Chapter I INTRODUCTION Maize, Zea mays (L.

) (Poaceae), ranking third in area after wheat and rice is one of the important cereal crops in the world. In India, maize was cultivated over an area of 8.55 million hectares, with a production and productivity of 21.73 million tones and 25.40 q ha-1, respectively in 201011 (Anonymous 2011). The maize production in India has increased more than 12 times from a mere 1.73 million tonnes in 195051 to 21.73 million tonnes at present. However, during 201011 it was grown in 133 thousand hectares with a production of 491 thousand tonnes in the Punjab (Anonymous 2012). The demand for maize will touch 42 million tonnes by 2025, of which 2021 per cent will be used for human consumption, more than 60 per cent as poultry and livestock feed and remaining 12 13 per cent for industrial raw material (Anonymous 2011). The generation of new agricultural technology in India during last 50 years has not only helped to increase production of maize but also has given rise to new cropping patterns. This has led to round the year cropping of maize in one or the other region of Indian union (Panwar 1998) and offers a bright prospect in crop diversification in the Punjab, where its cultivation is recommended in kharif, rabi and recently in spring season too (Anonymous 2006). The cultivation of spring maize is becoming popular because of its higher yield potential. The productivity level of maize in the Punjab after the recommendation of cultivation of spring maize in the state, has gone up by 34.9 per cent in 2011 with productivity of 36.93 q ha-1 than that of 27.38 q ha-1 in 2005 (Anonymous 2006, 2012). The average productivity of maize in India (2 t/ha) was quite low in comparison to the major maize producers in Asia (Sharma et al 2006). The attack of insect pests at various crop growth stages poses serious limitations in full manifestation of yield potential of maize during different seasons. Although 139 insect pests are reported to cause varying degree of damage to maize crop from sowing to harvest, but only about a dozen are quite serious and require control measures (Siddiqui and Marwaha 1994). However, according to Mathur (1987), the total number of insect and mite pests exceeds 250 in India. The insect pest complex changes in time and space and their incidence in maize has increased due to its large scale cultivation as sole crop and widespread use of pesticides for pest control. The insects hitherto unknown to attack the maize have become problems (Kumar et al 2005, Kumar and Kanta, 2011). The continuous planting of maize has led to the invasion of polyphagous shoot flies species as the key pests on spring sown maize in the northern India (Sarup et al 1984). The incidence and extent of losses by insect pests complex to the newly recommended cultivation of maize during spring season in the Punjab is not available. Among the different species of shoot fly prevalent in north India, Atherigona naqvii Steyskal is a serious pest of spring sown maize in the Punjab (Sandhu and Kaushal 1976, Singh and Kanta, 2004). The

maggots of shoot fly attack the emerging seedlings and feed on the whorl leaves causing deadhearts in small and slowly growing plants and resulting in curled and distorted leaves in bigger plants (Panwar 2005, Kanta et al 2006). The severity of the pest can be judged from the fact that up to 85.8 per cent deadhearts incidence was reported from spring-sown maize in the Punjab (Sajjan and Sekhon 1985) and even the 15 days delay in adoption of control practices may result in yield losses to the tune of 32.4 per cent (Gagandeep and Kanta 2007a). As the adult flies oviposit on the emerging seedlings and in cracks and crevices around the seedlings, the soil application of carbofuran 3 G @ 12.5 kg and phorate 10 G @ 10.0 kg per hectare at sowing time (Sajjan and Sekhon 1985, Kanta et al 2006) and seed treatment with imidacloprid 70 WS @ 5 g per kg seed and imidacloprid 600 FS @ 6 ml per kg seed one day before sowing was found effective and recommended for control of shoot fly in Punjab (Jindal and Hari 2008, 2011). However, the use of chemical pesticides is hazardous to the beneficial organisms and the environment; and the management strategy for any pest must involve more than one control tactics. Moreover, shoot fly, in general, is not easily accessible to insecticides sprayed on maize as the larvae feed inside the leaf whorls. The exploitation of host plant resistant also has a good potential for the management of shoot fly. The use of resistant cultivars is a realistic alternative to chemical control, if they are able to compete economically with the commonly used hybrids and varieties (Raina, 1985). In India, up till now, about 2000 maize germplasm lines have been screened against Atherigona species and number of resistant sources has been identified by various workers from different locations, but the levels of resistance are low to moderate (Siddiqui et al 1988, Panwar 2005, Jindal et al 2007, Kumar and Kanta 2011). Resistance to shoot fly in sorghum is expressed in terms of oviposition non-preference, antibiosis and tolerance (Dhillon et al 2005a). However, the factors influencing these mechanisms have not been identified in the promising maize cultivars for spring season in the Punjab. To develop crop cultivars with durable resistance to insect pests, it is important to identify germplasms with diverse combinations of factors associated with resistance to the target pests and then to combine identified

components/mechanisms of resistance in the same genetic background. Some of the factors associated with resistance to insects can be quantified or monitored easily in plant populations and such characters can be used as "marker traits" to screen and select for resistance to insect pests (Chamarthi 2008). There is a need to develop integrated pest management (IPM) strategies involving cultural methods of management of shoot fly spp. in spring season. But it requires a thorough knowledge of the life history, behaviour and ecology of the insect as well as of its plant or animal hosts. The most vulnerable stage (s) in the life cycle must be determined to develop practices to lessen its attack, to kill it, or to slow down its rate of reproduction (Lawani 1982). The principle of controlling insect pest populations by increasing the diversity (Solomon 1973) can successfully be employed by using sorghum

as a trap crop, which had been reported as a preferred host for egg lying by shoot fly (Ogwaro 1978, Sarup et al 1986). The low humidity increased the duration of egg development and decreased the egg survival; and continuous rainfall reduced shoot fly abundance (Raina 1981, Delobel 1983, Taneja et al 1986). These facts can be exploited by standardization of irrigation frequency and evaluation of sorghum as trap crop for management of shoot fly during spring season in the Punjab. Keeping in view the above facts, the present studies on incidence of insect pests and management of shoot fly, Atherigona spp. in spring sown maize were undertaken with the following objectives: Objectives:

a)

To study the populations build up of insect pests and their extent of losses in spring maize.

b) To establish the morphological and biochemical bases of resistance against shoot fly in different
promising cultivars.

c)

To formulate integrated approaches for management of shoot fly in spring sown maize.

Chapter-II REVIEW OF LITERATURE The literature of previous years recorded 160 insect and mite species on maize crop (Pant and Kalode 1964), but afterwards Mathur (1987) observed over 250 species of pests associated with maize in field and storage conditions. Siddiqui and Marwaha (1994) reported 139 insect pests causing varying degree of damage to maize crop from sowing to harvest. They further reported that only about a dozen of these are quite serious and require control measures. Insect pest complex changes in time and space. The crambid, Chilo partellus (Swinhoe) and muscids, Atherigona spp. are of major production constraint in maize cultivation in north India during different seasons (Kumar et al 2005). The present studies were undertaken on various aspects of shoot fly, Atherigona naqvii Steyskal to formulate appropriate strategies for its effective management and to estimate loses due to insect pests in spring sown maize in the Punjab. The relevant literature pertaining to different aspects of shoot fly, Atherigona spp. management and other insect pests has been reviewed under the following headings: 2.1 2.2 2.3 Prevalence of shoot flies, Atherigona spp. Geographic distribution and host range of shoot fly, A. naqvii Bioecology of shoot fly, A. naqvii 2.3.1 2.3.2 2.3.3 2.4 Nature of Damage of Atherigona spp. Ovipositional preference of Atherigona spp. under field conditions Effect of sowing time on the incidence of Atherigona spp.

Host plant resistance for the management of shoot flies, Atherigona spp. 2.4.1 2.4.2 Evaluation of germplasm against Atherigona spp. Mechanisms of host plant resistance to Atherigona spp. 2.4.2.1 2.4.2.2 2.4.2.3 Antixenosis for oviposition Antibiosis Tolerance

2.4.3 Bases of resistance to Atherigona spp. 2.4.3.1 2.4.3.2 2.5 Morphological plant traits Biochemical plant traits

Management of shoot fly, Atherigona spp. 2.5.1 2.5.2 Management of Atherigona spp. with insecticide Management of Atherigona spp. with non-chemical methods

2.6.

Other insect pests of spring maize

2.1

Prevalence of shoot flies, Atherigona spp. Shoot fly was first noticed infesting maize at Pantnagar in spring season during 1967 (Rathore et

al 1969) and the reared flies were identified as Atherigona orientalis Schiner. Atherigona naqvii Steyskal was observed attacking maize crop in the Punjab in 1972 (Sandhu and Kaushal 1976) with report of more than one type of eggs on maize plant. Sixteen shoot fly species namely, Atherigona bidens Henning, A. falcata Thomson, A. naqvii, A. orientalis, A. punctata Karl, A. soccata Rondani, A. oryzae Malloch, A. vorkif Deeming, Anacamptoneurum obliquum Becker, Anatricus erinaceus Loew, Aprometopis flavofacies Becker, Delia arambourgi (Seguy), Elachiptera scapularis (Adams), Lasiosina sp., Scoliophthalmus micantipennis Duda and S. trapezoids Becker have so far been recorded on maize in Africa and Asia (Panwar and Sarup 1985) including six species A. bidens, A. falcate, A. naqvii, A. orientalis, A. punctata and A. soccata are from India. A. orientalis occurred in Pantnagar (Rathore et al 1969). A. soccata is a serious pest of sorghum in kharif (monsoon) season (Jotwani et al 1970) whereas, A. naqvii attacks barley (Srivastava et al 1969) and wheat (Kundu and Kishore 1971). A. naqvii and A. soccata are serious pests in northern India (Sandhu and Kaushal 1976, Sarup et al 1984). A. soccata, A. bidens, A. falcata and A. punctata occur occasionally in the southern region (Seshu Reddy and Davies 1977, Davies and Seshu Reddy 1981). However, A. naqvii, invading spring sown maize, is of a serious concern for the farmers in the Punjab. 2.2 Geographic distribution and host range of shoot fly, A. naqvii A. naqvii has been reported from India (Delhi, Kerala, Uttar Pradesh, Rajasthan and Punjab), Ceylon, Australia, Nigeria, Pakistan, Ethiopia, Saudi Arabia and South Africa (Panwar 2005). Its host plants include Triticum aestivum Linn. ( Ramachandra Rao 1924, Srivastava and Pandey 1968, Pont 1972); Hordeum vulgare Linn. (Srivastava et al 1969); Arundinella metzii Hochst ex. Miq., Digitaria sanquinalis (Linn.) Scop and Pennisetum typhoides (Burm.) (Pont 1972); Zea mays Linn. (Deeming 1971, Pont 1972, Sandhu and Kaushal 1976) and Avena ludoviciana Linn. (Singh and Khan 1980). 2.3 Bioecology of shoot fly, A. naqvii A complex of Atherigona spp. cause severe losses in spring sown maize in northern India. Two major shoot fly species viz., A. soccata and A. naqvii produce similar type of dead hearts. However, the difficulty in identification of shoot fly species on the basis of type of damage can be overcome by its site of oviposition and distinguishing characters of eggs of both the species. The eggs of A. soccata are laid singly on the underside of sorghum leaf almost parallel to mid rib, whereas that of A. naqvii are laid either in soil in cracks or crevices (around maize seedlings) or at the base of stem/under surface of lower leaves of other cereals (Kishore 1991). In case of A. soccata, the egg is elongated with hexagonal sculpture

giving an appearance of reticulation whereas that of A. naqvii is cylindrical in shape and marked by reverse dichotomous lines running somewhat parallel (longitudinally) from one end to the other. The egg distribution by both A. naqvii and A. soccata is random or slightly aggregated rather than regular (Delobel 1981 and Hari et al 2013). The ovipositional behaviour of shoot fly species did not have any effect on its damage on varieties having different plant population levels in spring sown maize (Rao and Panwar 1994). The incubation period of A. naqvii eggs is 1 to 3 days. The freshly hatched larva is dirty white and full grown larva is yellow in colour. On hatching, the maggot feeds on leaf surface in funnel primarily as whorl maggot and attacks the growing point. The maggot is full grown in 15 -18 days. The pupal period lasts for 8-9 days. The adult longevity was 1-2 weeks when reared on yeast glucose dry food (Sandhu and Kaushal 1976).The total life cycle of A. naqvii on maize was completed in 17-33 days (Kumar and Chander 1987). 2.3.1 Nature of Damage of Atherigona spp. The Atherigona spp. causes damage at seedling stage of the crop. The newly emerged maggots enter the leaf whorl by crawling along the leaf sheath, move downwards by feeding on plant tissues which results in the browning of the central axes and gradual drying up of the growing points form the deadhearts in the young plants. However, the older plants do not show dead hearts but the damaged leaves of such plants get interwoven with central leaf and show curling and distortion. Irregular slits and deformed leaves are formed as a result of attack (Sarup et al 1984, Kanta et al 2006). Raina (1981) found that very little damage is done to the growing point by the first instar and the deadheart is caused by cutting the base of the central shoot. The first and the second instars are mainly involved in cutting of leaf tissues, whereas the third instar feeds on dead and decaying tissues. Sajjan and Sekhon (1985) reported that the shoot fly complex produced upto 85.8 per cent dead hearts in spring sown maize in Punjab. Gagandeep and Kanta (2007a) reported that under natural conditions the protected maize crop, in spring season of 2004, yielded 11.78 q per ha, with subsequent reduction of yield by 16.46, 22.41, 29.79, 34.88, 40.66, 44.22 per cent in the treatments plots with 5, 10, 20, 30, 40 and 50 per cent deadhearts incidence, respectively. They further reported that a delay of 15 days in control practices resulted into yield losses to the tune of 32.4 per cent.

2.3.2

Preferential plant age and site for oviposition by Atherigona spp. in the field The egg laying by A. soccata is random in field (Delobel 1981). The observations on preferential

plant age and site for oviposition by A. soccata and A. naqvii under field conditions at different locations for two years made by Sarup and Panwar in 1987 on spring maize revealed that in both the years, regardless of the site of oviposition, the maximum number of eggs were laid on 9 to 11 days old plants. Significantly more number of eggs were laid on/in the soil around each plant (in cracks and crevices) than those laid on the foliage, established the dominance of A. naqvii over the A. soccata. However, no significant difference either in the eggs laid or in the deadheart formation due to shoot fly species was obtained at various locations in the field. 2.3.3 Effect of time of sowing on the incidence of Atherigona spp. in spring sown maize The most practical control method of Atherigona spp. is the choice of a sowing date that enables the crop plants to pass the susceptible stage at a period when its density is low (Jotwani et al 1970). The investigation on the effect of date of sowing on shoot fly incidence, not only pinpoints the period of its maximum activity, but also helps in adjusting the sowing date of the crop to escape the period of high incidence of the pest in the field. During 1970-71 at Pantnagar, the number of shoot fly eggs as well as deadhearts were recorded as maximum in the crop sown in the 3rd week of February than that in the 1st and 2nd week of February. Thus it could be deduced that in terai area of Uttar Pradesh, the early sown maize crop i.e., sown during 1st and 2nd week of February, escapes shoot fly incidence. The delay in sowing might attract shoot fly incidence with consequential losses in yield (Sarup et al 1979). The peak period of activity of A. soccata and A. naqvii was reported from 3rd week of February to 1st week of March at Indian Agricultural research Institute, New Delhi, on the basis of number of eggs laid and deadhearts produced by them (Marwaha et al 1984). Two maize cvs Deccan 105 and Kiran were sown on six sowing dates viz., 8th, 15th, 22nd, 29th March, 5th and 12th April in 1993 at Delhi to observe the peak period of shoot fly activity during spring season. But in respect of egg laying, the shoot flies did not show any preference to any of the two varieties sown on different dates. The maximum and minimum deadhearts incidence of 17.9 and 6.6 per cent was observed irrespective of varieties, on the crop sown on 8th March and 6th April, respectively. However, maximum number of eggs (6.28/ plant) was observed in crop sown on 22nd March (Rao and Panwar 1995a). Singh and Kanta (2004) on the basis of two years study (in 1999 and 2000) at Ludhiana reported comparatively more incidence of shoot fly on crop sown in mid February than that sown in March and April. Gagandeep and Kanta (2007b) in another study on influence of date of sowing on incidence of A. naqvii on maize reported that A. naqvii produced maximum damage of 25.77 per cent dead hearts on 6th March sown crop. They further reported that among different dates the deadhearts incidence varied from 2.33 to 12.83, 2.63 to 12.67, 17.64 to 33.90

and 13.50 to 17.60 per cent under protected and unprotected conditions, on crop sown on 15th, 25th February, 6th and 16th March, respectively. The maximum pooled grain yield (9.46 q/ha) over protected/unprotected conditions was obtained from the crop sown on 15th February and mean yield of crops sown at different dates was significantly higher under protected conditions (11.36 q/ha) than that in unprotected conditions (4.75 q/ha). 2.4 Host plant resistance for the management of shoot flies, Atherigona spp. Host plant resistance to insects is one of the easiest and cheapest components of an integrated pest management programme. It is an environmentally safe method of pest management, and is compatible with other control strategies such as biological, cultural and chemical control (Smith 2004). Host plant resistance is one of the most effective means of keeping shoot fly population below Economic Threshold Levels, as it does not involve any cost input by the farmers. Plant resistance to shoot fly appears to be complex character and depends on the interplay of number of componential characters, which finally sum up in the expression of resistance to shoot fly (Dhillon 2004). 2.4.1 Evaluation of germplasm to Atherigona spp. Germplasm evaluation in maize for reaction to Atherigona spp. has been carried out under natural infestation only. The fish-meal technique (Nwanze 1997) is quite useful for increasing shoot fly abundance for screening the test material under field conditions. The moistened fish-meal is applied @ 50 g per m2 one day after seedling emergence by broadcasting in the field to screen maize germplasm (Jindal et al 2007). Among the various forms of shoot flies damage, the evaluation of germplasm is done by counting the deadhearts produced by them. The deadheart count is considered easy, quick and useful criteria for assessing the relative reaction of genotypes as it gives approximate idea about the total loss of plant population and yield (Sharma and Singh 1975). The 19 ratings scale for scoring shoot fly species damage in maize was given by converting percentages of the dead hearts due to shoot fly into the rating scale (Sharma et al 1992). The detail of rating and description is as 1 = < 10 %, 2 = 1020 %, 3 = 2130 %, 4 = 3140 %, 5= 4150 %, 6 = 5160 %, 7 = 6170 %, 8 = 7180 %, 9 > 80 % deadhearts. The plants having only leaf injury, generally, show remarkable recovery (Sekhon et al 1993). To confirm the resistance to shoot fly observed under field conditions and to study the resistance mechanisms involved in host plant resistance to sorghum shoot fly, A. soccata in sorghum a cagescreening technique was developed by Soto (1972). The technique was modified at International Crops Research Institute for the Semi-Arid Tropics (ICRISAT) to simulate field conditions more closely and has the advantage of requiring no artificial rearing of shoot flies (Taneja and Leuschner 1985). Shoot flies used for cage screening are collected from a trap baited with fishmeal. The cage screening technique can

be used for multiple as well as no- choice conditions. Such technique is not in use for screening of maize genotypes against A. naqvii in India. In India, up till now about 2000 maize germplasm lines have been screened against Atherigona spp. and number of resistant sources has been identified by various workers from different locations (Siddiqui et al 1988, Panwar 2005, Kumar and Kanta 2011). On the basis of deadhearts by shoot fly, maize germplasm with white semiflint br pop, white semiflint br pop-2 and D-37 were categorized as least susceptible while, early white composite and Romanian flint proved to be highly susceptible (Sharma and Singh 1975). Among forty two maize germplasms comprising of some locals, elite material and single cross hybrids tested under heavy natural incidence of shoot fly complex in spring season the percentage of deadhearts varied from 34.8 to 82.6, with Lakshmi composite from Bihar and Khevawal local 2 from Punjab were relatively less susceptible (Marwaha et al 1986). Sarup and Panwar (1986) reported tolerance to shoot fly in maize germplasm in different maturity groups. In another germplasm evaluation study Sharma and Panwar (1998) found that deadhearts percentages in different germplasms varied from 23.0 to 83.6 per cent and tolerant lines belong to early maturity group. The judicious utilization of less susceptible lines to both maize borer and shoot fly in resistance breeding can immensely help in developing maize cultivars resistant to pests possessing other desirable agronomic traits (Panwar and Sharma 1998). Continuing the process of identifying the resistant lines to shoot fly in maize Panwar (1995) reported that among the twenty seven elite maize germplasms tested, the deadhearts percent ranged from 5.0 to 24.2 and the four lines EH 40428, HYD 8275, SSM 5010 and MMH 41 were found to be tolerant for incorporation in future breeding programmes. Multiple pest resistance in cultivar Diara EV and in two inbred lines (IPA 34-10-13-3-1-1-#-2-1 and IPA 3-6-14-2-#-1) to the shoot fly species and maize borer was observed by Panwar and Sarup (1988) and Panwar et al (2000), respectively. Similarly multiple resistance to maize borer and shoot fly in four genotypes EV5098, EV 6098, Agaiti 2002 and EV 1098 at Sahiwal in Pakistan was reported by Shahzad et al (2006). Jindal et al (2007) reported maize hybrids PMH 1 (8.53 %) and JH 3459 (11.03 %) as less susceptible on the basis of deadhearts formation in comparison to Parkash (25.36 %) and PMH 2 (24.95 %) to shoot fly in spring maize in the Punjab. Among inbreds tested, CM 143 was found to be tolerant (16.24 %) while the highest incidence was highest (46.32 %) on CM 144. 2.4.2. Mechanisms of resistance All the three major mechanisms of resistance viz, oviposition non-preference (antixenosis), antibiosis, and recovery contribute to host plant resistance to sorghum shoot fly, A. soccata (Soto 1974, Raina et al 1981, Taneja and Leuschner 1985, Sharma et al 1997, Dhillon et al 2005a, 2006). However,

with respect to maize, a few reports are available on oviposition and tolerance but scanty information is available in the literature elucidating the mechanism of antibiosis for plant resistance to A. naqvii. 2.4.2.1 Antixenosis for oviposition Non-preference by insects is often projected as a property of the plant to render it unattractive for oviposition, feeding or shelter. Non preference for oviposition is considered to be a primary mechanism of resistance to shoot fly in sorghum (Singh and Jotwani 1980a, Singh et al 1995). Behavioral responses of shoot fly, A. soccata in sorghum have shown that initial choice of the host was random, but the females spent less time on the resistant cultivars (Raina et al 1984). Females laid eggs on non-preferred cultivars only after laying several eggs on the seedlings of susceptible cultivar in sorghum. However, according to Dhillon et al (2005a) this mechanism of resistance was not stable and tended to breakdown under nochoice conditions. The varying preference for oviposition by shoot fly (Atherigona spp.) was reported and different maize germplasms were grouped on the basis of number of eggs laid per plant at Pantnagar (Sharma and Singh 1975). However, no difference in discriminating plants of various maize germplasms for oviposition by Atherigona spp. was reported by subsequent workers (Sarup and Panwar 1986, Panwar and Sarup 1988, Rao and Panwar 1996a). Similarly, Rao and Panwar (2001a, 2001b) reported that none of the morphological and biochemical plant characters influence egg laying by shoot fly in field on maize. Seven maize cultivars viz., Antigua Gr.I, Deccan 103, Kiran, Ganga 11, Deccan 105, Pusa composite 1 and CM 300 were evaluated against Atherigona spp. under natural infestation conditions. Over number of years it was observed that Atherigona spp. did not discriminate for egg laying on 9-11 day old plants of different varieties. Jindal et al (2007) reported that among the different genotypes tested at Ludhiana, JH 3980 was the least preferred genotype for oviposition by shoot fly. However, there was no significant difference in egg laying in susceptible PMH 2 and moderately resistant hybrid JH 3459.

10

2.4.2.2 Antibiosis The resistance to sorghum shoot fly is a cumulative effect of non-preference and antibiosis (Raina et al 1981). Retardation of growth and development; prolonged larval and pupal periods; and poor emergence of adults on resistant genotypes provide an evidence of antibiosis to sorghum shoot fly (Sharma et al 1997). Sharma and Singh (1975) reported that some genotypes in maize, more preferred for oviposition, had lesser average number of surviving maggots per plant indicating the role of antibiosis in different germplasms. The shoot fly maggots fed on resistant sorghum cultivars i.e. SFCR 151, ICSV 705, SFCR 125 and IS 18551 exhibited longer larval period (10.111.0 days), lower larval survival (54.767.8 %) and adult emergence (46.752.2 %) than on susceptible check Swarna i.e.9.3 days, 73.3 and 60.6 %, respectively (Dhillon et al 2005a). Singh and Narayana (1978) reported that the fecundity of shoot

fly females was greater when raised on susceptible check variety Swarna, than that on moderately resistant varieties IS 2123 and IS 5604 However, reverse trend was observed by Dhillon et al (2005a). 2.4.2.3 Tolerance Seedling vigour and high rate of recovery of sorghum are important characteristics of resistant cultivars to shoot fly (Sharma et al 1977). Maize germplasms vigorous in growth in the early growth stage showed lower percentage of deadhearts and it might be due to quick recovery of plant from the injury by shoot fly maggot or coming out of maggots along with the extended leaves during the quick growth of plants (Sharma and Singh 1975). The differences in increase of deadhearts in subsequent observations among maize germplasms were due to their relative capacity to withstand or recover from the shoot fly damage (Pandey and Sharma 1980). Patel and Sukhani (1990) reported negative correlation of plant height and stem length with oviposition and deadhearts by shoot fly in sorghum, suggesting that the pest resistant plants grew faster and might therefore escape the damage. The significantly higher plant height in infested plants of less susceptible maize cultivar JH 3459 (116.9 cm) than the susceptible cultivar PMH 2 (76.3 cm) vis-a-vis non significant difference in height of healthy plants of these genotypes indicates tolerance in less susceptible hybrid. The lowest reduction in height (33.2 %) and grain weight per plant (38.7 %) in infested plants due to Atherigona spp. over healthy plant was also observed in hybrid JH 3459 (Jindal et al 2007). This supports the idea that the resistance to shoot fly in maize is primarily dependent on the degree of tolerance.

11

2.4.3

Bases of resistance to Atherigona spp. Plant resistance to Atherigona spp. is a complex character and depends on the interplay of number

of componential characters, so it is important to identify genotypes with different mechanisms to increase the levels and diversify the bases of resistance to this pest. A diverse array of plant characteristics may influence its resistance/susceptibility to Atherigona spp. 2.4.3.1 Morphological plant traits Plant morphology can have a strong impact on shoot fly population dynamics, especially in case of seedling characteristics that physically reduce feeding, oviposition and shelter. Faster seedling growth and toughness of the leaf sheath are associated with resistance to shoot fly in sorghum (Singh and Jotwani 1980b). Jayanthi et al (2002) showed that shoot fly resistant parental lines and their hybrids showed significantly high seedling vigour compared to susceptible parental lines and their hybrid groups. The negative correlation between seedling vigour with egg counts and deadheart formation was reported by Kamatar and Salimath (2003) in sorghum. Trichomes or plant hairs are common anatomical features on leaves, stem and/or reproductive structures in higher plants. Levin (1973) had described the role of trichomes in plant defense and pointed out that in numerous species, there is negative association between trichome density and insect feeding, oviposition responses and nutrition of larvae. The trichome density has a positive correlation with resistance to Atherigona spp. in maize. The Atherigona spp. resistant varieties of maize possessed higher trichome density on leaves (Rao K R and Panwar, 1995b). In sorghum also, number of eggs per plant and per cent plants with eggs were negatively correlated with trichome density when observed 14 days after emergence (Patel and Sukhani 1990, Dhillon et al 2005a). Karanjkar et al (1992) suggested that although there are highly significant negative correlation between the trichome density and shoot fly infestation, yet it seems that trichomes do not have a direct role in reducing the deadhearts, but are associated with reduced oviposition and can be used as a reliable selection criterion to select germplasm for resistance to shoot fly. The leaf glossiness (pale green and shiny leaves) at seeding stage probably has a strong influence on the orientation of shoot fly, A. soccata females due to reflection of light in sorghum. The genotypes with high leaf glossiness are resistant to shoot fly (Dhillon et al 2005 a,b). The leaf surface wetness (LSW) i.e. the role of morning dew in the movement of freshly hatched shoot fly, A. soccata larvae through the leaf sheath to the growing point in sorghum was first studied by Rivnay (1960). Nwanze et al (1990) reported that LSW is associated with shoot fly resistance in sorghum which was reported to be more in 10-day old seedlings than seedlings of other ages. Larval survival is

12

affected by the wetness of the central shoot and initial contact of larva with moisture enhances its movement and survival (Nwanze et al 1992). The taller varieties with more leaves are desirable for minimizing the shoot fly incidence. Rao and Panwar (2001a) reported that the leaf width and stem thickness were positively associated where as the number of leaves per plant and leaf length were negatively associated with shoot fly deadhearts in maize. However, there was no significant influence of these plant characters on the oviposition. Singh and Jotwani (1980b) reported that in sorghum, the leaves of resistant varieties are narrower and longer than the susceptible ones. The rapid seedling growth; and long and thin leaves during the seedling stages make plants less susceptible to shoot fly attack (Singh 1998). The oviposition by shoot fly in sorghum was negatively correlated with seedling height, leaf length and stem length, but positively correlated with number of leaves per plant, leaf width, stem girth, and panicle initiation. However, deadhearts produced by shoot fly were negatively correlated with seedling height, leaves per plant, leaf length, leaf width, and stem length and positively correlated with stem girth and panicle initiation (Verma and Singh 2000). The green colour leaves with rough surfaces were more preferred for egg laying by A. soccata (Raina 1982). Similarly, Patel and Sukhani (1990) reported the positive correlation between seedling weight at 20 DAG and leaf colour with shoot fly damage in sorghum and suggested that healthy and stout seedlings with dark green leaves are more preferred for egg laying resulting in higher deadhearts incidence. The plumule and leaf sheath pigmentation in sorghum were found to be associated with resistance to shoot fly (Dhillon 2004, Dhillon et al 2005 a, b). 2.4.3.2 Biochemical plant traits: The literature reviewed identified only a few deliberate screening programmes for identification of the role of biochemical components in imparting resistance to A. naqvii on maize. But, a lot of information is available on the role of biochemical constituents imparting resistant to related species A. soccata on sorghum. Plant resistance is attributed largely to secondary metabolites such as phenolics, nonprotein amino acids and alkaloids, which are documented to be deleterious to insect herbivores (Manuwoto and Scriber 1985, Wang et al 2006).Plants are known to produce certain chemical compounds in different quantities and proportions, which affect the behavior and biology of phytophagous insects (Painter 1958, Beck 1965, Schoonhoven 1968), and can be attractants (oviposition and feeding stimulants) or repellents (oviposition and feeding deterrents) or antibiotic (reduced survival, growth, and fecundity). Cultivars that lack these defense mechanisms are often too vulnerable to damage of insect pests. An important group of defense chemicals in maize host plant resistance to corn earworm is

13

consisted of secondary metabolites C-glycosyl flavone maysin and the phenylpropanoid product chlorogenic acid in silk. The first brood of the European corn borer (Ostrinia nubilalis) is controlled by high levels of the benzoxazinoid DIMBOA in seedlings and young plants (Klun et al 1970). These secondary metabolites are produced independent of the presence of the pest in a tissue and develop in a specific manner. In addition to secondary metabolites, ubiquitous phenolic acids, especially ferulic acid, may contribute to insect resistance in maize (McMullen et al 2009). Resistance to Atherigona spp. and C. partellus in maize (Rao C N and Panwar 2001b, 2002), and sorghum shoot fly in sorghum (Mote et al 1979, Kamatar et al 2003) is associated with low levels of proteins. Kabre and Ghorpade (1999) indicated that protein content was positively correlated with stem borer susceptibility in maize, while such relationship between free amino acid content and stem borer susceptibility was not evident. The maize plants with resistance to spotted stem borer, C. partellus had low amounts of sugar (Sekhon and Kanta 1994). Similarly Kabre and Ghorpade (1999) reported that total and reducing sugars were positively correlated with stem borer susceptibility in maize. The slight increase and decrease in reducing sugars between 17 and 20 days after seedling emergence in shoot fly resistant sorghum genotypes and susceptible varieties, respectively was reported by Bhise et al (1997). However, Singh et al (2004) reported that resistance to shoot fly is associated with low levels of reducing and total sugars in sorghum seedlings. Development of sugarcane aphid, Melanaphis sacchari (Zhent.), and delphacid, Peregrinus maidus (Ashm.) populations were more pronounced in varieties with higher sugar content in leaves (Mote and Shahane 1994). However, soluble sugar content had little influence on midge, Sitodiplosis mosellana (Gelin.) resistance in wheat (Shi ZhongLiang et al 2002). Tannins are polymers resulting from condensation of flavan-3-ols. The significant and negative correlation between tannin content and shoot fly damage was reported by Chamarthi (2008). Shi ZhongLiang et al (2002) reported that tannins are important secondary metabolites for induced resistance to blossom midge in wheat. Polyphenols are widely distributed in plants, but they are not directly involved in any metabolic process and therefore, are considered to be secondary metabolites. Phenolic compounds in maize are (E) ferulic and (E) -p -coumaric acid which are attached to hemicellulose through pentose sugars. The plant phenolics have attracted great interest in relation to their diversity in chemistry; and functionality in biology, chemistry, medicine, ecology and agriculture. In agriculture, they have long been recognized as allelochemicals (Rice 1984), and constitutive and induced plant defense mechanisms (Vidhyasekaran 1988).

