biology and management of the sugarcane aphid, melanaphis sacchari (zehntner) (homoptera:...

Crop Protection 23 (2004) 739–755

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Review

Biology and management of the sugarcane aphid, Melanaphissacchari (Zehntner) (Homoptera: Aphididae), in sorghum: a review

B.U. Singh*, P.G. Padmaja, N. Seetharama

National Research Centre for Sorghum, Rajendranagar, Hyderabad, Andhra Pradesh 500 030, India

Received 1 October 2003; received in revised form 3 December 2003; accepted 13 January 2004

Abstract

The sugarcane aphid, Melanaphis sacchari (Zehntner, 1897) is a key pest on sorghum and sugarcane in many areas of Africa, Asia,

Australia, the Far East, and parts of Central and South America. The status of research of its geographical distribution, host range,

nature of damage, extent of crop losses, and ecobiology in sorghum is summarized and research programs in different countries are

reviewed. Numerous germplasm accessions, A/B- and R-lines, agronomic elite lines, hybrids, and varieties, identified as sources of

resistance providing genetic diversity from different countries are listed. Studies on the components of resistance showed the

predominance of antixenosis for colonization/establishment on IS 1144C, IS 12664C, and TAM 428, and antibiosis was observed on

IS 12609C, IS 12664C, and TAM 428 for least number of days to reproduction, greater mortality, shorter longevity, and production

of no or fewer nymphs. The morpho-physiological traits and biochemical factors associated with resistance have been discussed.

There is a significant decline in diastase activity but increase in crude fiber and carbohydrates in the grain due to infestation by M.

sacchari. It is a vector of three persistent viruses (millet red leaf, sugarcane yellow leaf, and sugarcane mosaic viruses). Among the

control tactics, cultural practices, natural enemies, and chemical control together can prevent the sugarcane aphid from reaching the

economic threshold levels. Current progress has been reviewed and ideas for future research are suggested.

r 2004 Elsevier Ltd. All rights reserved.

Keywords: Biology; Management; Mechanisms of resistance,Melanaphis sacchari; Sorghum; Sugarcane aphid; Varietal resistance

troduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 740

eographical distribution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 740

ost range . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 740

ature of damage . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 742

rop losses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 742

cobiology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 743

arietal resistance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 743

irus vector . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 748

ultural practices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 748

atural enemies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 748

hemical control . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 751

ng author.

ss: [email protected] (B.U. Singh).

front matter r 2004 Elsevier Ltd. All rights reserved.

pro.2004.01.004

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12. Future research . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 751

Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 752

References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 752

Table 1

Geographical distribution of the sugarcane aphid, Melanaphis sacchari

(Zehntner)

Country Reference

Angola Mead (1978)

Argentina Delfino (1985)

Australia Passlow (1985)

Bhutan Agarwala (1985)

Botswana Flattery (1982)

Brazil Mead (1978)

China Wang (1961), Mead (1978), and Miao and Sunny

(1987)

Colombia Mead (1978)

Ecuador Mead (1978)

Egypt Mead (1978)

Ethiopia Megenasa (1982)

Florida Denmark (1988)

Haiti Mead (1978)

B.U. Singh et al. / Crop Protection 23 (2004) 739–755740

1. Introduction

The sugarcane aphid, Melanaphis sacchari (Zehntner,1897) (Homoptera: Aphididae) has been reportedvariously as Aphis sacchari (Zehntner) (Zimmerman,1948) and Longiunguis sacchari (Zehntner) (Eastop,1965). Roy Chaudhuri and Banerjee (1974) synony-mized both these species as M. sacchari (Zehntner).Eastop and Hille Ris Lambers (1976) reviewed theliterature on the nomenclature of sugarcane aphid, M.

sacchari, and listed that the genera, Geoktapia Mordo-vilko (1921), Longiunguis Van der Goot (1977), Masra-

phis Soliman (1938), Nevsikia Mordvilko (1932),Piraphis Borner (1932), Schizaphideilla Hille Ris Lam-bers (1939), Yezabura Matsumura (1917) as synonymsof the genus Melanaphis Van der Goot (1917).

Hawaii Mead (1978)

India Young (1970), Jotwani and Young (1972), Young

and Teetes (1977), Shuja-Uddin (1975), David and

Sandhu (1976), Alexander and Madhusudhanrao

(1977), Varma et al. (1978), Bapat (1981), Bhagat

(1981), Agarwala et al. (1983), Mote (1983), Mote

and Kadam (1984), Patil (1992), and Balikai (1997)

Indonesia Mead (1978)

Jamaica Edward (1937)

Japan Setokuchi (1973), Mead (1978), Hagio et al. (1985),

and Hagio and Ono (1986)

Kenya Le Pelley (1959), Nye (1960), and Starks (1969)

Nigeria Mead (1978)

Pakistan Hamid (1983)

Peru Mead (1978)

Philippines Rueda and Catling (1978), and Mead (1978)

South

Africa

Brain (1929), Muller and Scholl (1958), Matthee and

Oberholzer (1958), Matthee (1962), van Rensburg

and van Hamburg (1975), Anonymous (1981), van

Rensburg and Malan (1993), and van den Berg

(1999)

Sudan Schmutterer (1969), and Mead (1978)

Tabago Mead (1978)

Taiwan Chang (1981a, b), Chang and Fang (1984), and

Wilbrink (1922)

Tanzania Bohlen (1973)

Thailand Young (1970), Banzoger (1976), and Meksongsee

and Chawanapong (1985)

Trinidad Mead (1978)

Uganda Schmutterer (1969), and Mead (1978)

2. Geographical distribution

The geographical distribution of M. sacchari followsthe cultivation of sorghum (Sorghum bicolor (L.)Moench) and sugarcane (Saccharum officinarum (L.))worldwide covering Angola, Brazil, China, Colombia,Ecuador, Egypt, Ethiopia, Haiti, Hawaii, India, Indo-nesia, Japan, Jamaica, the middle East, Nigeria, Paki-stan, Peru, Philippines, Sudan, Thailand, Trinidad,Tabago, Uganda, and Venezuela (Eastop, 1955, 1965;Mead, 1978; CIE, 1981) (Table 1). It is considered as aneconomically important pest on sorghum in China(Wang, 1961), Taiwan (Chang, 1981a, b; Pi and Hsieh,1982a), Japan (Setokuchi, 1973), India (Young, 1970),South Africa (van Rensburg, 1973a), and Botswana,while it is common on cultivated sorghum in Zimbabwe(Flattery, 1982), where its economic importance has notbeen determined (Page et al., 1985). In North America,it occurs on sugarcane in Florida (Summers, 1978;Mead, 1978; Denmark, 1988), Hawaii (Zimmerman,1948; Pemberton, 1948), and Louisiana (White et al.,2001), and its economic status on sugarcane still remainsunclear (White et al., 2001).

