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Abstract

  1. Top of page
  2. Abstract
  3. Introduction
  4. Sorghum Insect Pests
  5. Life Cycles, Feeding, and Economical Significance of a Few Sorghum Insects
  6. Control and Management
  7. Acknowledgements
  8. References

Sorghum (Sorghum bicolor) has high levels of starch, sugar, and fiber and is one of the most important energy crops in the world. Insect damage is one of the challenges that impacts sorghum biomass production. There are at least 150 insect species that can infest sorghum varieties worldwide. These insects can complete several generations within a growing season, they target various parts of sorghum plants at developmental stages, and they cause significant biomass losses. Genetic research has revealed the existence of resistant genetics in sorghum and insect tolerant sorghum varieties have been identified. Various control methods have been developed, yet more effective management is needed for increasing sorghum biomass production. Although there are no transgenic sorghum products on the market yet, biotechnology has been recognized as an important tool for controlling insect pests and increasing sorghum production.

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Introduction

  1. Top of page
  2. Abstract
  3. Introduction
  4. Sorghum Insect Pests
  5. Life Cycles, Feeding, and Economical Significance of a Few Sorghum Insects
  6. Control and Management
  7. Acknowledgements
  8. References

Sorghum (Sorghum bicolor) is an important cereal crop worldwide that is widely cultivated for food, fiber, forage, ethanol, and sugar production (Li and Gu 2004; Liu et al. 2009). Sorghum fiber has great potential to be used as a low-cost feedstock for ethanol production (Thomason et al. 2009). Sweet sorghum (Sorghum bicolor Linn. Moench) contains high levels of sugar, grows tall and fast, and generates a large amount of biomass (Reddy et al. 2007; Liu et al. 2009). To use sorghum for biofuel generation, developments are needed to further increase and stabilize sorghum biomass production and to store it for a continuing supply. Sorghum's sweetness attracts not only people, but also pathogens and insects, so that effective pest management practices are essential. This review will focus on the insect pests and on management practices for increasing sorghum biomass production.

Sorghum Insect Pests

  1. Top of page
  2. Abstract
  3. Introduction
  4. Sorghum Insect Pests
  5. Life Cycles, Feeding, and Economical Significance of a Few Sorghum Insects
  6. Control and Management
  7. Acknowledgements
  8. References

At least 150 insect species have been reported as pests of sorghum worldwide (Harris 1995), and more than 100 of them occurred in Africa (Kruger et al. 2008). Twenty-nine important insect families were actively studied by entomologists in the last decade (Table 1).