14

The (E)-ferulic acid content of the grain of nine populations of land races of maize derived from CIMMYT's collections was found to be negatively correlated to susceptibility characteristics towards the maize weevil, Sitophilus zeamais Motsch. (Classen et al 1989). Shoot fly resistance has been reported to be associated with high amounts of phenolic compounds in sorghum seedlings (Khurana and Verma 1983, Kumar and Singh 1998). The moisture content was low in maize genotypes resistant to shoot fly damage (Rao and Panwar 2002). However, no relationship was observed between moisture content of sorghum seedlings and shoot fly resistance (Singh et al 2004). Low levels of nitrogen were earlier reported to be associated with the shoot fly resistance in sorghum (Singh and Narayana 1978, Singh and Jotwani 1980c, Khurana and Verma 1983, Chavan et al 1990). Maize germplasm with high level of resistance to spotted stem borer, C. partellus had high contents of silica and iron, but low contents of nitrogen, phosphorus and potassium (Sekhon and Kanta 1994). The susceptible variety (HY 4642) of maize to C. partellus had higher percentage of nitrogen and sugar than resistance variety (CM 137) having higher percentages of P, K, Fe and Si in stem tissues (Arabjafari and Jalali 2007). In maize, none of the biochemical plant factors had influenced the eggs laid by the shoot flies in the field. Significant positive correlation of nitrogen and crude protein content at 5, 10 and 15 DAE and carotenoid content in 10 DAE was observed with deadhearts percentage caused by Atherigona spp. It showed that resistant varieties had low carotenoid, nitrogen and crude protein contents as compared to susceptible maize cultivars having more nitrogen and protein favour the insect growth and development (Rao C N and Panwar 2001b). In sorghum, the analysis of various chemical constituents showed that percentages of nitrogen; reducing and total sugars; moisture; and chlorophyll of leaf in shoot fly susceptible hybrid CHS 1 were 6.15; 41.08 and 35.33; 1.11; and 26.62 times higher, respectively than in resistant varieties. The lysine content was also more in susceptible hybrid (Singh and Jotwani 1980c). The significant and positive correlations in percentage seedlings with eggs, number of eggs per seedling and per cent deadhearts were observed with nitrogen; potassium; phosphorus; reducing and total sugars at different DAE of sorghum seedlings (Singh et al 2004). The lower amount of chlorophyll in sorghum leaves turns the leaves yellowish green and probably make them unattractive for oviposition (Patil et al 2006). Pest attacks generally set in motion or accelerate a complicated series of metabolic disturbances in the host, rather than effecting a simple change in a unique process (Hewett 1977). The total phenols, glucose and total free sugars increased significantly in the samples infested with Ostrinia furnacalis Guenee in maize after 14 to 16 days of its infestation (Santiago and Mendoza 1983). Similar

15

alterations in biochemical constituents in mulberry after infestation with leaf roller, Diaphania pulverulentalis (Hampson) were reported by Mahadeva and Negaveni (2011). 2.5 2.5.1 Management of Atherigona spp. Management of Atherigona spp. with insecticide The just emerged maize seedlings suffer extensive damage leading to deadhearts formation by shoot fly species. Since, the adult of shoot flies oviposit on the emerging seedlings and in cracks and crevices around the seedlings (Kishore 1991), the plant protection measures at early seedling stage are essential to protect the crop from their severe infestations. Earlier application of a number of systemic granular insecticides in the furrows at the time of sowing has been reported to give good results against this pest (Chatterji et al 1973, Panwar et al 1999a). However, foliar sprays were not effective against shoot flies (Sharma et al 1979) in spring sown maize. The seed treatment with carbofuran 40F @ 10 g per 100 g of maize seed was found to be most effective for controlling A. naqvii at Ludhiana. The soil application of carbofuran 3 G @ 12.5 kg and phorate 10 G @ 10.0 kg per hectare at sowing time was also found to be effective against this pest at same place (Sajjan and Sekhon 1985, Kanta et al 2006). The seed treatment with 1.5 % carbofuran 35 ST or 25 ST or 1.0 % furathiocarb 40 SD or furrow application of carbofuran 3 G (0.375 kg a.i./ ha) and phorate 10 G (1.25 kg a.i./ ha) was most effective in reducing the deadheart formation due to Atherigona species in spring sown maize. Further more, the foliar application of synthetic pyrethroids and insect growth regulator chlorfluazuron (PP 145) also showed significantly less number of deadhearts vis--vis control but these were comparatively less effective than the above insecticides (Marwaha et al 1985, 1987). The effectiveness of dimethoate, monocrotophos, phosalone @ 0.1 % and @ 0.05 % chlorpyriphos reduced the formation of deadhearts due to shoot fly species. The spraying and dusting of maize crop at the seedling stage did not deter the shoot flies from ovipositing on the treated maize leaves or in the cracks and crevices in soil around the maize seedlings (Sinha 1989). Rao and Panwar (1992) evaluated the efficacy of carbofuran 3G, carbosulfan 25 STD, 3% neem kernel suspension, neem oil 20 EC, mohua oil 20 EC and phorate 10 G against shoot fly in spring sown maize and reported that average incidence of deadhearts was least in plots treated with phorate 10 G (8.82 %) and was highest in plots treated with neem oil (45.62 %). The plant products were not found effective to protect the crop against the ravages of shoot fly species in spring sown maize. The shoot fly did not discriminate between different treatments for egg laying. The effectiveness of maize treated seed with carbosulfan 25 STD @ 20 g and 40 g per kg seed, stored for different periods; and mixture of treated (60 %) and untreated (40%) maize seeds was also reported to be effective against shoot fly by Rao K R and Panwar (1995c, 1995d). Among different treatments i.e. application of carbofuran (slow release) 5 G @ 0.5 and 1.0 g m-1 row and carbofuran 3 G @ 1.0 g m-1 row; PEG-phorate

16

10 @ 0.25 g m-1 row and phorate 10 G @ 1 g m-1 row; and seed treatment with imidacloprid 70 WS @ 5.0, 7.5 and 10.0 g kg-1 seed significantly reduced deadheart formation in spring sown maize. However, shoot flies did not discriminate amongst the plants in different treatments for egg laying in the field (Rao K R and Panwar, 1996 b). The spot treatment by pinch application of Marshal 6 G (carbosulfan), Furadan 3 G (carbofuran) and Thimet10 G (phorate) proved effective in reducing incidence of A. naqvii in summer maize (Singh and Kanta 2006). The effectiveness of seed treatment with thiamethoxam 70 WS @ 2 g per kg seed of sorghum in reducing dead hearts incidence was reported by Kumar and Prabhuraj (2007). In spring sown maize at Ludhiana, Jindal and Hari (2008a) reported high effectiveness of seed treatment with imidacloprid 70 WS @ 5 g per kg seed in reducing dead heart incidence (3.42 %) by A. naqvii as compare to untreated control (70.03%) and carbofuran 3G application (13.59%). The foliar application of endosulfan 35 EC @ 200 ml per acre was also reported to be at par with granular insecticide applications in controlling the shoot fly (Hari and Jindal 2008). The improved formulation of imidacloprid (Gaucho) 600 FS @ 6 and 9 ml per kg seed was more effective in reducing deadhearts due to attack of A. naqvii (5.41 and 8.99 %, respectively), whereas the formulation of thiamethoxam (Cruiser) 70 WS @ 3.0 and 5.0 g per kg seed with (17.02 and 15.10 % deadhearts, respectively) was less effective (Jindal and Hari 2011). 2.5.2 Management of shoot fly with non chemical methods The more emphasis has been given on chemical control of shoot flies and is often being adopted by the farmers. The indiscriminate use of insecticides causes environmental pollution, residue problems, development of resistance, pest resurgence, secondary pest outbreaks and destruction of natural enemies of the pest. So, there is need to develop IPM strategies involving cultural methods of pest control for the management of shoot fly species in spring season. But it requires a thorough knowledge of life history, behaviour and ecology of insect as well as of its plant or animal hosts. The most vulnerable stage (s) in its life cycle must be determined and the practices be developed to prevent attack by the insect, to kill it, or to slow down its rate of reproduction (Lawani 1982). Inter-cropping of maize for the control of Atherigona spp. The principle of controlling insect pest populations is by increasing the diversity of an agroecosystem. It is crucial, however, that the right kind of diversity be established (Smith 1970). Under choice test for oviposition, sorghum shoot fly markedly preferred Sorghum bicolor than other graminaceous plants species i.e. Digitaria scallarum, Rottboellia exaltata, Setaria verticillata and Panicum maximum (Ogwaro 1978). Raina and Kibuka (1983) reported that in intercrop field trials 6 and 61 per cent eggs of shoot fly, A. soccata eggs, were received by maize and sorghum plants, respectively. Sarup et al (1984) reported higher per cent dead hearts in sorghum due to shoot fly species as compared to maize irrespective of sowing dates in spring season. The peak activity of shoot fly species exhibited in

17

maize sown on 16th march was at par with the lowest activity in sorghum planted on 30th March. Almost cent per cent deadhearts were produced in sorghum in the early three sowings implaying that peak activity of shoot fly in spring sown maize and sorghum was unrelated. The polyphagous nature of shoot fly species invading the spring sown maize led to suggestion of possible exploitation of this habit to control damage in maize fodder by providing a trap of most suitable host sorghum (Sarup et al 1986). Subsequent evaluation of Sorghum planting in different permutation and combinations (5-35 per cent) along with maize was unworkable in spite of varied intensities of shoot fly infestation. The per cent deadhearts in maize in different years (1984-86) varied from 1.1-2.4, 24.8-37.9 and 56.4-7.12 with corresponding figure of 68.0-76.0, 87.5-100.0 and 94.6-98.9 in sorghum. There was simultaneous occurrence of both A. soccata and A. naqvii in spring maize seedlings as against only A. soccata damaging sorghum seedlings perhaps masked the overall effect of preferred host sorghum. The oviposition by A. naqvii soon after the emergence of maize seedlings at 2-3 leaf stage restricted the success of sorghum as trap crop (Sarup et al 1986). Maize was inter-cropped with sorghum, bajra, cowpea and soybean for the control of shoot fly species in spring 1993 season at Delhi. There was no significant effect of egg laying or dead heart formation in maize due to shoot fly species when maize was inter-cropped with other millets and pulses. On the other hand, inter-cropped sorghum received significantly more eggs per plant and dead heart formation due to shoot fly species vis vis other inter cropped crops with no egg laying or deadheart formation in unrelated hosts i.e. soybean and cowpea. The lowest per cent deadhearts were recorded in maize + soybean intercropping. More deadhearts (24.7 %) formed in maize intercrop with sorghum proved that it was not useful in management of shoot fly (Rao K.R. and Panwar, 1995 e). Effect of weather factors on incidence of Atherigona spp. The temperature and humidity influenced the development of the eggs, 30oC and 90 per cent relative humidity (R.H.) being the best combination, for the development of shoot fly, A. Soccata as reported by Doharey et al (1977). On wheat, A. naqvii was more active from the second fortnight of October to end December; and again during February and March. These fluctuations appear to have a positive correlation with maximum and minimum temperature while the relative humidity and the rainfall did not have any influence on activity of A. naqvii (Khan and Singh 1980). The egg hatching in A. soccata coincides with the presence of moisture on the leaf (Raina 1981). Low humidity (30%) increased the
0 0

duration of egg development and decreased the egg survival. Temperatures above 35 C and below 18 C and continuous rainfall reduce the shoot fly abundance (Delobel 1983, Taneja et al 1986). Marwaha et al (1984) found that average maximum temperature ranging from 27 to 30oC and minimum 12-15oC coupled with higher relative humidity between 75-80 per cent and some rainfall in the last week of February with sporadic drizzling in March were conducive to Atherigona spp. activity in terms of higher oviposition and

18

deadhearts formation. These aspects can successfully be utilized for the management of shoot fly. Naitam and Sukhani (1989) reported that more soil moisture and temperature results into lesser shoot fly damage and increased sorghum. Sekhon et al (1986) reported that winter maize crop, sown on southern face of ridges, suffered significantly less damage than that sown as flat and on northern face of the ridge due to more rapid and vigorous growth of maize plants, which is attributed to the prevalence of relatively higher temperature during day time on southern side of ridges. Similar observation was made on spring sown maize by Jindal et al (2012). Nair et al (1995) reported significant positive and negative correlation of egg laying by A. soccata in sorghum with maximum and minimum temperature, respectively. Its correlation with morning and evening relative humidity was significantly negative and with sunshine hours was significantly positive. Effect of Nitrogen on incidence of Atherigona spp. The introduction of hybrids and high yielding varieties responsive to fertilizers has impact on insect pest abundance also. Sharma and Singh (1975) reported that more application of nitrogenous fertilizer in maize prompted growth and vigour of plants and helped in recovery of plants from shoot fly damage. Nitrogen treated sorghum plants were more preferred by shoot fly for egg laying than control plants but reduced incidence of shoot fly was observed with nitrogen application and it might be due to vigorous fast growth of plants by nitrogen (Mote and Ramshe 1987). However, Chamarthi (2008) reported that more vigorous plants were preferred by shoot fly for egg laying. Biological control of A. naqvii Three parasites namely, Tetrastichus nyemitawus Rohwer, Tetrastichus spp. and Neochrysocharis spp (Eulophidae) were reported parasitizing A. naqvii from Udaipur and Rajasthan (Kundu and Kishore 1972). Egg parasitoids i.e. Trichogramma spp. and Trichogrammatoidea spp. were reported from related species A. soccata (Sharma 1985). 2.6. Other insect pests infesting spring maize The availability of maize at one or more stages of growth throughout the year added new dimensions to the pestilence front (Kumar and Kanta, 2011). Insects hitherto unknown to attack the maize have become problems, as in present studies small brown planthopper Laodelphax striatellus (Falln) is being reported for the first time infesting spring sown maize in the Punjab. The brief description of insect pests prevalent in spring season is as under: Maize stem borer, Chilo partellus (Swinhoe) is a widely distributed insect pest in maize and sorghum agro-ecosystem in India. The pest is active from March to October and has 6 7 over

lapping generations (Kumar et al 2005). The major loss was caused in the early stages of the plant. The

19

freshly hatched tiny larvae began to feed on the tender leaves, particularly in the central whorl, which later emerged out in a severely fenestrated condition. Thereafter, the larvae bored into stem to form tunnel. In case of younger plants wherein sufficient stem formation had not taken place, the caterpillar cut the growing point of the crop and the base of the central whorl got badly damaged, resulting in the drying up of the central whorl and forming deadheart. This condition did not occur in the later stage when the stem was so thick that even a large number of larvae could continue to tunnel therein without leading to any external symptom. Right from germination till harvest, maize crop suffered from

depredation of this borer. All parts of the maize plant except roots were attacked (Panwar 2005). In peninsular India, the winter was not severe, therefore, the pest remains active around the year (Sharma et al 2006). The management of this pest involve use of resistant cultivars, manipulation of sowing dates, napier bajra hybrids as trap crop, intercropping of maize with pulses, ploughing up of field after the crop harvest, optimum seed and fertilizer rate, use of egg parasitoid Trichogramma chilonis and spraying of insecticides at early whorl stage of plant growth, spot application with granular insecticides (Hari and Jindal 2009, Jalali and Singh 2003; Kumar and Kanta 2011; Sarup et al 1978; Srinivas and Panwar 2003). The extent of damage of this pest in spring sown maize is not reported in literature. Army worm, Mythimna separata (Walker) is a polyphagous graminaceous pest. Srivastava and Khan (1961) observed that the army worm cause heavy losses to maize at two stages of its growth, viz., immediately after germination up to development of nodes and internodes; and secondly one month after sowing. The late infestation results in complete defoliation due to larval feeding from edge towards mid-rib. The pest attacks winter maize crop and incidence increases during March-April. The larva also attack maize silk in April-May. Corn earworm, Helicoverpa armigera (Hbner) is a polyphagous pest and noticed in summer and winter season maize in Punjab. There may be as many as eight generations in a year. The young larvae start damaging tassel and move downwards. Eat silk and may tunnel in the ears and damage grains, in the milky stage. The damage in the ear may form a channel like structure, which is seen after exposure of the grains. The attacked cobs show masses of moist excreta on them (Kumar and Kanta 2011). Other silk cutter, Euproctis virguncula Walker feeds on the freshly matured silk during September-October, but do not effect the pollination (Sandhu and Kaushal 1972, Sandhu et al 1974b). Cutworms, two species belonging to Agrotis, have been reported to attack maize. The caterpillars, hiding during day time in cracks in soil, damage maize seedlings at dusk. The older larvae kill the plants by cutting it just above the soil surface. At Pantnagar, the larvae number as high as 18 per meter length were recorded infesting spring sown maize (Kumar and Kanta 2011). The climbing cutworm, Rhyacia

20

herculea (Corti and Draudt) was reported to cause 20 to 40 per cent loss in winter sown maize crop in Bihar (Verma et al 1979). Grasshoppers are sporadic pests on maize crop. In favourable season they may prove harmful and leave nothing on plant except stem and mid-rib of leaves (Kumar and Kanta 2011). These are polyphagous pests, both nymphs and adults, generally feed on grasses before attacking the crop. Regular summer ploughing and destruction of bunds to expose insect eggs to bright sun is recommended along with mechanical control of gregarious (newly hatched) nymphs. Sugarcane leaf hopper or Pyrilla, Pyrilla perpusilla Walker was reported to cause heavy loss to maize crop during March-May (Kumar and Kanta 2011). It feeds on number of host plants including maize (Kumarasinghe and Wratten 1996). Both nymphs and the adults suck the plant sap from underside of the leaves. The infested plants turn pale and in severe attack may dry up completely and the black fungus develops on the leaves which hinders photosynthesis and growth of the plants. The maize jassid, Zygnidia manaliensis (Singh) is one of the major pests of spring sown maize in the Punjab. The nymphs and adults of this pest suck sap from leaves and produce whitish stippling, giving bad look to maize fodder and deteriorating its quality (Sajjan et al 1982). They further reported that malathion and phosphamidon gave complete control of this after 4 days of application. Economic losses of 32 per cent in fodder yield during spring season and 15 per cent loss in grain yield of winter crop in the Punjab have been reported by Singh (1991). The Jassid infestation on spring sown maize fodder was reported to be high by 3rd week of May (36/150 leaves) which declined later (Sodhi and Sekhon 1997). The cereal thrips, Anaphothrips sudanensis Trybom cause damage to maize seedlings in winter and spring seasons (Sandhu et al 1974a). The infestation has been reported to range between 29 to 86 per cent. Both adults and nymphs suck the sap from leaves, growing tips and gradually congregate within the leaf sheath to form colonies. As thrips start feeding on the oozed out sap, small silvery specks appear on the infested leaves, which enlarge to form brown patches. These patches increase in size spread all over the leaf which ultimately dries up. Heavy infestation at the seedling stage may cause complete loss of the crop. Its population reaches peak in February to April (Mathur 1988). Small brown planthopper, Laodelphax striatellus is being reported for the first time infesting spring sown maize in the Punjab in the present studies. L. striatellus commonly occurs in Taiwan, Japan, korea, China and the palearctic regions. Shukla (1979) first reported its prevalance in India on rice nursery (variety Jaya) in the farmers fields near Ludhiana, Punjab. Thus could be distinguished from white backed plant hopper by coloured scutellum as against white. It is polyphagous and its hosts include rice, maize, oats, tall oat grass , wheat , barley etc. It vectors a number of viral diseases like Barley yellow striate mosaic virus (BYSMV), Maize rough dwarf virus (MRDC), Northern cereal mosaic virus

21

(NCMV; including the Wheat rosette stunt virus - WRSV), Rice stripe tenuivirus (RSV), Rice blackstreaked dwarf virus (RBSDV) and Wheat chlorotic streak virus (WCSV) (Caciagli 1991). The life table studies on five varieties of rice, wheat and two weed speceis i.e. Echinocloa crusgalli and Leptochloa chinensis showed that nymphal duration on E. crusgalli was shortest (23 days). E. crusgalli was the most suitable host follwed by japonica rice variety and then wheat (Li et al 2009). High temperature determines the ups and downs of small brown planthopper population. The development duration, longevity of adults, pre oviposition and oviposition duartion decreased with the increase of temperature from 18oC to 27oC, whereas development of nymph became slow at 30oC and the developmental duration was delayed about 16 days. The survival rate was higher (81-88 %) at 21-27oC, but it decreased to 5 % as the temperature increased to 30oC. At 30oC the longevity of female adults was very short and they did not lay eggs (Zhang et al 2008). L. striatellus harbours intracellular yeast like symbiotes in the fat body, transmitting them to next generation through the female ovary. High temperature of 35oC destroyed the yeast like symbiotes in the mycetocutes. Under the continuous high temperature no adults were obtained (Noda and Saito 1979). The high temperature also influences the transovarial passage of rice stripe virus (RSV) in small brown planthopper. Most of the females at 17.5oC showed high transovarial passage i.e. 82.6 % of viruliferous females passed RSV to progeny at the rate of more than 90%. The percentage of females which showed high transovarial passage ( >90% ) was 60.9 % at 25oC and 12.5 % at 32.5oC and higher the rearing temperature, the lower was the percentage transovarial passage (Raga et al 1998). The fipronil, nitenpyram, chlorpyriphos are the few insecticides that provide effective control of L. striatellus in China and its populations have been discovered with resistance to organophosphate, carbamate and neonicotinoid insecticide classes (Endo and Tsurumachi 2000, Gao et al 2008). The information of infestation of this pest on maize is scanty. To sum up this review, the available literature reveals that a number of maize genotypes/ varieties/ lines have been screened against Atherigona spp. in spring sown maize, but detailed investigations regarding the bases of host plant resistance is required. A better understanding of the behaviour of shoot fly and interaction of environmental factors could serve as a basis for manipulation of the cultural practices, need based pesticide application to reduce the risk of high pest population build up and to minimize its damage to the host plant. In the era of changing climate and introduction of new intensive cropping pattern, the systematic information on insect pests incidence and extent of damage by insect pests complex needs to be updated for strengthening the IPM programmes in different crops including maize.

22

Chapter-III MATERIALS AND METHODS The studies on the Incidence of insect pests and management of shoot fly, Atherigona spp. in spring sown maize were conducted at the Research Farm and the Maize Entomology Laboratory, Department of Plant Breeding and Genetics, Punjab Agricultural University (PAU), Ludhiana, during spring 2011 and 2012. The university is situated at 30 45' N, 75 40' E with an altitude of 247 m.a.s.l. One of the field experiments was also carried out at the farmers field in village Chagra of district Hoshiarpur (Punjab). The biochemical analysis was carried out in the Quality Laboratory, Department of Plant Breeding and Genetics, PAU, Ludhiana. 3.1. Procurement of seed of maize/ sorghum The seed of different hybrids and inbred lines of maize was procured from Senior Maize Breeder, Maize Section, Department of Plant Breeding and Genetics and Director Seeds, PAU, Ludhiana for conducting the field experiments. The seed of sorghum variety SL 44 was procured from Incharge, Fodder Section, Department of Plant Breeding and Genetics, PAU, Ludhiana. The seeds were stored air tight at 40C in refrigerator for further use. 3.2. Test genotypes The eight maize genotypes viz., JH 3459, PMH 2, PMH 1, JH 3956, JH 31244, Parkash, LM 16 and CM 143 were tested in different experiments. These genotypes, recommended for cultivation in the state by PAU, were selected on the basis of their field reaction to shoot fly as observed in the previous years studies under All India Coordinated Research Project (AICRP) on maize. 3.3. Raising of the crop The maize crop was sown under flat and raised as per PAU, recommendations (Anonymous 2010) except any insecticide application in the untreated control treatments meant for making comparisons. The crop was sown @ two seeds per hill in each treatment plot of different experiments in the field. After completion of crop germination, the thinning was done to ensure one plant per hill in each treatment plot. 3.4. Population build up of insect pests The experiment on population build up of insect pests in spring sown maize was conducted at the farmers field in village Chagra of district Hoshiarpur under unprotected conditions. The maiz e cultivars JH 3459 and PMH 2 with two nitrogen level of 90 and 120 kg N ha-1 were flat sown in randomized block design (R.B.D.) factorial with 4 replications on February 21st during 2011 and 2012. The nitrogen was

23

applied in three splits as per PAU recommendations. Each treatment plot had 6 rows with 4 m row length. The row to row and plant to plant spacing was 60 and 15 cm, respectively and the trial was managed as per PAU recommendations (Anonymous 2010). The observations on the number of eggs laid by Atherigona spp. on the plant and nearby soil were recorded from 10 randomly selected plants in each plot at 7 ,14 and 21 days after germination (DAG) during both the test years. The leaf injury and deadhearts incidence due to attack of shoot fly were recorded from 7, 14, 21 and 28 days old seedlings from the inner 4 rows in each plot using standard procedures (Panwar 2005). From this data, the per cent leaf injury and deadhearts incidence was calculated separately as under: Leaf injury incidence (%) = (number of plants with leaf injury symptoms/ total number of plants) 100 Deadhearts incidence (%) = (number of plants with deadheart symptoms/ total number of plants) 100 Similarly, the population counts of small brown planthopper, Laodelphax striatellus (Falln), grasshopper nymphs; Sugarcane Pyrilla, Pyrilla perpusilla (Walker); coccinellids; and spiders were made from 10 randomly selected plants per plot at weekly intervals starting from seedling emergence stage onwards i.e. till the maturity of the crop. The damage by foliage feeder viz. maize stem borer, Chilo partellus (Swinhoe); army worm, Mythimna separata (Walker); silk cutter, Helicoverpa armigera (Hbner); and grasshopper spp. was recorded on the whole plot basis and was converted to per cent incidence. The two hybrids were compared for incidence and relative damage of insect pests under two nitrogen levels. The grain yield was also recorded from four inner rows after adjusting at 15 per cent moisture level and was converted to quintal per hectare. 3.5. Estimation of losses caused by insect pests in spring sown maize The experiment on estimation of losses caused by insect pests in spring sown maize was conducted at Research Farm, Department of Plant Breeding and Genetics, PAU, Ludhiana. The maize cultivars JH 3459 and PMH 2 comparatively tolerant and susceptible to shoot fly, respectively (Jindal et al 2007) were sown in three sets based on sowing dates i.e. first sown on 29th and 27th January; second on 11th and 14th February; and third on 26th and 28th February during spring 2011 and 2012, respectively. Three treatments viz. complete crop protection from shoot fly and other insect pests; crop protection from insect pests other than shoot fly and untreated control (no crop protection from any insect pest) were given to each of the above two hybrids sown under each of three sowing date sets during both the test years. The seed treatment with Gaucho (imidacloprid) 600 FS @ 6 ml/kg seed was done before sowing for crop protection from shoot fly attack. The spot application of carbofuran 3 G in 2nd week of April was

24

given to the crop against C. partellus in 2012 only. The experiment was conducted in R.B.D. factorial with 3 replications. Each treatment plot consisted of 5 rows with 3.6 m row length. The crop raising practices except the application of insecticides in untreated control plots were carried as per PAU recommendations (Anonymous 2010). The observations on the number of eggs laid by Atherigona spp. on the plant and nearby soil were made from 15 randomly selected plants in each treatment plot at 3, 7, 10 and 15 DAG in 2011 and at 7, 12 and 17 DAG in 2012 as no egg lying was observed at earlier observations. The leaf injury and deadhearts incidence due to shoot fly were recorded from 10, 15 and 20 days old plants on whole plot basis using standard procedures (Panwar 2005) to calculate per cent incidence. The population counts of small brown planthopper, L. striatellus; grasshopper nymphs; Sugarcane pyrilla, P. perpusilla; coccinellids and spiders from 10 randomly selected plants per plot were recorded at weekly intervals from seedling emergence stage onwards i.e. upto maturity of the crop in different sets during both the test years. In 2011, the population of L. striatellus was comparatively high and thus its population counts were recorded from two sweep catches of insect collection net from each treatment plots. The damage by foliage feeders viz. maize stem borer, C. partellus; army worm, M. separata; and grasshopper spp. was also recorded on whole plot basis and was converted to per cent incidence. The comparisons were made from the extent of losses caused by key insect pest i.e. shoot fly in two hybrids sown during different periods to identify most damaging crop growth period, most susceptible crop stage and to know whether early sown crop had any escape from this pest without any adverse effect on its grain yield. Similar observations were made for other insect pests also. The meteorological data was obtained from Department of Agricultural Meteorology, PAU, Ludhiana for making the correlations between incidence and damage of insect pests with weather parameters i.e. temperature, relative humidity, sunshine hours and evaporation prevailed during the period of studies. The grain yield was recorded from inner three rows in each treatment plot, adjusted at 15 per cent moisture level and was converted to quintal per hectare for estimation of comparative losses under protected and un-protected conditions sown under three sowing dates. 3.6. Role of various morphological and biochemical plant characteristics in resistance against shoot fly, Atherigona naqvii The experiments to study the bases of resistance against most prevalent species of shoot fly in spring sown maize in Punjab i.e. Atherigona naqvii in eight maize genotypes (JH 3459, PMH 2, PMH 1, JH 3956, JH 31244, Parkash, LM 16 and CM 143) were conducted during spring 2011 and 2012 at the Research Farm, the Maize Entomology Laboratory and Quality Laboratory, Department of Plant Breeding and Genetics, PAU, Ludhiana. The experiment to record incidence of shoot fly, A. naqvii and to study the

25

morphological basis of resistance in maize genotypes against A. naqvii was conducted under field conditions using fish-meal technique given by Nwanze (1997). The test genotypes were sown on 20th and 22nd February in 2011 and 2012, respectively in R.B.D. with 3 replications. Each treatment plot had 5 rows with 3 m row length. The crop raising practices were followed as per PAU recommendations (Anonymous 2010) except the application of insecticides. The optimum shoot fly population in field was ensured by broadcasting the moistened fish-meal at the rate of 50 gm per m2 (Jindal et al 2007). The correlation and regression coefficient for its infestation, biological attributes with morphological and biochemical parameters were also worked out. 3.6.1. The following observations were recorded on various parameters:

3.6.1.1. Incidence of shoot fly: The number of eggs laid per plant was recorded from 15 plants per treatment plot at 3 days interval from 3 to 15 DAG to assess antixenosis for oviposition in different genotypes. The leaf injury and deadhearts incidence was recorded on whole plot basis at 5, 10, 15 and 20 DAG to work out per cent infestation. The proportion of deadhearts incidence out of total shoot fly incidence (leaf injury and deadhearts) at 20 DAG was also worked out to calculate the recovery of seedlings for assessing the tolerance in different genotypes. 3.6.1.2. Expression of antibiosis to shoot fly, A. naqvii: To study antibiosis test genotypes were exposed to shoot flies adults under field conditions and were further used to study survival and development of shoot fly on different genotypes. The field plants along with 75 plants of each genotype in fish-meal baited small pots in 3 replications were sown and observed daily for egg laying and deadhearts formation. The plants with eggs were tagged after ensuring that the laid eggs were of shoot fly, A. naqvii species only. To quantify antibiosis deadhearts formed in tagged plants in field and pots were labelled on the day of their appearance to compute the larval period. Ten seedlings with deadhearts, six days after deadhearts formation in each genotype were taken from each replication and placed in 50 ml vials. These seedlings were dissected carefully to ensure the presence of the maggot; and on drying the seedling was replaced with the fresh one of the same age. A similar separate set of vials was also maintained for each genotype to ensure optimum number of pupae and adults for further studies. The observations were recorded on life cycle parameters i.e. larval and pupal periods; larval and pupal survival; pupal weight and fecundity (number of eggs laid per female) as per Dhillon et al (2005a). There were three replications for each genotype and the experiment was carried out in completely randomized design (C.R.D). Larval period and survival rate: The deadhearts with maggots (i.e. 6 days after deadheart formation) placed in glass vials were observed daily for their pupation. The days from deadheart appearance to pupation plus one day (because it takes one day for deadheart realization after egg hatching) was recorded as larval duration (Meksongsee et al 1981). The larval period was recorded separately for each larvae per

26

replication; and the mean larval period and survival rate was calculated on the basis of surviving larvae out of 10 larvae per replication. Pupal period and survival rate: The time taken from pupation to adult emergence was recorded as pupal period. The pupal period was recorded separately for each individual per replication and the mean pupal period and survival rate was calculated from the pupae which emerged as adults. The additional pupae of same age were obtained from the culture maintained in separate vials for each genotype . The number of pupae survived or emerged as adults were recorded and expressed as percentage pupal survival. Pupal survival = (Number of adult emerged / Total number of pupae) x 100. Pupal weight: Pupal weight (in mg) was recorded for individual pupa on an electronic balance, within 24 h of its pupation. The pupae were sorted into males and females; and their individual weights were recorded separately in each replication comprising of three pupae of each sex. After weighing, the pupae were placed in respective jars on moist sand to avoid their mortality because of desiccation for further studies. Fecundity: Three pairs of adults emerging from larvae reared on each genotype were released in each of three replications i.e. in glass chimneys (19 cm height), each having its mouth covered with piece of white muslin cloth and tied with rubber band. The bottom end of the glass chimneys were firmly adhered to the soil inside the pots (12 cm dia.) to prevent any exit route left for the shoot flies to escape. The cotton swab dipped in 20 per cent sucrose solution and brewers yeast + glucose in the ratio of 1:1 was kept in a Petri dish under each chimney to provide food to the adult flies. Two one week old maize seedlings (planted in pots of 12 cm dia.) of the same genotype, on which the larvae were fed, were provided to the shoot flies as an oviposition substrate. The seedlings were changed with fresh ones after every 2 days and the data on number of eggs laid in each case were recorded. 3.6.1.3. Evaluation of Morphological characteristics of maize genotypes: The observations on various morphological characters were recorded at fifth leaf stage of seedlings. The data on seedling vigour, leaf glossiness, trichomes, pigmentation, leaf length, leaf width, leaf area and stem girth were recorded on the test genotypes grown under field conditions, while the data on leaf surface wetness was recorded in seedlings grown under greenhouse conditions (Dhillon et al 2005a). There were three replications for each genotype. Seedling vigour: The seedling vigour (in terms of plant height, leaf expansion, plant growth, robustness and adaptation) was recorded on a 1 to 5 rating scale. The three observations were recorded for each

27

genotype per replication. The ranking of genotypes was done as per following rating scale (Dhillon et al 2005a): 1 = highly vigorous (plants showing maximum height, more number of fully expanded leaves, good adaptation and robust seedlings). 2 = vigorous (good plant height, good number of fully expanded leaves, and good adaptation and seedling growth). 3 = moderately vigorous (moderate plant height with moderate number of fully expanded leaves and fairly good seedling growth). 4 = 5 = less vigorous (less plant height with poor leaf expansion and poor adaptation). poor seedling vigour (plants showing poor growth and weak seedlings).