Uruguay Delfino (1985)

Venezuela Mead (1978), Sanchez and Cermeli (1987), and

Aponte et al. (1988)

Zimbabwe Sithole et al. (1987)
3. Host range
Although M. sacchari (Zehntner) (Homoptera: Aphi-didae) is a minor pest on several crops, its pest status hasincreased rapidly since the early 1970s. The genusMelanaphis has 20 species associated with Gramineae

(Blackman and Eastop, 1984). The host range of M.

sacchari is largely restricted to the species of the genera:Saccharum, Sorghum, Oryza, Panicum, and Pennisetum

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Table 2

Host range of the sugarcane aphid, M. sacchari (Zehnt.) reported from different countries

Scientific name Common name Country from

which reported

Reference

Cynodon dactylon (L.) Bermuda grass, Taiwan Wilbrink (1922)

Burmagrass,

Common stargrass,

Devilgrass,

Dhubgrass

Miscanthus sinensis, (L.) Ornamental grass, Japan Setokuchi (1973), and Kawada (1995)

Japanese silvergrass

Oryza sativa (L.) Paddy, China, Miao and Sunny (1987)

Rice USA (Florida) Denmark (1988)

Panicum colonum Barnyard grass, USA (Florida), Denmark (1988)

(Syn: Echinochloa colonum) Jungle rice, Taiwan Wilbrink (1922)

Tufted annual grass

Panicum maximum Jacq. Hamilgrass, Botswana, van Rensburg (1973a)

Jacquin Guineagrass Zimbabwe van Rensburg (1973a)

Paspalum sanguinale Lamarck Hairy crabgrass USA (Florida) Wilbrink (1922)

(Syn: Digitaria sanguinalis)

Pennisetum sp. USA (Florida) Denmark (1988)

Saccharum officinarum Sugarcane Argentina, Delfino (1985)

USA (Florida), Mead (1978), and Denmark (1988)

USA (Hawaii), Pemberton (1948)

India (Sikkim), Agarwala et al. (1983)

India (Tamil Nadu), Alexander and Madhusudhanrao (1977)

India (Uttar Pradesh), Shuja-Uddin (1975), and Varma et al. (1978)

India (West Bengal), Agarwala et al. (1983)

Jamaica, Edward (1937)

USA (Louisiana), White et al. (2001)

Pakistan, Hamid (1983)

Philippines, Rueda and Catling (1978)

Taiwan Wilbrink (1922)

Setaria italica (L.) Beauv. Boar millet, South Africa, van Rensburg (1973a, b)

Foxtail millet, USA (Florida) Wilbrink (1922)

German millet,

Hay millet,

Italian millet,

Nunbank setaria

Sorghum bicolor (L.) Moench Sorghum Argentina, Delfino (1985)

USA (Florida), Wilbrink (1922), and Denmark (1988)

India (Karnataka), Patil (1992), and Balikai (1997)

India (Kashmir), Bhagat (1981)

India (Maharashtra), Mote (1983), and Mote and Kadam (1984)

India (Punjab), David and Sandhu (1976)

Japan, Setokuchi (1973), Hagio et al. (1985), and Hagio

and Ono (1986)

South Africa, van Rensburg and van Hamburg (1975)

van Rensburg and Malan (1983)

Taiwan, Chang (1981a, b), and Chang and Fang (1984)

Thailand, Banzoger (1976)

Uruguay, Delfino (1985)

Venezuela Sanchez and Cermeli (1987), and Aponte et al. (1988)

Sorghum halepense (L.) Pers. Aleppo grass, Japan, Kawada (1995)

Aleppo millet grass, South Africa van Rensburg (1973a)

Cuba grass,

Johnson grass

Sorghum verticilliflorum (Steud.)

Stapf.

Wild Sudangrass South Africa van Rensburg (1973a), and van Rensburg and

van Hamburg (1975)

Zea mays (L.) Maize Bhutan Agarwala (1985)

B.U. Singh et al. / Crop Protection 23 (2004) 739–755 741

(Denmark, 1988) (Table 2). Brain (1929) recorded M.

sacchari as a potential pest of sugarcane in coastal areasof Natal; and Muller and Scholl (1958), Matthee and

Oberholzer (1958), and Matthee (1962) found it as aregular pest on sorghum in Northern Transvaal in SouthAfrica. In addition, its host range spreads to Setaria

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italica (L.) Beauv. in the USA (Wilbrink, 1922) andSouth Africa (van Rensburg, 1973a), and Zea mays (L.)in Bhutan (Agarwala, 1985), including species of grassessuch as Cynodon dactylon (L.) in Taiwan (Wilbrink,1922), Miscanthus chinensis (L.) in Japan (Setokuchi,1973; Kawada, 1995), Paspalum sanguinale Lamarck inthe USA (Wilbrink, 1922), Sorghum halepense (L.) Pers.in South Africa (van Rensburg, 1973a) and Japan(Kawada, 1995), and Sorghum verticilliflorum (Steud.)Stapf. in South Africa (van Rensburg, 1973a; vanRensburg and van Hamburg, 1975). A list of the hostplants from which the sugarcane aphid has beenreported is provided in Table 2. The sugarcane aphidis anholocyclic throughout most of its range in tropicaland subtropical regions, but some sexual oviparae havebeen observed on Gramineae in Punjab, India (Davidand Sandhu, 1976). In addition, monoecious holocyclicforms have also been reported on sorghum (David andSandhu, 1976) and sugarcane (Yadava, 1966). Both alateand apterous virginoparous adults showed a strongertendency of preference to Sorghum bicolor and S. halepense

than Japanese silvergrass, Miscanthus sinensis (Anders)(Kawada, 1995), but had a strong preference for M.

sinensis over sugarcane (S. officinarum L.).

4. Nature of damage

Damage to sorghum by the sugarcane aphid dependson a number of factors including aphid density andinfestation duration. Sorghum is typically infested soonafter plant emergence, but significant infestationsusually occur during late growth stages, and in dryperiods (van Rensburg, 1973a). Sorghum responses toM. sacchari injury include purple leaf discoloration ofseedlings followed by chlorosis, necrosis, stunting, delayin flowering, and poor grain fill, including quality andquantity yield losses. The sugarcane aphid feeds on theabaxial surface of older sorghum leaves. Leaves belowthe infected ones are often covered with sooty moldswhich grow on the honeydew produced by the aphid(Narayana, 1975). Plant stress due to drought mayintensify damage to sorghum by the sugarcane aphid.The importance of M. sacchari as a pest on sorghum

results from its colonization when plants are 2–3 weeksold. Despite early colonization, notable increases inaphid numbers take place only after panicle exsertion(van Rensburg, 1973a). Therefore, the sugarcane aphidcolony has a relatively narrow time window withinwhich to increase population buildup. Physiological andbiological changes taking place during sorghum plantdevelopment can cumulatively affects the exponentialgrowth rates during early and mid-season, reaching asmany as 30 000 aphids on a single plant (Setokuchi,1977). However, the aphid densities decline quickly in2–3 weeks after peak abundance, and the factors

influencing decline are alate dispersal induced by aphiddensity as well as the poor host condition (vanRensburg, 1973b).There is a significant increase in population of M.