Table 1. Sorghum insect species and distributions
Family NameLatin nameCommon nameDistributionReferences
AcrididaeDichroplus elongatusGrasshopperArgentinaBulacio et al. 2005
AcrididaeOrphulella punctataShort-horned grasshoppersArgentinaBulacio et al. 2005
AphididaeSchizaphis graminumGreenbugBrazil, Egypt, and USAFigueira et al. 2005; Nagaraj et al. 2005; Slman et al. 2006; Radchenko and Zubov 2007; Ayyanath et al. 2008; Wu and Huang 2008; Burd and Puterka 2009
AphididaeMelanaphis sacchariSorghum aphidAfrica, Australia, China, India, and USADiarisso et al. 2005; Chang et al. 2006; Li et al. 2006; Sharma et al. 2006a, 2006b; Tiwari and Bhamare 2006; Ibrahim et al. 2008; Wang et al. 2009
AphididaeRhopalosiphum maidisCorn leaf aphidBrazil, China, and USAFonseca et al. 2006; Sun and Dong 2006; Michels and Matis 2008
BlissidaeBlissus occiduusChinch bugUSAAnderson et al. 2006
BostrichidaeRhizopertha dominicaGrain borerIndia and USAGetchell and Subramanyam 2008; Kudachi and Balikai 2009.
BruchidaeCallosobruchus chinensisPulse beetleIndiaHampanna et al. 2006
CecidomyiidaeStenodiplosis sorghicolaSorghum midgeAustralia, Burkina, Mali, and USADiarisso et al. 2005; Sharma et al. 2005; Franzmann et al. 2006; Michels and Burd 2007; Radchenko and Zubov 2007; Damte et al. 2009
ChrysomelidaeDiabrotica speciosaAmerican corn rootwormUSAWalsh 2007
ChrysomelidaeDiabrotica virgiferaWestern Corn RootwormItalySaladini et al. 2009
CicadellidaeDalbulus elimatusMexican corn leafhopperMexicoMoya-Raygoza 2007
CicadellidaeDalbulus maidisCorn leafhopperMexicoMoya-Raygoza 2007
CoccinellidaeHippodamia convergensLady beetleCzechMichaud and Jyoti 2007
CoccidaePulvinaria tenuivalvataRed-striped soft scale insectEgyptEl-Shazly et al. 2005
CoreidaeLeptoglossus phyllopusLeaf-footed bugUSAProm and Perumal 2008
CrambidaeDichocrocis punctiferalisYellow peach mothChinaWang et al. 2004; Sun and Dong 2006; Zhao et al. 2008; Lu et al. 2009a; Wang et al. 2009
CurculionidaeAnthonomus grandisBoll weevilsUSAShowler 2006
CurculionidaeSitophilus zeamaisMaize weevilNigeriaBabarinde et al. 2008
CurculionidaeSitophilus oryzaeRice weevilCameroon, India, Nigeria, and USASunilkumar et al. 2005; Bamaiyi et al. 2007; Getchell and Subramanyam 2008; Hampanna et al. 2006; Haile 2006; Ladang et al. 2008
CydnidaeCyrtomenus bergiThe subterranean burrower bugDenmarkRiis et al. 2005
DelphacidaePeregrinus maidisShoot bug and corn planthopperIndia, the Caribbean Islands, and Pacific OceansKumar and Prabhuraj 2006; Singh and Nadoor 2008; Balikai and Bhagwat 2009; Balikai et al. 2009
FormicidaeAcromyrmex lundiLeaf-cutting antArgentinaDans et al. 2009
GelechiidaeSitotroga cerealellaAngoumois grain mothEritreaHaile 2006
HetrodinaeAcanthoplus discoidalis MalawiMviha et al. 2007
HomopteraPyrilla perpusillaSugarcane leaf hopperIndiaSingh et al. 2007
HodotermitidaeAnacanthotermes turkestanicusTurkestan termiteCentral AsiaKhamraev et al. 2007
Miridae/LygaeidaeNysius natalensisThe LygaeidAfrica and ChinaWan and Hao 1992; Kruger et al. 2007, 2008
Miridae/LygaeidaeLygaeidaeSeed bugsAfricaKruger et al. 2008
MiridaeEurystylus oldiHead bugAfrica, Ghana, Mali, Nigeria, and NigerDeu et al. 2005; Diarisso et al. 2005; Showemimo 2006; Tanzubil et al. 2007; Kruger et al. 2007, 2008
MiridaeCalocoris angustatusSorghum earhead bugsIndiaDalip et al. 2007
MiridaeCampylomma sp.Mullein bugAfrica and GhanaTanzubil et al. 2007; Kruger et al. 2007, 2008
MiridaeMirid sp.Mirid bugSouth AfricaKruger et al. 2007
MiridaeCreontiades pallidusThe Shedder bugAfrica and GhanaTanzubil et al. 2007; Kruger et al. 2008
MiridaeTaylorilygus sp.The avocado bugGhanaTanzubil et al. 2007
MiridaeMegacoelum apicaleNotFindGhanaTanzubil et al. 2007
MiridaeSthenaridea suturalisPlant bugsAfricaKruger et al. 2008
MiridaeStenotus rubrovittatusSorghum plant bugAfrica and JapanKashin et al. 2009; Kawasaki et al. 2009; Kichishima et al. 2009; Kruger et al. 2008
MuscidaeAtherigona soccataShoot flyAfrica, Europe, Egypt, India, Iraq, Kenya, and PakistanBalikai et al. 2008, Sharma et al. 2006a, 2006b; Kumar and Prabhuraj 2007; Tahir et al. 2005; Al-Karboli and Al-Nakhli 2008; Biradar et al. 2008; Gomashe et al. 2008; Kumar et al. 2008; Obonyo et al. 2008; Salman and Abdel Moniem 2008; Aruna and Padmaja 2009; Satish et al. 2009; Shrinivas et al. 2009
Noc NoctuidaeSesamia creticaGreater sugar cane borersEgypt, CameroonAboubakary et al. 2008; Ezzeldin et al. 2009
NoctuidaeMythimna separataArmywormIndiaSpurthi et al. 2009.
NoctuidaeBusseola fuscaAfrican stem borersub-Saharan and AfricaCalatayud et al. 2008; Juma et al. 2008
NoctuidaeSesamia calamistisPink stem borerAfrica and SudanOkrikata and Anaso 2008; Ong’amo et al. 2008
NoctuidaeHelicoverpa zeaCorn earwormUSAChilcutt and Matocha 2007
NoctuidaeHelicoverpa armigeraCotton bollwormAustralia, India, and ChinaHerde 2005; Sun and Dong 2006; Vijaykumar et al. 2007; Jaspreet et al. 2008
NoctuidaePectinophora gossypiellaPink bollwormIndiaJaspreet et al. 2008
NoctuidaeEarias spp.Spiny bollwormIndiaJaspreet et al. 2008
NoctuidaeSpodoptera frugiperdaFall ArmywormBrazil and USACosta et al. 2006; Zeledon and Pitre 2007
NoctuidaeSesamia nonagrioidesCorn stalk borerGreeceDimou et al. 2007
PemphigidaeTetraneura nigriabdominalisThe root aphidTaiwanKuo et al. 2006
PentatomidaeNezara viridulaSouthern green stink bugAfrica and USAKruger et al. 2007, 2008; Smith et al. 2008
PentatomidaeAcrosternum hilareGreen stink bugUSASmith et al. 2008
PentatomidaeEuschistus servusBrown stink bugUSASmith et al. 2008
PentatomidaeOebalus pugnaxRice stink bugUSAChilcutt and Matocha 2007
PyralidaeSciropohaga excerptalisTop shoot borersIndiaShukla et al. 2006
PyralidaeChilo partellusStem borerAsia, Africa, Ethiopia, and IndiaJiang and Schulthess 2005; Kandalkar and Men 2006; Marulasiddesha et al. 2007; Kumar et al. 2006; Sharma et al. 2006a, 2007; Asmare 2008; Chand and Deepthi et al. 2008; Reddy et al. 2009; Spurthi et al. 2009
PyralidaeChilo sacchariphagusSorghum strip borerChinaHuang et al. 2000, 2001; Xu et al. 2002; Wang et al. 2004; Sun and Dong 2006
PyralidaeMampava bipunctella ChinaWang et al. 1989; Huang et al. 1991; Xu et al. 2003; Wang et al. 2004; Sun and Dong 2006; Zhao et al. 2008
PyralidaeOstrinia furnacalisAsia corn borerChinaWang et al. 2004; Sun and Dong 2006; Zhao et al. 2008; Wang et al. 2009
ScarabaeidaePachnoda interruptaThe sorghum chaferEthiopiaBengtsson et al. 2009
ScarabaeidaePhyllophaga crinitaWhite grubsMexicoRodríguez-del-Bosque and Salinas-García 2008
TenebrionidaeTribolium confusumThe confused flour beetleEritrea and IndiaHaile 2006; Vanita et al. 2006
ThripidaeThrips palmiThrips palmiIndiaLokesh et al. 2008