Leaf glossiness: Leaf glossiness (plants with pale green, shiny, narrow and erect leaves) was evaluated on a 1 to 5 rating in the early morning hours when there was maximum reflection of light from the leaf surfaces (Dhillon et al 2005a). The three observations were recorded for each genotype per replication. The rating of genotypes was done as under: 1 = 2 = 3 = 4 = 5 = highly glossy (light green, shining, narrow and erect leaves). glossy (light green, less shining, narrow and erect leaves). moderate glossy (fair green, light shining, medium leaf width and less drooping leaves). moderate non-glossy (green, pseudo-shine, broad, and drooping leaves). non-glossy (dark green, dull, broad and drooping leaves).

Pigmentation: Pink pigment on leaf sheath was assessed on a 1 - 5 rating scale (Dhillon et al 2005a). The three observations were recorded for each genotype per replication. The rating of genotypes was done as under: 1 = 2 = 3 = 4 = 5 = leaf sheath with dark pink pigment. leaf sheath with fair pink pigment. leaf sheath with light pink pigment. leaf sheath with very light pink pigment. leaf sheath with green colour.

Leaf surface wetness: The test genotypes were planted in small pots (12 cm dia). The observations on leaf surface wetness were recorded between 7.00 to 7:30 A.M. The seedlings at the 5 th leaf stage were brought to the laboratory; the central whorl leaf was opened and mounted on a slide with a sticky tape. Water droplets on the leaf surface were observed under the microscope (10 x magnification). The three observations were recorded for each genotype per replication. Leaf surface wetness was rated on a 1 to 5 scale i.e. 1 = leaf blade without water droplets, 2 = leaf blade with sparsely placed few water droplets, 3 =

28

leaf blade near mid rib covered with water droplets, 4 = water droplets spread all over the leaf blade and 5= entire leaf blade densely covered with water droplets (Sharma et al 1997). Leaf length: The leaf length (cm) was recorded from fully formed 5th leaf (from the base) taken at random in each genotype. The linear leaf length was recorded with measuring scale. The three observations were recorded for each genotype per replication. Leaf width: The leaf width (cm) was recorded from fully formed 5th leaf (from the base) taken at random in each genotype. The width was recorded from upper, middle and lower portion of leaf with measuring scale to calculate mean width. There were three observations for each genotype per replication. Leaf area: For studying the leaf area (cm2) of different maize genotypes 5 leaf (from the base) taken at random was detached from the plant and the leaf area was determined using Cl-203 Laser Leaf Area Meter (CID, Inc, USA). Three plants were selected from each genotype per replication.
th th

Stem girth (Circumference): The stem girth (cm) was recorded from seedling at 5 leaf stage. The mean diameter (d) of the seedling was recorded from two sides using vernier caliper. The three observations were recorded for each genotype per replication. Stem girth: 2 (3.14) r or (3.14) d
th

; as d=2r

Trichomes: The presence and number of trichomes was observed on the abaxial surface central portion of the 5 leaf (from the base) taken at random. The following parameters were recorded: Number of trichomes: The number of trichomes was determined per 200 m linear length using Scanning Electron Microscope (SEM) as per standard protocol (Bozzola and Russell 1999). Imaging and counting was performed from three regions of single leaf per genotype. Trichome length: The trichome length was determined from the same leaf region selected for counting the trichomes of each genotype and was imaged using SEM as per standard protocol. Imaging and measurement was performed from three regions of single leaf per genotype. Trichome angle: The angle of trichome was also determined from the same leaf region selected for recording the trichome number and length in each genotype and was imaged using Scanning Electron Microscope (SEM) as per standard protocol. Imaging and measurement was performed from three regions of single leaf per genotype. Standard protocol for imaging leaf samples under SEM The leaf hairiness parameters, viz. trichome number, trichome length and angle of insertion of trichome were analyzed and imaged under Scanning Electron Microscope (SEM) at the Electron Microscopy and Nanoscience (EMN) Laboratory, College of Agriculture, PAU, Ludhiana as per standard protocol given by Bozzola and Russell (1999). Fresh leaves of each test genotype were collected and immediately immersed in individual vials containing 2.5 per cent glutaraldehyde solution for primary

29

fixation and kept overnight at a temperature of 4C. The leaf specimens were then washed thrice with distilled water. For secondary fixation, the specimens were immersed in 4 per cent osmium tetraoxide solution for a period of 2-4 hours at 4C. After post-fixation, the specimens were again washed thrice (each washing of 5 to 10 minute duration) using distilled water. Dehydration of the specimen discs was performed using different grades of ethanol (25, 50, 70, 95 and 100%) each for a period of 20 minutes whereas the final dehydration (with 100% ethanol) was performed for 30 minutes. The specimens were dried to critical point in CO2 at 5oC and mounted on aluminium stub using double-sided carbon tape. Each specimen leaf disc was mounted with its lower surface up allowing the lower epidermal surface of each leaf to be examined. The mounted leaf specimens were sputter-coated with a thin layer of gold using an automated sputter coater. Finally, the specimens were examined and imaged using Hitachi S-3400N Scanning Electron Microscope operated at an accelerating voltage of 15 KV using secondary electron detector. 3.6.1.3. Biochemical composition of maize seedlings in relation to infestation by shoot fly, A. naqvii: For studying the biochemical basis of resistance, seedling samples representing 5th leaf stage were used in each replication. A set of test genotypes was grown in screen house to obtain shoot fly infestation free plants for comparison of biochemical parameters (proteins, phenol, tannins, reducing sugars and free amino acids) between healthy and infested plants. The 5th leaf stage maize seedlings, collected from the field and screen house, were dried at 60oC in oven. The dried seedlings were cut and powdered in a Willey mill using a 0.5 pore size blade??? to obtain a fine powder for estimating different biochemical parameters. The leaf biochemical investigations were carried out in the Quality lab, Department of Plant Breeding and Genetics, PAU, Ludhiana for the following parameters: Estimation of total chlorophyll: Chlorophyll estimation from 5th leave of maize genotypes was performed using hand-held SPAD-502 Plus Chlorophyll Content Meter (Konica Minolta sensing, inc. Japan). The meter calculates a unit less Chlorophyll Content Index (CCI) value. Three non-overlapping measurements were taken in each replication per genotype and the mean of three CCI values was obtained. Each measurement consists of 10 observations. Moisture content: Ten seedlings at 5th leaf stage were cut at the base and immediately weighed to record
0

fresh weight of the seedlings. The seedlings were then kept in an oven at 60 C till observed for no further change in weight and then the weight of the oven dried material was recorded. The moisture percentage was calculated as follows: Moister content (%) = [(fresh weight dry weight) / (fresh weight)] 100.

30

Proteins: The protein content was calculated from nitrogen content of plant tissue. The estimation of nitrogen content was carried from plant samples each weighing 0.5 g by macro-kjeldahl method following the procedure described by the Association of Official Agricultural Chemists (AOAC 2000). Estimation of Phenols: The 100 mg of powdered plant tissue was dipped in 25 ml of 80 per cent aqueous methanol for 24 hours and then boiled on hot plate for 2 hours to evaporate methanol. The material was filtered and the volume was made to 15 ml with distilled H2O. This extract was used for estimation of phenols as per Swain and Hills (1959). (A) Reagents: i) Folin denis reagent: To prepare Folin Denis reagent 100 gm of sodium tungstate (Na Wo 2H O), 20
2 4 2

g phosphomolybdic acid, and 50 ml phosphoric acid were dissolved in 750 ml of water. The mixture was refluxed for 2 hours, and the final volume was made to 1litre by adding distilled water. ii) Saturated solution of sodium carbonate (35.0 g/100 ml H 2O): To prepare saturated sodium carbonate solution 35 g anhydrous sodium carbonate was dissolved in 100 ml of water at 70 80 C, and allowed to cool. (B) Procedure: Methanolic extract (0.1 ml) was made to 7.0 ml with 6.9 ml of distilled water. To this 0.5 ml of folin denis reagent was added and shaken thoroughly. After 3 minutes, 1 ml of saturated solution of sodium carbonate was added and volume was made to 10.0 ml with distilled water. The black colour formed was read after 1 hour at 725 nm against the blank in a spectrophotometer. The standard curve was prepared by using gallic acid in the range of 1080 g. Tannins: One gram of the powdered plant tissue was dipped in 25 ml of 80 per cent aqueous methanol for 24 hours and then boiled on hot plate for 2 hours to evaporate methanol. The material was then filtered and the volume was made to 25 ml with distilled water. This extract was used for estimation of tannins as per Sadasivam and Manickam (1992). (A) Reagents: i) Folin Denis reagent ii) Saturated solution of sodium carbonate (35.0 g/100 ml H2O) (B) Procedure: Methanolic extract (0.1 ml) was made to 7.5 ml with 7.4 ml of dH2O. To this 0.5 ml of folin denis reagent was added and shaken thoroughly. After 3 minutes, 1 ml of saturated solution of sodium carbonate was added and black colour formed was read at 760 nm against the blank. The amount of tannin was calculated from the standard curve using tannic acid (10-100 g) as standard.
0

31

Free amino acids: The 500 mg of the powdered plant tissue was dipped in 25 ml of 80 per cent aqueous methanol and incubated for 24 hours. The mixture was boiled on hot plate for 2 hours to evaporate methanol and the volume was made to 30 ml with distilled water. This extract was used for estimation of amino acids as per Elahi and Khan (1973). (A) Reagents: i) 0.5 M citrate buffer (a): Sol. A: 52.52 g of citric acid in 500 ml of distilled water Sol. B: 73.52 g of sodium citrate in 500 ml of distilled water 0.5 M citrate buffer was prepared by mixing 14.85 ml (Sol. A) and 35.15 ml (Sol. B ) ; and made to 100 ml with distilled water, pH was adjusted at 5.5. ii) 1.0 % ninhydrin (b): Dissolved 400 mg ninhydrin in 40 ml of 0.5 M citrate buffer just before use. iii) Pure glycerol (c) Prepared working ninhydrin solution just before use by mixing 1.0 % ninhydrin (b): pure glycerol (c): 0.5 M citrate buffer (a) in the ration of 5: 12: 2. (B) Procedure: To 0.5 ml of the appropriately diluted sample, 5.0 ml of ninhydrin solution was added. The contents were mixed well and 4.5 ml of distilled water was added to make the volume 10 ml. The mixture after boiling for 12 minutes was immediately cooled in ice water. The intensity of brown colour developed was read at 570 nm. The amount of protein in samples was calculated from the standard curve prepared from different concentrations of glycine (10-100 g). Reducing sugars: The 500 mg of the powdered plant tissue was dipped in 25 ml of 70 per cent aqueous ethanol and incubated for 24 hours. The mixture was boiled on hot plate for 2 hours to evaporate ethanol and the volume was made to 100 ml with distilled water. This extract was used for estimation of reducing sugars as per Nelson (1944). (A) Reagents: i) Solution 1 : 2.5 g of sodium carbonate , 2.5 g sodium potassium tartrate, 2.0 g of sodium bi carbonate and 20 gm of sodium sulphate was dissolved in distilled water and the final volume was made to 100 ml. ii) Solution 2: Two drops of sulphuric acid were added in 15 g of copper sulphate and the volume was made to 100 ml. Solution 1 and 2 were mixed in the ration of 25:1 just before the use. iii) Solution 3: A. 5 g of ammonium molybdate was dissolved in 4 ml of sulphuric acid and 90 ml of distilled water.

32

B. Mixed 600 mg of disodium arsenate in 5 ml of distilled water. Mixed A and B fractions to get the solution 3. (B) Procedure: 0.1 ml of ethanolic extract was used in estimation of reducing sugars from healthy plants and 0.5 ml from infested plants. The volume was made to 1.0 ml by adding required amount of distilled water. To this 1.0 ml of mixture of solution 1 and 2 was added and boiled for 20 minutes. The mixture was then cooled in ice water and 1 ml of solution 3 was added along with 3.5 ml of distilled water. The bluish green colour formed was read at 620 nm against the blank. The amount of reducing sugars was calculated from the standard curve using glucose (10-100 g) as standard. The amount of phenol, tannins, reducing sugars and free amino acids was calculated as: = (OD of test/ OD of standard) x concentration of standard x Volume made/ volume taken x 1/ weight of tissue. The information was also subjected to percentages of changes (decrease/ increase) in the infested and healthy plants and was calculated as per (Mahadeva and Nagaveni 2011):

% decrease/ increase =

(Values of infested plants - Values of healthy plants) 100 (Values of healthy plants)

3.7. Cultural and chemical control of shoot fly, Atherigona spp in spring maize. The experiments to develop IPM for shoot fly, Atherigona spp. in spring sown maize were conducted with different cultural and chemical control strategies at Research Farm, Department of Plant Breeding and Genetics, PAU, Ludhiana. The maize cultivar PMH 2 comparatively susceptible to shoot fly, was flat sown on 14th February in 2011 and twice on 14th February and 6th March in 2012. Each treatment plot was of 5 rows with 3 m row length. In 2 nd experiment in 2012 plots was of 4 rows with 3 meter row length. The experiments were laid in R.B.D. with 3 replications as per the eleven treatments replicated thrice during 2011 and 2012 as given below: T1 T2 T3 T4 T5 thiamethoxam 30 FS @ 4 g per kg seed one day before sowing thiamethoxam 30 FS @ 6 g per kg seed one day before sowing Single row of unprotected sorghum as trap crop Two rows of unprotected sorghum as trap crop Single row of protected sorghum as trap crop

33

T6 T7 T8 T9 T10 T11 -

Two rows of protected sorghum as trap crop First irrigation at emergence First irrigation after one week of emergence carbofuran 3 G @ 12.5 kg per ha at sowing (standard) imidacloprid 600 FS @ 6 ml per kg seed one day before sowing (standard) Untreated control The sorghum variety SL 44 was used for raising the trap crop and protected with imidacloprid 600

FS @ 5 ml per kg seed. The irrigation was given one day after emergence except in T 8. The crop raising practices except chemical control in untreated treatment plots were followed as per PAU recommendations (Anonymous 2010). Only the four selected treatments (T 6, T8, T10 & T11) based on earlier experimentation was evaluated against untreated control in 2nd experiment conducted during 2012. The different treatments were observed for any oviposition differentiation by shoot fly, Atherigona spp. by counting the number of eggs laid on the plant and nearby soil from 10 randomly selected plants in each treatment plot at 3, 6, 9, 15 and 18 DAG. In 2 nd experiment of 2012 eggs were counted at 5 and 10 DAG. The egg count was also recorded from all the tillers of 10 randomly selected plants of sorghum also. The leaf injury and deadhearts incidence due to shoot fly were recorded from 7, 14 and 21 days old maize seedlings on whole plot basis to calculate per cent incidence. In 2nd experiment in 2012, it was recorded from 5, 10 and 15 days old maize seedlings. The comparison was made for the damage and extent of losses caused by shoot fly in different treatments to identify most promising treatments for formulating effective IPM strategy for shoot fly in spring sown maize. The grain yield was recorded from inner three rows in each treatment plot and on whole plot in 2nd experiment of 2012, adjusted at 15 per cent moisture level and was converted to quintal per hectare for comparative estimation of effectiveness of different treatments. 3.8. Recording of weather data: The requisite meteorological data i.e. temperature, relative humidity, sunshine hours prevailing during these studies were obtained from Department of Agricultural Meteorology, PAU, Ludhiana. 3.9. Statistical analysis: The data on infestation and biological parameters of shoot fly, A. naqvii population counts of insect pests, morphological, biochemical parameters of plants and grain yield was analyzed using

ANOVA and the different treatment means were separated by least significant differences test (LSD) at

34

p=0.05 (Gomez and Gomez 1984). The correlation and regression coefficients for oviposition, biological parameters and damage by shoot fly with morphological, biochemical leaf characteristics of maize cultivars and weather factors with were also worked out.

35

Chapter-IV RESULTS AND DISCUSSION The results of investigation on the Incidence of insect pests and management of shoot fly, Atherigona spp. in spring sown maize have been presented and discussed below: 4.1. Population build up of insect pests of spring maize at Hoshiarpur: The experiment on population build up of insect pests of spring maize on two maize hybrids JH 3459 and PMH 2 at two nitrogen levels of 120 and 90 kg ha-1 under unprotected conditions was conducted at Hoshiarpur during spring 2011 and 2012 and the data for the same have been presented in Table 1-4 and Annexure I-VI. 4.1.1. Oviposition by shoot flies, Atherigona spp.: The number of eggs laid by shoot fly per plant were on par during different observations i.e. 7,14 and 21 days after germination (DAG) in both the test hybrids and at different nitrogen level during 2011 and 2012 (table 1). The interaction of hybrids and nitrogen levels was also non significant during both the test years. The egg count was comparatively higher during 2012 than that in 2011. The data on egg counts in different treatments revealed that mean egg count under two different levels of nitrogen irrespective of hybrids was same after 14 (0.31/ plant) and 21 (0.20/ plant) DAG and was numerically more under lower level of application of nitrogen (0.09/ plant) than that under higher level of application of nitrogen (0.08/ plant) at 7 DAG during 2011 (Table 1) However, it was numerically high in higher level of nitrogen (120 kg/ ha) application than that the lower level of nitrogen to the crop when observed at 7, 14 and 21 DAG during 2012. Mote and Ramshe (1987) also reported that nitrogen treated sorghum plants were more preferred by shoot fly, A. soccata for egg laying (1.11/ plant) than control plants (0.95/ plant). Chamarthi (2008) also reported that preference by shoot fly, A. soccata for egg laying was more on vigorous plants of sorghum. The non significant difference in the present studies might be due to differences in maize sowing seasons, host plant, predominant species of shoot fly. Though the mean egg counts made from randomly selected plants of both the hybrids i.e. JH 3459 and PMH 2, irrespective of nitrogen levels, did not differ significantly, yet numerically more numbers of eggs were laid in hybrid PMH 2 than JH 3459 during all the observations in both the years i.e. 2011 and 2012 except at 21 DAG in 2012 when it was same (2.63 eggs/ plant) on both the hybrids. Similar to present studies, the non discrimination of plants of various maize germplasms for oviposition by shoot flies were reported earlier also by Sarup and Panwar (1986), Panwar and Sarup (1988); and Rao and Panwar (1996a).

36

Table 1. The oviposition (eggs/ plant) by shoot fly, Atherigona spp. in different treatments during spring 2011 and 2012 at Hoshiarpur. Mean number of eggs per plant at 2011 7 DAG Nitrogen Hybrid JH PMH 2 3459 0.50 0.10 (1.02) (1.05) 0.08 0.10 (1.04) (1.05) 0.06 0.10 (1.03) (1.05)
Nitrogen (A) = NS, Hybrid (B) = NS, A x B = NS

14 DAG Mean 0.08 (1.04) 0.09 (1.04) Hybrid JH PMH 3459 2 0.27 0.35 (1.13) (1.16) 0.30 0.32 (1.14) (1.15) 0.29 0.34 (1.13) (1.16)
Nitrogen (A) = NS, Hybrid (B) = NS, A x B = NS

21 DAG Mean 0.31 (1.14) 0.31 (1.14) Hybrid JH PMH 3459 2 0.20 0.20 (1.09) (1.09) 0.18 0.22 (1.08) (1.11) 0.19 0.21 (1.09) (1.10)
Nitrogen (A) = NS, Hybrid (B) = NS, A x B = NS

Mean 0.20 (1.09) 0.20 (1.09)

N120 N90 Mean CD (p=0.05) 2012

7 DAG Nitrogen Hybrid JH PMH 2 3459 0.40 0.52 (1.18) (1.22) 0.45 0.35 (1.20) (1.16) 0.43 0.44 (1.19) (1.19)
Nitrogen (A) = NS, Hybrid (B) = NS, A x B = NS

14 DAG Mean 0.46 (1.20) 0.40 (1.18) Hybrid JH PMH 3459 2 0.40 0.50 (1.18) (1.22) 0.35 0.43 (1.16) (1.19) 0.38 0.46 (1.17) (1.21)
Nitrogen (A) = NS, Hybrid (B) = NS, A x B = NS

21 DAG Mean 0.45 (1.20) 0.39 (1.18) Hybrid PMH JH 3459 2 0.28 0.32 (1.13) (1.15) 0.25 0.20 (1.11) (1.09) 0.26 0.26 (1.12) (1.12)
Nitrogen (A) = NS, Hybrid (B) = NS, A x B = NS

Mean 0.30 (1.14) 0.22 (1.10)

N120 N90 Mean CD (p=0.05)

DAG: Days after germination 4.1.2. Incidence of shoot fly, Atherigona spp.: The mean leaf injury and deadhearts incidence, irrespective of hybrids, among test nitrogen levels was non significant in all the observations during 2011 and 2012 except for deadhearts incidence at 21 and 28 DAG during 2011 where it differed significantly (table 2 and 3). The mean leaf injury incidence, irrespective of nitrogen levels, differed significantly among test hybrids at 14 DAG during 2011; and at 14 and 21 DAG during 2012. The mean deadhearts incidence, irrespective of nitrogen levels differed among test hybrids at 21 and 28 DAG during 2011; and at 14, 21 and 28 DAG in 2012. However, the interaction of test hybrids with test nitrogen levels for mean leaf injury and deadhearts incidence were non significant during both the test years. The leaf injury incidence caused by shoot fly was comparatively more during 2012 (ranging from 5.03- 39.26 %) than that in 2011 (ranging from 1.49-24.75 %). At 7

37

DAG, it was numerically more (2.25 and 7.33 %) at higher nitrogen level than that at lower nitrogen level (1.99 and 5.70 %) during the respective years. Similarly, the leaf injury incidence among hybrids did not differ significantly during both the test years at this observation and it was numerically more in PMH 2 (2.49 and 7.21 %) than that in JH 3459 (1.75 and 5.83 %). At 14 DAG also leaf injury, irrespective of hybrids was numerically more at higher nitrogen level than that in lower nitrogen level during 2011, however, it was lower at this nitrogen level than the lower one during 2012. But, irrespective of nitrogen level, it was significantly was more in hybrid PMH 2 (17.17 and 28.87 %) than that in JH 3459 (13.02 and 14.68 %) during 2011 and 2012, respectively. At 21 DAG again it was less at higher nitrogen level (23.14 and 31.97 %) than that at lower one (23.19 and 33.04 %) during both the test years. The mean leaf injury, irrespective of nitrogen levels differed significantly among hybrids during 2012 only being more in PMH 2 (39.09 %) than that in JH 3459 (25.91%). The non significant interaction of hybrids and nitrogen level for leaf injury incidence also implies that susceptible hybrid PMH 2 had higher incidence than less susceptible JH 3459 at both the nitrogen levels at different observations during both the test years. As in case of leaf injury incidence, the mean deadhearts incidence was also comparatively more during 2012 (ranging from 5.40-26.48 %) than that in 2011 (ranging from 2.98-18.31 %). At 14 DAG, the mean deadhearts incidence, irrespective of hybrids being on par, was numerically more (4.50 and 8.73 %) at higher nitrogen level than that at lower nitrogen level (3.75 and 8.33 %) during the respective years. However, the same , irrespective of nitrogen levels was significantly more in PMH 2 (11.33 %) than JH 3459 (5.73 %) during 2012 only. At 21 DAG, irrespective of hybrids, significantly more deadhearts were formed at higher nitrogen level (12.90 %) than the lower one (11.01 %) during 2011 only whereas, the hybrid PMH 2 (14.41 and 24.92 %) irrespective of nitrogen levels, had more deadhearts incidence than JH 3459 (9.50 and 17.06 %) during 2011 and 2012, respectively. At 28 DAG also, during 2011 irrespective of hybrids, significantly more deadhearts were formed at nitrogen level of 120 kg ha -1 (14.65 %) than that in 90 kg ha-1 (11.99 %) the mean deadhearts incidence, irrespective of nitrogen levels was significantly more in PMH 2 (16.40 and 25.58 %) than that in JH 3459 (10.24 and 16.40 %) during 2011 and 2012. This infestation level thus, confirmed the key pest status of shoot fly in spring sown maize in the Punjab. The non significant interaction of hybrids and nitrogen levels for deadhearts incidence again implies that hybrid PMH 2 had higher incidence than JH 3459 at both the nitrogen levels at different observations during both the test years. However, the numerical increase in deadhearts under nitrogen level of 120 kg ha-1 than that in 90 kg ha-1 was more pronounced in susceptible hybrid PMH 2 with 3.82 per cent increase than that of 1.50 per cent in moderately tolerant hybrid JH 3459 during 2011. In 2012, deadhearts incidence differed non significantly at different nitrogen levels and had a decrease of 0.34 per cent in JH 3459 and slight increase of 1.81 per cent in hybrid PMH 2 at higher nitrogen level. This might

38

be due to higher incidence of shoot fly during 2012 as compared to 2011. However, reduction in deadhearts at higher dose of nitrogen in sorghum due to A. soccata incidence, might be due to vigorous fast growth of plants was reported by Mote and Ramshe (1987). They further reported that in susceptible variety CSH-8R there was increase in incidence of shoot fly at 30 and 60 kg h-1, while it reduced at 90 kg h-1 as compared to control; and in moderately tolerant variety SPV-86 and M.35-1 its incidence was reduced with increase in nitrogen level which is in line with the present findings that deadhearts incidence reduced at higher nitrogen level in least susceptible hybrid JH 3459 during 2012 but marginally increased during 2011. Sharma and Singh (1975) also reported that application of nitrogenous fertilizer in some of the germplasms promoted growth and vigour in the plants thus helping in recovery of shoot fly infestation in plants. Table 2. The shoot fly, Atherigona spp. incidence in different treatments during spring 2011 at Hoshiarpur. Mean leaf injury and deadhearts at 7 DAG LI (%) Hybrid JH PMH 2 3459 2.01 2.48 1.49 2.50 1.75 2.49 14 DAG LI (%) Hybrid JH PMH 3459 2 13.53 17.54 12.51 16.81 13.02 17.17 DH (%) Hybrid JH PMH Mean 3459 2 3.50 5.50 4.50 2.98 4.51 3.75 3.24 5.01

Nitrogen

Mean 2.25 1.99

Mean 15.53 14.66

N120 N90 Mean

CD (p=0.05)

Nitrogen (A) = NS Hybrid (B) = NS A x B = NS

Nitrogen (A) = NS Hybrid (B) = 2.81 A x B = NS

Nitrogen (A) = NS Hybrid (B) = NS A x B = NS

21 DAG Nitrogen LI (%) Hybrid JH PMH 2 3459 21.54 24.75 22.54 23.83 22.04 24.29 DH (%) Hybrid JH PMH Mean 3459 2 10.01 15.79 12.90 8.99 13.04 11.01 9.50 14.41

28 DAG DH (%) Hybrid JH PMH Mean 3459 2 10.99 18.31 14.65 9.49 14.49 11.99 10.24 16.40

Mean 23.14 23.19

N120 N90 Mean

CD (p=0.05)

Nitrogen (A) = NS Hybrid (B) = NS A x B = NS

Nitrogen (A) = 1.52 Hybrid (B) = 1.52 A x B = NS

Nitrogen (A) = 1.78 Hybrid (B) = 1.78 A x B = NS

LI: Leaf injury, DH: Deadhearts; DAG: Days after germination

Table 3. The shoot fly, Atherigona spp. incidence in different treatments during spring 2012 at Hoshiarpur.

39

Mean leaf injury and deadhearts at 7 DAG Nitrogen LI (%) Hybrid JH PMH 2 3459 6.63 8.04 5.03 6.38 5.83 7.21 LI (%) Hybrid JH PMH 3459 2 14.77 26.41 14.59 31.33 14.68 28.87 14 DAG DH (%) Hybrid JH PMH Mean 3459 2 5.40 12.05 8.73 6.05 10.61 8.33 5.73 11.33

Mean 7.33 5.70

Mean 20.59 22.96

N120 N90 Mean CD (p=0.05)

Nitrogen (A) = NS Hybrid (B) = NS A x B = NS LI (%) Hybrid JH PMH 2 3459 25.01 38.93 26.82 39.26 25.91 39.09

Nitrogen (A) = NS Hybrid (B) = 7.29 A x B = NS 21 DAG DH (%) Hybrid JH PMH Mean 3459 2 17.12 25.80 21.46 17.01 24.05 20.53 17.06 24.92

Nitrogen (A) = NS Hybrid (B) = 3.04 A x B = NS 28 DAG DH (%) Hybrid JH PMH Mean 3459 2 17.39 26.48 21.94 17.73 24.67 21.20 17.56 25.58

Nitrogen

Mean 31.97 33.04

N120 N90 Mean CD (p=0.05)

Nitrogen (A) = NS Hybrid (B) = 4.17 A x B = NS LI: Leaf injury, DH: Deadhearts; DAG: Days after germination

Nitrogen (A) = NS Hybrid (B) = 7.36 A x B = NS

Nitrogen (A) = NS Hybrid (B) = 4.05 A x B = NS

The grain yield (q/ ha) differed significantly among the genotypes as well as under different nitrogen levels during both the years (table 4), but their interactions were non significant during both the years. It was significantly more in hybrid PMH 2 (46.11 and 42.29 q/ ha) than that in JH 3459 (42.45 and 38.28 q/ ha) in 2011 and 2012, respectively. The yield was also significantly higher at nitrogen level of 120 kg ha-1 (46.29 and 41.58 q/ ha) than that of 90 kg ha-1 (42.26 and 38.28 q/ ha) during both the test years, respectively. The differences in yield, even at varied infestation levels of insect pests might be due to the inherent capacity of the tested genotypes with positive response to more nitrogen application.