sacchari on sorghum from the boot to the soft doughstage (40–70 days after planting) in the spring, andheading to harvesting (60–100 days after planting) inautumn (Fang, 1990). Waghmare et al. (1995) observedpopulation increase and peaks during January, when thepost-rainy sorghum crop was between flowering andmilk stage, and declined thereafter till maturity. Plantgrowth stage and temperature had significant effects onthe population buildup of the sugarcane aphid, anddispersal occurs within 6–10 days at a temperatureregime of 15.1�C and 31.0�C (Balikai, 2001), 16.0�C and29.0�C (Mote and Kadam, 1984), 19.5�C and 34.7�C(Narayana et al., 1982), 22.5�C and 32.5�C, and at 84%RH (AICSIP, 1979–2003) and 18.0–31.0�C (van Re-nsburg, 1973a, b), but the population died at 35.0�C(Behura and Bohidar, 1983). In addition to thetemperature, cloudy weather together with increasinghumidity can result in aphids colonies completelycovering the abaxial surface of all leaves of sorghumplants (Mote, 1983). The highest rate of populationbuildup was at 94% and 43% RH and at 11.4�C and30.0�C temperature in the morning and afternoon,respectively (Waghmare et al., 1995). Aphid density wasgreater under irrigation than in unirrigated conditions,and its occurrence on sorghum at milk stage did notaffect the grain yields severely, but the fodder qualitydeteriorated (Balikai, 2001).

5. Crop losses

The sugarcane aphids remove the plant sap fromxylem tissues of leaves, and in large densities they causephysiological losses such as wilting/curling of leaves,and also result in chlorosis. However, the aphidnumbers necessary to cause yield reductions in sorghumvary based on the plant stage, interval between, andduration of infestation. The degree of plant moisturestress under which sorghum is grown as well as theinduction of stress due to aphid infestation also plays asignificant role in the amount of aphid injury that can betolerated.Sorghum yield losses ranging from minor to severe

have been reported in Botswana (Anonymous, 1974;Flattery, 1982), Zimbabwe (Page et al., 1985) and India(Mote and Kadam, 1984; Mote et al., 1985). In SouthAfrica, grain yield losses reached 60% (Matthee, 1962),and 46–78%, without insecticide control (van Rensburgand van Hamburg, 1975; van Rensburg, 1979; van denBerg, 2002). There are few direct and indirect estimatesof the sugarcane aphid damage in sorghum andsugarcane. In sorghum, the losses varied between

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12–26% and 10–31% with an overall loss of 16% and15% for grain yield and fodder yield, respectively(Balikai, 2001). Sugarcane aphid infested sorghum grainwas significantly associated with the poor preparation ofbeverages (Pi and Hsieh, 1982b), and reduction indiastatic activity, malt, and abrasive hardness index (vanden Berg et al., 2003) as well as causing grain yieldreduction and poor quality of forage sorghums (Seto-kuchi, 1979) similar to decrease in crude protein andincrease in neutral detergent fiber (NDF) by the yellowsugarcane aphid, Sipha flava (Fukumoto and Mau,1989). There was a significant reduction of 40.2% and39.1% in grain and fodder yields, respectively, wheninfested at 60 DAE, compared to 70 and 80 DAE with23% and 28% loss in grain yield, and 15% and 17%reduction in forage yields, respectively (Balikai, 2001).

6. Ecobiology

The earliest detailed study of the biology and lifehistory of the sugarcane aphid has been on sorghum inSouth Africa (van Rensburg, 1969, 1973a, b, 1976) andto a limited extent on sugarcane in Japan (Setokuchi,1980, 1988) and sorghum in India (Varma et al., 1978).The sugarcane aphid forms colonies of lemon-yellowapterae and alate individuals on the abaxial surface ofbasal leaves of a sorghum plant. Some alates havepatterned black markings along the dorsal scleritis(Eastop, 1955; Roy Chaudhuri and Banerjee, 1974;Blackman and Eastop, 1984). Reproduction is predo-minantly asexual with adults being either apterae oralate viviparous females. Sexual reproduction is alsoknown to occur on sorghum (David and Sandhu, 1976),however, the environmental conditions under whichsexual reproduction takes place have not been reported.It has four nymphal stadia, which are completed in 4.3–12.4 (Chang et al., 1982) or in 5 days (Manthe, 1992).Adults normally survive for 10–16 (Meksongsee andChawanapong, 1985), 14–37 (Chang et al., 1982), or 28days (van Rensburg, 1973a), and produce up to 68nymphs female�1 with an average of 34 (Meksongseeand Chawanapong, 1985), 45–89 (Chang et al., 1982), or96 at 18.0–31.0�C (van Rensburg, 1973a). Alatesproduce fewer nymphs and have a shorter life expec-tancy (van Rensburg, 1973a). It develops 51–61 genera-tions, averaging 56 generations annually underscreenhouse conditions (Chang et al., 1982). The lifespan of each generation is shorter in summer than inwinter (Chang et al., 1982). The number of days fornymphal development was increased with a decrease inlongevity and fecundity in aphids reared on sorghum at25.0�C and 16 h photoperiod (Kawada, 1995). There isalso evidence that the performance of the sugarcaneaphid varies in its adaptation when reared on sugarcanealone, while those reared on sorghum are adapted to

both sorghum and sugarcane (Setokuchi, 1988). Thepopulation density is influenced by variable tempera-tures and rainfall pattern (Chang et al., 1982).It overwinters parthenogenetically on ratoon sorghum

and wild alternate hosts such as S. verticilliflorum, S.

halepense (L.), Panicum maximum Jacq., and Setaria

spp. (van Rensburg, 1973a). The dispersal of alatesthroughout the year ensures that young cultivatedsorghum is infested soon after germination (vanRensburg, 1973a). The seasonal history of the sugarcaneaphid has been described in two phases: (i) fastexponential increase in numbers during the early seasonis terminated by heavy dispersal in the mid-season, andresponsible for considerable loss in yield; and (ii) latesummer phase, wherein natural enemies regulate theaphid populations at sub-economic levels for rest of theseason (van Rensburg and van Hamburg, 1975). Sincehighly significant correlations occur between the popu-lation development of wheat aphid and the sugarcaneaphid, the developmental parameters of the former hasbeen suggested to predict the similar population trend ofM. sacchari on sorghum (Niu, 1987).