Africa is one of the main sorghum production areas. Forty-two panicle-feeding pests have become serious pests of sorghum in West, Central, and South Africa. Thirty-nine herbivorous Hemipteran species were found in South Africa, with the most abundant family being Miridae and Lygaeidae (Kruger et al. 2008). Greenbug (Schizaphis graminum), sorghum midge (Stenodiplosis sorghicola), fall armyworm (Spodoptera frugiperda), and corn borers (Ostrinia furnacalis) are the major pests in North America (Munson et al. 1993; Wu and Huang 2008; Damte et al. 2009). Grasshoppers (Dichroplus elongatus) are important sorghum pests in South America. Sorghum shoot fly (Atherigona soccata), corn rootworm (Diabrotica speciosa), and corn borers are important sorghum pests in Asia, Africa, and Europe (Table 1). Sorghum aphid (Melanaphis sacchari) and sorghum midge significantly impact sorghum production in Australia.

In China, 22 sorghum insect species have been reported. These pests include soil insects (i.e. mole cricket Scapteriscus vicinus, black cutworm Agrotis ipsilon, and wireworm Agriotes lineatus), foliage pests (i.e. fall armyworm S. frugiperda), stalk pests (i.e. Asian corn borer O. furnacalis and sorghum strip borer Chilo sacchariphagus), and panicle pests (i.e. cotton bollworm Helicoverpa armigera and yellow peach moth Dichocrocis punctiferalis). Sorghum panicle pests are particularly important because they can cause complete yield losses (Sun and Dong 2006). Sorghum aphids (M. sacchari Zehntner) are widely distributed in China and are one of the most important sorghum pests (Wang et al. 2009).

