RK (2008) Evaluation of biorational pesticides for the management of stem borer, Chilo partellus Swinhoe in sweet sorghum. Karnataka J. Agr. Sci. 21, 293–294. Diarisso NY, Diourte M, Pendleton BB ´ (2005) Use of local plants to control sorghum insect pests in the field. Int. Sorghum Millets Newsl.46, 60–62. Spurthi GS, Shekarappa, Patil RK, Puttanavar MS, Ramegowda GK(2009) Effect of intercropping on the incidence of stem borer and armyworm in sorghum. J. Entomol. Res. 33, 89–92. Wang LJ, Xu XD, Jiang Y, Dong HY, Li LK, Luan SY (2009) Control technology for major sorghum pests. Bull. Agr. Sci. Technol. Sin.12, 159–160. Qing J, Chen C, Wang K, Huang R (1990) Study on fluctuation,harmfulness and the techniques of sex pheromone mass-trapping for forecasting spotted sugarcane borer. J. Southwest Agr. Univ.Sin. 12, 30–32. Chilcutt CF, Matocha JE (2007) Effects of crop rotation, tillage, and fertilizer applications on sorghum head insects. J. Ecol. Entomol.100, 88–94. Balikai RA, Raju A, Biradar BD, Sajjanar GM, Hundekar AR (2009) Loss estimation due to shoot bug, Peregrinus maidis (Ashmead) in rabi sorghum under field conditions. J. Entomol. Res. 33, 77–83. Tiwari RB, Bhamare VK (2006) Bioefficacy of different insecticides against sorghum aphid (Melanaphis sacchari Zehnter). Int. J. Plant Sci. (Muzaffarnagar) 1, 113–115. Satish K, Srinivas G, Madhusudhana R, Padmaja PG, Nagaraja Reddy R, Murali Mohan S, Seetharama N (2009) Identification of quantitative trait loci for resistance to shoot fly in sorghum[Sorghum bicolor (L.) Moench]. Theor. Appl. Genet. 119, 1425–1439.
  • 17. Sharma HC, Reddy BVS, Dhillon MK, Venkateswaran K, Singh BU, Pampapathy G, Folkertsma RT, Hash CT, Sharma KK (2005) Host plant resistance to insects in sorghum: present status and need for future research. J. SAT Agr. Res. 1, 1–8. Singh BU, Padmaja PG, Seetharama N (2004) Biology and management of the sugarcane aphid, Melanaphis sacchari (Zehntner) (Homoptera: Aphididae), in sorghum: A review. Crop Prot. 23, 739–755. Sharma HC, Dhillon MK, Pampapathy G (2006a) Multiple-resistance to sorghum shoot fly, spotted stemborer and sugarcane aphid in sorghum. Int. J. Tropical Insect Sci. 26, 239– 245. Sharma HC, Dhillon MK, Reddy BVS (2006b) Expression of resistance to Atherigona soccata in F1 hybrids involving shoot flyresistant and susceptible cytoplasmic male-sterile and restorer lines of sorghum. Plant Breed. 125, 473–477.