7. Varietal resistance

Aphid density and plant damage under naturalinfestations have been used to select resistant sorghumgenotypes in the greenhouse and field conditions(Setokuchi, 1976; Pi and Hsieh, 1982a; Hagio andOno, 1986). Seedling and mature plant evaluationsdisplayed similar results (Pi and Hsieh, 1982a; Hagioand Ono, 1986), however, seedlings were preferred foreasy handling and control of infestation levels (Teetes,1980).

Sources and mechanisms of resistance: Several sources,levels, and mechanisms of resistance in sorghum to thesugarcane aphid comprising germplasm accessions,parental lines (A/B and R lines), agronomic elite lines,hybrids, varieties, and locals have been reported fromdifferent countries (Tables 3–6). Resistance to insectshas been characterized into three components asantixenosis (nonpreference), antibiosis, and tolerance(Painter, 1951; Horber, 1980; Smith, 1989; Smith et al.,1994). However, antixenosis and antibiosis resistance toM. sacchari in sorghum do not differ with plant age(Teetes, 1980). Both alate and apterous virginoparousadults showed a stronger tendency of preference to S.

bicolor and S. halepense rather than M. sinensis.Tolerance to the sugarcane aphid injury in sorghumincreases greatly with slight increase in plant height, andhas inherent advantage over antibiosis and antixenosisin that it does not impose selection pressure on aphidpopulations, and thus may have greater permanence.Setokuchi (1988) reported that M. sacchari failed toestablish on resistant lines under field conditions

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Table 3

Germplasm accessions of sorghum resistant to the sugarcane aphid, M. sacchari (Zehnt.) reported from different countries

Germplasm accession Origin Classification Resistance Country from

which reported

Reference

Level Mechanism

IS 44 USA Bicolor R India Balikai (1993)

IS 84 Mexico Durra/Kaura MR Japan Hagio and Ono (1986)

IS 718 USA Bicolor R India AICSIP (1988)

IS 1063 India Durra R India AICSIP (1988)


IS 1133C (SC 202) India Durra HR Botswana, Manthe (1992) and Teetes et al. (1995)

Zimbabwe Manthe (1992) and Teetes et al. (1995)





IS 1144C (SC 451) India Durra HR Ax, Ab Botswana, Manthe (1992) and Teetes et al. (1995)


IS 1366C (SC 210) India Durra R Ax Botswana, Manthe (1992) and Teetes et al. (1995)


IS 1598C (SC 214) India Dochna HR Ax, Ab Botswana, Manthe (1992) and Teetes et al. (1995)


IS 1840 R India Mote and Shahane (1988, 1993)

IS 2312 USA Durra R India AICSIP (1997)

IS 3796 USA Caffrorum MR Japan Hagio and Ono (1986)

IS 4657 India Durra R India Balikai (1993)

IS 5188C (SC 245) India Durra/Dochna HR Ab Botswana, Manthe (1992) and Teetes et al. (1995)



IS 5887C (SC 248) India Roxburghii HR Ax Botswana, Manthe (1992) and Teetes et al. (1995)


IS 6389C (SC 489) India Nandyal HR Botswana, Manthe (1992) and Teetes et al. (1995)


IS 6416C India Nandyal HR Ax Botswana, Manthe (1992) and Teetes et al. (1995)


IS 6426C (SC 497) India Nandyal HR Ax Botswana, Manthe (1992) and Teetes et al. (1995)


IS 6962C India Caudatum R Botswana, Manthe (1992) and Teetes et al. (1995)


IS 8100C Japan Caudatum/

Nigricans

HR Japan Hagio (1987, 1992) and Hagio and Ono

(1986)

IS 12158C (SC 984) Ethiopia Bicolor HR Ax, Ab Botswana, Manthe (1992) and Teetes et al. (1995)


IS 12551C (SC 31) Ethiopia Caudatum HR Ax, Ab Botswana, Manthe (1992) and Teetes et al. (1995)


IS 12599C (SC 90) Congo Guinea HR Ax, Ab Botswana, Manthe (1992) and Teetes et al. (1995)


IS 12608C (PI 257 595,

SC 108-14-E)

Ethiopia Caudatum HR Ax, Ab Botswana, Manthe (1992) and Teetes et al. (1995)


IS 12609C (SC 109) Ethiopia Caudatum R Botswana, Manthe (1992) and Teetes et al. (1995)


IS 12610C (SC 110-14) Ethiopia Caudatum R Botswana, Manthe (1992) and Teetes et al. (1995)

Taiwan Chang (1981a, b)


IS 12611C (SC 111 or

SC 170)

Ethiopia Bicolor R Botswana, Manthe (1992) and Teetes et al. (1995)


IS 12612C Ethiopia Bicolor R Japan, Hagio and Ono (1986), Hagio et al. (1985),

and Hagio (1987, 1992)

South Africa Wenzell et al. (1998)

IS 12637C (SC 146) Ethiopia Bicolor R Ab Botswana, Manthe (1992) and Teetes et al. (1995)


IS 12645C Ethiopia Bicolor HR Ax, Ab Botswana, Manthe (1992) and Teetes et al. (1995)



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Table 3 (continued)

Germplasm accession Origin Classification Resistance Country from

which reported

Reference

Level Mechanism

IS 12661C Ethiopia Bicolor HR Ax, Ab Botswana, Manthe et al. (1984), Manthe (1992), and

Teetes et al. (1995)

Zimbabwe Manthe et al. (1984), Manthe (1992), and

Teetes et al. (1995)

IS 12664C Ethiopia Bicolor HR Ax, Ab Botswana, Manthe (1992) and Teetes et al. (1995)


IS 12667C Ethiopia Bicolor R Botswana, Manthe (1992)

Zimbabwe Manthe (1992)

IS 14048 Malawi Guinea R India Balikai (1993)

IS 23250 (Sima) Caudatum–

Bicolor

R South Africa Peterson (2002)

IS 33843 Durra R India Balikai (1993)

Ax, antixenosis; Ab, antibiosis; HR, highly resistant; MR, moderately resistant; and R, resistant.


although resistant sources served as suitable hosts inconfinement studies, which indicate the involvement ofboth antixenosis and antibiosis mechanisms. Antixeno-sis for adult colonization was noticed in TAM 428, IS1144C, IS 1366C, IS 1598C, IS 6416C, IS 6426C, IS12661C, and IS 12664C. Among them, IS 1144C and IS12664C were preferred less than the resistant check,TAM 428. High levels of antibiosis expressed in TAM428, IS 1144C, IS 5188C, IS 12609C, and IS 12664C forleast number of days to reproduction; TAM 428, IS12609C, and IS 12664C for greater mortality andshorter longevity of adults, and production of fewer orno nymphs (Teetes et al., 1995).