Many of the sorghum pests can damage crops other than sorghum, such as corn, cotton, and millet. Therefore, the host ranges of these sorghum pests need to be carefully considered when rotating crops for managing the insects. Further complicating management practices, several insect species can occur simultaneously in the same field. Except for the differences in host ranges, the target tissues and plant developmental target stages vary significantly among the pests.

Life Cycles, Feeding, and Economical Significance of a Few Sorghum Insects

  1. Top of page
  2. Abstract
  3. Introduction
  4. Sorghum Insect Pests
  5. Life Cycles, Feeding, and Economical Significance of a Few Sorghum Insects
  6. Control and Management
  7. Acknowledgements
  8. References

Insect life cycle means the period from the egg to the adult. The length of a life cycle varies under different environmental conditions and for different insect species. The main environmental factors affecting cycle time include temperature and humidity. Knowledge of insect life cycles can improve the control of insect damage. A few stalk insects (Asian corn borer and sorghum strip borer), panicle pests (cotton bollworm, yellow peach moth, and honeydew moth), and sorghum aphid are described below as examples to illustrate their life cycles and impacts on sorghum production.

Asian corn borer (Ostrinia furnacalis)

Late stage larvae overwinter in the stalks and/or ear shank of crops. In the spring, the larvae pupate, and the adults emerge in May. After eclosion, the adults can lay eggs on the back of the leaves by the leaf veins. The eggs then hatch into larvae 3–5 d later and the larvae feed on the plants. The early stage larvae (1st–3rd instars) mainly damage new leaves, tassels, bracts, and filaments. After the 4th instar, the larvae can enter into stalks to feed and pupate there. The larvae are able to move and are attracted by light, moisture, and sugars. The life cycle of this pest is 31–39 d long, depending on the temperature and humidity in the field. One female can lay 300–700 eggs, which begin the next generation (Duan et al. 2008; Li 2009).

Asian corn borer can reproduce two to three generations per year in the Yellow River valley of China, and four to seven generations in South China, depending on the environmental conditions. Overlapping generations are often observed under high temperatures and humidity. Asian corn borer can cause 10–30% sorghum yield losses annually (Duan et al. 2008; Li 2009). Yield losses caused by this pest result from the damage to leaves and stems, which reduces photosynthesis and the nutrient supply to the grains, and increases lodging of the sorghum plants.

Sorghum strip borer (Chilo sacchariphagus)

The sorghum strip borer is also called the sugarcane stem borer. The adults lay eggs on the back of leaves near the petiole or on the stalk. One female can lay 200–250 eggs. Early stage larvae feed on the young leaves of sorghum, while later instars drill and tunnel into the stalk. The larvae cause damage to the stalk up until pupation. Late stage larvae overwinter in sorghum or other crop stalks. The larvae pupate at the end of May and emerge 10–15 d later. The number of instars is variable from four to nine, but is usually six to seven. It takes 42–60 d to complete one life cycle.

Sorghum strip borers can reproduce two generations annually in North China and three to five generations in South China. The newly hatched larvae gather on young leaves where they feed, causing damage to the plant. After the 3rd instar, the larvae drill into the stalks and feed on the pith. The larvae will also feed on the tassel of the sorghum. Yield losses caused by this pest can be 10–40% (Qing et al. 1990; Li 2009) due to reduced photosynthesis and increased stalk lodging.

Cotton bollworm (Helicoverpa armigera)

The larvae pupate in the soil about 5–10 cm underground in the late fall where they will overwinter. The moths emerge in May–June when the temperature is above 15 °C. The adults mate and oviposit during the night, and female bollworms lay eggs on sorghum plants. The eggs hatch 3–5 d later and larvae feed on sorghum heads or cotton leaves. The young larvae (the 1st and 2nd instars) cause little damage to crops. However, the larger larvae (the 3rd to 5th instars) infest the leaves, tears, buds, bolls or heads of sorghums. The larvae pupate in the soil 15–20 d later and moths emerge 10–20 d after pupation. The length of its life cycle is 35–50 d, depending on the field environmental conditions.

Cotton bollworm can occur in three to four generations in North China and four to five generations in South China per year. It likes high temperatures and high humidity. The female adults hide on the bottom leaves, and lay eggs on the ear and the glumes of sorghum at night. One female can produce 1000–3000 eggs. The newly hatched larvae feed on the flowers and ear, then enter into tassel and feed on the grain, sometimes eating the whole panicle (Yu 1989). Cotton bollworms can cause 50–70% sorghum yield losses.