40

Table 4. The grain yield (q/ha) in different treatments during spring 2011 and 2012 at Hoshiarpur Mean grain yield (q/ha) Nitrogen 2011 Hybrid JH 3459 PMH 2 44.27 48.32 40.62 43.89 42.45 46.11 Mean 46.29 42.26 2012 Hybrid JH 3459 PMH 2 39.45 43.71 37.11 40.88 38.28 42.29 Mean 41.58 38.99

N120 N90 Mean CD (p=0.05)

Nitrogen (A) = 3.11 Hybrid (B) = 3.11 A x B = NS

Nitrogen (A) = 2.26 Hybrid (B) = 2.26 A x B = NS

4.1.3. Population build up of other insect pests and beneficials at Hoshiarpur: The population counts per ten plants and damage on whole plot basis of different insect pests recorded at weekly intervals was presented in Annexure I-VI. The comparative lower infestation/ population levels of different insect pests showed that spring sown maize apart from its key pest i.e. shoot fly in general is comparatively free from the attack of other insect pests of maize. The Chilo partellus (Swinhoe) incidence was relatively more during 2011 and it differed significantly at different nitrogen levels and among hybrids in some observations during 2012 and 2011, respectively (Annexure I). However, the interaction of these factors for C. partellus incidence was non significant at all the observations during both the test years. Moreover, no deadhearts formation due to C. partellus was observed during the study period, indicated that its infestation did not occur in a serious form on February sown spring maize crop. The leaf injury incidence due to C. partellus showed that the pest appeared during first week of April, it increased progressively and minor damage was observed up to mid May during both the years. The highest incidence was observed during last week of April (15.75-21.36 %) and first week of May (10.60- 12.73 %) during 2011 and 2012, respectively. Though, irrespective of hybrids, it was numerically more at higher nitrogen level during this period in both the test years, yet it differed significantly during 4th week of April and 2nd week of May during 2012 only. Irrespective of the hybrids, the incidence was more (12.55 and 4.14 %) at higher nitrogen level than the lower one (9.48 and 2.36 %) at these observations. Similarly, Panwar et al (1997 &1999b) also reported that the nitrogen fertilizer application enhances the damage of C. partellus on maize in kharif season also. The incidence, irrespective of nitrogen levels, was significantly lower in hybrid JH 3459 (5.92, 8.17, 14.76 and 15.00 %) than PMH 2 (7.83, 10.33, 17.20 and 19.29 %) when observed on 2nd, 3rd & 4th week of April; and 1st week of May, respectively during 2011 only. Similar to these observations, Jindal and Hari (2008b) also reported that hybrid JH 3459 have some tolerance to C. partellus in Kharif season. The incidence of C.

41

partellus in end April on spring sown maize did not manifest any economic damage as observed by slight leaf injury only on mature leaves in the present studies. The incidence of other lepidopteran insect pests viz. corn earworm (Helicoverpa armigera) and army worm (Mythimna separata) was sporadic and their incidence differed non significantly at different nitrogen levels and among hybrids during 2011 and 2012 (Annexure II). Similarly, the interaction of these factors for incidence of these pests was non significant during both the test years. The incidence of corn earworm on silks was very low and observed during 4th and 2nd 3rd week May during 2011 and 2012, respectively. At different observation times it varied from 1.47 to 7.58 per cent during spring 2011; and from 1.28 to 4.01 per cent during spring 2012. Similarly, lower incidence of army worm was observed during April and May during both the test years. It was maximum in 1 st week of May (3.31-5.75 %) and last week of April (3.25-5.91 %) during 2011 and 2012, respectively. Thus, these insect pests did not cause any economic damage in the present studies, but corn earworm being a polyphagous pest with a tendency to feed on developing grains and may a pose threat to end February to early March sown spring maize. Among sucking insect pests, minor incidence of pyrilla (Pyrilla perpusilla) and newly recorded small brown planthopper (Laodelphax striatellus) were observed during both the test years (Annexure III). Both the sucking insect pests differed non significantly at different nitrogen levels and among hybrids (except for few observations) during 2011 and 2012. The mean population of pyrilla was also low and its maximum population counts were observed in last week of April (ranging from 0.15 to 0.20/ plant) and 1st week of May (ranging from 0.17 to 0.25/ plant) in the respective tests years. However, irrespective of nitrogen levels, its incidence in 1st week of May, 2011 was significantly more on hybrid JH 3459 (0.15/ plant) than that on PMH 2 (0.09/ plant). The occurrence of small brown planthopper, L. striatellus is being reported on spring maize for the first time in the Punjab, but its prevalence at this location was low as compared to Ludhiana during 2011. The mean population counts of L. striatellus were highest during 3rd week of April during 2011 (ranging from 0.97 to 1.75/ plant) and 2012 (ranging from 0.52 to 0.85/ plant). However, irrespective of nitrogen levels, the counts were significantly more in hybrid PMH 2 (0.87 and 0.80/ plant) as compared to JH 3459 (0.60 and 0.55/ plant) during 2nd and 3rd week of 2011 and 2012, respectively. Thus, again revealing that population of small brown planthopper had not been influenced by interactions of genotypes and nitrogen levels. The low incidence of these insect pests observed did not cause any economic damage to spring sown maize. The incidence of grasshopper spp. was observed during May in both the test years. (Annexure IV). The nymphal counts of grasshopper and per cent plant damage differed non significantly in different treatments during both the test years except in 3rd week of May for nitrogen levels during 2011. The mean nymphal counts per plant and per cent plant damage was significantly more (0.74 and 5.71) at higher

42

nitrogen level than that of lower one (0.46 and 4.60). The nymphal counts and plant damage varied from 0.25 to 0.80 & 0.10 to 0.20; and from 2.52 to 9.16 & 0.25 to 4.23 per cent during 2011 and 2012, respectively. The grasshopper population and its damage were found to be very low and did not cause economic damage, as the crop was already reaching near maturity and no damage was observed on developing cobs. The population of beneficial arthropods i.e. coccinellids and spiders was observed during the month of April and May (Annexure V and VI). The population of coccinellids i.e. Coccinella septempunctata (Linnaeus) appeared during the April and it declined after first week of May during both the test years, however it differed significantly among nitrogen levels and hybrids when observed on April 15, 22 & 29; and May 6, 2011 but differed non significantly in all the observations during 2012. It was higher (ranging from 0.10 to 0.68/ plant) in 2011 than that of (ranging from 0.03 to 0.22/ plant), which might be due to comparatively higher population of small brown planthopper prevalent at that time. The mean population, irrespective of hybrids, was significantly more (0.50 and 0.31/ plant) at higher nitrogen level than that in lower one (0.36 and 0.23/ plant) in 3rd week of April and 1st week of May, respectively during 2011 only. Similarly, irrespective of nitrogen levels, it was more on hybrid PMH 2 (0.29, 0.59, 0.40 & 0.36/ plant) than JH 3459 (0.12, 0.27, 0.17 & 0.17/ plant) in 2nd, 3rd & 4th week of April and 1st week of May, respectively. However, the interactions of nitrogen levels with hybrids for population counts of coccinellids was non significant. During 2011, the higher counts in different treatments were corresponding to the higher counts of small brown planthopper in those treatments. Similarly, the counts of spiders also differed non significantly among different treatments during both the tests years. The spider counts ranged from 0.08 0.20 in May, 2011 and 0.08 0.23 in April - May, 2012 in the present studies, emphasizes their minor importance in spring sown maize. To sum up the findings of the present studies, the sporadic incidence of other insect pests and beneficials was observed on spring sown maize at Hoshiarpur and further it needs more emphasis for the management of its key pest i.e. shoot fly using integrated approaches.

43

4.2.

Estimation of losses caused by insect pests in spring sown maize: The experiment on insect pests incidence and estimation of losses caused by them in spring sown

maize was conducted at Punjab Agricultural University Ludhiana during spring 2011 and 2012. These studies were conducted on hybrid JH 3459 and PMH 2 sown on end January, mid and end February under protected/ unprotected conditions against shoot fly. The data of these experiments have been presented in Table 5-15 and Annexure VII-XII. 4.2.1. Oviposition by shoot fly, Atherigona spp. The number of eggs laid by shoot fly per plant was recorded at 3, 7, 10, 15 DAG during 2011 and 7, 12, 17 DAG in 2012 (table 5 and 6). The egg count differed non significantly at different observations in both the test hybrids under protected as well as unprotected conditions during 2011, whereas the differences in egg count per plant among hybrids irrespective of crop protection condition differ significantly in Set 1 (end January sown crop); and among crop protection conditions and among hybrids in Set 3 (end February sown crop) at 3 DAG and 17 DAG , respectively. However, the interaction between hybrids and crop protection conditions for egg laying were non significant during all the observations in both the test years. The egg count in end January sown crop (Set 1) was observed to be 0.02 per plant at 3 DAG which reached to maximum counts of 0.19 per plant at 10 DAG in 2011 (fig 1), whereas during 2012 (fig 2) the egg count was observed to be 0.06 per plant at 7 DAG which reached to maximum level of 0.15 per plant at 17 DAG. At maximum egg laying period also the differences for egg count among different hybrids as well as protection conditions and their interaction were non significant during both the test years. During 2011 at 10 DAG, the mean egg count, irrespective of crop protection conditions was 0.19 and 0.20 per plant in least susceptible JH 3459 and susceptible PMH 2, while they same, irrespective of hybrids was 0.20 and 0.17-0.22 per plant under protected and unprotected conditions, respectively. In 2012, at 17 DAG the mean egg count, irrespective of crop protection conditions was 0.16 and 0.15 per plant in least susceptible JH 3459 and susceptible PMH 2, however, the same irrespective hybrids was numerically lower under protected conditions (0.12/ plant) than that under unprotected conditions (0.17-0.19/ plant). At 7 DAG in 2012, the susceptible hybrid PMH 2 (0.10/ plant) received significantly more eggs than least susceptible hybrid JH 3459 (0.03/ plant). The mid February sown crop (Set 2) had higher egg counts than end January sown crop (set 1) during both the test years (fig 1 and 2), but it differed non significantly in all the observations among different hybrids and crop protection conditions; and the interaction among these factors was also non significant. Numerically, the maximum mean egg counts in different treatments in set 2 were observed at 10 and 7 DAG during 2011 and 2012, respectively (table 5 and 6). The egg counts increased progressively from 3 DAG (0.01/ plant) to 10 DAG (0.27/ plant) and then decreased to 0.12 eggs per plant at 15 DAG (fig 1). At maximum egg laying period at 10 DAG in 2011, the mean egg counts,

44

irrespective of crop protection conditions were 0.27 and 0.28 per plant in least susceptible JH 3459 and susceptible PMH 2, while the same, irrespective of hybrids, were 0.27 and 0.26-0.31 per plant in protected and unprotected conditions, respectively. In 2012, the egg counts were higher than that in 2011 which decreased from 0.79 per plant at 7 DAG, to 0.26 per plant at 12 DAG and then to a minimum of 0.12 per plant at 17 DAG (fig 2). At 7 DAG, JH 3459, irrespective of crop protection conditions, received numerically less eggs (0.67/ plant) than that in PMH 2 (0.90/ plants) and the same, irrespective of hybrids were also lower in protected (0.72/ plant) than that in unprotected condition (0.76 - 0.87 per plant). The non significant interaction between hybrids and protection conditions indicated that these factors did not influence the egg laying behavior of shoot fly in present study. The end February sown crop (set 3) had higher egg count than end January (set 1) and mid February sown crops (set 2) during both the years except at 15 DAG in set 1 during 2011 and at 15 and 7 DAG in set 2 during 2011 and 2012, respectively (Fig.1 and 2). The egg counts differed non significantly among hybrids and crop protection conditions at all the observations during both the test years, except at 12 and 17 DAG in 2012. The counts, irrespective of hybrids, differed among crop protection conditions at 12 DAG and the same, irrespective of plant protection conditions, among hybrids at 17 DAG. However, the interaction among these factors for egg counts was non significant in all the observations during both the test years. Numerically, the egg counts were maximum in all the treatments at 10 and 12 DAG during 2011 and 012, respectively (table 5 and 6). During 2011, the mean egg counts irrespective of crop protection conditions and hybrids increased progressively from 3 DAG (0.16/ plant) to 10 DAG (0.52/ plant) and then decreased to 0.02 eggs per plant at 15 DAG (fig 1). At 10 DAG i.e. at maximum egg count period, the mean egg counts, irrespective of crop protection conditions were numerically high in hybrid PMH 2 (0.54/ plant) than that in JH 3459 (0.50/ plant), while the same irrespective of hybrids were numerically lower in protected conditions (0.34/ plant) than unprotected conditions (0.56-0.68/ plant). In 2012, the egg count was higher than that during 2011, and it increased from 0.48 per plant at 7 DAG to 0.65 per plant at 12 DAG and then decreased to 0.27 per plant at 17 DAG (fig. 2). At 12 DAG, the mean egg counts, irrespective of crop protection conditions being on par with each other , were 0.67 per plant and 0.64 per plant in JH 3459 and PMH 2, while the same, irrespective of hybrids were significantly lower in protected conditions (0.46/ plant) than that in unprotected conditions (0.72 0.78 per plant). However, at 17 DAG the egg count differed significantly in hybrids as well as under protection/ unprotection conditions. Table 5. Oviposition by shoot fly, Atherigona spp in crops sown at different times during 2011 at Ludhiana Egg count per plants 2011: Set 1 3 DAG 7 DAG Condition Mean Condition

Hybrid

Mean

45

JH 3459 PMH 2 Mean CD (p=0.05) Hybrid JH 3459 PMH 2 Mean CD (p=0.05)

Hybrid JH 3459 PMH 2 Mean CD (p=0.05) Hybrid JH 3459 PMH 2 Mean CD (p=0.05)

FP OT UT FP OT UT 0.02 0.02 0.02 0.02 0.07 0.04 0.07 0.06 (1.01) (1.01) (1.01) (1.01) (1.03) (1.02) (1.03) (1.03) 0.02 0.00 0.05 0.02 0.04 0.00 0.11 0.05 (1.01) (1.00) (1.02) (1.01) (1.02) (1.00) (1.05) (1.03) 0.02 0.01 0.04 0.06 0.02 0.09 (1.01) (1.00) (1.01) (1.03) (1.01) (1.04) Hybrid (A) = NS, Condition (B) = NS Hybrid (A) = NS, Condition (B) = NS A x B = NS A x B = NS 10 DAG 15 DAG Condition Condition Mean Mean FP OT UT FP OT UT 0.20 0.18 0.20 0.19 0.05 0.07 0.05 0.05 (1.09) (1.08) (1.09) (1.09) (1.02) (1.03) (1.02) (1.03) 0.20 0.15 0.25 0.20 0.04 0.05 0.09 0.06 (1.09) (1.07) (1.11) (1.09) (1.02) (1.02) (1.04) (1.03) 0.20 0.17 0.22 0.05 0.06 0.07 (1.09) (1.08) (1.10) (1.02) (1.03) (1.03) Hybrid (A) = NS, Condition (B) = NS Hybrid (A) = NS, Condition (B) = NS A x B = NS A x B = NS Egg count per plants 2011: Set 2 3 DAG 7 DAG Condition Condition Mean Mean FP OT UT FP OT UT 0.03 0.00 0.03 0.02 0.11 0.20 0.13 0.15 (1.01) (1.00) (1.01) (1.01) (1.05) (1.10) (1.06) (1.07) 0.04 0.00 0.00 0.00 0.14 0.13 0.11 0.13 (1.00) (1.00) (1.00) (1.00) (1.07) (1.06) (1.05) (1.06) 0.01 0.00 0.01 0.13 0.17 0.12 (1.01) (1.00) (1.01) (1.06) (1.08) (1.06) Hybrid (A) = NS, Condition (B) = NS Hybrid (A) = NS, Condition (B) = NS A x B = NS A x B = NS 10 DAG 15 DAG Condition Condition Mean Mean FP OT UT FP OT UT 0.27 0.25 0.31 0.27 0.09 0.13 0.18 0.13 (1.13) (1.12) (1.14) (1.13) (1.04) (1.06) (1.08) (1.06) 0.27 0.27 0.31 0.28 0.11 0.09 0.09(1.04) 0.10 (1.13) (1.12) (1.14) (1.13) (1.05) (1.04) (1.05) 0.27 0.26 0.31 0.10 0.11 0.13 (1.13) (1.12) (1.14) (1.05) (1.05) (1.06) Hybrid (A) = NS, Condition (B) = NS Hybrid (A) = NS, Condition (B) = NS A x B = NS A x B = NS

46

Table 5. contd Egg count per plants 2011: Set 3 Hybrid JH 3459 PMH 2 Mean CD (p=0.05) Hybrid JH 3459 PMH 2 Mean CD (p=0.05) 3 DAG Condition Mean FP OT UT 0.07 0.16 0.27 0.16 (1.03) (1.07) (1.12) (1.07) 0.02 0.18 0.29 0.16 (1.01) (1.08) (1.13) (1.08) 0.05 0.17 0.28 (1.02) (1.08) (1.13) Hybrid (A) = NS, Condition (B) = NS A x B = NS 10 DAG Condition Mean FP OT UT 0.20 0.71 0.60 0.50 (1.09) (1.30) (1.26) (1.22) 0.47 0.64 0.51 0.54 (1.21) (1.28) (1.23) (1.24) 0.34 0.68 0.56 (1.15) (1.29) (1.24) Hybrid (A) = NS, Condition (B) = NS A x B = NS 7 DAG Condition Mean FP OT UT 0.11 0.38 0.22 0.24 (1.05) (1.17) (1.10) (1.11) 0.27 0.18 0.35 0.27 (1.12) (1.08) (1.16) (1.12) 0.19 0.28 0.29 (1.09) (1.13) (1.13) Hybrid (A) = NS, Condition (B) = NS A x B = NS 15 DAG Condition Mean FP OT UT 0.00 0.04 0.02 0.02 (1.00) (1.02) (1.01) (1.01) 0.00 0.02 0.09 0.03 (1.00) (1.01) (1.04) (1.02) 0.00 0.03 0.06 (1.00) (1.02) (1.03) Hybrid (A) = NS, Condition (B) = NS A x B = NS

Set 1: end January sown crop, Set 2: mid February sown crop, Set 3: end February sown crop FP: Protection from shoot fly and other insect pests, OT: Protection from insect pests other than shoot fly, UT: No protection for insect pests

47

Table 6. Oviposition by shoot fly, Atherigona spp in crops sown at different times during 2012 at Ludhiana
Egg count per plants 2012: Set 1 Hybrid JH 3459 PMH 2 Mean CD (p=0.05) 7 DAG Condition FP OT UT 0.00 0.09 0.00 (1.00) (1.04) (1.00) 0.07 0.11 0.11 (1.03) (1.05) (1.05) 0.03 0.10 0.06 (1.02) (1.05) (1.03) Hybrid (A) = 0.03, Condition (B) = NS A x B = NS 7 DAG Hybrid JH 3459 PMH 2 Mean CD (p=0.05) Hybrid JH 3459 PMH 2 Mean CD (p=0.05) Condition FP OT UT 0.71 0.50 0.80 (1.30) (1.22) (1.34) 0.73 1.02 0.93 (1.32) (1.42) (1.39) 0.72 0.76 0.87 (1.31) (1.32) (1.37) Hybrid (A) = NS, Condition (B) = NS A x B = NS 7 DAG FP OT UT 0.26 0.44 0.38 (1.12) (1.20) (1.17) 0.42 0.65 0.71 (1.19) (1.28) (1.30) 0.35 0.55 0.55 (1.16) (1.24) (1.24) Hybrid (A) = NS, Condition (B) = NS A x B = NS Mean 0.67 (1.29) 0.90 (1.38) 12 DAG Condition FP OT UT 0.02 0.09 0.07 (1.03) (1.01) (1.04) 0.05 0.09 0.09 (1.02) (1.04) (1.04) 0.04 0.10 0.08 (1.04) (1.02) (1.04) Hybrid (A) = NS, Condition (B) = NS A x B = NS Egg count per plants 2012: Set 2 12 DAG Condition Mean FP OT UT 0.33 0.18 0.22 0.15 (1.07) (1.15) (1.08) (1.13) 0.29 0.29 0.29 (1.13) 0.29 (1.13) (1.14) (1.14) 0.31 0.22 0.23 (1.10) (1.14) (1.11) Hybrid (A) = NS, Condition (B) = NS, A x B = NS Egg count per plants 2012: Set 3 Mean 0.36 (1.17) 0.59 (1.26) FP 0.49 (1.22) 0.42 (1.29) 0.46 (1.21) Hybrid (A) = NS 12 DAG OT UT 0.71 0.80 (1.34) (1.31) 0.74 0.75 (1.32) (1.32) 0.72 0.78 (1.33) (1.31) NS, Condition (B) = 0.09, Mean 0.67 (1.29) 0.64 (1.28) 17 DAG FP OT UT Mean 0.02 0.30 0.22 0.33 (1.15) (1.01) (1.14) (1.10) 0.13 0.47 0.35 (1.16) 0.32 (1.06) (1.21) (1.15) 0.08 0.38 0.34 (1.16) (1.04) (1.17) Hybrid (A) = 0.03, Condition (B) = 0.04 A x B = NS 17 DAG Condition FP OT UT 0.11 0.20 0.18 (1.09) (1.05) (1.09) 0.13 0.18 0.15 (1.07) (1.06) (1.08) 0.12 0.19 0.17 (1.08) (1.06) (1.09) Hybrid (A) = NS, Condition (B) = NS A x B = NS 17 DAG Condition FP OT UT 0.15 0.11 0.11 (1.07) (1.05) (1.05) 0.16 0.15 0.09 (1.07) (1.07) (1.04) 0.16 0.13 0.10 (1.07) (1.06) (1.05) Hybrid (A) = NS, Condition (B) = NS A x B = NS Mean 0.12 (1.06) 0.13 (1.06)

Mean 0.03 (1.01) 0.10 (1.05)

Mean 0.06 (1.02) 0.08 (1.04)

Mean 0.16 (1.08) 0.15 (1.07)

AxB=

Set 1: end January sown crop, Set 2: mid February sown crop, Set 3: end February sown crop FP: Protection from shoot fly and other insect pests, OT: Protection from insect pests other than shoot fly, UT: No protection for insect pests

48

Figure 1 & 2

49

The mean egg counts, irrespective of crop protection conditions, were lower in JH 3459 (0.22/ plant) than PMH 2 (0.32/plant) and similar to egg count at 12 DAG, the same, irrespective of hybrids were significantly lower in protected conditions (0.08/ plant) than that in unprotected conditions (0.34 0.38/ plant). The non significant interaction between hybrids and crop protection conditions indicated that the similar reduction occurred in egg counts in both the hybrids under protected and unprotected conditions. The maximum egg counts were observed at 10 DAG irrespective of sowing date in 2011 and at 17, 7 and 12 DAG crop sown in end January, mid February and end February, respectively during 2012 (fig. 1 and 2) was in agreement with findings of Sarup and Panwar (1987) who also reported that regardless of the site of oviposition, maximum number of eggs were laid on 9 to 11 days old plants (3.08-3.22 eggs/ plant). The non consistent correlation between stages of the crop sown at different times and oviposition, in the present studies indicated the egg laying preference of shoot fly towards the stage of crop was up to 10 DAG and 12 DAG in 2011 and 2012, respectively. The period of maximum egg laying depends upon the abundance of shoot flies and the prevailing environmental factors at a particular time and place, for which continuous monitoring is required to strengthen the management of this key pest in spring sown maize in the Punjab. However, the non significant differences of egg counts in hybrids under different crop protection conditions except for few observations in 2012 observed in the present studies are in line with the findings of Rao and Panwar (1992, 1996a). The correlation of egg counts of shoot fly with weather factors is depicted in Table 7. The egg counts had positive but non significant correlation with maximum and mean air temperature in crops sown at different time during both the test years; except for the end January sown crop during 2012, the correlation of egg counts was significant and positive with mean temperature (r= 0.99*); and was non significant and negative in the same crop during 2011. The increase in minimum temperature had a positive influence on egg laying by shoot fly, though the correlation of egg count and minimum temperature was negative (r= - 0.63) in end January sown crop in 2011 also. It was positive in February sown crops (r= 0.97 and 0.78) during 2011. It was positive and significant in end January and mid February sown crops (r= 0.99* each) during 2012. Taneja et al (1986) also reported that most important factors affecting the number of eggs laid by shoot fly A. soccata were adult population density and temperature in sorghum during kharif season. They further reported the negative and significant correlation of maximum (r= - 0.48 to -66) and

50

minimum temperature with egg counts (r = - 0.24 to - 0.58) in sorghum crops sown during different years in kharif season. However, highly significant positive and negative correlation of egg laying with maximum (r = 0.72) and minimum temperature (r = - 0.91), respectively in kharif sorghum was reported by Nair et al (1995). The differences in correlations of egg counts in relation to temperature in the present studies might be due to prevalent temperature range, host plant and predominant shoot fly species in different studies. In the present studies, the maximum, minimum and mean temperature range in 2011 and 2012 during oviposition period ranged from (16.6 31.2 oC and 19.0 32.0oC), (7.0 oC 19.4oC and 4.8 20.6oC) and (13.1 25.0 oC and 12.9 23.4oC), respectively as compared to maximum and minimum temperature range of 23.4oC 33.0 oC and 18.2 oC 23.6 oC prevalent during period of studies by Nair et al (1995). So, it can be concluded that in spring sown maize under the Punjab conditions, the shoot fly oviposition increases with the rise of temperature especially during the period of mid February to mid March. The relative humidity (R.H.) of air had non significant correlation with egg laying by shoot fly during both the test years (table 22) except in end February sown crop during 2011 where it was significant (r = 0.99*) with evening R.H. The correlation between egg laying and evening R.H. was negative with end January and positive with February sown crops during both the test years. However, the correlation of morning and mean R.H. in crop sown at different times did not showed any trend during both the test years. Nair et al (1995) also reported significant and negative correlation (r= -0.84) of egg laying by shoot fly with evening humidity only in sorghum. While, the positive correlation of oviposition with mean R.H. (r=0.54 to 0.68) was reported by Taneja et al (1986). Again the differences of correlation of egg counts in relation to R.H. in the present studies might be due to prevalent weather conditions, host plant and predominant shoot fly species. The morning, evening and mean R.H during period of observations for egg counts during 2011 and 2012 ranged from 72.0 100.0 & 72.0 97.0 %, 36.0 91.0 & 24.0 66.0 %; and 61.5 93.0 & 55.0 81.0 %, respectively as compared to morning and evening R.H range of 88 97 % and 38 84 % reported by Nair et al (1995) during period of their studies. Marwaha et al (1984) found that R.H. between 75.0 80.0 % was favourable for shoot fly infestation in spring maize. In the present studies, egg laying by shoot fly at wider range of mean R.H (55.0 93.0 %) indicated that oviposition is more influenced by crop growth stage and minimum air temperature. Thus, emphasizing the need of further detailed studies at micro level in field conditions for ascertaining the role of R.H. in influencing the oviposition by shoot fly.

51

Table 7.The correlations of egg count with weather parameter in spring 2011 and 2012. 2011 Air temperature (C) Max. Min. Mean 0.23 -0.63 -0.29 0.09 0.97 0.59 0.29 0.78 0.64 2012 SET 1 SET 2 SET 3 0.87 0.81 0.30 0.99* 0.99* 0.69 0.99* 0.96 0.52 -0.58 -0.04 0.87 - 0.54 0.76 0.75 - 0.56 0.85 0.75 Relative humidity (%) Mor. Eve. Mean 0.53 -0.56 0.09 0.13 0.49 0.49 -0.03 0.99* 0.92

SET 1 SET 2 SET 3

* Significant at 5 % level. Set 1: end January sown crop, Set 2: mid February sown crop, Set 3: end February sown crop 4.2.2. Incidence of shoot fly, Atherigona spp.: The leaf injury and deadhearts incidence differed significantly for few observation in the test hybrids; and in almost all the observations for protected and unprotected conditions in crop sown at different periods during both the test years (table 8 and 9). However, the interaction of hybrids and crop protection conditions for deadhearts incidence was significant for later observations only in crop sown at different times, whereas, the interaction for leaf injury between these factors was non significant for all the observations during the study periods. The leaf injury and deadhearts incidence was comparatively lower in end January sown crop (set 1) than February sown crops during 2011 and 2012 (fig.3, 4, 5 and 6). In January sown crop at 10 DAG, the mean leaf injury incidence in hybrids did not differ significantly during both the test years. Irrespective of crop protection conditions, it was 7.89 and 2.46 per cent in JH 3459; and 10.72 and 2.68 per cent in PMH 2 during, respectively 2011 and 2012. In 2011, the mean leaf injury, Irrespective of hybrids, was significantly lower under protected (1.47 %) conditions than that in unprotected conditions (13.14 -13.31 %) whereas in 2012, it differed non significantly and ranged from 2.24 - 2.91 per cent. The mean deadhearts incidence, Irrespective of crop protection conditions differed significantly in the hybrids during 2012 only, being lower in JH 3459 (1.71 %) than that in PMH 2 (4.04 %) Irrespective of hybrids, it was significantly lower in protected condition (0.17 and 0.96 %) than that in unprotected conditions (7.76 and 3.66 %) during both the test years, respectively. At 15 DAG, the mean leaf injury incidence, irrespective of crop protection conditions was significantly low in JH 3459 (5.60 %) than PMH 2 (8.23 %) in 2012, while it was on par in 2011. Irrespective of hybrids, it was significantly lower in protected conditions (2.00 and 3.03 %) than that in unprotected conditions (17.70 and 8.80 %) during 2011

52

and 2012, respectively. Similar trend was observed for deadhearts incidence during both the test years with significantly lower deadhearts in JH 3459 (4.16 and 3.71 %) as compared to PMH 2 (7.61 and 9.07 %). In protected conditions again it was lower (0.17 and 1.28 %) than unprotected conditions (9.23 and 9.12 %) during both the test years. The interaction of hybrids and crop protection conditions for deadhearts incidence was also significant during 2011. However, both the test hybrids did not differ significantly under protected conditions, thus indicating the importance of seed treatments based plant protection to manage the shoot fly in spring sown maize. At 20 DAG, the mean leaf injury incidence, irrespective of crop protection conditions, in JH 3459 (16.56 and 9.97 %) and PMH 2 (14.29 and 13.70 %) was on par with each other during both the test years. However, irrespective of hybrids, it was significantly lower in protected conditions (4.52 and 4.66 %) than that in unprotected conditions (22.08 and 14.53 %) during the respective years. The interaction between hybrids and crop protection conditions was non significant indicating equal leaf injury by shoot fly on susceptible and least susceptible hybrid during both the test years. The deadhearts incidence at this stage differed significantly in hybrids, under protected/ unprotected conditions and in their interactions during both the test years. Irrespective of crop protection conditions, the hybrid JH 3459 had significantly lower mean deadhearts incidence (5.03 and 8.18 %) as compared to PMH 2 (10.07 and 13.22 %) in the respective years. Irrespective of hybrids, the deadhearts incidence was significantly lower in protected conditions (0.50 and 3.83%) than that in unprotected conditions (11.65 and 14.03 %) during 2011 and 2012, respectively. The interaction between hybrid and crop protection conditions was significant and the mean deadhearts incidence was significantly lower in both the hybrids in protected conditions than that in unprotected conditions indicating deadhearts incidence as the real indicator of shoot fly susceptibility in different genotypes. However, the hybrids did not differ significantly in deadhearts incidence under protected conditions again emphasizing the importance of protection measures even in end January sown crop. The leaf injury and deadhearts incidence in set 2 i.e. mid February sown crop was comparatively higher than that in set 1 i.e. end January sown crop during both the test years (Table 8 and 9). At 10 DAG, the mean leaf injury incidence, irrespective of crop protection conditions was significantly higher in PMH 2 (7.77 %) than JH 3459 (4.67 %) during 2012, whereas, in 2011 it was on par in both the hybrids (8.66 and 8.17 %, respectively). Irrespective of hybrids, it was significantly low under protected (1.32 and 1.41 %) than unprotected conditions (11.77and 8.13 %) during the respective years. Similar to leaf injury incidence the mean

deadhearts incidence, irrespective of crop protection conditions, was also significantly higher in PMH 2 (4.68 %) than JH 3459 (3.15 %) in 2012 whereas, both were on par with each other

53

during 2011. The mean deadhearts incidence, irrespective of hybrids, was again low under protected condition (0.33 and 1.13 %) than that in unprotected conditions (6.31 and 5.27 %) during 2011 and 2012, respectively. The interaction between hybrids and crop protection conditions was non significant during both the test years. In 2011 at 15 DAG, the mean leaf injury incidence, irrespective of crop protection conditions was significantly more in JH 3459 (23.85 %) than that in PMH 2(16.89 %) where as in 2012, both were on par with each other. However, the mean leaf injury, irrespective of hybrids was significantly lower under protected conditions (1.64 and 3.79 %) than that in unprotected conditions (28.15and 21.39 %). The mean deadhearts incidence, irrespective of crop protection conditions also differed significantly only in hybrids during 2011, being higher in PMH 2 (12.64 %) than that in JH 3459 (7.34 %) while both were on par with each other in the test hybrids (15.87 and 14.93 %) during 2012. Irrespective of hybrids, it was again significantly lower in protected condition (0.17 and 2.85 %) than that in unprotected conditions (16.24 and 23.68 %) during 2011 and 2012, respectively. At this observation also, the interaction between the hybrids and crop protection conditions for deadhearts incidence was non significant during both the test years. At 20 DAG, as in the end January sown crop (Set 1) the test hybrids differed significantly for deadhearts incidence only again revealing the importance of deadhearts in assessment of damage due to shoot fly in spring sown maize. Moreover, the mean leaf injury incidence, irrespective of crop protection conditions, was variable among the test hybrids being numerically higher in JH 3459 (24.38 %) than that in PMH 2 (21.20 %) in 2011 and also in PMH 2 (14.75 %) than that in JH 3459 (11.70 %) in 2012. However, the same, irrespective of hybrids, was significantly lower under protected (11.17 and 8.86 %) than that in unprotected conditions (30.83 and 16.24 %) during the respective years. The mean deadhearts incidence, irrespective of crop protection conditions, differed significantly in the hybrids, being lower in JH 3459 (9.18 and 15.95 %) than that in PMH 2 (15.45 and 25.28 %) during 2011 and 2012. The mean deadhearts incidence, irrespective of hybrids, was also significantly lower under protected (1.97 and 5.55 %) than unprotected conditions (17.98 and 28.83 %) during 2011 and 2012, respectively. The interaction between hybrids and crop protection conditions was also significant for deadhearts incidence only during both the test years, however, the mean deadhearts were significantly lower in both the hybrids, being on par with each other, under protected conditions, than that under unprotected conditions as already reported under the end January sown crop, emphasizing the indispensability of seed treatment for protection of the crop from shoot fly. The leaf injury and deadhearts incidence was higher in set 3 i.e. end February sown crop (Table 8 and 9) than earlier sown crops during both the test years. At 10 DAG, the leaf injury

54

differed non significantly among the hybrids during both the test years. However, the mean leaf injury incidence, irrespective of crop protection conditions was higher during 2011 in JH 3459 (36.42 % ) than that in PMH 2 (33.99%) as compare to 2.86 and 3.00 per cent incidence, respectively in 2012. As in other sowing dates, the mean leaf injury, irrespective of hybrids, was significantly lower under protected conditions (3.30 and 1.10 %) than that under unprotected conditions (48.07and 3.43 %) in 2011 and 2012, respectively. In 2011, the mean deadhearts incidence, irrespective of crop protection conditions was significantly lower in JH 3459 (8.34 %) than that in PMH 2 (13.71 %) and both the hybrids were on par with each other in 2012. Irrespective of hybrids, it was again significantly lower under protected (1.05 and 0.75 %) than that in unprotected conditions (14.33and 2.89 %) during 2011 and 2012, respectively again indicating that incidence was higher at earlier stage during 2011. The interaction between hybrids and crop protection conditions for deadhearts incidence was non significant during both the test years. At 15 DAG, the mean leaf injury incidence differed non significantly in both the test years, irrespective of crop protection conditions being 33.36 and 16.63 per cent in JH 3459; and 30.08 and 19.86 per cent in PMH 2 during 2011 and 2012, respectively. Irrespective of hybrids, it was significantly lower under protected (2.73 and 2.22 %) than that in unprotected conditions (44.13 and 26.56 %) during the respective years. The mean deadhearts incidence, irrespective of crop protection conditions, was significantly lower in JH 3459 (10.48 and 10.49 %) than that in PMH 2 (17.11 and 18.40 %) in 2011 and 2012, respectively. Irrespective of hybrids, it was also significantly lower under protected (1.73 and 1.10 %) than that in unprotected conditions (18.73 and 20.71 %) during 2011 and 2012, respectively. The interaction between hybrids and crop protection conditions was significant for deadhearts occurrence during both the test years. However, the mean deadhearts incidence in both the hybrids was significantly lower under protected conditions under unprotected conditions. At 20 DAG as in end January and mid February sown crops, the hybrids in end February sown crop also the mean leaf injury incidence, irrespective of hybrids was significantly lower under protected (2.22 and 4.77 %) than that in unprotected conditions (39.87 and 19.98 %) during the respective years. The mean deadhearts incidence, irrespective of crop protection conditions was significantly lower in JH 3459 (11.17 and 18.00 %) than that in PMH 2 (18.33 and 27.43 %) during 2011 and 2012, respectively. Irrespective of hybrids, it was also significantly lower under protected conditions (2.07 and 2.69 %) than that under unprotected conditions (19.87 and 33.63 %) during 2011 and 2012, respectively. The interaction between hybrids and crop protection conditions was also significant for deadhearts incidence during both the test years.