Morpho-physiological traits: In sorghum, some of themorpho-physiological traits such as genotypes withsmall, narrow, and fewer leaves, and low leafbending at the seedling stage (Mote and Kadam,1984), greater plant height and greater distance betweentwo leaves and the presence of waxy lamina (Mote andShahane, 1994), and epicuticular wax on the ventralsurface of the leaves were associated with reducedsusceptibility to the sugarcane aphid (Pi and Hsieh,1982a, b).

Biochemical factors: Host suitability to variousphloem-feeding Homoptera has frequently been relatedto nitrogen levels in host plants. Hsieh (1988) stated thatthe presence of p-hydroxybenzaldehyde during HCNrelease from the sorghum leaves due to aphid biting, atthe seedling stage, may be important to repel furtherattack. The mean HCN content of F1 hybrids producedfrom crosses between high and low HCN contentgenotypes usually had correlations intermediate betweenthose of the parents or lines were closer to the parentwith a high HCN content. The development of aphidpopulations and leaf sugary exudation was morepronounced in sorghum genotypes having higher nitro-

gen, sugar, and chlorophyll content of leaves (Mote andJadhav, 1993; Mote and Shahane, 1994). The varietiesICSV 9, BTP 28, IS 1640, ICSV 148, and Swati (SPV504) with higher contents of phosphorus, potassium,and polyphenols were less preferred by the sugarcaneaphids and also showed less development of leaf sugaryexudation (Mote and Shahane, 1994).There is a significant reduction in nitrogen, phos-

phorus, potash, total sugars, and chlorophyll content insorghum due to infestation by the sugarcane aphid. Incontrast, there is a significant reduction in polyphenolsin resistant over the susceptible sorghum genotypes(Balikai, 2001). In India, planting in the third week ofSeptember caused a reduction in protein, total minerals,and fat content in the grain due to infestation by thesugarcane aphid. Similarly, there is also a decline indiastase activity in sorghum grain, while there is anincrease in crude fiber and carbohydrate content wheninfested by M. sacchari (van den Berg et al., 2003). Insorghum fodder, there was a loss of 10% and 7.0% incrude protein and crude fiber content, respectively(Balikai, 2001).

Genetic basis of resistance: Greenhouse and fieldstudies with the crosses between PI 257595 (highlyresistant), 129-3A (moderately resistant), and RTx 430(susceptible) have shown that resistance is monogenicand controlled by a single dominant gene (Hsieh and Pi,1982; Pi and Hsieh, 1982b; Tan et al., 1985). Studieshave also indicated that PI 257595 and 129-3A have thesame gene for resistance, although the resistance gene of129-3A has modifiers (Pi and Hsieh, 1982a). Althoughdominant and additive gene actions are involved,additive gene action accounts for the resistance expres-sion (Hsieh and Pi, 1988). The cross between RTx430� 129-3A indicated the presence of complimentarygene action (Chang and Fang, 1984).

ARTICLE IN PRESS

Table 4

Parental lines (A/B- and R-lines) of sorghum resistant to the sugarcane aphid, M. sacchari (Zehnt.) reported from different countries

Parental line Resistance Country from which reported Reference

Level Mechanism

A/B-line

2B R India Balikai (2001)

9B R India Balikai (2001)

27A R India AICSIP (1998)

33B R India Patil (1992)

42B R India AICSIP (1990, 1998), Patil (1992), and

Balikai (1993)

53B R India Patil (1992)

104A/B R India AICSIP (1998) and Balikai (2001)

116A/B R India AICSIP (1991, 1998) and Patil (1992)

117B R India Patil (1992) and AICSIP (1996)

129-3A R Taiwan, Pi and Hsieh (1982a)


205B R India Patil (1992) and AICSIP (1996)

296B R India Kishore (2000)

2077A/B R India Balikai (2001)

2219A/B R India Patil (1992) and Balikai (1993)

4692A MR Japan Hagio and Ono (1986)

A 36227B R India AICSIP (1991)

AB 31 MR China Chang (1981a, b)

AKMS 14A R India AICSIP (1997)

ICSB 53 R India AICSIP (1991)

ICSB 70 R India Patil (1992)




ICSB 2730 HR Ax, Ab India AICSIP (2003)

ICSB 2731 HR Ax, Ab India AICSIP (2003)



R-line

2R R China, Chang (1981a, b)


5R R China Chang (1981a, b)

47R MR China Chang (1981a, b)

57R MR China Chang (1981a, b)

C 43 HR Ax, Ab India AICSIP (2003)

C 81 R India Balikai (1993)

ICSR 160 R India AICSIP (1991)

ICSR 161 R India Patil (1992)


ICSR 165 R India AICSIP (1991) and Patil (1992)




ICSR 38111 HR Ax, Ab India AICSIP (2003)



M 148-138 R India AICSIP (1990)

R 128 R China Chang and Fang (1984)



R 1413 R India Balikai (1993)

R 354 R India Kishore (2000)

RS 29 R Ax, Ab India Patil (1992), Balikai (1993),

Kishore (2000), and AICSIP (2003)

RS 67 R India Patil (1992), Balikai (1993),

Kishore (2000), and AICSIP (2003)

RS 291 R India AICSIP (1997)



RTAM 428 HR Ax, Ab China, Xu (1982)


SB 1085 R India AICSIP (1997)



ARTICLE IN PRESS

Table 5

Agronomic elite lines/experimental varieties of sorghum resistant to the sugarcane aphid, M. sacchari (Zehnt.) reported from different countries

Agronomic elite line/

experimental variety

Resistance Country from

which reported

Reference

Level Mechanism

Agr 4S2560 Shallu HR Taiwan Pi and Hsieh ( 1982a)

7511 HR Ax, Ab China Wang et al. (1991)

BPT 28 R India Mote and Shahane (1988, 1993)

CE 151 R South Africa Peterson (2002)

E 108 R India AICSIP (1982)



Ent. 62SADC R South Africa Peterson (2002)

FGYQ 336 R South Africa Peterson (2002)

FGYQ 353 R South Africa Peterson (2002)

Hami R India Mote et al. (1985)

HB 37 R China Chang (1981a, b)

ICSV 9 R India Mote and Shahane (1988, 1993)

ICSV 148 R India AICSIP (1988) and Mote and Shahane (1988, 1993)

ICSV 197 R India Sharma (1993)



ICSV 89013 R India AICSIP (1991)










NK 266 MR Japan Hagio and Ono (1986)

NR 349 R India AICSIP (1990)

PE 954 177 HR Ax, Ab Japan Hagio et al. (1985), Hagio and Ono (1986), and

Hagio (1992)

PNR 8537 HR Ax Botswana, Teetes et al. (1995)

Zimbabwe Teetes et al. (1995)

PVR 10 R India Patil (1992)

SA 967 R South Africa Wenzell et al. (1998)

SDSL 89426 R South Africa Peterson (2002)

SPV 97 R India AICSIP (1980)