Yellow peach moth (Dichocrocis punctiferalis)

This pest overwinters in late stages of larvae in the straw of sorghum and other crops. The larvae pupate and emerge in May and June. Then the adults mate, lay eggs on the peach at night, and hatch a week later. One female can deposit as many as 700 eggs in a lifetime. The larvae drill into the peach and damage it. There are five stages of instars and the larvae live for 15–20 d. The adults emerge in August and may lay eggs on the late-maturing peach or transfer to sorghum, sunflower, and late-maturing corn to complete their 2nd or 3rd generations. The yellow peach moth adults lay eggs on the branches of ears and glumes. The larvae drill into panicles and damage the seeds at the milky maturity stage of the sorghum. After the 3rd instar, the larvae feed and excrete on the ear to cause ear mold and decrease the sorghum quality.

Yellow peach moth can complete its life cycle within 34 to 43 d, reproduce two to three generations in North China and four to five generations in South China. It is becoming an important pest for corn in the Yellow River Valley. It can cause 30–40% yield losses of summer sorghum (Zhang 1969; Wang et al. 2009).

Honeydew moth (Cryptoblabes gnidiella)

They overwinter in a form of late stage larva in the ear or sheath of the sorghum at the beginning of October. Then the larva pupates in the middle of the following June and emerges in July. One female can produce ∼200 eggs on average. The eggs hatch 3–5 d later, and the larvae can damage the ear till pupation. The pupae may last 7–8 d before becoming adults (Li 2009).

Honeydew moth is widely distributed in China. It can complete two to three life cycles in North China and four to six generations in South China each year, depending on the weather conditions. It can occur in drought fields under proper temperatures. The Honeydew moth adults lay eggs on the ear or glumes of sorghum. The larvae will feed on the ear and the seeds, and cause yield losses of about 40% on average (Li 2009).

Sorghum aphids (Melanaphis sacchari)

Aphids are small soft-bodied pests that have piercing mouth parts for sucking juice from plant tissues. These aphids usually attack newly emerged sorghum leaves. Yellow sugarcane aphids are lemon yellow in color (Munson et al. 1993). Sugarcane aphid is an important sorghum pest in China and many other countries (Chen 1999; Singh et al. 2004).

An aphid life cycle is illustrated in Figure 1. Aphids cause damage throughout the sorghum lifecycle by both the adult and the nymph. The aphids first occur at the bottom leaves and then disperse to the upper leaves and stalks. Piercing and sucking of the sorghum juice by the sorghum aphids slows down the growth of plants and even causes plant death. The sorghum aphids are also vectors of maize dwarf mosaic virus, which impacts the development of sorghum (Munson et al. 1993; Wang and Liu 1999; Wang et al. 2009).

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Figure 1. The illustration of sorghum aphid life cycle. The length of a life cycle varies with environmental conditions (temperature and humidity). Aphids overwinter by eggs in winter hosts. The eggs hatch into the apterous type and give birth to nymph until the third generation, which is alate virginoparae and transfer to the summer hosts. Their reproduction rate is surprisingly high, about 5 d per generation on the summer host. During this period, the aphids accumulate, migrate, and badly damage the summer plants. At the end of fall, the alate virginoparae migrates back to the winter hosts, and produces the female generation as sexual forms. The females mate with the alate males which are from the summer host, and then lay eggs on the winter hosts. Alate, aphid with wings; Apterous, aphid without wings.

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Sorghum aphids reproduce more than 10 generations per year. In Liaoning Province, 19–20 generations may occur, depending on weather conditions. A female aphid may produce 50–80 offspring within 3–4 weeks. The newly born aphids may complete their growth and development in 1–2 weeks, depending on the temperature. This explains how large populations of aphids can develop in a short period of time, and can result in severe sorghum yield losses (Munson et al. 1993; Singh et al. 2004).

Insect rearing in the laboratory is required for developing insect control technology, such as insecticide discovery and biotechnology (see next section). Our laboratory has successfully developed rearing methods for Asian corn borer (Figure 2), sorghum strip borer, cotton bollworm, and yellow peach moth. These insects all go through eggs, larvae, pupae, and adults/oviposition stages in their life cycles under the laboratory conditions as summarized in Table 2. Since controlled conditions (temperature, humidity, and light) and artificial diets were used, the developmental stage of the insects is more uniform compared with the situation in the field. The life cycles of these pests are on average relatively shorter in the laboratory than in the field (Table 2). These rearing features facilitate insect larva supply for developing pest controlling technologies.