55

Table 8. Incidence of shoot fly, Atherigona spp in crops sown at different times during 2011 at Ludhiana Mean leaf injury and deadhearts at Set 1-2011 LI (%) at 10 DAG Hybrid FP JH 3459 PMH 2 Mean CD (p=0.05) 1.65 1.30 1.47 A x B = NS LI (%) at 15 DAG Condition Hybrid FP JH 3459 PMH 2 Mean CD (p=0.05) 2.37 1.64 2.00 OT 15.47 21.82 18.65 UT 17.67 17.73 17.70 11.84 13.73 Mean FP 0.35 0.00 0.17 OT 5.28 11.22 8.25 UT 6.86 11.60 9.23 4.16 7.61 Condition Mean OT 9.62 16.99 13.31 UT 12.41 13.86 13.14 7.89 10.72 FP 0.35 0.00 0.17 A x B = NS DH (%) at 15 DAG Condition Mean OT 4.63 10.60 7.62 UT 5.22 10.30 7.76 3.40 6.97 DH (%) at 10 DAG Condition Mean

Hybrid (A) = NS, Condition (B) = 5.07,

Hybrid (A) = NS, Condition (B) = 4.89,

Hybrid (A) = NS, Condition (B) = 5.64, A x B = NS LI (%) at 20 DAG

Hybrid (A) = 1.67, Condition (B) = 2.04, A x B = 2.89 DH (%) at 20 DAG Condition

Hybrid FP JH 3459 PMH 2 Mean CD (p=0.05) 4.44 4.60 4.52

Condition Mean OT 21.42 17.95 19.68 UT 23.83 20.32 22.08 16.56 14.29 FP 0.67 0.33 0.50

Mean OT 6.92 14.09 10.51 UT 7.51 15.79 11.65 5.03 10.07

Hybrid (A) = NS, Condition (B) = 3.51, A x B = NS

Hybrid (A) = 2.52, Condition (B) = 3.09, A x B = 4.37

Set 1: end January sown crop; LI: Leaf injury, DH: deadhearts FP: Protection from shoot fly and other insect pests, OT: Protection from insect pests other than shoot fly, UT: No protection for insect pests

56

Table 8. contd

Mean leaf injury and deadhearts at Set 2-2011 LI (%) at 10 DAG Hybrid FP JH 3459 PMH 2 Mean 1.31 1.33 1.32 Condition Mean OT 11.61 12.71 12.16 UT 11.59 11.95 11.77 8.17 8.66 FP 0.33 0.34 0.33 OT 5.65 6.33 5.99 UT 6.29 6.33 6.31 4.09 4.33 DH (%) at 10 DAG Condition Mean

CD Hybrid (A) = NS, Condition (B) = 2.10, Hybrid (A) = NS, Condition (B) = 0.96, (p=0.05) A x B = NS A x B = NS LI (%) at 15 DAG Hybrid FP JH 3459 PMH 2 Mean 2.30 0.98 1.64 Condition Mean OT 37.17 25.47 36.34 UT 32.08 24.22 28.15 23.85 16.89 FP 0.33 0.00 0.17 OT 9.52 17.59 13.56 UT 12.15 20.32 16.24 7.34 12.64 DH (%) at 15 DAG Condition Mean

CD Hybrid (A) = 6.08, Condition (B) = Hybrid (A) = 3.34, Condition (B) = 4.09, A (p=0.05) 7.44, A x B = NS x B = NS LI (%) at 20 DAG Hybrid FP JH 3459 PMH 2 Mean 12.57 9.76 11.17 Condition Mean OT 28.53 24.2 26.37 UT 32.04 29.62 30.83 24.38 21.20 FP 1.98 1.95 1.97 OT 12.15 21.83 16.99 UT 13.40 22.57 17.98 9.18 15.45 DH (%) at 20 DAG Condition Mean

CD Hybrid (A) = NS, Condition (B) = 6.61, (p=0.05) A x B = NS

Hybrid (A) = 2.87, Condition (B) = 3.51, A x B = 4.97

Set 2: mid February sown crop; LI: Leaf injury, DH: deadhearts FP: Protection from shoot fly and other insect pests, OT: Protection from insect pests other than shoot fly, UT: No protection for insect pests

57

Table 8. contd Mean leaf injury and deadhearts at Set 3-2011 LI (%) at 10 DAG Hybrid FP JH 3459 PMH 2 Mean 3.77 2.82 3.30 Condition Mean OT 59.11 49.39 54.25 UT 46.39 49.75 48.07 36.42 33.99 FP 0.69 1.40 1.05 OT 13.74 21.68 17.71 UT 10.60 18.05 14.33 8.34 13.71 DH (%) at 10 DAG Condition Mean

CD Hybrid (A) = NS, Condition (B) = 7.70, Hybrid (A) = 3.76, Condition (B) = 4.61, (p=0.05) A x B = NS A x B = NS LI (%) at 15 DAG Hybrid FP JH 3459 PMH 2 Mean 4.44 1.03 2.73 Condition Mean OT 52.60 43.99 48.30 UT 43.04 45.22 44.13 33.36 30.08 FP 2.06 1.40 1.73 OT 16.11 25.75 20.93 UT 13.28 24.19 18.73 10.48 17.11 DH (%) at 15 DAG Condition Mean

CD Hybrid (A) = NS, Condition (B) = 6.06, Hybrid (A) = 3.64, Condition (B) = 4.46, (p=0.05) A x B = NS A x B = 6.31 LI (%) at 20 DAG Hybrid FP JH 3459 PMH 2 Mean 3.75 0.69 2.22 Condition Mean OT 50.59 38.93 44.76 UT 39.04 40.69 39.87 31.13 26.77 FP 2.40 1.74 2.07 OT 17.49 27.11 22.30 UT 13.61 26.13 19.87 11.17 18.33 DH (%) at 20 DAG Condition Mean

CD Hybrid (A) = NS, Condition (B) = 5.55, (p=0.05) A x B = NS

Hybrid (A) = 3.77, Condition (B) = 4.62, A x B = 6.53

Set 3: end February sown crop ; LI: Leaf injury, DH: deadhearts FP: Protection from shoot fly and other insect pests, OT: Protection from insect pests other than shoot fly, UT: No protection for insect pests

58

Table 9. Incidence of shoot fly, Atherigona spp in crops sown at different times during 2012 at Ludhiana Mean leaf injury and deadhearts at Set 1-2012 LI (%) at 10 DAG Hybrid FP JH 3459 PMH 2 Mean CD (p=0.05) 1.89 2.60 2.24 A x B = NS LI (%) at 15 DAG Hybrid FP JH 3459 PMH 2 Mean CD (p=0.05) 3.16 2.91 3.03 Condition Mean OT 6.94 10.89 8.91 UT 6.71 10.88 8.80 5.60 8.23 FP 0.63 1.93 1.28 OT 5.09 12.47 8.78 UT 5.43 12.82 9.12 3.71 9.07 Condition Mean OT 3.27 2.56 2.91 UT 2.23 2.89 2.56 2.46 2.68 FP 0.63 1.28 0.96 A x B = NS DH (%) at 15 DAG Condition Mean OT 1.95 6.08 4.02 UT 2.55 4.77 3.66 1.71 4.04 DH (%) at 10 DAG Condition Mean

Hybrid (A) = NS, Condition (B) = NS,

Hybrid (A) = 1.73, Condition (B) = 2.11,

Hybrid (A) = 2.32, Condition (B) = Hybrid (A) = 2.93, Condition (B) = 3.59, 2.84, A x B = NS A x B = NS LI (%) at 20 DAG DH (%) at 20 DAG Condition Mean FP OT 14.63 18.01 16.32 UT 12.76 16.31 14.53 9.97 13.70 FP 2.83 4.84 3.83 OT 11.85 16.62 14.24 UT 9.86 18.20 14.03 8.18 13.22 Mean

Hybrid

Condition

JH 3459 PMH 2 Mean CD (p=0.05)

2.52 6.80 4.66

Hybrid (A) = NS, Condition (B) = Hybrid (A) = 1.90, Condition (B) = 2.33, 7.98, A x B = NS A x B = 3.30

Set 1: end January sown crop; LI: Leaf injury, DH: deadhearts FP: Protection from shoot fly and other insect pests, OT: Protection from insect pests other than shoot fly, UT: No protection for insect pests

59

Table 9. contd Mean leaf injury and deadhearts at Set 2-2012 LI (%) at 10 DAG Hybrid FP JH 3459 PMH 2 Mean 1.88 0.95 1.41 Condition Mean OT 6.89 11.36 9.13 UT 5.25 11.00 8.13 4.67 7.77 FP 1.34 0.93 1.13 A x B = NS DH (%) at 15 DAG Condition Mean FP JH 3459 PMH 2 Mean 4.74 2.85 3.79 OT 16.55 20.37 18.46 UT 19.45 23.33 21.39 13.58 15.52 FP 4.11 1.60 2.85 A x B = NS DH (%) at 20 DAG Condition Mean FP JH 3459 PMH 2 Mean 6.60 11.11 8.86 OT 12.70 16.45 14.57 UT 15.79 16.69 16.24 11.70 14.75 FP 4.74 6.36 5.55 A x B = 7.24 OT 19.95 34.99 27.47 UT 23.17 34.49 28.83 15.95 25.28 Mean OT 18.31 21.01 19.66 UT 22.37 24.99 23.68 14.93 15.87 Mean OT 4.16 6.51 5.34 UT 3.97 6.58 5.27 3.15 4.68 DH (%) at 10 DAG Condition Mean

CD Hybrid (A) = 2.51, Condition (B) = 3.07, (p=0.05) A x B = NS LI (%) at 15 DAG Hybrid Condition

Hybrid (A) = 0.56, Condition (B) = 0.69,

CD Hybrid (A) = NS, Condition (B) = 6.04, (p=0.05) A x B = NS LI (%) at 20 DAG Hybrid Condition

Hybrid (A) = NS, Condition (B) = 7.74,

CD Hybrid (A) = NS, Condition (B) = 5.15, (p=0.05) A x B = NS

Hybrid (A) = 4.18, Condition (B) = 5.12,

Set 2: mid February sown crop; LI: Leaf injury, DH: deadhearts FP: Protection from shoot fly and other insect pests, OT: Protection from insect pests other than shoot fly, UT: No protection for insect pests

60

Table 9. contd Mean leaf injury and deadhearts at Set 3-2012 LI (%) at 10 DAG Hybrid FP JH 3459 PMH 2 Mean CD (p=0.05) 0.96 1.24 1.10 A x B = NS LI (%) at 15 DAG Hybrid FP JH 3459 PMH 2 Mean CD (p=0.05) 2.89 1.55 2.22 A x B = NS LI (%) at 20 DAG Hybrid FP JH 3459 PMH 2 Mean CD (p=0.05) 5.49 4.06 4.77 A x B = NS Condition Mean OT 19.97 21.53 20.75 UT 20.11 19.86 19.98 15.19 15.14 FP 2.56 2.81 2.69 OT 25.31 38.38 31.84 UT 26.13 41.12 33.63 18.00 27.43 Condition Mean OT 25.19 27.94 26.56 UT 21.82 30.08 25.95 16.63 19.86 FP 0.96 1.23 1.10 OT 15.75 27.30 21.53 UT 14.76 26.66 20.71 10.49 18.40 Condition Mean OT 4.59 4.84 4.71 UT 3.02 3.83 3.43 2.86 3.30 FP 0.70 0.79 0.75 A x B = NS DH (%) at 15 DAG Condition Mean OT 3.31 2.50 2.90 UT 2.97 2.81 2.89 2.33 2.03 DH (%) at 10 DAG Condition Mean

Hybrid (A) = NS, Condition (B) = 2.02,

Hybrid (A) = NS, Condition (B) = 0.60,

Hybrid (A) = NS, Condition (B) = 8.23,

Hybrid (A) = 2.50, Condition (B) = 3.06, A x B = 4.33 DH (%) at 20 DAG Condition Mean

Hybrid (A) = NS, Condition (B) = 4.01,

Hybrid (A) = 4.02, Condition (B) = 4.93, A x B = 6.97

Set 3: end February sown crop ; LI: Leaf injury, DH: deadhearts FP: Protection from shoot fly and other insect pests, OT: Protection from insect pests other than shoot fly, UT: No protection for insect pests

61

Figure 3 & 4

62

Figure 5 & 6

63

The mean incidence of deadhearts being on par with each other was significantly low under protected conditions in both the hybrids i.e. JH 3459 and PMH 2 than that under unprotected conditions. But the infestation level of 13.61 - 17.49 and 25.31 26.13 per cent in 2011 and 2012, respectively under unprotected conditions even in moderately tolerant hybrid JH 3459 again emphasized the need of plant protection measures in February sown crop. The pooled mean over the cultivars and the protected/ unprotected conditions for shoot fly infestation at 20 DAG revealed that the mean leaf injury and deadhearts incidence was higher in end February followed by mid February sown crop and least in end January sown crop during both the test years (Table 10). The mean leaf injury incidence, irrespective of hybrids was lower in protected condition (5.97 and 5.56 %) than that in unprotected condition (30.60 and 15.02 %) during 2011 and 2012, respectively. The mean leaf injury incidence was significantly lower in end January (12.70 and 6.16 %) than that in mid February (19.89 and 12.13 %) and end February (22.27 and 12.57 %) sown crops during 2011 and 2012, respectively. However, both the sowing times in February did not differ significantly in this respect. The significant interaction between sowing times and crop protection conditions showed that leaf injury incidence was significantly lower in protected conditions than that under unprotected conditions at all the three sowing times during both the test years and it increased progressively with delay in sowing under unprotected conditions. The mean leaf injury incidence, irrespective of hybrids, was maximum and

minimum in mid February (11.17 %) and end February sown crop (2.22 %) under protected condition; and in end February (42.32 %) and end January sown crop (20.88 %) under unprotected conditions, respectively during 2011. Similarly in 2012, the maximum incidence was observed in mid (8.86 %) and end February (20.37 %); and minimum incidence was observed in end January sown (3.04 and 9.27 %) crop under protected and unprotected conditions,

respectively. The mean deadhearts incidence, irrespective of hybrids was also lower in protected conditions (1.52 and 4.03 %) than that in unprotected conditions (16.55 and 25.01 %) during 2011 and 2012, respectively. It was significantly lower in end January sown crop (5.79 and 8.98 %) than that in mid February (9.73 and 16.85 %) and end February sown crops (11.58 and 17.72 %) during 2011 and 2012, respectively. As in case of leaf injury incidence, both the February sown crops did not differ significantly for deadhearts incidence also during both the test years. The interaction between times of sowing and crop protection conditions was significant. The mean deadhearts incidence, irrespective of hybrids was lower in protected conditions than that under unprotected conditions at different sowing times during both the test years, again emphasized the role of plant protection against shoot fly in spring sown crop. Similar to leaf injury incidence, deadhearts also increased progressively with delay in sowing under unprotected

64

conditions. During 2011, the mean deadhearts incidence, irrespective of hybrids, was maximum in end February (2.08 %) and minimum in end January sown crop (0.50 %) under protected conditions; and in end February (21.09 %) and end January sown crop (11.08 %) under unprotected conditions, respectively. While in 2012, the deadhearts incidence was maximum in mid February (5.55 %) and minimum in end February (2.69 %) under protected conditions; and it was maximum in end February (32.74 %) and minimum in end January sown (14.13 %) crop under unprotected conditions. Similarly, the higher shoot fly incidence and more losses in yield with delayed sowing i.e. in the 3rd week of February as compared to 1st and 2nd week of February sown crop in terai area of Uttar Pradesh was reported by Sarup et al (1979). Marwaha et al (1984) also reported that irrespective of maize varieties, the sowing dates manifested significant differences in deadhearts formation due to shoot fly species (A. soccata and A. naqvii). According to them, the incidence was higher in mid February (63.99 %) and end February (26.7 %) sown crop during 1981 and 1983, respectively at Delhi. Similarly, Rao and Panwar (1995a) also reported that irrespective of varieties, maximum deadhearts incidence due to shoot flies (17.9 %) with delayed sowing of maize on 8th March at Delhi. At Ludhiana, Gagandeep and Kanta (2007a) reported maximum damage of 25.77 per cent deadhearts due to shoot fly infestation on 6 th March sown crop as compared to 7.53 and 7.65 per cent in 15th and 25th February sown crop, respectively. They further reported that among different dates deadhearts incidence varied from 2.33 to 17.64 and 12.67 to 33.90 per cent under protected and unprotected conditions, respectively. The appraisal of the data indicated that though the early sown crop in end January recorded lower shoot fly infestation but seed treatment with Gaucho (imidacloprid) 600 FS @ 6 ml per kg seed further reduced the damage. The seed treatment in end January, mid February and end February sown crops is must. The effectiveness of imidacloprid seed treatments in reducing shoot fly infestation was reported earlier also (Rao and Panwar 1996b, Jindal and Hari 2008a, 2011). To study the most vulnerable stage for shoot fly incidence and to ascertain the window for application of spray treatment to manage shoot fly if any, the infestation data was pooled. The overall mean for the leaf injury and deadhearts incidence at various observations in crops sown at different times during both the test years has been depicted in fig. 3, 4, 5 and 6. In 2011, the leaf injury incidence was maximum at 20 DAG on end January sown crop (15.43 %) and mid February sown crop (22.79 %) while it was maximum at 10 DAG on end February sown crop (fig. 3). In 2012 (fig. 4), it was again maximum at 20 DAG in end January (11.84 %) and at 15 DAG for both mid and end February sown crops (14.60 and 18.25 %, respectively). The increase in leaf injury on crop sown in recommended time i.e. during mid February was 8.42 to

65

20.37 and 6.22 to 14.60 per cent from 10 to 15 DAG in 2011 and 2012, respectively. Thus, indicating that there exists a window to minimize the damage by the application of insecticides at this time. Similarly the increase in deadhearts incidence (fig 5 and 6) was higher from 10 to 15 DAG in 2011 (4.21 to 9.99 %) and 2012 (3.91 to 15.40 %), in the recommended mid February sown crop. Thus again supporting the fact that the damage can be minimized if suitable control measures are applied at the first appearance of shoot fly infestation or after one week of crop emergence. However, the further detailed studies are needed in future in this regard.

Table 10. Incidence of shoot fly, Atherigona spp pooled over the cultivars in protected and
unprotected conditions at Ludhiana
Mean leaf injury and deadhearts during 2011 Condition LI (%) Sowing time DH (%) Sowing time Mean

Protected Un protected Mean CD (p=0.05)

Set 1 4.52 20.88 12.70

Set 2 11.17 28.60 19.89

Set 3 2.22 42.32 22.27

Mean

5.97 30.60

Set 1 0.50 11.08 5.79

Set 2 1.97 17.49 9.73

Set 3 2.08 21.09 11.58

1.52 16.55

Condition (A) = 1.98, Sowing time (B) = 2.43, A x B = 3.44

LI (%) Sowing time

Condition (A) = 1.59, Sowing time (B) = 1.95 A x B = 2.76

2012 DH (%) Sowing time

Condition

Protected Un protected Mean CD (p=0.05)

Set 1 3.04 9.27 6.16

Set 2 8.86 15.40 12.13

Set 3 4.77 20.37 12.57

Mean

5.56 15.02

Set 1 3.84 14.13 8.98

Set 2 5.55 28.15 16.85

Set 3 2.69 32.74 17.72

Mean 4.03 25.01

Condition (A) = 2.23, Sowing time (B) = 2.73, A x B = 3.86

Condition (A) = 2.29, Sowing time (B) = 2.80, A x B = 3.96

Set 1: end January, Set 2: mid February, Set 3: end February sown crops ; LI: Leaf injury, DH: deadhearts

The leaf injury and deadhearts incidence was positively correlated with maximum air temperature during both the test years in crops sown at different times, except in January sown crop in 2012, where deadhearts incidence was negatively correlated with it (table 11). This correlation was significant (r=0.99*) for deadhearts incidence in end February sown crop during 2011 only. The correlations of leaf injury and deadhearts incidence were also positive and significant with minimum air temperature in end January and end February sown crops during

66

both the test years except for deadhearts incidence in end January sown crop during 2012 (r= 0.88). In contrast, it was positive and significant (r=0.99*) on the crop sown at same time during 2011, indicating climatic factors vis--vis shoot fly abundance influence the ultimate degree of damage to the crop sown at different times. In mid February sown crop, the correlation of leaf injury and deadhearts incidence with minimum temperature was negative but non significant, this might be due to prevalent temperature range, abundance of shoot fly and the stage of the crop at that time. The leaf injury and deadhearts incidence was also positively correlated with mean air temperature in crop sown at different times during both the test years except for leaf injury in mid February and deadhearts in end January sown crop during 2012. However it was significant for deadhearts in end January (r=0.99*) and for leaf injury in end February (r=1.00*) sown crops in 2011 and 2012, respectively. Overall, during the period of study the incidence (leaf injury and deadhearts) increased with rise of minimum air temperatures in maize sown at different times. Khan and Singh (1980) based on mean of four seasons also reported that incidence of A. naqvii on wheat increased with the rise of air temperature. According to Taneja et al (1986) formation of deadhearts due to shoot fly in kharif sorghum was highly dependent on adult population density, temperature and minimum humidity. They further reported that the correlation of deadhearts with maximum and minimum temperature was significant and negative (r= 0.57 to 0.83 and 0.23 to 0.43, respectively) during different years, while it was positive and significant (r=0.43) in one year with minimum temperature also. Singh and Verma (1986) also reported that deadhearts formation was negatively correlated with temperature (r=0.71) in sorghum. The negative correlation of shoot fly incidence with temperature as reported by these workers in contrast to positive correlation in the present findings. This might be due to difference in weather conditions, host plant and predominant shoot fly species during kharif season than that in spring season crop. However Nair et al (1995) also reported that maximum and minimum temperature; morning and evening R.H.; and bright sunshine hours had week correlation with deadheart formation and inferred that the meteorological parameters indirectly affected deadhearts formation. The correlations of shoot fly incidence (leaf injury + deadhearts) with morning R.H. either positive or negative but non significant in crop sown at different times during both the test years except for leaf injury incidence for mid February (r= 0.99*) and end February (r= -0.99*) sown crop during 2011 and 2012, respectively (table 11). The correlation of leaf injury and deadhearts incidence was negative with evening R.H. for all sowings times in 2012 and for mid February sown crop during 2011. However, it was significant for mid February sown crop for leaf injury (r= -0.98*) and deadhearts (r= -0.99*) during 2011 and 2012; and only for deadhearts incidence (r= -0.98) in end February sown crop during 2012 also. Similarly, mean R.H. had

67

significant correlations for leaf injury (r= -0.98) and deadhearts (r= -0.99) incidence in only mid February sown crop during 2011 whereas, deadhearts incidence was significantly correlated in end January and end February crop (r= -0.99 and -1.00) during 2012. The correlations of shoot fly incidence with mean R.H. were non significant for other sowing dates. Thus, overall R.H. was not strongly correlated with shoot fly damage and it may have indirect effect as R.H. decrease with increase of temperature. Moreover, the wide range of mean R.H. (61.5-90.5 and 49.0-73.0 % in respective years) prevailed during the shoot fly infestation observation period indicated that microclimatic factors might have more influence on the shoot fly abundance and crop damage. However, positive and significant correlation of deadhearts due to shoot fly with R.H. as reported by Taneja et al (1986) and Singh and Verma (1986) in sorghum might be due to coincidence of peak activity period of A. soccata on sorghum with rainy season. Table 11. The correlations of shoot fly incidence with weather parameter in spring 2011 and 2012. 2011 Air temperature (C) Leaf injury Set 1 Set 2 Set 3 Deadhearts Set 1 Set 2 Set 3 Leaf injury Set 1 Set 2 Set 3 Deadhearts Set 1 Set 2 Set 3 -0.65 0.50 0.97 -0.88 -0.31 0.52 -0.80 0.08 0.82 -0.44 0.59 -0.75 -0.45 -0.99* -0.98* -0.99* -0.59 -1.00* 0.55 0.06 0.93 0.56 -0.71 0.91 0.56 -0.36 1.00* 0.73 0.98 0.99* 0.99* -0.32 0.82 0.99* 0.91 0.89 2012 0.82 0.17 -0.99* -0.45 -0.98* -0.71 0.51 -0.89 -0.82 -0.33 0.97 -0.85 0.75 -0.99* 0.12 0.44 -0.99* -0.53 Max. 0.91 0.95 0.86 Min. 0.91 -0.43 0.84 Mean 0.93 0.85 0.87 Mor. -0.60 0.99* -0.60 Eve. 0.51 -0.98* 0.21 Mean 0.14 -0.98* -0.16 Relative humidity (%)

Set 1: end January sown crop, Set 2: mid February sown crop, Set 3: end February sown crop

68

4.2.3. Population builds up of other insect pest: The population counts and damage of other insect pests infesting spring sown maize on different dates at Ludhiana was recorded at weekly intervals during both the test years and data has been presented in Table 12-14 and Annexure VII-XII. Similar to observation in the first experiment at Hoshiarpur, the damage and population level of other insect pests was low at Ludhiana also in maize sown at different time. However, apart from incidence of shoot fly (key insect pest of spring maize) the high incidence of small brown planthopper, Laodelphax striatellus (Falln) was also observed for first time in spring sown maize in the Punjab in spring 2011. This insect pest was reported for the first time in India on rice nursery (variety Jaya) in the farmers fields near Ludhiana by Shukla (1979). The pest appeared on the spring sown maize crop in the second week of April and its population started declining in the second week of May during both the test years (fig 7 and 8). The highest population counts of this pest were observed in third week of April and fourth week of April on crop sown at different times during 2011 and 2012, respectively (fig 7 and 8). However, the pest population differed non significantly on both the test hybrids and in different crop protection conditions irrespective of spring maize sown on different dates during both the test years. The interaction between hybrids and crop protection conditions for the abundance of L. striatellus was also non significant during both the test years. In 2011, on end January sown crop, the pest population was observed on JH 3459 and PMH 2 i.e. 1.85 & 2.02 hoppers per plant; 86.22 & 105.89, 116.33 & 145.78; 40.28 & 35.67 and 1.00 & 0.72 hoppers per sweep during second, third & fourth week of April; first and second week of May, respectively (table 12). In the corresponding period the pest population was numerically lower (1.63 hoppers/ plant; 80.0, 113.58, 33.58 and 1.00 hoppers/ sweep) in plots having protection from shoot fly than unprotected conditions (2.07-2.12 hoppers/ plant; 100.42-107.75, 137.08-142.50, 34.42-45.92, 0.75-0.83 hoppers/ sweep). The interaction of these factors was non significant and the population counts were lower in protected conditions than that under unprotected conditions in both the test hybrids during all the observations. However, during 2012 the incidence of L. striatellus was low in end January sown crop but it again differed non significantly in hybrids and crop protection conditions (table 13). The pest counts on JH 3459 and PMH 2 were 0.10 & 0.13; 0.16 & 0.16; and 0.48 & 0.39 hoppers per plant, during third and fourth week of April and first week of May, respectively. The counts in protected and unprotected conditions varied from 0.13-0.43 and 0.10-0.52 hoppers per plant, respectively during the period of different observations. The hopper counts in mid February sown crop were also higher during 2011 and the maximum counts were observed in third and last week of April during 2011 and 2012, respectively (fig. 7 & 8). However, it differed non significantly among treatments at different observation times

69

during both the test years (table 12 and 13). In 2011, the counts of L. striatellus on JH 3459 and PMH 2 were 3.37 & 3.10 hoppers per plant; 183.11 & 189.05, 116.50 & 125.39, 51.78 & 51.28 and 1.56 &0.83 hoppers per sweep during second, third & fourth week of April; first and second week of May, respectively (table 12). As in end January sown crop the pest population was also

comparatively lower (2.92 hoppers/ plant; 188.17, 114.42, 50.08 and 1.25 hoppers/ sweep) in plots with protection from shoot fly as compare to unprotected conditions (3.32-3.47hoppers/ plant; 181.82-188.17, 122.17-126.25, 50.42-54.08 and 1.08-1.25 hoppers/ sweep). The interaction of these factors was non significant and the population counts were lower in protected conditions than that under unprotected conditions in both the test hybrids during all the observations. During 2012, as in end January sown crop, the incidence of L. striatellus was also low in mid February sown crop and it differed non significantly in hybrids and crop protection conditions. The hopper counts per plant on JH 3459 and PMH 2 were 0.37 & 0.42, 0.21 & 0.29, 1.10 & 1.20, 0.49 & 0.79 and 0.06 in each during second, third & fourth week of April; and first & second week of May, respectively. The hopper counts in protected and unprotected conditions varied from 0.07-1.00 and 0.05-1.32 per plant, respectively during the period of different observations. The population counts in end February sown crop were comparatively lower than in mid February sown crop during 2011 (fig. 7). The highest pest population was observed during third & fourth week of April during 2011 and 2012 respectively. As in end January and mid February sown crops, the pest counts differed non significantly among hybrids as well as crop protection conditions during both the test years in this sowing date also (table 12 and 13). In 2011, the population of small brown planthopper on JH 3459 and PMH 2 was 2.25 & 2.10 hoppers per plant; 146.28 & 145.11, 47.27 & 48.17, 25.50 & 28.11 and 0.29 & 0.28 hoppers per sweep during second, third & fourth week of April; first and second week of May, respectively. The pest population during

corresponding period in protected plots was 1.78 hoppers per plant; 148.67, 45.17, 28.42 and 0.28 hoppers per sweep in comparison to 2.28-2.47 hoppers per plant; 133.83-154.58, 47.58-50.42, 25.3326.67 and 0.27-0.30 hoppers per sweep in unprotected conditions. The interaction of these factors was non significant and the population counts were variable in protected conditions in both the test hybrids during all the observations. In 2012 the incidence of L. striatellus was again low in this crop also and differed non significantly between hybrids and crop protection conditions. The hopper counts per plant on JH 3459 and PMH 2 were 0.18 & 0.19, 0.20 & 0.22, 1.74 & 1.24, 0.49 & 0.51 and 0.03 & 0.09 during second, third & fourth week of April; and first & second week of May, respectively. The hopper counts in protected and unprotected conditions varied from 0.07-1.55 and 0.03-1.48 per plant, respectively during the period of different observations.