SPV 224 R India AICSIP (1980, 1982)










SPV 422 MR India Mote and Kadam, 1984)


SPV 504 R India Mote and Shahane (1988, 1993)

SPV 625 MR India Mote and Kadam, 1984)

SPV 969 R India Patil (1992)








ARTICLE IN PRESS

Table 5 (continued)

Agronomic elite line/

experimental variety

Resistance Country from

which reported

Reference

Level Mechanism


SPV 1155 R India Kishore (2000)

SPV 1178 R India Balikai (1993)











8. Virus vector

The sugarcane aphid is a vector and transmits threepersistent viruses namely, millet red leaf virus (Black-man and Eastop, 1984), and sugarcane yellow leaf viruson sorghum and sugarcane (Schenck, 2000), andsugarcane mosaic virus (SMV) on sorghum (Bhargavaet al., 1971; Kondaiah and Nayudu, 1984; Setokuchiand Muta, 1993). Yang (1986) demonstrated that boththe sugarcane aphid and corn leaf aphid (Rhopalosiphum

maidis) were efficient transmitters of SMV betweensorghum, corn, and sugarcane. Latent periods of 30 and20 days were observed for SMV on corn and sweetsorghum, respectively. However, the disease symptomscolonized by M. sacchari takes a longer time to expresson sweet sorghum than on corn or sugarcane (Yang,1986).

9. Cultural practices

Among the cultural practices, early planting results inthe crop escaping from aphid attack (van Rensburg,1974) and high plant density promotes low plant vigorand concomitantly reduces aphid abundance (Flattery,1982; van Rensburg, 1979; Setokuchi, 1975). Cutting offorage sorghum before the first week of aphid abun-dance prevents not only the damage but also regulatessubsequent increase in aphid density on a ratoon crop(Setokuchi, 1977). Since the sugarcane aphid over-winters on ratoon sorghum, S. verticilliflorum, S.

halepense, P. maximum, and Setaria spp., their destruc-tion before the sorghum crop is planted reduces thecarryover of the pest. Mulching with rice/wheat straw isanother practice effective in reducing colonization byaphids. Sanchez and Cermeli (1987) used yellow trapswith water to capture the migrant aphids in the fields ofsorghum to forecast their migratory pattern andpopulation dynamics in Venezuela.

10. Natural enemies

Over 47 species of natural enemies attack M. sacchari

worldwide. Although unable to prevent the buildup ofdamaging numbers of the sugarcane aphid even in theseason, natural enemies play a very important role (vanRensburg and van Hamburg, 1975), and often maintainthe sugarcane aphid populations below the economicthreshold levels (ETLs) in sorghum (van Rensburg,1973b; Anonymous, 1978; Chang, 1981a, b; Meksongseeand Chawanapong, 1985) (Table 7).Zimmerman (1948) reported Aphelinus maidis para-

sitizing the sugarcane aphid in Hawaii. Similarly, theparasitoids Enrischia comperei Ashm. in Australia (Gil-strap, 1980), Exochonus concavus (Fursch), Leucopus

spp., and Lioadalia flavomaculata (DeGeer) in SouthAfrica (van Rensburg, 1973b) have been recorded. Theaphid parasite, Lysiphlebus testaceipes (Cresson) (Hy-menoptera: Braconidae) attacks the sugarcane aphid inHawaii (Zimmerman, 1948) and indigenous L. dehliensis

Zehntner is credited with biological control on sugar-cane in India (Varma et al., 1978). Low levels ofparasitism by unidentified parasitoids have been ob-served in sorghum in South Africa, but generally areconsidered less important in suppressing the aphidpopulation abundance (Anonymous, 1978; van Re-nsburg, 1973b).Emphasis has been placed on predators, primarily

ladybeetles (Coleoptera: Coccinellidae), lacewings (Neu-roptera: Chrysomelidae and Hemerobiidae), and hover-flies (Diptera: Syrphidae) as species causing greatestmortality to the sugarcane aphid populations. Numer-ous predators suppressed sugarcane aphid abundanceon sorghum in South Africa (Anonymous, 1978; vanRensburg, 1973a, b), Tanzania (Bohlen, 1973), andSudan (Schmutterer, 1969). In southern Africa, impor-tant predator species are ladybeetles, Cheilomenes

propinqua (Muls.) var. quadrilineata (Muls.), C. lunata

(Fabr.), C. sulphurea (Fabr.), L. flavomaculata (De Geer),

ARTICLE IN PRESS

Table 6

Hybrids, varieties, and locals of sorghum resistant to the sugarcane aphid, M. sacchari (Zehnt.) reported from different countries

Hybrid/variety/local Resistance Country from

which reported

Reference

Level Mechanism

Hybrid

272 x 3122 R South Africa Wenzell et al. (1998)




1470 derivative R South Africa Wenzell et al. (1998)


(6BRON161/(7E0366�Tx2783)-HG54)�CE151)-CG-3-BGBK


(CE1518BDM499)-LD17-BD1 R South Africa Peterson (2002)

(Macia�TAM 428)-1-1-2 R South Africa Peterson (2002)

(MR112B-92M2�Tx 2880)-SM3-

SM1-ML 52


7511 x Kang 60-P341 R South Africa Wang et al. (1991)

7511 x Kang60-P400 R South Africa Wang et al. (1991)

CSH 13K&R HR Ax, Ab India Balikai (1993)

SPH 634 R India AICSIP (1996)

Variety

CSV 8R R India Balikai (1993, 2001)

Sel 3 R India Kishore (2000)

Local

Boushi Kaoliang R Japan Hagio (1987)

Dabor HR Ax, Ab China Xu (1982)

Dagadi R India Mote et al. (1985)

Dairyukoku R Japan Hagio (1967)

DJ 6514 R India AICSIP (1982, 1987)

Gyushinpaku R Japan Hagio (1987)

Hakubai Kaoliang R Japan Hagio (1987)

Jogri R India Mote et al. (1985)

Koumairou HR Ax, Ab Japan Hagio (1992)

Kuchkuchi R India Mote et al. (1985)

Lakadi R India Mote et al. (1985)

Ludende HR Ax, Ab Botswana, Teetes et al. (1995)

Zimbabwe Teetes et al. (1995)

Minasel R Japan Hagio (1987)

SA 1429 R South Africa Manthe et al. (1992)

SC 110-14 R Taiwan Chang (1981a, b)

PJ 4R R India Mote et al. (1985)

Senkinshiro HR Japan Setokuchi (1976), Hagio et al. (1985),

Hagio and Ono (1986), and Hagio (1992)

Setokho-2-gou MR Japan Hagio and Ono (1986)

Shoukoubai R Japan Hagio (1987)

Shoumai Kaoliang HR Ax, Ab Japan Hagio (1992)

Suzuho HR Ax, Ab Japan Hagio et al. (1985), Hagio and Ono (1986), and

Hagio (1992)