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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).

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Table 2. Life cycle information of four sorghum insect pests under laboratory conditions
ItemsInsects
Asian corn borerSorghum strip borerCotton bollwormYellow peach moth
Life cycle (d)27–3045–5529–3432–37
Egg (d)4–54–53–45–7
Larva (d)12–1428–3514–1612–15
Pupa (d)3–47–105–77–8
Adult/oviposition (d)6–86–86–86–8
No. of instars64–965
Light: dark (h)16:816:816:816:8
Temperature (°C)27 ± 127 ± 128 ± 126 ± 1
Relative humidity (%)70–8080–9060–70>70

Control and Management

  1. Top of page
  2. Abstract
  3. Introduction
  4. Sorghum Insect Pests
  5. Life Cycles, Feeding, and Economical Significance of a Few Sorghum Insects
  6. Control and Management
  7. Acknowledgements
  8. References

Insect resistant varieties

A prevalent strategy for controlling sorghum insects is to breed insect resistant varieties. Sorghum geneticists and breeders have identified a number of quantitative trait loci (QTLs) for insect resistant breeding programs. Subudhi et al. (2002) and Sharma et al. (2005) have summarized and intensively reviewed the progress in this area.

Shoot fly (A. soccata) is one of the most destructive pests at the seedling stage for sorghum. Sharma et al. (2006a, 2006b) evaluated a number of male-sterile and maintainer lines, restorer lines, and their F1 hybrids against three sorghum shoot fly species, shoot fly, spotted stem borer (Cerambycidae partellus), and sugarcane aphid (M. sacchari). Satish et al. (2009) reported QTLs for resistance to sorghum shoot fly. They discovered 29 QTLs by multiple QTL mapping. IS18551 contributed resistant alleles for most of the QTLs, and the related QTLs were co-localized, indicating they may be tightly linked genes. Interestingly, the insect resistant QTLs are located in syntenic maize genomic regions, showing conservation of insect resistance loci between maize and sorghum.

Radchenko and Zubov (2007) carried out intensive research on the genetic diversity of sorghum in greenbug (S. graminum) resistance. They included sorghum germplasm from Africa (the primary centre of sorghum origin), USA, India, Latin American countries, and China in their study. Sweet sorghum is more susceptible to greenbug, while grain sorghum, broomcorn sorghum, and grass sorghum (Sudan grass) are relatively resistant to greenbug. Nagaraj et al. (2005) carried out linkage group mapping studies using single-sequence repeat (SSR) markers with sorghum recombinant inbred lines for sorghum greenbug resistance. They identified three QTLs associated with biotype I and five QTLs associated with biotype K of the greenbug. These QTLs account for 9.0–19.6% of phenotypic variation. Wu and Huang (2008) carried out mapping experiments to dissect genetic resistance to greenbug using 118 SSR markers. They discovered two QTLs on chromosome 9 for the greenbug resistance. These two QTLs account for approximately 55–80% and 1–6% of the resistance to greenbug feeding, respectively.

Marulasiddesha et al. (2007) developed a screening method for sorghum resistance to the stem borer C. partellus (Swinhoe). They reported that genotype SSV-7073 was the most stem borer resistant variety among the 23 grain and sweet sorghum lines they evaluated in the field.

Chang et al. (2006) reported that the aphid (M. sacchari) resistance is controlled by a single dominant gene. They have mapped an SSR maker which is located on linkage group 9 and linked with this resistance gene.

Deu et al. (2005) mapped head-bug resistant QTLs in an F2 population from a cross of sorghum cultivar Malisor 84–7 (head-bug resistant) and S 34 (head-bug susceptible). They built a genetic map including 92 loci distributed over 13 linkage groups. Three significant and seven putative QTLs were identified in this study.

Shukla et al. (2006) compared 14 sweet sorghum genotypes in a field experiment and reported that ASSVT-8, ASSVT-9 and ASSVT-10 showed significant tolerance to various borers compared to other genotypes. Gupta et al. (2007) reported that RSSV-9 and NSSH-104 were relatively tolerant to top shoot borer compared to SSV-84 in India. RSSV-9 and NSSH-104 also exhibited superior stalk strength.