70

Table 12. The small brown planthopper, Laodelphax striatellus (Falln) population counts per plant* or per sweeps** in spring 2011 at Ludhiana
The populations count of small brown planthopper, L. striatellus during 2011: Set 1 Hybrid JH 3459 PMH 2 Mean CD (p=0.05) 14.04.11* 21.04.11** Condition Mean Condition Mean FP OT UT FP OT UT 1.53 1.93 2.10 1.85 78.16 89.00 91.50 86.22 1.73 2.30 2.03 2.02 81.83 126.50 109.33 105.89 1.63 2.12 2.07 80.00 107.75 100.42 Hybrid (A) = NS, Condition (B) = NS Hybrid (A) = NS, Condition (B) = NS A x B = NS A x B = NS 28.04.11** 07.05.11** Condition Condition Mean Mean FP OT UT FP OT UT 112.67 111.50 124.83 116.33 31.83 35.17 53.83 40.28 114.50 173.50 149.33 145.78 35.33 33.67 38.00 35.67 113.58 142.50 137.08 33.58 34.42 45.92 Hybrid (A) = NS, Condition (B) = NS Hybrid (A) = NS, Condition (B) = NS A x B = NS A x B = NS 13.05.11** Condition Mean FP OT UT 1.33 0.83 0.83 1.00 0.67 0.83 0.67 0.72 1.00 0.83 0.75 Hybrid (A) = NS, Condition (B) = NS, A x B = NS 2011: Set 2 16.04.11* 23.04.11** Condition Condition Mean Mean FP OT UT FP OT UT 2.87 3.63 3.60 3.37 207.00 187.33 155.00 183.11 2.97 3.30 3.03 3.10 169.33 176.50 221.33 189.05 2.92 3.47 3.32 188.17 181.82 188.17 Hybrid (A) = NS, Condition (B) = NS Hybrid (A) = NS, Condition (B) = NS A x B = NS A x B = NS 29.04.11** 6.05.11 ** Condition Condition Mean Mean FP OT UT FP OT UT 113.17 115.00 121.33 116.50 52.50 54.50 48.33 51.78 115.67 137.50 123.00 125.39 47.67 53.67 52.50 51.28 114.42 126.25 122.17 50.08 54.08 50.42 Hybrid (A) = NS, Condition (B) = NS Hybrid (A) = NS, Condition (B) = NS A x B = NS A x B = NS 13.05.11** Condition Mean FP OT UT 1.50 1.50 1.67 1.56 1.00 1.00 0.50 0.83 1.25 1.25 1.08 Hybrid (A) = NS, Condition (B) = NS, A x B= NS

Hybrid JH 3459 PMH 2 Mean CD (p=0.05)

Hybrid JH 3459 PMH 2 Mean CD (p=0.05)

Hybrid JH 3459 PMH 2 Mean CD (p=0.05)

Hybrid JH 3459 PMH 2 Mean CD (p=0.05)

Hybrid JH 3459 PMH 2 Mean CD (p=0.05)

71

Table 12 Contd 2011: Set 3 Hybrid JH 3459 PMH 2 Mean CD (p=0.05) FP 1.60 1.97 1.78 16.04.11* Condition OT UT 2.63 2.53 2.3 2.47 2.03 2.28 25.04.11** Condition FP OT UT 138.67 155.67 144.5 158.67 148.67 112.00 133.83 164.67 154.58

Mean 2.25 2.10

Mean 146.28 145.11

Hybrid (A) = NS, Condition (B) = NS A x B = NS 30.04.11**

Hybrid (A) = NS, Condition (B) = NS A x B = NS 7.05.11 ** Condition Mean 25.50 28.11

Hybrid FP JH 3459 PMH 2 Mean CD (p=0.05) Hybrid JH 3459 PMH 2 Mean CD (p=0.05) 48.33 42.00 45.17

Condition OT 48.33 52.50 50.42 UT 45.17 50.00 47.58 Mean 47.27 48.17 FP 27.83 29.00 28.42

OT 25.33 28.00 26.67

UT 23.33 27.33 25.33

Hybrid (A) = NS, Condition (B) = NS A x B = NS

Hybrid (A) = NS, Condition (B) = NS A x B = NS 13.05.11

FP 0.30 0.27 0.28

Condition OT 0.27 0.27 0.27

UT 0.3 0.3 0.3

Mean 0.29 0.28

Hybrid (A) = NS, Condition (B) = NS, A x B = NS

Set 1: end January sown crop, Set 2: mid February sown crop, Set 3: end February sown crop; LI: Leaf injury, DH: deadhearts; FP: Protection from shoot fly and other insect pests, OT: Protection from insect pests other than shoot fly, UT: No protection for insect pests

72

Table 13. The small brown planthopper, Laodelphax striatellus (Falln) population counts per plant in spring 2012 at Ludhiana
The populations count of small brown planthopper, L. striatellus during 2012: Set 1 16.04.12 Condition Mean FP OT UT 0.13 0.03 0.13 0.10 (1.06) (1.02) (1.06) (1.05) 0.13 0.20 0.07 0.13 (1.06) (1.09) (1.03) (1.06) 0.13 0.12 0.10 (1.06) (1.06) (1.05) Hybrid (A) = NS, Condition (B) = NS A x B = NS 01.05.12 Condition FP OT 0.47 (1.21) 0.40 (1.18) 0.30 (1.14) 0.40 (1.18) 23.04.12 Condition Mean FP OT UT 0.17 0.10 0.20 0.16 (1.08) (1.05) (1.09) (1.07) 0.13 0.13 0.20 0.16 (1.06) (1.06) (1.09) (1.07) 0.15 0.12 0.20 (1.07) (1.06) (1.09) Hybrid (A) = NS, Condition (B) = NS A x B = NS Mean 0.48 (1.21) 0.39 (1.18)

Hybrid JH 3459 PMH 2 Mean CD (p=0.05)

Hybrid JH 3459 PMH 2 Mean CD (p=0.05)

UT 0.67 (1.28) 0.37 (1.17) 0.52 (1.22)

0.43 (1.19) 0.35 (1.16) Hybrid (A) = NS, Condition (B) = NS, A x B = NS 2012: Set 2

Hybrid JH 3459 PMH 2 Mean CD (p=0.05)

Hybrid JH 3459 PMH 2 Mean CD (p=0.05) Hybrid JH 3459 PMH 2 Mean CD (p=0.05)

11.04.12 16.04.12 Condition Condition Mean Mean FP OT UT FP OT UT 0.33 0.37 0.40 0.37 0.07 0.33 0.23 0.21 (1.15) (1.17) (1.18) (1.17) (1.04) (1.15) (1.11) (1.10) 0.30 0.57 0.37 0.42 0.37 0.27 0.23 0.29 (1.14) (1.24) (1.17) (1.18) (1.16) (1.22) (1.11) (1.13) 0.32 0.47 0.38 0.22 0.30 0.23 (1.15) (1.21) (1.17) (1.10) (1.14) (1.11) Hybrid (A) = NS, Condition (B) = NS Hybrid (A) = NS, Condition (B) = NS A x B = NS A x B = NS 23.04.12 01.05.12 Condition Condition Mean Mean FP OT UT FP OT UT 1.13 1.43 0.73 1.10 0.47 0.47 0.53 0.49 (1.46) (1.54) (1.31) (1.44) (1.21) (1.21) (1.24) (1.22) 0.87 1.20 1.53 1.20 0.50 0.73 1.13 0.79 (1.36) (1.48) (1.59) (1.48) (1.22) (1.31) (1.45) (1.33) 1.00 1.32 1.13 0.48 0.60 0.83 (1.41) (1.51) (1.45) (1.22) (1.26) (1.34) (A) = NS, (B) = NS, A x B = NS (A) = NS, (B) = NS, A x B = NS 10.05.12 Condition Mean FP OT UT 0.07 (1.03) 0.03 (1.02) 0.07 (1.03) 0.06 (1.03) 0.07 (1.03) 0.07 (1.03) 0.03 (1.02) 0.06 (1.03) 0.07 (1.03) 0.05 (1.02) 0.05 (1.02) Hybrid (A) = NS, Condition (B) = NS, A x B = NS

73

Table 13 contd.. 2012: Set 3 11.04.12 Hybrid JH 3459 PMH 2 Mean CD (p=0.05) Condition FP OT UT 0.13 0.17 0.23 (1.06) (1.08) (1.11) 0.27 0.23 0.07 (1.12) (1.11) (1.03) 0.20 0.20 0.15 (1.09) (1.09) (1.07) Hybrid (A) = NS, Condition (B) = NS A x B = NS 23.04.12 Condition FP OT UT 1.90 1.67 1.67 (1.70) (1.62) (1.63) 1.20 1.23 1.30 (1.48) (1.49) (1.51) 1.55 1.45 1.48 (1.59) (1.55) (1.57) Hybrid (A) = (0.14), Condition (B) = NS, A x B = NS Mean 0.18 (1.08) 0.19 (1.09) 16.04.12 Condition FP OT UT 0.53 0.00 0.07 (1.22) (1.00) (1.03) 0.10 0.20 0.37 (1.05) (1.09) (1.16) 0.32 0.10 0.22 (1.14) (1.05) (1.10) Hybrid (A) = NS, Condition (B) = NS A x B = NS 01.05.12 Condition FP OT UT 0.47 0.57 0.43 (1.21) (1.25) (1.20) 0.40 0.60 0.53 (1.18) (1.25) (1.23) 0.43 0.58 0.48 (1.20) (1.25) (1.22) Hybrid (A) = NS, Condition (B) = NS A x B = NS Mean 0.20 (1.09) 0.22 (1.10)

Hybrid JH 3459 PMH 2 Mean CD (p=0.05)

Mean 1.74 (1.65) 1.24 (1.49)

Mean 0.49 (1.22) 0.51 (1.22)

08.05.12 Condition Hybrid JH 3459 PMH 2 Mean CD (p=0.05) FP 0.03 (1.02) 0.10 (1.05) 0.07 (1.03) OT 0.07 (1.03) 0.10 (1.05) 0.08 (1.04) UT 0.00 (1.00) 0.07 (1.03) 0.03 (1.02) Mean 0.03 (1.02) 0.09 (1.04)

Hybrid (A) = NS, Condition (B) = NS, A x B = NS

Figures in parentheses are n+1 transformed values. Set 1: end January sown crop, Set 2: mid February sown crop, Set 3: end February sown crop; LI: Leaf injury, DH: deadhearts; FP: Protection from shoot fly and other insect pests, OT: Protection from insect pests other than shoot fly, UT: No protection for insect pests

74

Figure 7 & 8

75

The maximum temperature had been reported to influence the population of small brown planthopper especially above 35oC due to killing of intracellular yeast like symbiotes (Noda and Saito 1979). The correlation of population counts of small brown planthopper with weather factors were also worked out in the present studies (table 14). The correlation of population counts of L. striatellus was negative with maximum (r= - 0.14 to - 0.52) and mean air temperature (r= - 0.13 to - 0.57) in different sowing times during both the test years except in end January sown crop in 2012, which was might be due to its very low population. The negative correlation was also observed with minimum air temperature during the high population year i.e. 2011 and population decreased as the minimum temperature increased during the month of May. However, in 2012 this correlation was not conclusive may be due to lower population abundance. The present findings were in corroboration with earlier reports of decreased survival of small brown plant hopper with increase in temperature. Zhang et al (2008) also reported that survival of this pest was higher (81-88 %) at 21-27oC, but it decreased to 5 per cent as the temperature increased to 30o C. Moreover, at 30oC the longevity of female adults reduced and they did not lay eggs. The population decline in the present studies in end April and first week of May during 2011 and 2012, respectively might be due to rise of temperature (40o C). The correlation of pest population was positive with R.H. during both the years except in end January sown crop during 2012 (table 14). However, it was significant (r=0.94*) with evening R.H. in end February sown crop during 2011 only, thus indicating the indirect effect of R.H. as it decreased with rise in temperature, which is deleterious to survival of small brown planthopper. The variations in its population over the years at different locations revealed that its occurrence is sporadic and it could be a limiting factor in spring maize cultivation in future, because it is a vector of number of viruses on cereal crops including Maize rough dwarf virus (Caciagli 1991). Thus this pest requires due attention during planning of insect pest management strategies in spring maize. Moreover, detailed studies on its life and seasonal history, host range, carry over, damage potential and influence of biotic and abiotic factors on population dynamics are required in future. Maize stem borer, Chilo partellus (Swinhoe) is a serious threat to maize cultivation worldwide. In India, it was reported to cause 27-85 per cent damage in different agro-climatic zones during kharif season (Jindal and Hari 2008b). However, its incidence was found to be relatively low in spring sown maize in the present studies. The incidence of borer also differed non significantly among hybrids (except a few observations) and crop protection condition at different sowing times during both the test years (Annexure VII).

76

Table 14. The correlations of small rice plant hopper, Laodelphax striatellus (Falln) population counts with weather parameter in spring 2011 and 2012. 2011 Air temperature (C) Set 1 Set 2 Set 3 Max. -0.18 -0.49 -0.52 Min. -0.26 -0.53 -0.60 Mean -0.22 -0.52 -0.57 2012 0.27 -0.30 -0.13 Relative humidity (%) Mor. 0.23 0.41 0.45 Eve. 0.37 0.78 0.94* Mean 0.31 0.57 0.61

0.44 0.49 -0.95 -0.62 0.95 Set 1 -0.28 0.15 0.62 0.33 0.51 Set 2 -0.14 -0.01 0.50 0.16 0.37 Set 3 * Significant at 5 % level. Set 1: end January sown crop, Set 2: mid February sown crop, Set 3: end February sown crop The leaf injury incidence due to C. partellus appeared during the first week of April and its minor damage was observed up to first week of May during both the years in all the sowing times. The adult emergence from carry over population of C. partellus was reported to occur in the fourth week of March (Singh et al 1985), which might have laid eggs on the spring sown maize crop and caused leaf injury symptoms. On end January sown crop the leaf injury incidence of C. partellus differed non significantly among hybrids and protection conditions during both the test years and it was highest during third and second week of April during 2011 and 2012, respectively. During this period it was 3.66 & 1.39 per cent in JH 3459 and 3.10 & 1.86 per cent during 2011 and 2012, respectively. The incidence was numerically lower in shoot fly protected (2.48 and 1.21 %) as compared to unprotected treatment plots (4.33 and 1.92 %), during 2011 and 2012. The non significant interaction of these factors for leaf injury incidence was observed for both the test years. In mid February sown crop again the leaf injury incidence differed non significantly for hybrids and crop protection conditions except in second week of April during 2012, where the differences, irrespective of crop protection conditions was significant among the hybrids, being higher in PMH 2 (2.52 %) than JH 3459 (1.14 %). However, highest incidence was observed in third week of April during both the test years. During this period it was 3.21 & 3.90 per cent in JH 3459 and 2.77 & 3.88 per cent during 2011 and 2012, respectively. The incidence was numerically lower in protected conditions (2.75 %) as compared to unprotected conditions (5.79 %) during 2012, whereas it was 2.99 per cent each in protected/ unprotected conditions in 2011. The interaction of these factors for leaf injury incidence was again non significant for both the test years.

77

In end February sown crop also the leaf injury incidence differed non significantly for hybrids and crop protection conditions except in fourth week of April and first week of May during 2011, where the difference among the hybrids was significant being higher in JH 3459 (5.99 and 3.61 %) than PMH 2 (3.85 and 1.88 %), in respective weeks. Similar to mid February sown crop, the highest incidence was recorded in third week of April during both the test years. During this period it was 9.35 & 2.97 per cent in JH 3459 and 7.80 & 2.75 per cent during 2011 and 2012, respectively. The incidence was numerically lower in protected conditions (2.37 %) as compared to unprotected conditions (3.64 %) during 2012, whereas it was higher (11.65 %) in protected conditions than unprotected conditions (6.42 %) during 2011. The interaction of these factors for leaf injury incidence was again non significant for both the test years. There was no deadhearts formation observed due to C. partellus infestation during study periods revealed that this pest did not cause any economic loss on spring sown maize in the Punjab. The incidence of the other lepidopteran insect pest army worm, Mythimna separata (Walker) was also low and differed non significantly in hybrids and crop protection conditions in crop sown at different times during both the test years (Annexure VIII). Its incidence was observed in second and first fortnight of April during 2011 and 2012, respectively. The incidence varied from 1.00 4.32 & 0.33 1.06 per cent in end January, 0.66 2.71 & 1.04 3.10 per cent in mid February and 0.61 3.20 & 0.34 1.37 per cent in end February sown crop in different treatments during 2011 and 2012, respectively. Thus, occurrence of minor incidence of this pest suggested that armyworms did not seems to pose any potential threat to spring maize cultivation in Punjab. The minor incidence of sucking pest pyrilla, Pyrilla perpusilla Walker was also observed during both the test years (Annexure IX). It differed non significantly in hybrids and crop protection conditions as well as in their interactions in crops sown at different times. Its incidence was observed in second fortnight of April during both the test years. The population count per plant varied from 0.03 0.17 & 0.03 0.13 in end January, 0.03 0.23 & 0.07 0.13 in mid February and 0.03 0.13 & 0.00 0.10 in end February sown crop in different treatments during 2011 and 2012, respectively. The incidence of grasshopper spp. was observed during May in all the crops sown at different times during both the test years (Annexure X). The nymphal counts and per cent plants damage differed non significantly among hybrids and crop protection conditions in both the test years. The nymphal count per plant varied from 0.00 0.10 & 0.03 0.13 in end January, 0.03 0.13 & 0.07 0.10 in mid February and 0.07 0.17 & 0.07 0.17 in end February sown crop in different treatments during 2011 and 2012, respectively. Similarly, the plant damage was also low

78

and it varied from 0.34 1.67 & 0.71 1.75 in end January, 1.33 3.73 & 2.04 2.41 in mid February and 2.42 3.58 & 2.06 3.09 per cent in end February sown crop in different treatments during 2011 and 2012, respectively. The observed lower level of grasshopper population and its minor damage that too on maturing leaves was observed which did not cause any economic loss in spring sown maize crop in the present studies. 4.2.4. Population builds up of beneficials: The population of beneficial insect, coccinellids appeared during the April and their population declined after first week of May during both the test years (Annexure XI). However, it differed non significantly among hybrids and crop protection conditions during both the test years. Different species of family coccinellidae are known for their predatory role against a variety of insect pests, particularly the aphids and sucking pests (Frazer 1988). In the present studies when, the higher counts of coccinellids were observed during 2011, comparatively higher population of small brown planthopper was also prevalent at that time. Two predominant species of coccinellids in Punjab i.e. Coccinella septempunctata (Linnaeus) and Cheilomenes sexmaculatus (Fabricius) were found to prey on small brown planthoppers. In 2011, C. septempunctata population on end January sown crop ranged from 0.43-0.53, 1.13-1.53, 2.003.16, 0.60- 0.73, 0.07- 0.23 per plant in second, third & fourth week of April; and first & second week of May, respectively. At highest population density in last week of April, it was 2.70 and 2.45 per plant in JH 3459 and PMH 2; and it was numerically higher in protected conditions (3.00 per plant) than unprotected conditions (2.35-2.38 per plants). The interaction of these factors was non significant and higher population counts were observed in protected conditions in both the test hybrids. However, the population of C. sexmaculatus was observed only in last week of April. It was 0.78 and 0.62 per plant in JH 3459 and PMH 2, respectively. Similar to C. septempunctata, its population was numerically higher in protected conditions (0.85 per plant) than unprotected conditions (0.58-0.65 per plants). The interactions of hybrids and protected conditions for C. sexmaculatus abundance was non significant. In 2012, only C. septempunctata was observed during the month of April and May. Its population counts were 0.17-0.30, 0.070.17, 0.00-0.17, 0.01-0.10 and 0.07-0.17 during first, second, third & fourth week of April; and first week of May, respectively. In this year, the lower population of coccinellids was in correspondence with lower incidence of small brown planthopper. On mid February sown crop also the population of coccinellids was higher during 2011 and again differed non significantly for hybrids and crop protection conditions. The counts of C. septempunctata ranged from 0.30-0.40, 0.83-1.13, 2.03 2.60, 0.43 - 0.63, 0.07- 0.20 per plant in second, third & fourth week of April; and first & second week of May, respectively. At highest

79

population density in last week of April, it was 2.27 and 2.44 per plant in JH 3459 and PMH 2; and but it was numerically lower in protected conditions (2.28 per plant) than unprotected conditions (2.32-2.47 per plants). The interaction of these factors was also non significant for the abundance of C. septempunctata. Similar to end January sown crop, the population of C. sexmaculatus was observed only in the last week of April. It was 0.47 and 0.50 per plant in JH 3459 and PMH 2, respectively. Similar to C. septempunctata, its population was numerically lower in protected conditions (0.40 per plant) than unprotected conditions (0.52-0.53 per plants). The interactions of hybrids and crop conditions for C. sexmaculatus abundance was non significant. In 2012, only C. septempunctata was observed during the month of April and May. Its population counts were 0.00-0.07, 0.10-0.20, 0.23-0.60 and 0.17-0.37 and 0.03-0.10 during second, third & fourth week of April; and first & second week of May, respectively. In this sowing date again the lower population of coccinellids was in correspondence with lower incidence of small brown planthopper. On end February sown crop again the populations of coccinellids was higher during 2011 and differed non significantly for hybrids and crop protection conditions. The counts of C. septempunctata ranged from 0.17-0.27, 1.08-2.03, 0.47 0.97, 0.23 - 0.63, 0.33- 0.67 per plant in second, third & fourth week of April; and first & second week of May, respectively. At highest population density in third week of April, it was 1.97 and 1.73 per plant in JH 3459 and PMH 2; and it was numerically lower in protected conditions (1.73 per plant) than unprotected conditions (1.83-1.98 per plants). The interaction of these factors was also non significant for the abundance of C. septempunctata. Similar to end January and mid February sown crops, the population of C. sexmaculatus was observed only in the last week of April. It was 0.35 and 0.29 per plant in JH 3459 and PMH 2, respectively. However, its counts were numerically higher in protected conditions (0.35 per plant) than unprotected conditions (0.32-0.30 per plants). The interaction of hybrids and protected conditions for C. sexmaculatus abundance was non significant. In 2012, only C. septempunctata was observed during the month of April and May. Its population counts were 0.03-0.07, 0.10-0.17, 0.27-0.43 and 0.13-0.23 and 0.03-0.13 during second, third & fourth week of April; and first & second week of May, respectively. Similar to other sowing dates, again the lower population of coccinellids was observed in correspondence with lower incidence of small brown planthopper. The decline in population of coccinellids was due to decline in prevalent population of its host i.e. small brown planthopper at that time. Further more in the present studies the coccinellids were able to manage the population of small brown planthopper without any insecticidal application. The present studies, highlights the need of detailed studies

80

on insect biodiversity and their interaction with environmental factor to devise management strategies in future. The population of spiders was observed during April in all the crops sown at different times during study period (Annexure XII). The counts per plant differed non significantly in hybrids and crop protection conditions in both the test years. It varied from 0.17 0.23 & 0.07 0.10 in end January, 0.03 0.10 & 0.07 0.30 in mid February and 0.07 0.10 & 0.03 0.17 in end February sown crop in different treatments during 2011 and 2012, respectively. The very low population of spiders observed in the present studies, indicated that these are of minor importance for management of insect pests in spring sown maize in Punjab. Thus, the study on insect pest incidence on maize sown with different sowing dates in spring season revealed that the shoot fly is the major limiting factor in spring maize cultivation but sporadic incidence of insect pest like small brown planthopper laid the emphasis on regular monitoring of insect pests and their natural enemies in the subsequent years. 4.2.5. Losses in grain yield due to insect pests: In the present studies the incidence of shoot fly, maize stem borer, armyworm, pyrilla, grasshopper and small brown planthopper were recorded, but shoot fly was observed to be the major yield reducing factor during both the test years. Though the higher incidence of small brown planthopper in 2011 was recorded, but it did not cause any economic damage to the crop sown with different sowing times. The grain yield (q/ha) differed significantly under protected/ unprotected conditions against shoot fly during both the test years and among the hybrids in mid February sown crop during 2012 only (table 15). However, the interaction among these two factors was non significant during both the years. During 2011, in end January sown crop the grain yield was 62.24 and 63.10 q/ ha in JH 3459 and PMH 2, respectively. It was significantly higher in protected conditions (77.16 q/ ha) than that in unprotected conditions (54.78-55.04 q/ ha). The interaction of these factors was non significant and both the hybrids performed better under protected conditions. Similarly, in 2012 both the hybrids JH 3459 (50.69 q/ ha) and PMH 2 (52.09 q/ ha) were statistically on par in grain yield. It was again significantly higher under protected conditions (68.79 q/ ha) than that in unprotected conditions (41.57 43.82 q/ ha). The interaction between hybrids and crop protection conditions was non significant. In mid February sown crop during 2011, the grain yield differed non significantly in hybrids and was 57.78 and 57.96 q/ha in JH 3459 and PMH 2, respectively. It was significantly higher in protected conditions (77.67 q/ ha) than that in unprotected conditions (45.30-49.64 q/ ha).

81

Table 15. The grain yield (q/ ha) in different cultivars under protected/unprotected conditions in crops sown at different times at Ludhiana Mean grain yield (q/ ha) 2011 Hybrid JH 3459 PMH 2 Mean CD (p=0.05) FP 75.10 79.22 77.16 Set 1 Condition OT UT 55.04 57.10 55.56 52.43 55.04 54.78 Set 2 Condition OT UT 51.44 48.35 47.83 44.24 49.64 45.30 Set 3 Condition OT UT 43.72 41.67 37.55 35.49 40.64 38.58

Mean 62.24 63.10

FP 73.56 81.79 77.67

Mean 57.78 57.96

FP 63.63 66.36 64.99

Mean 49.67 46.47

Hybrid (A) = NS, Condition (B) = 8.81 A x B = NS Set 1

Hybrid (A) = NS, Condition (B) = 10.65 A x B = NS 2012 Set 2

Hybrid (A) = NS, Condition (B) = 5.02 A x B = NS Set 3

Hybrid JH 3459 PMH 2 Mean CD (p=0.05) FP 65.59 71.98 68.79

Condition OT 44.21 43.43 43.82 UT 42.29 40.84 41.57

Mean 50.69 52.09 FP 68.92 78.85 73.88

Condition OT 46.26 46.73 46.49 UT 42.69 45.69 44.19

Mean 52.63 57.09 FP 62.25 64.22 63.24

Condition OT 37.81 32.52 35.17 UT 36.41 32.25 34.33

Mean 45.49 43.00

Hybrid (A) = NS, Condition (B) = 6.19 A x B = NS

Hybrid (A) = 4.46, Condition (B) = 5.46 A x B = NS

Hybrid (A) = NS, Condition (B) = 4.64 A x B = NS

Set 1: end January sown crop, Set 2: mid February sown crop, Set 3: end February sown crop FP: Protection from shoot fly and other insect pests, OT: Protection from insect pests other than shoot fly, UT: No protection for insect pests

82

Similar to end January sown crop, the interaction of hybrids and crop protection conditions was non significant with both the hybrids performing better under protected conditions. However, in 2012 the grain yield was significantly higher in hybrids PMH 2 (57.09 q/ ha) than that in JH 3459 (52.63 q/ ha). It was also significantly higher under protected conditions (73.88 q/ ha) than that in unprotected conditions (44.19 46.49 q/ ha). However, the interaction between hybrids and crop protection conditions was non significant. In end February sown crop, the grain yield differed non significantly among hybrids during both the test years. In 2011, it was 49.67 and 46.47 q/ ha in JH 3459 and PMH 2, respectively. It was significantly higher in protected conditions (64.99 q/ ha) than that in unprotected conditions (38.58 - 44.64 q/ ha). Similar to earlier observations, the interaction of hybrids and crop protection conditions was non significant. In 2012, the grain yield was 45.49 and 43.00 q/ ha in JH 3459 and PMH 2, respectively. It was significantly higher under protected conditions (63.24 q/ ha) than that in unprotected conditions (34.33 35.17 q/ ha). However, the interaction between hybrids and crop protection conditions was again non significant. The analysis of pooled data of grain yield performed for cultivars under protected/ unprotected conditions against shoot fly attack with different times of sowing revealed that it was maximum in end January and mid February sown crop during 2011 and 2012, respectively (table 16). The gain yield was significantly higher under protected conditions than that under unprotected conditions during both the test years. It was 73.28 & 68.66 q/ ha in protected and 47.75 & 40.93 q/ ha in unprotected conditions during 2011 and 2012, respectively. In 2011, the grain yield being on par with mid February sown crop (62.83 q/ ha) was significantly higher in end January sown crop (66.38 q/ ha) than that in end February sown crop (52.34 q/ ha). Accordingly, the losses were minimum in end January sown crop (27.93 %) followed by mid February sown crop (38.21 %) and were maximum in end February crop (38.99 %). In 2012, the grain yield was significantly more in mid February crop (59.65 q/ ha) than that in end January sown crop (55.76 q/ ha) which was significantly better than end February crop (48.99 q/ ha). As in 2011, the grain yield losses were minimum in end January sown crop (37.97 %) followed by mid February sown crop (38.66 %) and maximum in end February crop (45.04 %). Overall the grain yield losses were 34.84 and 40.39 per cent during 2011 and 2012, respectively. The interaction between crop protection conditions and different sowing times was non significant and the yield was higher in protected conditions in all the crops during both the test years. Similarly, higher grain yield (9.46 q/ ha) in maize crop sown on 15th February as compared to end February (8.28 q/ ha) and March sown crops (5.93- 8.56 q/ ha) was reported by Gagandeep and Kanta (2007b). They further reported that pooled yield over sowing dates was also significantly more in protected (11.36 q/ ha) than that in unprotected conditions (4.75 q/ ha) against shoot fly. Thus, the tune of losses due to shoot fly in present studies emphasizes the need for timely plant protection measures against this pest in spring sown maize.