E. concavus (Furch), Chilocorus nigritus (F.), andScymnus morelleti (Muls.) (Coleoptera: Coccinellidae)in South Africa (van Rensburg, 1973b), S. babar Sassjiin China (He et al., 1987), and Diomus terminatus (Say)in the USA (Hall, 1987). He et al. (1987) also reportedthat adults of S. babar overwintered in the leaf sheathsof Juncellus serotinus, Acorus calamus L., Typha

latifolia, and Boiboschnoenus mantinus. They become

active in mid-April and migrated to the sorghum fieldsin mid-June to predate upon M. sacchari and then intomaize fields in early August to feed on R. maidis and R.

padi, and complete one and two generations on sorghumand maize, respectively.Among the dipteran predators, syrphids also play an

important role on the sugarcane aphid viz., Allograpta

exotica (Wiedemann, 1830) in the USA (Florida)

ARTICLE IN PRESS

Table 7

Natural enemies of the sugarcane aphid, M. sacchari (Zehnt.) reported in different countries

Natural enemy Order Family Country Reference

Pathogen

Verticillium lecanii (Zimm.) Viegas Hypocreales Hypocreaceae USA (Florida) Hall (1987)

Parasitoid

Aphelinus maidis Timberlake Hymenoptera Aphelinidae USA (Hawaii) Zimmerman (1948) and Gilstrap (1980)

Enrischia comperei ashm. Hymenoptera Elasmidae Australia Gilstrap (1980)

Bracon sp. Hymenoptera Braconidae India Hussain (1991)

Lioadalia flavomaculata (DeGeer) Hymenoptera Aphelinidae South Africa van Rensburg (1973)

Lysiphlebus dehliensis Zehntner Hymenoptera Braconidae India Varma et al. (1978)

Lysiphlebus testaceipes (Cresson) Hymenoptera Braconidae USA (Florida), Hall (1987)

USA (Hawaii) Zimmerman (1948)

Predator

Allograpta exotica (Wiedemann, 1830) Diptera Syrphidae USA (Florida) Hall (1987)

Brumus suturalis (Fabr.) Coleoptera Coccinellidae India Seshu Reddy and Davies (1979)

(Syn: Brumoides suturalis (Fabr.))

Ceratomegilla (Megilla) inonata Muls. Coleoptera Coccinellidae Puerto Rico Gilstrap (1980)

Chilochorus nigritus (F.) Coleoptera Coccinellidae India, Gilstrap (1980)

South Africa van Rensburg (1973)

Chrysoperla sp. Neuroptera Chrysomelidae India Seshu Reddy and Davies (1979)

Chrysoperla basalis Walk. Neuroptera Chrysomelidae Thailand Meksongsee and Chawanapong (1985)

Chrysoperla collaris Schneider Neuroptera Chrysomelidae Puerto Rico Gilstrap (1980)

Chrysoperla externa (Hagan) Neuroptera Chrysomelidae USA (Florida) Hall (1988)

Coelophora inaequalis (Fabricius) Coleoptera Coccinellidae Australia, Gilstrap (1980)

USA (Hawaii) Gilstrap (1980)

Coleomegilla maculata Deg. Coleoptera Coccinellidae Dutch Guiana Gilstrap (1980)

Coleomegilla maculata fuscilabris

(Mulsant)

Coleoptera Coccinellidae USA (Florida) Hall (1988)

Cycloneda sanguinea (L.) Coleoptera Coccinellidae Dutch Guiana, Gilstrap (1980)

Puerto Rico, Gilstrap (1980)

USA (Florida) Hall (1988)

Diomus terminatus Say Coleoptera Coccinellidae USA (Florida) Hall (1987)

Exochomus concavus (Furch) Coleoptera Coccinellidae South Africa van Rensburg (1973)

Geocoris sp. Hemiptera Lygaeidae India Seshu Reddy and Davies (1979)

Hippodamia convergens Guerin Coleoptera Coccinellidae USA (Florida) Hall (1988)

Illeis indica Timberlake Coleoptera Coccinellidae India Seshu Reddy and Davies (1979)

Leucopus spp. Diptera Chamaemyiidae South Africa van Rensburg (1973b)

Menochilus (Cheilomenes) lunata

(Fabr.)

Coleoptera Coccinellidae South Africa van Rensburg (1973b)

Menochilus (Cheilomenes) propinqua

(Muls.) var quadrilineata

Coleoptera Coccinellidae South Africa van Rensburg (1973b)

Menochilus (Cheilomenes) sexmaculatus

(Schall.)

Coleoptera Coccinellidae India, Seshu Reddy and Davies (1979) and

Gilstrap (1980)

Thailand Meksongsee and Chawanapong (1985)

Menochilus (Cheilomenes) sulphurea

(Fabr.)

Coleoptera Coccinellidae South Africa van Rensburg (1976)

Micraspis discolor (Fabr.) Coleoptera Coccinellidae Thailand Meksongsee and Chawanapong (1985)

Micromus subanticus (Walker) Neuroptera Hemerobiidae USA (Florida) Hall (1988)

Nacoleia (Omiodes) accepta Btlr. Diptera Cecidomyiidae USA (Hawaii) Gilstrap (1980)

Ola v-nigrum Mulsant Coleoptera Coccinellidae USA (Florida) Hall (1988)

Phaenobremiameridionalis Felt. Diptera Cecidomyiidae USA (Hawaii) Gilstrap (1980)

Prospaltella transvena Timb. Hymenoptera Aphelinidae USA (Hawaii) Zimmerman (1948) and Gilstrap (1980)

Scoloposcelis parallelus Motsch. Hemiptera Anthocoridae Puerto Rico Gilstrap (1980)

Scymnodes (Platynomus) lividigaster

(Mulsant)

Coleoptera Coccinellidae Australia Gilstrap (1980)

Scymnus babar Sassji Coleoptera Coccinellidae China He et al. (1987)

Scymnus loewi Muls. Coleoptera Coccinellidae Puerto Rico Gilstrap (1980)

Scymnus morelletti (Muls.) Coleoptera Coccinellidae South Africa van Rensburg (1976)

Scymnus nubilus Mulsant Coleoptera Coccinellidae India Gilstrap (1980)

Scymnus roseicollis Muls. Coleoptera Coccinellidae Puerto Rico Gilstrap (1980)

Syrphus balteatus (DeGeer) Diptera Syrphidae Thailand Meksongsee and Chawanapong (1985)

Xanthogramma aegyptium (Wied.) Diptera Syrphidae USA (Louisiana) White et al. (2001)

South Africa van Rensburg (1976)

Xanthogramma scutellaris Diptera Syrphidae India Seshu Reddy and Davies (1979)