Sorghum hybrids have been developed using cytoplasmic male-sterility (CMS), maintainer, and restorer lines. Some of the hybrids are not only high-yielding, but also more resistant to insect pests (Sharma et al. 2005). For example, ICSA88019 hybrid showed 10.9% of midge damage, whereas the corresponding susceptible hybrid had 22.2% damage (Sharma et al. 2005).

Insecticides

A number of insecticides have been developed and used to control sorghum insect pests (Buutin 2009; Wang et al. 2009). Tiwari and Bhamare (2006) studied a number of insecticides against aphids. Among the insecticides they evaluated, Dimethoate 30 EC at 0.03% and imidacloprid 17.8 SC at 0.009% were the most effective, and treatments could decrease the aphid population to 1.17 and 1.84 aphids per cm2 leaf, respectively. In these experiments, Dimethoate treatment protected the sorghum plant well and gave the highest grain yield (2 205.75 kg/ha).

The efficiency of insecticide treatments depends on the dosage, insect stage, sorghum developmental stage, and application method. Higher levels of insecticides may increase costs. However, low levels of chemical applications may not be effective. Applying chemicals at early larvae and eclosion stages may be more effective in controlling insects compared to other stages. For the sorghum plants, the flowering stage is the most critical period for controlling insect damage.

Seed treatments

Seed treatment is another common method to control insects (Buutin 2009; Wang et al. 2009). Balikai and Bhagwat (2009) carried out seed treatment experiments against shoot fly, shoot bug (Peregrinus maidis), and aphid in rabi sorghum. Their results indicated that treatment with thiamethoxam 70 WS at 3 g/kg seed was one of the effective methods in controlling these three insects.

Wang and Liu (1999) reported that imidacloprid is a very effective insecticide for controlling aphids by seed treatment. This treatment had no negative impact on seed germination and seedling growth. The protective effects can extend to the heading stage. In addition, this seed treatment can control black cutworm and increase yield.

Crop rotation and other managements

Crop rotation is a common practice, since it effectively reduces the build up of sorghum insects in the same field. Chilcutt and Matocha (2007) conducted a 3 year field study and reported the effects of crop rotation, tillage, and fertilizer applications on sorghum head insects. Rotation of sorghum with cotton could reduce Helicoverpa zea, although with increased density of Oebalus pugna x. Reducing tillage may decrease O. pugnax in some instances.

Spurthi et al. (2009) reported that sorghum hybrid CSH-14 intercropped with pigeon peas (red gram, Cajanus cajan) or soybean (Glycine max) could significantly reduce the infestation of stem borer (C. partellus) and increase yields.

Many sorghum insects overwinter in soil or in previous-crop residues. Therefore, conventional tillage and the removal of previous-plant residue (crop and weeds) can reduce damage by insects, since these reduce the pest sources. Early or late sowing can be effective in some areas, depending on the insect species. Alternating the planting time in order to create a mismatch between insect larvae stages and plant developmental stages can effectively reduce insect damage.

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.

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 (Cerambycidae partellus) in sweet sorghum. Their treatments include endosulfan, carbofuran, neem seed kernel extract, Metarhizium anisopliae, nimbecidine, Bacillus thuringiensis (Bt), plant mixture, and Vitex negundo. Bt treatment gave the best leaf protection and less dead heart. 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 sorghum stem borer.

Biotechnology

The commercialization and cultivation of transgenic Bt-cotton and Bt-corn are significantly increasing agricultural productivity. More and more evidence demonstrates that transgenic Bt-crops are safe for human food chain (Whalon and Wingerd 2003; Zehnder et al. 2007). Also, the World Health Organization (WHO) (2002) concluded that genetically modified (GM) foods “currently available on the international market have passed risk assessments and are not likely to present risks for human health” and it is well accepted that individual transgenic crops and their safety should be assessed on a case-by-case basis.

A wide range of insecticidal genes from Bt strains have been used in insect control through biotechnology methods. Bt genes that are currently used in Bt-crops exhibit significant efficacy against Lepidoptera. The sucking insects, such as aphids, are not sensitive to normal Bt proteins. Many of the sorghum insects (i.e. Asian corn borer and cotton bollworm) are Lepidoptera insects. Therefore, Bt genes used in corn and cotton crops could be used to control Lepidoptera insects in transgenic sorghum plants.