83

Table 16. The grain yield (q/ ha) pooled over cultivar for comparison of protected and unprotected conditions in crops sown at different times at Ludhiana Mean grain yield (q/ ha) during 2011 Condition Protected Un protected Mean CD (p=0.05) % change (FP-UP/ FP x 100) Condition Protected Un protected Mean CD (p=0.05) % change (FP-UP/ FP x 100) Set 1 77.16 55.61 66.38 Set 2 77.67 47.99 62.83 Set 3 65.02 39.67 52.34 Mean 73.28 47.75

Condition (A) = 5.08, Sowing time (B) = 6.23, A x B = NS 27.93 Set 1 68.82 42.69 55.76 38.21 2012 Set 2 73.93 45.35 59.65 38.99 Set 3 63.23 34.75 48.99 34.84 Mean 68.66 40.93

Condition (A) = 2.43, Sowing time (B) = 2.98, A x B = NS 37.97 38.66 45.04 40.39

Set 1: end January sown crop, Set 2: mid February sown crop, Set 3: end February sown crop FP: Protection from shoot fly, UP: No protection from shoot fly 4.3: Relative susceptibility of different genotypes to shoot fly damage: The experiment for screening of eight maize genotypes i.e. six hybrids (JH 3459, PMH 2, JH 31244, JH 3956, PMH 1 and Parkash) and two inbred lines (LM 16 and CM 143) against Atherigona naqvii was conducted to determine the role of various morphological and biochemical plant characteristics in resistance against shoot fly, Atherigona naqvii during spring 2011 and 2012 and the data for the same have been presented in Table 17-31. 4.3.1 Oviposition by shoot fly, A. naqvii: Oviposition non-preference of shoot fly towards the crop cultivars is one of the important components of host plant resistance. Except at 3 DAG 2011, the mean number of eggs per plant in the test genotypes differed non significantly during all the observations recorded at 3, 6, 9, 12 and 15 DAG during both the test years (table 17). During 2011, the overall mean numbers of eggs per plant were comparatively more at 3 DAG (0.57) than all the remaining observations. At 3 DAG it was maximum in susceptible LM 16 (0.67) which was also on par with remaining susceptible cultivar i.e. Parkash and PMH 2 (0.64). It was minimum in intermediate JH 31244 and least susceptible JH 3956 (0.49) which were also on

84

par with intermediate CM 143 (0.51). However, intermediate PMH 1 (0.53) and least susceptible JH 3459 (0.58) were on par with both the groups. In subsequent observations, the mean egg counts per plant decreased progressively in all the test genotypes and were comparatively low at 15 DAG. Among different genotypes at 6 DAG, 9 DAG, 12 DAG and 15 DAG, it ranged from 0.24 to 0.51, 0.20 to 0.38, 0.18 to 0.36, and 0.11 to 0.18 eggs per plant, respectively. Numerically, on the basis of overall mean of different observations on different test cultivars maximum and minimum number of eggs were observed on genotype LM 16 (0.42) and JH 31244 (0.28), respectively. In 2012, though the differences were non significant, yet the egg counts were comparatively more on all the genotypes irrespective of different observations as compared to 2011. However, the maximum egg counts per plant were observed at 6 DAG (1.33) as against 3 DAG during 2011, which decrease progressively in subsequent observations and again it was minimum at 15 DAG. The egg counts at 3 DAG, 6 DAG, 9 DAG, 12 DAG and 15 DAG, ranged 0.22 to 0.29, 1.04 to 1.53, 0.60 to 0.84, 0.22 to 0.31 and 0.02 to 0.11 respectively. Numerically, on the basis of overall mean of different observations on different test cultivars maximum and minimum number of eggs were laid on genotype on JH 31244 (0.59) and JH 3459 (0.42), respectively as against LM 16 and JH 31244 in 2011. The perusal of data indicated that at peak oviposition time (i.e. at 3 and 6 DAG during 2011 and 2012, respectively) the least susceptible genotypes JH 3956 and JH 3459 received comparatively less eggs per plant (0.49 - 1.49) than susceptible genotypes Parkash and PMH 2 (0.64 - 1.53 per plant). The non significant differences in oviposition of A. naqvii on different test genotypes might be due to high shoot fly pressure (indicated with a maximum of 0.57 and 1.33 eggs/ plant during 2011 and 2012, respectively irrespective of test genotypes) in fish meal baited treatment plots. Dhillon et al (2005 b) also reported that the mechanism of antixenosis for oviposition in sorghum to A. soccata was not stable and tends to breakdown under no-choice conditions. However, the varying preference for oviposition by shoot fly (Atherigona spp.) was reported on maize at Pantnagar and different germplasms were grouped on the basis of number of eggs laid per plant (Sharma and Singh 1975). The observations of Sarup and Panwar (1986), Panwar and Sarup (1988) and Rao and Panwar (1996a) are in line with the present studies as they also reported the non significant differences for oviposition by shoot flies on various maize germplasms. In the present studies, among different morphological and biochemical plant characteristics of different test genotypes with oviposition by A. naqvii, only reducing sugars showed (table 29) positive and significant correlation (r = 0.81*). Singh and Jotwani (1980c) and Singh et al (2004) also reported the positive correlation between oviposition by shoot fly, A. soccata and reducing sugars in sorghum. However, in the present studies, the other morphological and biochemical

85

Table 17. Oviposition by shoot fly, Atherigona naqvii on the different test genotypes in field conditions. Mean egg count per plant during 2011 Genotype 3 DAG JH 3459 PMH 2 JH 31244 JH 3956 PMH 1 PARKASH LM 16 CM 143 Mean CD (p=0.05) Genotype 3 DAG JH 3459 PMH 2 JH 31244 JH 3956 PMH 1 PARKASH LM 16 CM 143 Mean CD (p=0.05) 6 DAG 9 DAG 0.29 (1.13) 0.31 (1.14) 0.27 (1.12) 0.20 (1.09) 0.24 (1.11) 0.22 (1.10) 0.38 (1.17) 0.22 (1.10) 0.27 NS 12 DAG 0.27 (1.12) 0.33 (1.15) 0.18 (1.08) 0.27 (1.12) 0.20 (1.09) 0.27 (1.12) 0.36 (1.16) 0.29 (1.13) 0.27 NS 15 DAG 0.13 (1.06) 0.18 (1.08) 0.11 (1.05) 0.16 (1.07) 0.11 (1.05) 0.13 (1.06) 0.18 (1.09) 0.11 (1.05) 0.14 NS 0.58 (1.26) 0.42 (1.19) 0.64 (1.28) 0.42 (1.19) 0.49 (1.22) 0.36 (1.16) 0.49 (1.22) 0.24 (1.12) 0.53 (1.24) 0.36 (1.16) Mean 0.34 0.38 0.28 0.27 0.29 0.33 0.42 0.30

0.64 (1.28) 0.38 (1.17) 0.67 (1.29) 0.51 (1.23) 0.51 (1.23) 0.36 (1.16) 0.57 0.38 (0.05) NS

Mean egg count per plant during 2012 6 DAG 9 DAG 0.47 (1.21) 0.69 (1.30) 0.84 (1.35) 12 DAG 0.27 (1.12) 0.27 (1.12) 0.31 (1.14) 15 DAG 0.09 (1.04) 0.07 (1.03) 0.11 (1.05) Mean 0.42 0.56 0.59 0.54 0.53 0.57 0.50 0.46

0.22 (1.10) 1.04 (1.13) 0.24 (1.11) 1.53 (1.58) 0.24 (1.11) 1.47 (1.56) 0.27 (1.13) 1.49 (1.57) 0.24 (1.12) 1.44 (1.56) 0.29 (1.13) 1.38 (1.54) 0.29 (1.13) 1.16 (1.47) 0.29 (1.13) 1.11 (1.45) 0.26 NS 1.33 NS

0.69 (1.30) 0.22 (1.10) 0.02 (1.01) 0.73 (1.31) 0.82 (1.35) 0.73 (1.31) 0.60 (1.26) 0.70 NS 0.22 (1.10) 0.27 (1.13) 0.27 (1.13) 0.24 (1.11) 0.26 NS 0.02 (1.01) 0.09 (1.04) 0.07 (1.03) 0.07 (1.03) 0.07 NS

Figures in parentheses are n+1 transformed values; DAG: days after germination

86

parameters of test genotypes were non significantly correlated with the number of eggs laid by shoot fly indicating that these plant characters did not influence the oviposition pattern of A. naqvii (table 29). Rao and Panwar (2001a, b) are also in line with the present findings as they also reported that none of the morphological and biochemical characters influence egg laying by shoot fly, Atherigona spp. on maize at New Delhi. 4.3.2. Incidence of shoot fly, Atherigona spp. The per cent leaf injury and deadhearts differed significantly among the test cultivars at 5, 10, 15 and 20 DAG during both the test years i.e. 2011 and 2012 except the observations recorded at 5 DAG where incidence of deadhearts was on par among different cultivars during both the test years (table 18). At 5 DAG, the per cent leaf injury in different cultivars ranged from 2.36 to 12.33 and from 3.78 to 9.29 per cent in 2011 and 2012, respectively. It was maximum in Parkash (12.33 and 9.29 %, respectively) during 2011 and 2012, being on par with JH 31244 (11.05 %) and PMH 1(10.38 %) in 2011; and PMH 2 (8.61 %) and LM 16 (8.28 %) in 2012. During 2011 and 2012, minimum leaf injury was observed on CM 143 (2.36 %), being on par with LM 16 during 2011 (3.34 %) and in 2012 (3.78 %) which during 2012 was also on par with JH 3956 (4.27 %) followed by JH 3459 (4.79 %), PMH 1 (5.10 %) and JH 31244 (5.70 %). However, the remaining cultivars formed a medium group being on par with either of the two above groups. The deadhearts incidence was on par in all the genotypes and it ranged from 0.00 to 3.08 per cent at 5 DAG during both the test years. At 10 DAG, the per cent leaf injury ranged from 6.23 to 24.44 and 5.31 to 10.91 per cent in 2011 and 2012, respectively. In 2011, JH 31244 (24.41%) being on par with PMH 1(24.41 %) and Parkash (23.68%) had maximum leaf injury incidence. It was lowest in CM 143 (6.23 %), which was further on par with LM 16 (8.96 %). However, during 2012, though the leaf injury was maximum in Parkash (10.91%) but except CM 143 was on par with all the remaining genotypes (7.83-10.14 %). It was minimum in CM 143 (5.31 %), being on par with LM 16 (7.07 %) and further on par with JH 3459. The deadhearts incidence ranged from 5.37 to 12.68 and 2.92 to 7.38 per cent in 2011 and 2012, respectively. In 2011, the deadhearts incidence was maximum in Parkash (12.68 %), being on par with PMH 2 (11.99 %) and LM 16 (10.75 %), and it was minimum in JH 3956 (5.37 %), being on par with JH 3459 (6.35 %) and PMH 1(7.35 %), followed by JH 31244 (7.98 %) and CM 143 (8.14 %). During 2012, the deadhearts incidence was maximum in Parkash (7.38 %), being on par with PMH 2 (7.00 %), which were further on par with PMH 1 (5.88%). It was minimum in CM 143 (2.92 %) which was on par with JH 3956 (3.99 %). However, the remaining genotypes formed a medium group with 4.85 to 5.02 per cent incidence of deadhearts. The data in the table 18 revealed that in general the deadhearts incidence was not in direct proportion to the incidence of leaf injury in the test genotypes during different observations and despite significant differences occurred in infestation i.e. leaf injury and deadhearts incidence in the test genotypes, no clear

87

cut manifestation were found to draw conclusions upto 10 DAG. However, the differential reaction in test genotypes to deadhearts incidence (occurred due to shoot fly incidence) started appearing on 15 DAG and clear cut differential reaction of the test genotypes in terms of deadhearts incidence became evident at 20 DAG. At 15 DAG, the leaf injury incidence ranged from 15.96 to 29.59 and 23.94 to 41.92 per cent in 2011 and 2012, respectively. During 2011, the leaf injury was maximum in JH 31244 (29.59 %) which was further on par with PMH 2 (27.23 %) and Parkash (26.01%). It was minimum in JH 3459 (15.96 %), which was also on par with JH 3956 (22.06%) and further being on par with LM 16 (19.37 %) and CM 143 (17.83 %). However, PMH 1 (23.98 %) except JH 3459 (15.96 %) was on par with both the above groups. During 2012, the maximum leaf injury incidence was in Parkash (41.92 %) being on par with PMH 2 (40.1 %) and JH 31244 (39.27%) further being on par with PMH 1(35.96 %) and it was minimum in CM 143 (23.94 %) being on par with LM 16 (26.52%). However, JH 3459 (30.75 %) and JH 3956 (31.94 %) were on par with both the above groups. The deadhearts incidence ranged from 5.66 to 15.27 and 12.30 to 28.45 per cent in 2011 and 2012, respectively. In 2011, the deadhearts incidence was maximum in PMH 2 (15.27 %), being on par with Parkash and LM 16 (each with 14.65 %), and was followed by JH 31244 (10.98 %) which was further on par with CM 143 (9.77 %) and PMH 1(9.36 %). It was minimum in JH 3956 (5.66 %), which was further on par with JH 3459 (7.34 %). The JH 31244 was on par with other remaining cultivars except PMH 2, Parkash and LM 16 (in first group); and JH 3459 and JH 3959 (in second group). During 2012, the maximum deadhearts incidence occurred in Parkash (28.45 %), being on par with LM 16 (23.08 %) and PMH 2 (22.87 %), were further on par with PMH 1 (18.96 %), and JH 31244 (18.55 %). It was minimum in JH 3459 (12.30 %), which was on par with JH 3956 (13.43%) further being on par with CM 143 (17.75 %). The differences occurred in shoot fly infestation in test genotypes was clear but some genotypes showed differential reaction during different test years. At 20 DAG, the leaf injury incidence ranged from 16.61 to 28.01 and from 19.37 to 38.13 per cent in 2011 and 2012, respectively. During 2011, it was maximum in Parkash (28.01 %) further, being on par with JH 31244 (25.61 %) and PMH 2 (23.56 %) and minimum in JH 3459 (16.61 %), being on par with CM 143 (16.99 %), was further also on par with JH 3956 (19.39 %), LM 16(21.01 %) and PMH 1(21.72 %). Similarly in 2012, per cent leaf injury was maximum in Parkash (38.13 %), being on par with PMH 2 (35.67 %) and JH 31244 (32.51 %), was further also on par with PMH 1(30.23 %). It was minimum in CM 143 (19.37 %) which was also on par with JH 3459 (24.29 %)

88

Table 18. Incidence of shoot fly, Atherigona naqvii on the different test genotypes in field conditions during 2011 and 12 2011 Genotype JH 3459 PMH 2 JH 31244 JH 3956 PMH 1 PARKASH LM 16 CM 143 CD (p=0.05) 2012 Genotype JH 3459 PMH 2 JH 31244 JH 3956 PMH 1 PARKASH LM 16 CM 143 CD (p=0.05) 4.79 8.61 5.70 4.27 5.10 9.29 8.28 3.78 3.53 6.06 5.97 11.05 7.99 10.38 12.33 3.34 2.36 2.72 5 DAG LI (%) DH (%) 0.00 0.00 0.00 0.00 0.20 0.42 0.65 0.22 NS 5 DAG LI (%) DH (%) 0.97 3.08 1.33 2.21 1.17 3.01 1.97 1.10 NS 10 DAG LI (%) 14.09 16.45 24.44 16.65 24.41 23.68 8.96 6.23 6.04 LI (%) 7.83 10.14 9.41 8.60 9.50 10.91 7.07 5.31 3.27 DH (%) 6.35 11.99 7.98 5.37 7.35 12.68 10.75 8.14 3.03 DH (%) 5.02 7.00 4.89 3.99 5.88 7.38 4.85 2.92 1.85 15 DAG LI (%) 15.96 27.23 29.59 22.06 23.98 26.01 19.37 17.83 6.48 LI (%) 30.75 40.14 39.27 31.94 35.96 41.92 26.52 23.94 11.70 DH (%) 7.34 15.27 10.98 5.66 9.36 14.65 14.65 9.77 2.83 DH (%) 12.30 22.87 18.55 13.43 18.96 28.45 23.08 17.75 5.99 20 DAG LI (%) 16.61 23.56 25.61 19.39 21.72 28.01 21.01 16.99 5.77 LI (%) 24.29 35.67 32.51 24.30 30.23 38.13 25.26 19.37 7.37 DH (%) 7.98 17.27 12.32 6.99 10.73 15.65 16.33 11.49 3.32 DH (%) 12.99 25.76 21.08 14.91 19.98 30.49 26.64 18.84 5.00

10 DAG

15 DAG

20 DAG

LI: leaf injury, DH: deadhearts; DAG: days after germination

89

and JH 3956 (24.30 %). However, the leaf injury incidence in LM 16 (25.26 %), except PMH 2 (35.67 %) and Parkash (38.13 %) was on par with remaining cultivars. The deadhearts incidence ranged from 6.99 to 17.27 and from 12.99 to 30.49 per cent in 2011 and 2012, respectively. In 2011, PMH 2 with maximum deadhearts (17.27 %), being on par with LM 16 (16.33 %) and Parkash (15.65), formed a susceptible group. The minimum incidence was observed in JH 3956 (6.99 %) which was on par with JH 3459 (7.98 %) formed least susceptible group. The hybrid PMH1 (10.73 %), CM 143 (11.49 %) and JH 31244 (12.32 %) were intermediate in reaction. During 2012, similar grouping was observed, Parkash (30.49 %) had maximum incidence which was on par with LM 16 (26.64 %) and PMH 2 (25.76 %) which was also on par with JH 31244(21.08 %) form the susceptible group. The hybrid JH 3459 (12.99 %) having minimum deadhearts incidence was on par with JH 3956 (14.91 %) and formed the least susceptible group. The inbred CM 143 (18.84 %) and PMH 1(19.98 %) being on par with each other formed intermediate class. In pooled analysis of 2011 & 2012, the leaf injury and deadhearts incidence at 20 DAG ranged from 18.18 to 33.07 and from 10.49 to 23.57 per cent, respectively (table 19). The maximum leaf injury incidence was in Parkash (33.07 %) followed by PMH 2 (26.62 %) which was further on par with JH 31244 (26.06 %), PMH 1(25.97 %) and LM 16 (23.13%), all being on par with each other. The minimum leaf injury incidence was recorded in CM 143 (18.18 %), which was further on par with JH 3459 (20.45 %) and JH 3956 (21.84 %). The later two cultivars were also on par with LM 16 (23.13 %) which except Parkash (33.07 %) was also on par with the remaining genotypes. The data recorded on leaf injury incidence revealed that except for the maximum susceptible hybrid Parkash, all the remaining genotypes did not give any clear cut differential reaction but it was overlapping in different genotypes. However, the clear cut differences were found in deadhearts incidence in this regard. The deadhearts incidence was maximum in Parkash (23.57 %), being on par with PMH 2 (21.52 %) and LM 16 (21.48 %), they formed the susceptible group. It was minimum in JH 3459 (10.49 %), being on par with JH 3956 (10.95 %), they formed the least susceptible group. The other genotypes i.e. CM 143, PMH 1 and JH 31244 (15.16 - 16.70 %), formed the intermediate group. In the present studies, the overlapping results with respect to incidence of leaf injury might be due to inherent capacity of different cultivars to recover from the leaf injury. However, the differences in deadhearts formation were significant among different cultivars to shoot fly attack. Sharma and Singh (1975) observed that the deadhearts count is easy, quick and helpful in assessing the relative reaction of genotypes as it gives approximate idea about the total loss of plant population and yield. A large number of resistant sources to shoot fly in maize have been identified and categorized by various workers from different locations (Sarup and Panwar 1986, Panwar and Sarup 1986, 1988, Siddiqui et al 1988, Panwar 1995, 2005; Sharma and Panwar, 1998, Jindal et al 2007) to serve as resistant gene (s)

90

against the shoot fly complex. The moderately susceptible reaction of CM 143 might be helpful in resistance breeding as it has been demonstrated that the higher resistance level in inbreds is likely to result in higher level of resistance in the resultant variety/ hybrid (Siddiqui et al 1995). In the present studies also the hybrids JH 3459 and JH 3956 showed relatively less susceptibility to shoot fly infestation i.e. leaf injury and deadhearts incidence and their potential can be exploited in formulation of strategies for the management of this pest for spring sown maize in the Punjab. 4.3.3. Recovery of seedling or tolerance: The seedling survival in sorghum after infestation is related to faster seedling growth rate that provides a better chance to escape deadheart formation (Doggett et al 1970). In the present studies, the per cent total incidence of shoot fly (leaf injury + deadhearts ) differed significantly among genotypes and it ranged from 24.59 to 43.66 and 37.28 to 69.62 per cent in 2011 and 2012, respectively (table 19). During 2011, Parkash had maximum (total) incidence (43.66 %) which being on par with PMH 2 (40.84 %), was also on par with JH 31244 (37.93) and LM 16 (37.34 %). It was minimum in JH 3459 (24.59 %) which, being on par with JH 3956 (26.38 %) and CM 143 (28.49 %), was also on par with PMH 1 (32.15 %). PMH 1, in turn was also on par with LM 16 and JH 31244. Similarly in 2012 incidence was also maximum in Parkash (69.62 %) which was again on par with PMH 2 (61.43%). It was minimum in JH 3459 (37.28 %), being on par with CM 143 (38.21 %) and JH 3956 (39.21%). The incidence was moderate in PMH 1, LM 16 and JH 31244 (50.21- 53.60 %) and all being on par with each other, however, JH 31244 also being on par with PMH 2. The proportion of deadhearts out of total incidence in different genotypes was also worked out for both the test years i.e. 2011 and 2012 to determine the differential levels of tolerance in test genotypes. It ranged from 26.73 to 43.74 and 34.53 to 51.12 per cent in 2011 and 2012, respectively (table 19). In 2011, the proportion of deadhearts out of total incidence being maximum in LM 16 (43.74 %) which was also on par with PMH 2 (42.38 %) and CM 143 (40.76 %) which was further on par with Parkash (35.74 %). It was minimum in JH 3956 (26.73 %) being on par with JH 3459, JH 31244 and PMH 1 (32.49- 32.89 %) which was further on par with Parkash. Parkash was also on par with JH 31244 and JH 3459 which were further on par with JH 3956. During 2012, again the incidence was maximum in LM 16 (51.12 %), being on par with CM 143 (49.42 %), which were further on par with susceptible hybrid Parkash (45.24 %). It was minimum in JH 3459 (34.53 %) which was further on par with JH 3956 (38.17 %), which in turn was on par with JH 31244 (39.21%), PMH 1 (40.06 %) and PMH 2 (42.12 %) which were further on par with Parkash. The pooled data of 2011 & 2012 revealed that proportion of deadhearts out of total incidence was maximum in LM 16 (47.43 %), which was on par with CM 143 (45.09 %) and PMH 2 (42.25 %) showing least tolerance to shoot fly infestation. Whereas, it was minimum in JH 3956 (32.45 %) and being

91

on par with JH 3459 (33.51 %) followed by JH 31244 (35.90 %) and PMH 1(36.47 %) indicating comparative tolerance in these genotypes. The Parkash (40.49 %) being on par with JH 31244, PMH 1, PMH 2 and CM 143 was comparatively intermediate in tolerance to attack of shoot fly. The relative tolerance to A. naqvii infestation in terms of proportion of deadhearts out of total incidence of shoot fly among different genotypes was almost in line with the already grouping i.e. JH 3459 and JH 3956 (least susceptible); CM 143, PMH 1 and JH 31244 (intermediate); and Parkash, PMH 2 and LM 16 (susceptible) made for susceptibility of different cultivars to this pest based on deadhearts incidence. Similar to present studies, Pandey and Sharma (1980) also reported that the differences in increase of deadhearts in subsequent observations among maize germplasms were due to their relative capacity to withstand or recover from the damage of the shoot fly. The per cent increase in deadhearts in subsequent observations was lower in least susceptible genotypes in comparison to susceptible hybrids. They also reported that percentage of deadhearts did not have any relationship with either oviposition preference or maggot counts supports the idea that the resistance to shoot fly is not preferential but is dependent primarily on the degree of tolerance i.e. the ability of the plants to recover from the pest injury. Table 19. The shoot fly, Atherigona naqavii infestation parameters (recorded 20 days after germination) on the different test genotypes in field conditions Pooled mean of 2011 and 2012 LI (%) JH 3459 PMH 2 JH 31244 JH 3956 PMH 1 PARKASH LM 16 CM 143 CD (p=0.05) 20.45 26.62 26.06 21.84 25.97 33.07 23.13 18.18 4.47 DH (%) 10.49 21.52 16.70 10.95 15.35 23.57 21.48 15.16 2.87 Per cent total incidence (LI + DH) 2011 24.59 40.84 37.93 26.38 32.15 43.66 37.34 28.49 7.65 2012 37.28 61.43 53.60 39.21 50.21 69.62 51.89 38.21 9.20 Proportion of dead hearts out of total incidence 2011 2012 Pooled 32.49 42.38 32.59 26.73 32.89 35.74 43.74 40.76 6.74 34.53 42.12 39.21 38.17 40.06 45.24 51.12 49.42 6.26 33.51 42.25 35.90 32.45 36.47 40.49 47.43 45.09 5.19

Genotypes

LI: leaf injury, DH: deadhearts; DAG: days after germination

92

In the present studies, also egg counts of A. naqvii had a non significant correlation (r = 0.43) with per cent deadhearts out of total incidence, but the correlation with leaf injury (r = 0.75*) and deadhearts (r = 0.86*) was positive and significant (table 30). Similar positive and significant correlation between egg laying by A. soccata and deadhearts incidence in sorghum was also reported by Singh and Jotwani (1980a), Patel and Sukhani (1990) and Singh et al (2004). In present studies, the deadhearts incidence was also positively correlated with per cent deadhearts out of total incidence (r = 0.70*), but non significant correlation between leaf injury and per cent deadhearts out of total incidence again proved that different genotypes had differential tolerance to withstand the infestation of A. naqvii. The per cent deadhearts out of total incidence had negative and significant correlation with larval period and adult longevity (r = -0.84* and -0.71*, respectively) but positive and significant with female pupal weight (r = 0.83*) indicating cultivars with short larval period, adult longevity and with more female pupal weight had more deadhearts formation (table 30). It had positive and significant correlation with seedling score (r = 0.84*) thus revealing that more deadhearts formation in less vigorous genotypes (table 29). Seedling vigour and high rate of recovery were reported to be important characteristics of resistant cultivars of sorghum also by Sharma et al (1977) and Dhillon et al (2005b). According to Sharma and Singh (1975) also maize germplasms vigorous in growth in the early stage of growth had less percentage of deadhearts due to quick recovery of plant from the injury by shoot fly. 4.3.4. Expression of antibiosis component of resistance to shoot fly, Atherigona naqvii: Antibiosis component of resistant to shoot fly offers exciting possibilities of exerting pressure against insect feeding and development, resulting in lower survival of larvae on the resistant cultivars (Soto 1974). The studies on antibiosis component of resistance to shoot fly, Atherigona naqvii in the test genotypes after exposing to shoot flies under field conditions were conducted in Maize Entomology Laboratory. The larval and pupal periods; adult longevity ; per cent larval and pupal survival; male and female pupal weights; and fecundity on eight maize genotypes ranged from 17.16 18.17 and 7.28 7.88 ; 6.32 8.44 days; 46.39 57.57 and 60.28 73.22 per cent; 3.00 3.97 and 3.99 4.68 mg; and 5.89 7.67 eggs per female, respectively in 2011; and from 17.19 18.62 and 7.43 8.80 ; 6.76 8.62 days; 46.94 63.87 and 61.80 71.69 per cent; 2.85 4.14 and 3.95 4.79 mg; and 8.00 9.78 eggs per female, respectively in 2012 (table 20). Larval period: The larval period differed significantly when reared in the laboratory on different genotypes during both the test years. In 2011, it was significantly prolonged on least susceptible hybrid JH 3459 (18.17 days) which was also on par with the other least susceptible hybrid JH 3956 (17.89 days); two intermediate JH 31244 (17.78 days) and PMH 1; and one susceptible hybrid PMH 2 (17.72 days in each). It was minimum in susceptible LM 16 (17.16 days) which was also on par with susceptible Parkash (17.22 days) and

93

intermediate CM 143 (17.33 days). The hybrid Parkash and CM 143 were also on par with PMH 1 and PMH 2. During 2012, larval period was maximum on JH 31244 (18.62 days) being on par with JH 3459 (18.57 days) and both were further on par with PMH 1(18.24 days), JH 3956 (18.14 days). and PMH 2 (18.10 days). It was minimum in CM 143 (17.19 days) which was also on par with LM 16 (17.48 days) and Parkash (17.52 days). However, PMH 2 was on par with all the cultivars. Though the larval periods were overlapping in most of the cultivars during both the test years, yet in general, the larval period was comparatively short in susceptible hybrid Parkash and susceptible inbred LM 16 and intermediate CM 143; and it was longer in least susceptible JH 3459 suggesting differential larval development in cultivars with varied susceptibility. Pupal period: The pupal period differed non significantly among test genotypes in both the test years, being maximum on PMH 1 (7.88 and 8.80 days) and minimum on CM 143 (7.28 and 7.43 days), in the respective years. Adult longevity: The adult longevity differed significantly only in 2011. The flies lived for significantly longer period on least susceptible hybrid JH 3459 (8.44 days). It was minimum in susceptible LM 16 (6.32 days) being on par with intermediate CM 143 (6.45 days) and both were further on par with susceptible Parkash (6.77 days), intermediate PMH 1 (6.91 days) and susceptible PMH 2 (6.95 days). The adult longevity, being on par with each other was intermediate in JH 3956 (7.32 days) and JH 31244 (7.50 days) which was further on par with Parkash, PMH 1 and PMH 2. Though in 2012, the differences were non significant yet numerically it was more on same least susceptible hybrid JH 3459 (8.62 days) and less on intermediate CM 143 (6.76 days). Thus again suggesting that in general the adult longevity was more on least susceptible hybrid JH 3459 but fecundity was more in this cultivar. Larval survival: Though the larval survival differed non significantly during both the test years, yet it was numerically maximum (57.57 and 63.87 %) in susceptible hybrid Parkash during both the test years and minimum in moderately tolerant hybrid JH 3956 (46.39 & 52.92 %) and JH 3459 (47.17 & 46.94 %) in 2011 and 2012, respectively. The other susceptible genotypes i.e. PMH 2 (51.04 and 55.41%) and LM 16 (54.31 and 59.23 %) also had more larval survival than the moderately tolerant genotypes in respective years. Pupal survival: The pupal survival differed non significantly, indicating no specific interaction with host plant reaction of different test genotypes. It was maximum in JH 31244 (73.22 and 71.69 %) and minimum in PMH 1 (60.28 %) and JH 3459 (61.80 %) in 2011 and 2012, respectively. Pupal weight: The female pupae of A. naqvii were heavier than the male pupae on all genotypes but differed non significantly in weight during both the test years. However, the male pupal weight differed significantly during both the test years. In 2011, male pupal weight was maximum in susceptible hybrid PMH2 (3.97 mg) being on par with PMH 1 (3.92

94

mg) and both were further on par with susceptible LM 16 (3.79 mg) and intermediate CM 143 (3.63 mg). Male pupal weight was minimum in Parkash (3.00 mg), being on par with JH 3956 (3.08 mg) and both were further on par with least susceptible JH 3459 (3.25 mg). JH 31244 (3.52 mg) was on par with some of the treatments of both the groups. Similarly in 2012, it was maximum in susceptible PMH 2 (4.14 mg) being on par with susceptible LM 16 (4.02 mg) and minimum insusceptible Parkash (2.85 mg). However, it was intermediate on genotypes JH 3459, PMH 1, CM 143 and JH 31244 (3.44-3.63 mg), being on par with each other. The female pupal weight was numerically more on susceptible genotypes LM 16 (4.68 and 4.79 mg) followed by PMH 2 (4.44 and 4.74 mg) during both the test years. However, It was comparatively less on least susceptible hybrids JH 3459 (3.99 mg) and JH 3956 (3.95 mg) in 2011 and 2012, respectively. Thus provides an evidence of antibiosis in spring maize cultivars to A. naqvii. Fecundity: The fecundity per female reared on different genotypes differed significantly during 2011 alone. The female laid maximum eggs when reared on susceptible cultivar Parkash (7.67) which was also on par with JH 3956 (7.33) and PMH 2 (7.45) and minimum on PMH 1 (5.89) which was on par with CM143 (6.11). However the remaining cultivars formed intermediate group. In 2012, though the differences in egg laying were non significant, yet numerically almost the same trend was observed in different test cultivars. It was comparatively more on susceptible genotypes PMH 2, Parkash and LM 16 (9.00-9.78) and comparatively less on intermediate JH 31244, PMH 1, CM 143 and least susceptible JH 3459 (8.00-8.11). Retardation of larval development, prolongation of larval and pupal periods; and reduction in larval/ pupal survival on the resistant genotypes provides an evidence of antibiosis to shoot fly A. soccata in sorghum (Singh and Jotwani 1980d, Raina et al 1981, Sharma et al 1997, Dhillon et al 2005a). Singh and Jotwani (1980d) also observed prolonged larval and pupal periods on the resistant varieties viz., IS 1054, IS 5469 and IS 5490 than that on susceptible variety Swarna in sorghum. They further reported that larval weight, larval length, pupal weight, pupal length and weight of the flies were higher on susceptible variety as compared to resistant ones. Similarly, resistant genotypes SFCR 151, ICSV 705, SFCR 125 and IS 18551 exhibited longer larval period (10.111.0 days as compared to 9.3 days on Swarna); and lower larval survival and adult emergence (54.767.8 and 46.752.2% as compared to 73.3 and 60.6% on Swarna, respectively) than the susceptible check in sorghum as reported by Dhillon et al (2005 a). In the present studies also, in general the susceptible genotypes, Parkash, LM 16 and PMH 2 had less larval period, more larval survival, more male and female pupal weight, more fecundity thus indicating the existence of antibiosis

95

Table 20. Expression of antibiosis component of resistance to shoot fly, A.naqvii in the different test genotypes under laboratory conditions 2011 Genotypes Larval period (days) 18.17 17.72 17.78 17.89 17.72 17.22 17.16 17.33 0.62 Larval period (days) 18.57 18.10 18.62 18.14 18.24 17.52 17.48 17.19 0.91 Pupal period (days) 7.50 7.66 7.61 7.50 7.88 7.55 7.39 7.28 NS Pupal period (days) 8.10 7.76 8.24 8.05 8.80 7.87 7.76 7.43 NS Adult longevity (days) 8.44 6.95 7.50 7.32 6.91 6.77 6.32 6.45 0.77 Adult longevity (days) 8.62 7.83 7.14 7.31 7.39 7.26 6.92 6.76 NS Larval survival (%) 47.17 51.04 47.28 46.39 50.49 57.57 54.31 50.12 NS Larval survival (%) 46.94 55.41 50.93 52.92 51.39 63.87 59.23 55.87 NS Pupal survival (%) 65.22 62.23 73.22 72.59 60.28 66.67 64.46 65.49 NS Pupal survival (%) 61.80 62.64 71.69 65.28 63.54 65.56 68.50 67.37 NS Male pupal wt. (mg) 3.25 3.97 3.52 3.08 3.92 3.00 3.79 3.63 0.38 Male pupal wt. (mg) 3.44 4.14 3.63 3.15 3.46 2.85 4.02 3.55 0.28 Female pupal wt. (mg) 3.99 4.44 4.18 4.02 4.30 4.16 4.68 4.39 NS Female pupal wt. (mg) 4.13 4.74 4.46 3.95 4.09 4.15 4.79 4.28 NS Fecundity