(Hall, 1987), Syrphus balteatus (DeGeer) in Thailand(Meksongsee and Chawanapong, 1985), Xanthogramma

aegyptium in the USA (Louisiana) (White, 1987), SouthAfrica (van Rensburg, 1973a, b), and X. scutellarius inIndia (Seshu Reddy and Davies, 1979); and Leucopus

spp. (Chamaemyiidae) in South Africa (van Rensburg,1973b). Hall (1988) reported Micromus subanticus

(Walker) (Neuroptera: Hemerobiidae) as feeding onthe sugarcane aphid in the USA (Florida). Otherchrysopids have also been recorded such as Chrysoperla

basalis Walker, C. collaris Schneider, and C. externa

(Hagan) in Thailand (Meksongsee and Chawanapong,1985), Puerto Rico (Gilstrap, 1980) and the USA(Florida) (Hall, 1988).Hall (1987) found that the entomogenous fungus,

Verticillium lecanii (Zimm.) Viegas is an importantbiological control agent in the USA (Florida). However,ants are notorious in interfering with the beneficialactivities of aphid predators and/or parasites and thesugarcane aphid may sometimes benefit from a symbio-tic association with certain ant species.

11. Chemical control

Soil application of systemic insecticides such asdisulfoton, disyston, and phorate (van Rensburg andvan Hamburg, 1976; Mote, 1977; Denmark, 1988) wereeffective, with phorate superior to disulfoton (Mote,1977). Foliar sprays of demeton-S-methyl, dimethoate,endosulfan, and parathion (van Rensburg, 1979; Mote,1985); diazinon, malathion, metasystox, and phosdrin(Denmark, 1988); quinalphos (Chaudhari et al., 1994);and carbofenthion and carbofuran were effective whenthe populations reach 70 and 155 aphids plant�1 at 50and 80 days after planting in spring and 60 and 90 daysafter planting in autumn, respectively (Fang, 1990).Spraying extracts of leaves and kernels of Vinca rosea,Pongamia pinnata, Azadirachta indica, and Vitex negun-

do took 10 days to cause mortality of aphids:Dimethoate, chlorpyriphos, endosulfan, alfamectin,and malathion not only reduced aphid populations butpromoted grain and fodder yields, besides increase inkernel weight (Balikai, 2001). Currently, the onlyeffective method to control the M. sacchari populationsis the use of insecticides, but thus use is not consideredeconomically feasible for small-scale farmers in devel-oping countries (van Rensburg and van Hamburg, 1975;Young and Teetes, 1977; Page et al., 1985). This isbecause, in the absence of natural enemies, aphidinfestations may resurge, so a second or even a thirdinsecticide application may be required for control (vanRensburg, 1978, 1979). Systemic insecticides are effec-tive against the sugarcane aphid, but do not kill thebeneficial arthropods (van Rensburg et al., 1978; vanRensburg, 1980). However, application of demeton-S-

methyl, parathion, and monocrotophos resulted inmortality of coccinellids and syrphids while endosulfanand pirimicarb were less toxic to predators (vanRensburg and van Hamburg, 1975).

12. Future research

Several factors affect the plant responses to M.

sacchari injury such as the time of infestation (seasonand plant growth stage), duration of the infestation,environmental stresses (especially drought) and nutri-tional status of the host plant, which cumulatively affectthe yield loss. The sugarcane aphid feeding at the basalleaves had a strong influence on mobility of mineralnutrients, amino compounds, and carbohydrates in thephloem. By feeding at these sites, M. sacchari alters thecarbohydrate-partitioning patterns of sorghum, suggest-ing that infestation might alter sink-source relationshipswithin the plant. Very few studies have investigated theimpact of environmental factors on M. sacchari damagesymptom development, which mostly depends on theinitial aphid density colonized as well as the inherentplant resistance. Thus, the impact of abiotic factors suchas temperature and photoperiod on the life history traitsof M. sacchari needs to be investigated. Further, thecontribution of weeds and the native vegetation to thesugarcane aphid population dynamics is not clearlyknown, although considered as a major factor influen-cing sorghum infestation. Diversity of natural enemiesmay vary within and among cultivar communitiesdepending on a variety of factors including microcli-mate, prey density, annual changes in host plant speciesabundance, the timing and rate of migration anddispersal, and geographic location. The epidemiologyof transmission of three persistent viruses transmitted bythe sugarcane aphid has not been substantially investi-gated. Emphasis should be given on the populationcolonization at the seedling stage of sorghum becausethey may have greater impact affecting the growth anddevelopment of the crop.The impact of natural enemies on sugarcane aphid

populations in sorghum is not well understood. Quanti-fication and predicting the impact of natural enemies, asa group or as individuals, has been difficult because ofthe many species involved. The natural enemies have agreater effect early in the season, since the increase pergeneration of the sugarcane aphid population early inthe season is slow compared with late season increases.Comprehensive information needs to be generated onthe population dynamics of M. sacchari and the factorsthat lead to rapid population buildup. Environmentalfactors are largely responsible for the initial decline inaphid levels following a summer outbreak (Hall, 1987).Quantification of development rates of immatures, andpotential population growth rates of the sugarcane


aphid under different temperature regimes similar tothose occurring in the post-rainy season sorghum nichein India would greatly aid in forecasting models.Drought and sugarcane aphid infestation are twofactors that individually cause yield reduction insorghum, but when they occur in succession results ina synergistic interaction between aphid infestation andpredisposing the crop to drought stress. Field observa-tions have indicated that aphid damage is greater whendrought also occurs as a stress factor. The magnitude ofplant damage to sorghum in relation to the density ofM. sacchari as well as the stage of plant growth is notknown. It may be that sorghum can recover andcompensate for damage due to M. sacchari as has beenobserved by the feeding injury caused by S. flava insorghum. Insect density relationships with yield loss arenot available. It is not known, whether the sugarcaneaphid injects a phytotoxin during feeding that degradeschloroplasts or the damage results from the plant’sresponse to mechanical injury. As little is known abouthow aphids elicit these changes and the physiologicalmechanisms associated with injury, improving ourunderstanding of how M. sacchari affects the sorghumplant physiology will be important in identifying newtargets for aphid resistance in sorghum.Cultural practices such as elimination of volunteer

sorghum plants from the previous crop and thedestruction of susceptible weedy hosts needs greateremphasis. Search for new sources of resistance isessential to diversify the genetic background. Identifica-tion of resistant sources in elite background as well as inparental lines in the development of hybrids, andknowledge on the mechanisms would greatly assist indeveloping cultivars with stable sources of resistance tothe sugarcane aphid, under high yield background.Combinations of different categories have effects thatmay be more beneficial and compatible in trophicinteractions than the effect of individual componentsof resistance.

Acknowledgements

We are greatly indebted to Dr. Mangala Rai, DirectorGeneral, Indian Council of Agricultural Research, NewDelhi for his constant encouragement and keen interest.

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