Although there is no transgenic sorghum product on the market yet, sorghum genomics and biotechnology have been significantly developed (Liu et al. 2009). The sorghum genome sequence has been reported and used (Paterson et al. 2009; Gui et al. 2010), and many genes and molecular markers have been developed (Liu et al. 2009). To generate insect resistant transgenic sorghum plants, efficient genetic transformation technology, promoters, and terminators are needed along with the insect resistance genes. Various genetic transformation methods have been developed for sorghum (reviewed by Girijashankar and Swathisree 2009). Among the methods, Agrobacterium-mediated transformation is the most efficient and has been improved using various treatments (Verma et al. 2008; Gurel et al. 2009). Lu et al. (2009b) have developed an Agrobacterium-mediated transformation system that can generate marker-free transgenic sorghum plants in a public line [P898012]. Eight co-cultivation conditions were examined for their effects on transformation. The average transformation frequencies were 0.4–0.7%. Visarada et al. (2008) have developed a simple system for selecting and advancing transgenic progeny of sorghum. Various sorghum promoters have been identified, including stem-specific (Potier et al. 2008), sucrose synthase gene (Sivasudha and Kumar 2008), meristem-specific (Verma and Kumar 2005), and wound-inducible (Girijashankar et al. 2005) promoters.

Transgenic sorghum plants have been generated and characterized for improving nutrient utilization (Burla and Podile 2009) and expressing γ-kafirin (Bansal et al. 2008). Interestingly, Girijashankar et al. (2005) developed transgenic sorghum plants expressing a synthetic cry1Ac gene under a wound-inducible promoter mpiC1. The Bt-transgenic sorghum plants showed partial tolerance against first instars of the spotted stem borer (C. partellus). Further optimization of the insect resistant genes and promoters can lead to better insect control in sorghum.

Bt-crops have become a very important tool for insect pest management worldwide. In 2009, a total of 134 million hectares of biotech crops were planted and more than 30% of them are Bt-crops (James 2009). Bt-crops, such as Bt-cotton, not only control the targeted insect pest well, but also reduce the same pest's impact on other host crops (Wu et al. 2008). However, it has been found that insect pests can develop resistance in the Bt-crops (Tabashnik et al. 2009) and nontarget pests may become a major insect problem for the Bt-crops (Lu et al. 2010). For example, mirid bugs (Heteroptera, Miridae) were only minor insects in Northern China, and have increased 12-fold since 1997 and become a main pest problem in the region for cotton and other crops (Lu et al. 2010) due to the Bt-cotton. Strategy for overcoming these issues with Bt technology, such as using multiple Bt genes for insect resistance management, has been proposed (Bates et al. 2005; Lu et al. 2010). An integrated insect management system covering the whole life of sorghum and having long-term benefits is required for durable management of sorghum insect pests and increasing its biomass production.

Insects are one of the most important biofactors impacting sorghum biomass production for food, feed, fiber, and fuel worldwide. Although many insect controlling methods have been developed, they have limitations in terms of effectiveness, cost, or safety. Resistant sorghum varieties exhibit advantages in control of insect damages over other methods. Biotechnology and Bt genes have great potential for developing insect resistant sorghum varieties in combination with traditional breeding programs for increasing sorghum biomass production. One method cannot solve all pest problems in a crop and different methods should be used based on insect pest management (IPM) principles in each specific program.

(Co-Editor: Hai-Chun Jing)

Acknowledgements

  1. Top of page
  2. Abstract
  3. Introduction
  4. Sorghum Insect Pests
  5. Life Cycles, Feeding, and Economical Significance of a Few Sorghum Insects
  6. Control and Management
  7. Acknowledgements
  8. References

The authors wish to thank Beth Arnold for literature search and collection. Thanks to Drs Barbara Mazur, Nina Schmidt, and Beth Arnold for editing and critical review of the manuscript. The original larvae of Asian corn borer and cotton bollworm were provided by Dr Zhenying Wang and Dr Jie Zhang at the Institute of Plant Protection, Chinese Academy of Agricultural Sciences. Sorghum strip borer and yellow peach moth were originally collected from the sorghum plants in the experimental field of Dr Ruiheng Du at the Millet Institute, Hebei Academy of Agricultural Sciences. This work was supported by Pioneer Hi-Bred International Inc., A DuPont Company.

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  2. Abstract
  3. Introduction
  4. Sorghum Insect Pests
  5. Life Cycles, Feeding, and Economical Significance of a Few Sorghum Insects
  6. Control and Management
  7. Acknowledgements
  8. References
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