JH 3459 PMH 2 JH 31244 JH 3956 PMH 1 PARKASH LM 16 CM 143 CD (p=0.05) 2012 Genotypes

6.44 7.33 7.00 7.45 5.89 7.67 6.67 6.11 1.01 Fecundity

JH 3459 PMH 2 JH 31244 JH 3956 PMH 1 PARKASH LM 16 CM 143 CD (p=0.05)

8.11 9.78 8.00 8.78 8.00 9.33 9.00 8.11 NS

96

mechanism in addition to tolerance in least susceptible cultivars i.e. JH 3459 and JH 3956. The higher larval survival in susceptible group i.e. LM 16, PMH 2 and Parkash might have resulted in comparative higher deadhearts formation and especially in higher proportion of deadhearts out of total incidence in these genotypes. More larval period, less larval survival, less pupal weight and lower fecundity was also observed in least susceptible cultivars JH 3459 and JH 3956. Further more the percent deadhearts also exhibited the positive and significant correlation with larval survival (r = 0.86*) indicating more larval survival related with more deadhearts formation (table 30). However, it showed non significant but negative correlation with larval period, pupal period, adult longevity and pupal survival; and the correlation was also positive with fecundity, male and female pupal weight, respectively. The observations of Singh and Jotwani (1980d) are in line with the present studies as they have also reported the negative correlation of deadhearts with larval (r = - 0.88*) and pupal period (r = - 0.75*) for A. soccata in sorghum i.e. these periods were short in the susceptible variety but per cent dead hearts were high. The above findings indicated that the shoot fly resistance observed under field conditions might be due to combined effects of at least two mechanisms. However, the factors influencing differential development of shoot fly on different genotypes needs to study in more details in more diverse arrays of maize genotypes indicating different levels of resistance to shoot fly. 4.3.5. Variation in morphological traits in relation to expression of resistance to shoot fly, A. naqvii: The various morphological traits of the seedlings i.e. leaf glossiness, seedling vigour, pink pigmentation of leaf sheath, leaf surface wetness; and length, angle and number of trichomes were recorded at 5th leaf stage during 2011 and 2012.The role of variation in morphological traits in different genotypes in relation to resistance to shoot fly, A. naqvii is discussed as under: Leaf glossiness: Leaf glossiness score ranging from 2.33 to 3.50 and 2.50 to 3.33 differed non significantly during 2011 and 2012, respectively (table 21). The leaf glossiness was reported to be associated with shoot fly resistance in sorghum (Dhillon et al 2005 a), the leaf glossiness (pale green and shiny leaves) at seeding stage probably has a strong influence on the orientation of shoot fly, A. soccata females due to reflection of light in sorghum. But the present studies on maize are not in line with the above finding as the correlation of glossiness with egg laying was negative and non significant (r= -0.14) and both were unrelated as least glossy hybrid JH 31244 received minimum and maximum number of eggs in 2011 and 2012, respectively (table 29). This might be due to differences in the host crop and oviposition sites of A. soccata and A. naqvii as A. soccata lays eggs on leaves of plants but A. naqvii lays eggs in cracks and crevices and on the basal portion of the plant stem. Seedling vigour: Seedling vigour scores ranging from 1.33 to 3.67 and from 2.00 to 3.83 differed significantly during 2011 and 2012, respectively (table 21). During 2011, the

97

seedling vigour score was minimum in JH 3956 (1.33) and on par with Parkash (1.67), both were further on par with JH 31244 (1.83), JH 3459 and PMH 1 (2.33). However, it was maximum in CM 143 (3.67) which was on par with LM 16 (3.17) and PMH 2 (2.83); which in turn were also on par with JH 3459 and PMH 1 both being on par with the remaining cultivars also. In 2012, minimum score was observed in JH 3459 (2.00) which was on par with JH 3956 (2.17), JH 31244, PMH 1 and Parkash (2.33). It was maximum in LM 16 (3.83), which was further on par with CM 143 (3.50). However, PMH 2 (2.83) was on par with both the above groups. Among the hybrids, the higher seedling vigour score i.e. low seedling vigour in susceptible hybrid PMH 2 during both the test years indicated that seedling vigour is related to tolerance in maize genotypes against shoot fly. The per cent deadhearts out of total incidence had positive and significant correlation with seedling vigour score (r = 0.84*) i.e. negatively correlated with seedling vigour (table 29). A negative correlation between seedling vigour with egg counts and deadhearts formation has also been reported by Kamatar and Salimath (2003) in sorghum. However, preference for oviposition and higher deadheart incidence in the genotypes with high seedling vigour were reported by Dhillon et al (2005a) and Chamarthi et al (2011) in sorghum. This may be due to differences in crop geometry, growing season and different environment under study. Leaf sheath pigmentation: The leaf sheath pigmentation scores ranging from 1.5 to 3.50 and 2.50 to 3.33 during 2011 and 2012, respectively differed significantly in 2011 only (table 21). During 2011, the pigmentation score was minimum in JH 3459 (1.50) which was further on par with PMH 2 (2.33) and JH 3956 (2.50). It was maximum in LM 16 (3.50) which was on par with JH 31244 (3.33), PMH1, Parkash, CM 143 (3.00 each) and JH 3956 (2.50). JH 3956 was also on par with JH 3459. In 2012, differences were non significant but the leaf sheath pigmentation score was minimum in JH 3459 (2.50) and maximum in CM 143 (3.33). The JH 3459 had more leaf sheath pigmentation whereas the other least susceptible hybrid JH 3956 had comparatively higher pigmentation score i.e. have less leaf sheath pigmentation. Moreover, in the present studies the correlation between oviposition and pigmentation in the test genotypes was also non significant. Therefore, the pigmentation did not influence egg laying by A. naqvii in the present studies. However, according to Dhillon et al (2005a) the leaf sheath pigmentation in sorghum was found to be associated with its resistance to shoot fly, A. soccata with correlation coefficient of 0.76 with respect to pigmentation scores and oviposition. This might be due to the differences in the sorghum crop physiology i.e. reflection of light from the leaf surface and leaf surface chemicals that influence the predominant shoot fly, A. soccata preferring leaf surface for oviposition on sorghum plant. Leaf surface wetness: Leaf surface wetness (LSW) ranging from 2.17 to 4.17 and 2.17 to 3.67 differed significantly during 2011 and 2012, respectively (table 21). In 2011, the LSW score was minimum in LM 16 and CM 143 (2.17 each), being on par with Parkash (2.50). It

98

was maximum in PMH 2 (4.17), being on par with all the remaining hybrids (3.67-3.83). During 2012, among the hybrids, it was significantly less in Parkash (2.17), which was on par with inbred lines LM 16 (2.33) and CM 143 (2.67). The inbred line CM 143 was also on par with hybrid JH 3459 (3.17), which in turn was on par with all the remaining hybrids (3.333.67). The PMH 2 and Parkash, both being susceptible to A. naqvii, but higher and lower LSW score indicated that the LSW of genotypes did not contributes towards resistance to A. naqvii. Furthermore, in the present studies the LSW score was non significantly correlated with deadhearts and their proportion out of total incidence (table 29). However, the leaf surface wetness was reported to be positively associated with shoot fly susceptibility in sorghum and had significantly positive correlation (r= 0.85) with deadhearts percentage (Nwanze et al 1992, Dhillon et al 2005a, Chamarthi et al 2011). The differences in results might be due to the predominant shoot fly species and crop growth season prevailing during the period of study. Leaf trichome: The maize genotypes possessed unbranched, slender, non-glandular trichomes with pointed tips that occurred singly on the leaf surface. The data pertaining to leaf trichome characteristics of different maize genotypes presented in table 21 revealed that the mean trichome length ranging from 35.33 m to 102.83 m and 38.30 m to 90.27 m differed significantly during 2011 and 2012, respectively. In 2011, JH 31244 (35.33 m) possessed shortest mean trichome length, being on par with PMH 2 (36.24 m), they were further on par with JH 3459 (51.28 m). It was maximum in Parkash (102.83 m) which was further on par with JH 3956 (86.14 m). JH 3956 was further on par with LM 16 (84.26 m) and PMH 1 (83.88 m). During 2012, shortest trichome length was observed in susceptible genotypes PMH 2 (38.30 m) which was on par with an intermediate hybrid, JH 31244 (40.88 m) but was significantly more in another susceptible hybrid i.e. Parkash (90.27 m). Thus, the trichome length of genotypes (with variable susceptibility levels) was not found associated with their reaction to A. naqvii. The correlation of trichome length with shoot fly oviposition and damage was also found non significant in the present studies (table 29).

The mean trichome angle among different genotypes ranging from 49.67 to 74.75 and 51.33 to 72.33 also varied significantly during 2011 and 2012, respectively (table 21). In 2011, the susceptible genotype Parkash was found to have minimum trichome angle (49.67), which was further on par with JH 31244 (53.89 ), CM 143 (57.33) and LM16 (64.00). JH 3956 (74.75 ) had maximum trichome angle being on par with PMH 1 (73.33), PMH 2 (71.25 ) both further being on par with JH 3459 (64.58) and all being on par with LM 16 also. In 2012, the trichome angle was again minimum in Parkash (51.33 ) being on par with CM 143 (53.33) and both were on par with JH 31244 (56.83). The trichome angle was maximum in JH 3956 (72.33 ) which was further on par with PMH 1 (68.00 ) and PMH 2 (67.67 ). As observed for trichome length, the trichome angle in different genotypes was

99

Table 21. Variation in morphological traits of the test maize genotypes at 5th leaf stage under field conditions during 2011 and 2012 2011 Genotypes Leaf glossiness (1-5)* 3.00 2.33 3.50 2.83 2.83 3.00 3.00 3.33 NS Leaf glossiness (1-5)* 2.83 2.50 3.33 2.67 3.00 3.33 3.17 3.00 NS NS Seedling vigour (1-5)* 2.33 2.83 1.83 1.33 2.33 1.67 3.17 3.67 1.15 Seedling vigour (1-5)* 2.00 2.83 2.33 2.17 2.33 2.33 3.83 Leaf sheath pigmentation (1-5)* 1.50 2.33 3.33 2.50 3.00 3.00 3.50 3.00 1.07 Leaf sheath pigmentation (1-5)* 2.50 2.83 3.00 3.00 2.67 2.83 3.17 3.50 0.79 Leaf surface wetness (1-5)* 3.67 4.17 3.67 3.67 3.83 2.50 2.17 2.17 0.65 Leaf surface wetness (1-5)* 3.17 3.50 3.67 3.33 3.50 2.17 2.33 2.67 0.62 Trichome Trichome length angle (m) 51.28 64.58 36.24 35.33 86.14 83.88 102.83 84.26 62.97 18.16 71.25 53.89 74.75 73.33 49.67 64.00 57.33 14.56 No. of Trichome per 200 m length 4.67 5.00 3.00 3.67 2.67 1.67 2.00 3.00 0.94 No. of Trichome per 200 m length 4.00 3.00 3.33 3.00 2.67 2.67 2.67 3.33 0.81

JH 3459 PMH 2 JH 31244 JH 3956 PMH 1 PARKASH LM 16 CM 143 CD (p=0.05) 2012 Genotypes

JH 3459 PMH 2 JH 31244 JH 3956 PMH 1 PARKASH LM 16 CM 143 CD (p=0.05)

Trichome Trichome length angle (m) 53.13 64.33 38.30 40.88 64.07 80.97 90.27 73.97 58.73 8.06 67.67 56.83 72.33 68.00 51.33 63.33 53.33 7.30

*Rating as per Dhillon et al ( 2005 a)

100

also not found to be associated with their reaction to A. naqvii. Thus, the present findings rules out the utility of trichome angle as morphological marker for the selection of resistance genotypes to shoot fly in breeding programmes. Non significant correlation of trichome angle with A. naqvii oviposition and damage was (table 29). The number of trichome per 200 m linear length ranging from 1.67 to 5.00 and 2.67 to 4.00 varied significantly among the test genotypes during 2011 and 2012, respectively (table 21). In 2011, the susceptible genotype Parkash possessed minimum trichome number 1.67 per 200 m linear length which was further on par with LM 16 (2.00/ 200 m linear length). It was maximum in PMH 2 (5.00/ 200 m linear length), being on par with JH 3459 (4.67 per 200 m linear length). Similarly in 2012, it was minimum in Parkash, PMH 1 and LM 16 (2.67/ 200 m linear length each) being on par with JH 3956, PMH 2 (3.00/ 200 m linear length each), which was further on par with JH 31244 and CM 143 (3.33 each). However, it was maximum in JH 3459 (4.00) which was further on par with JH 31244 and CM 143. The correlation of trichome number with A. naqvii oviposition and damage was also non significant (table 29). But, trichome density was reported to had a positive correlation

with resistance to shoot fly in maize. According to Rao C N and Panwar (2001a) the shoot fly (A. naqvii & A. soccata) resistant varieties of maize possessed higher trichome density on leaves with maximum and minimum of 3.21 and 2.14 trichomes per mm2 in resistant Antigua Gr. 1 and CM 300, respectively at New Delhi. However, they did not mention the relative abundance of the two species. In sorghum also, number of eggs per plant and per cent plants with eggs were negatively correlated with trichome density when observed 14 days after emergence (Maiti and Gibson 1983, Patel and Sukhani 1990, Dhillon et al 2005a). In present studies, PMH 2 (susceptible) and JH 3459 (least susceptible) possessing comparatively higher number of trichomes indicated that trichome density did not relate their susceptibility to A. naqvii. This might be due to predominance of A. naqvii which lays eggs on soil in cracks and crevices near the plant and on the basal stem portion. Leaf length: Significant variation was recorded in the leaf length of plants of different maize genotypes in healthy and infested conditions (table 22). The mean leaf length of different genotypes during 2011 ranged from 11.24 to 19.02 cm and 7.53 to 14.24 cm in healthy and infested plants, respectively. The leaf length of healthy plant was significantly less in inbred CM 143 (11.24 cm) than that in other susceptible inbred LM 16 (14.86 cm). Among least susceptible hybrids it was minimum in least susceptible JH 3459 (15.60 cm), which was further on par with JH 3956 (16.64 cm) which was in turn was on par with JH 31244 (16.80 cm) and PMH 1 (17.36 cm). It was maximum in susceptible hybrid PMH 2 (19.02 cm) being on par with other susceptible hybrid Parkash (18.82 cm). The leaf length of infested plants was minimum in susceptible inbred LM 16 (7.53 cm) with maximum per cent reduction of 49.33 per cent over healthy ones. Among hybrids leaf length of infested plants was minimum

101

in susceptible hybrid Parkash (10.13 cm) with maximum per cent reduction of 46.17 per cent over healthy ones. Leaf length of infested plants was maximum in least susceptible hybrid JH 3956 (14.24 cm) with minimum per cent reduction (14.42) over healthy plants. In 2012, the mean leaf length of different genotypes ranging from 11.56 to 17.42 cm and 8.55 to 12.40 cm in healthy and infested plants, respectively differed significantly. As in 2011, the length of healthy leaves was lower in inbred CM 143 (11.56 cm) than the susceptible inbred LM 16 (14.76 cm). Among hybrids again it was minimum in the second least susceptible hybrid JH 3956 (15.85 cm), being on par with PMH 1 (15.93 cm), JH 3459 (16.27 cm). However, again as in 2011, it was maximum in susceptible hybrid PMH 2 (17.42 cm) being on par with other susceptible hybrid Parkash (17.36 cm). The mean leaf length of infested plant was minimum in inbred LM 16 (8.55 cm) with maximum per cent reduction in length (42.07) over the healthy plant. Among hybrids, the infested leaf length was minimum in one susceptible hybrid PMH 2 (10.50 cm) with maximum per cent reduction (39.72) over healthy plant. Infested plant length was maximum in one least susceptible hybrid JH 3956 (12.40 cm) with minimum per cent reduction of 21.77. The lowest per cent reduction in leaf length, under infested conditions, in case of least susceptible cultivar JH 3956 during both the test years provided an evidence of its tolerance to A. naqvii infestation. The more leaf length in healthy plants of A. naqvii susceptible hybrids Parkash and PMH 2, than that in above least susceptible hybrids during both test years revealed that leaf length also seems to be linked with the susceptibility in these cultivars. Leaf width: The leaf width of maize genotypes also showed significant variation under healthy and infested conditions (table 22). The mean leaf width of different genotypes during 2011 ranged from 1.68 to 2.61 cm and from 0.82 to 1.90 cm in healthy and infested plants, respectively. The leaf width in healthy plants was minimum in less susceptible inbred CM 143 (1.68 cm) than the susceptible inbred LM 16 (1.98 cm). Among hybrids it was minimum in one least susceptible JH 3459 (2.11 cm) which was further on par with JH 3956 (2.24 cm). It was maximum in one susceptible hybrid Parkash (2.61 cm) being on par with the remaining susceptible hybrid PMH 2 (2.59 cm). The mean leaf width of infested plant was minimum in inbred LM 16 (0.82 cm) with maximum per cent reduction (58.59) in leaf width over the healthy one. Among hybrids, the leaf width of infested plants was minimum in one

susceptible hybrid Parkash (1.28 cm) with per cent reduction of 44.40 over the healthy one. Infested plant width was maximum in least susceptible hybrid JH 3956 (1.90 cm) with minimum per cent reduction (15.18). In 2012, the mean leaf width of different genotypes ranged from 1.76 to 2.50 cm and 1.20 to 1.91 cm in healthy and infested plants, respectively. The leaf width of healthy plants was minimum in inbred CM 143 (1.76 cm) being on par with the remaining susceptible inbred LM 16 (1.87 cm). Among hybrids as in 2011, it was

102

minimum in least susceptible JH 3956 (2.12 cm), being on par with JH 3459 (2.18 cm) and both further being on par with PMH 1 (2.29 cm). It was maximum, again as in 2011, in susceptible hybrid Parkash (2.50 cm) which was further on par with JH 31244 (2.45 cm) and PMH 2 (2.39 cm). As in 2011, the leaf width of infested plants was again minimum in the inbred LM 16 (1.20 cm) with maximum per cent reduction (35.83) in length over the healthy ones. Among hybrids infested leaf width was minimum in susceptible hybrid PMH 2 (1.64 cm), being on par with all the hybrid (on par with each other 1.65-1.73 cm) expect JH 31244 (1.91 cm) where it was significantly more. However, as in 2011, the per cent reduction was again maximum in the susceptible hybrid Parkash (34.00%) and minimum in JH 3956 (20.75 %). The lowest per cent reduction in leaf width, under infested conditions, in case of least susceptible cultivars JH 3956 and JH 3459 during both the test years again provided an evidence of their tolerance to A. naqvii infestation. The more leaf width in healthy plants of A. naqvii susceptible hybrid Parkash, than that in above least susceptible hybrids during both test years revealed that leaf width also linked with the susceptibility in this cultivars. Leaf area: The leaf area of healthy and infested plants of different maize genotypes varied significantly during 2011 and 2012 ( table 22). In 2011, the mean leaf area of different genotypes ranged from 17.50 to 33.49 cm2 and 7.93 to 16.84 cm2 in healthy and infested plants, respectively. The leaf area of healthy plants of inbred line CM 143 (17.50 cm2) was significantly lower than the remaining inbred LM 16 (21.59 cm2). Among hybrids least susceptible JH 3956 (25.93 cm2) has minimum leaf area being on par with JH 3459 (26.16 cm2). The hybrid Parkash (33.49 cm2) had maximum leaf area than all the remaining hybrids (28.73- 31.04 cm2). The leaf area of infested plants was minimum in inbred line LM 16 (7.93 cm2) with maximum per cent reduction (63.27) over the healthy one. Among hybrids the leaf area of infested plants was minimum in susceptible Parkash (10.96 cm2) with maximum 67.27 per cent reduction over the healthy one. Leaf area of infested plants was maximum in least susceptible hybrid JH 3956 (16.84 cm2) with minimum per cent reduction (35.06) over healthy ones. In 2012 the leaf area ranged from 16.97 to 32.77 cm2 and 9.31 to 15.75 cm2 in healthy and infested plants, respectively. As in 2011, the leaf area was minimum in less susceptible inbred line CM 143 (16.97 cm2) than the remaining susceptible inbred LM 16 (21.10 cm2). Among hybrids the leaf area was minimum in least susceptible hybrid JH 3956 (27.43 cm2), being on par with JH 3459 (27.85 cm2). The leaf area was maximum in susceptible hybrid Parkash (32.77 cm2) which was further on par with other susceptible hybrid PMH 2 (31.97 cm2). As in 2011, the leaf area of infested plant was again minimum in inbred LM 16 (9.31 cm2) with per cent reduction of 55.88 over the healthy one. Among hybrids leaf area of infested plants was minimum in PMH 2 (13.87 cm2) with maximum reduction of 56.62 per cent over healthy one. However, as in 2011 the leaf area was again

103

maximum in least susceptible hybrid JH 3956 (15.75 cm2) with minimum per cent reduction (42.58) over healthy ones. The lowest per cent reduction in leaf area, under infested conditions, in case of least susceptible cultivars JH 3956 during both the test years again provided an evidence of its tolerance to A. naqvii infestation. The higher per cent reduction in leaf area of susceptible hybrid Parkash and PMH 2 indicated that more leaf area is related to susceptible reaction of these genotypes to shoot fly infestation. Stem Girth: The stem girth (circumference) of healthy and infested plants differed significantly among test maize genotypes during 2011 and 2012 (table 22). In 2011, the mean stem girth of different genotypes ranged from 1.87 to 2.83 cm and 1.54 to 2.51 cm in healthy and infested plants, respectively. The stem girth of healthy plants was minimum in inbred CM 143 (1.87 cm) which was also on par with LM 16 (1.96 cm). Among hybrids, it was minimum in least susceptible hybrid JH 3956 (2.04 cm), being on par with JH 3459 (2.07 cm). It was maximum in susceptible hybrid PMH 2 (2.83 cm) followed by PMH 1 (2.61). It was intermediate and on par with each other in hybrids JH 31244 (2.34 cm) and Parkash (2.44 cm). The stem girth of infested plants was minimum in susceptible inbred LM 16 (1.54 cm) with per cent reduction of 21.43 over healthy ones. Among hybrids stem girth of infested plants was minimum in susceptible hybrid Parkash (1.85 cm) with maximum per cent reduction of 24.18 over healthy ones. Stem girth of infested plants was maximum in other susceptible hybrid PMH 2 (2.51 cm) with per cent reduction of 11.31 over healthy plants. During 2012, stem girth of different genotypes ranged from 1.95 to 2.67 and 1.63 to 2.23 cm in healthy and infested plants, respectively. The stem girth of healthy plants of both the inbreds i.e. LM 16 (1.95 cm) and CM 143 (1.97 cm) was on par with each other. Among the hybrids it was minimum in least susceptible hybrid JH 3956 (2.13 cm) and maximum in susceptible hybrid PMH 2 (2.67 cm). It was intermediate and on par with each other (2.35 2.41cm) in the remaining hybrids. The stem girth of infested plants was minimum in CM 143 (1.63 cm) with per cent reduction of 17.26 over healthy ones. Among hybrids stem girth of infested plants was minimum in susceptible hybrid Parkash and least susceptible hybrid JH 3956 (1.99 cm each) with per cent reduction of 17.43 and 6.57, respectively over healthy ones. Stem girth of infested plants was maximum in other least susceptible hybrid JH 3459 (2.23 cm) with least per cent reduction of 5.11 over healthy plants. The data revealed that susceptible hybrids had more stem girth. Similarly, Rao and Panwar (2001c) reported that resistant variety of maize Antigua Gr. 1 (5.57 mm) possessed less stem girth as compared to susceptible CM 300 (8.68 mm) and ascribed that thick stem in susceptible varieties containing juicy parenchymatous tissues might be providing better medium for development of shoot fly maggots. Patel and Sukhani (1990) also reported positive correlation of stem thickness with shoot fly, A. soccata infestation in sorghum.

104

Table 22. Variation in leaf and stem traits of the test genotypes at 5th leaf stage under field conditions during 2011 and 2012 2011 Leaf length (cm) Leaf width (cm) Leaf area (cm2) Stem girth (cm) Cultivar JH 3459 PMH 2 JH 31244 JH 3956 PMH 1 PARKASH LM 16 CM 143 CD (p=0.05) 2012 Cultivar JH 3459 PMH 2 JH 31244 JH 3956 PMH 1 PARKASH LM 16 CM 143 CD (p=0.05) H 16.27 17.42 16.72 15.85 15.93 17.36 14.76 11.56 1.04 H 15.60 19.02 16.80 16.64 17.36 18.82 14.86 11.24 1.15 I 10.24 12.63 13.40 14.24 12.63 10.13 7.53 9.06 1.42 Leaf length (cm) I 11.74 10.50 11.70 12.40 11.37 11.38 8.55 9.14 1.44 % cha. -32.02 -39.72 -30.02 -21.77 -28.63 -34.45 -42.07 -20.93 H 2.18 2.39 2.45 2.12 2.29 2.50 1.87 1.76 0.20 % cha. -34.36 -33.60 -20.24 -14.42 -27.25 -46.17 -49.33 -19.40 H 2.11 2.59 2.35 2.24 2.35 2.61 1.98 1.68 0.19 I 1.62 1.44 1.76 1.90 1.54 1.28 0.82 1.39 0.17 % cha. -23.22 -44.40 -25.11 -15.18 -34.47 -50.96 -58.59 -17.26 H 26.16 30.72 31.04 25.93 28.73 33.49 21.59 17.50 2.19 I 13.95 13.94 15.20 16.84 13.87 10.96 7.93 10.67 1.56 Leaf area (cm2) H 27.85 31.97 30.83 27.43 30.19 32.77 21.10 16.97 1.36 I 15.20 13.87 15.54 15.75 15.10 15.03 9.31 10.06 1.49 % cha. -45.42 -56.62 -49.59 -42.58 -49.98 -54.13 -55.88 -40.72 H 2.35 2.67 2.37 2.13 2.39 2.41 1.95 1.97 0.13 % cha. -46.67 -54.62 -51.03 -35.06 -51.72 -67.27 -63.27 -39.03 H 2.07 2.83 2.34 2.04 2.61 2.44 1.96 1.87 0.11 I 2.18 2.51 2.28 1.96 2.14 1.85 1.54 1.74 0.11 % cha 5.31 -11.31 -2.56 -3.92 -18.01 -24.18 -21.43 -6.95

Leaf width (cm) I 1.72 1.64 1.91 1.68 1.73 1.65 1.20 1.38 0.16 % cha. -21.10 -31.38 -22.04 -20.75 -24.45 -34.00 -35.83 -21.59

Stem girth (cm) I 2.23 2.16 2.19 1.99 2.19 1.99 1.75 1.63 0.10 % cha. -5.11 -19.10 -7.59 -6.57 -8.37 -17.43 -10.26 -17.26

H: Healthy, I: Infested; Per cent change in infested condition (% cha.) = I-H /H x 100

105

The oviposition, leaf injury and deadhearts incidence were positively correlated with leaf length, leaf width, leaf area and stem girth (table 29). However, only the leaf injury showed significant and positive correlation with leaf length (r = 0.79*), leaf width (r = 0.89*), leaf area (r = 0.85*) and stem girth (r = 0.76*) indicating that more infestation occurred on cultivars with seedlings having broad and long leaves with more stem girth. Thus susceptibility in Parkash and PMH 2 can be linked to these putative morphological markers in resistance breeding programmes against this pest. Rao C N and Panwar (2001a) also reported that leaf width had positive correlation with percentages of deadhearts by shoot fly in maize but leaf length was negatively correlated. Thus, according to them maize varieties with more leaf length and less leaf width showed resistant reaction to shoot fly. However, the results of present studies indicated that susceptible genotypes had more leaf length. The positive correlation between leaf length and A. soccata susceptibility in sorghum reported by Sandhu et al (1986) are in line with the present studies. 4.3.6. Variation in biochemical traits in relation to expression of resistance to shoot fly, Atherigona naqvii: The various biochemical traits i.e. chlorophyll, moisture content, proteins, tannins, phenols, amino acids and reducing sugars were estimated at 5th leaf stage of the seedling during 2011 and 2012. Chlorophyll: The data obtained for chlorophyll content of leaves of different maize genotypes have been given in table 23. The chlorophyll content index (CCI) differed significantly among the test genotypes during 2011 only and it varied from 25.40 to 31.07. It was maximum in JH 31244 (31.07), which was further on par with JH 3956 (29.50) and PMH 1 (29.20); and minimum in LM 16 (25.40), being on par with CM 143 (25.80) which were further on par with PMH 2 (27.53), JH 3459 and Parkash (27.57 each), which was also on par with JH 3956 and PMH 1. Though, during 2012 the differences were non significant among the test genotypes yet numerically, it was minimum in PMH2 (27.70) and maximum in JH 3459 (29.82). The results showed that the resistance of genotypes to A. naqvii did not have any consistent correlation with chlorophyll content of their leaves. The correlation (r= -0.10) between chlorophyll content and oviposition was non significant revealing that egg laying by A. naqvii was not related to chlorophyll content of the leaves of test genotypes (table 29). Similarly Rao C N and Panwar (2001b) have also reported the non significant correlation between oviposition by Atherigona spp. and chlorophyll content in maize. Singh et al (2004) further reported that there was no relationship between chlorophyll content and per cent seedlings with eggs of A. soccata in sorghum also. However, Patil et al (2006) reported that the lower amount of chlorophyll in sorghum leaves turns them yellowish green and probably make them unattractive for oviposition by A. soccata. These differences might be due to difference in egg laying preference of prevalent Atherigona spp. in different studies.

106

Table 23. Variation in chlorophyll content index (CCI) and moisture content of the test genotypes at 5th leaf stage under field conditions during 2011 and 2012 Chlorophyll content index (CCI) 2011 JH 3459 PMH 2 JH 31244 JH 3956 PMH 1 PARKASH LM 16 CM 143 CD (p=0.05) 27.57 27.53 31.07 29.50 29.20 27.57 25.40 25.80 3.13 2012 29.82 27.70 29.47 28.03 29.23 29.47 28.20 28.03 NS Moisture content (%) 2011 85.45 85.86 85.27 87.01 87.14 88.10 89.30 86.63 1.85 2012 84.47 85.69 85.62 86.11 86.56 86.73 85.32 85.04 0.84

Genotypes

Moisture content: There were significant differences in moisture content of plants of test genotypes in both the test years and it varied from 85.27 to 89.30 and 84.47 to 86.73 per cent during 2011 and 2012, respectively (table 23). During 2011, it was maximum in susceptible LM 16 (89.30 %), which was further on par with another susceptible hybrid Parkash (88.10 %) which was further also on par with PMH 1, JH 3956 and CM 143 (86.63 to 87.14 %). It was minimum in JH 31244 (85.27 %) which was further on par with genotypes JH 3459, PMH 2, JH 3956 and CM 143 (85.45-87.01 %), which were also on par with Parkash. During 2012, it was maximum in Parkash (86.73 %), being on par with PMH 1 (86.56 %) and both were further on par with JH 3956 (86.11 %) which was also on par with LM 16, JH 31244 and PMH 2 (85.32 to 85.69 %), which was on par with CM 143. It was minimum in JH 3459 (84.47 %) which was again on par with CM 143 (85.04 %). The resistance of the test genotypes to A. naqvii again did not have any consistent correlation with the moisture content of plants. Rao C N and Panwar (2001b) reported lower moisture content in shoot fly maize resistant genotype Antigua Gr. 1 (74.7 %) as compared to susceptible CM 300 (82.8 %). However, no relationship was reported between moisture content of sorghum seedlings and shoot fly resistance by Singh et al (2004). In present studies also moisture content was non significantly correlated (table 29) with deadhearts incidence (r= 0.23).

107

Proteins: The data pertaining to the protein content (%) on dry weight basis of shoot fly infested and non infested maize genotypes during spring 2011 and 2012 have been presented in table 24. The protein content increased under infested plants as compared to healthy plants but it differed significantly among test genotypes in 2012 only. During 2011, it was maximum in CM 143 (19.04 %) and minimum in JH 31244 (15.85) in healthy plants; and maximum in PMH 2 (25.56 %) and minimum in CM 143 (21.98 %) infested plants. The percentage change of protein content in infested over healthy was maximum in susceptible hybrid PMH 2 (49.91%) followed by JH 31244 (42.52 %) and minimum in CM 143 (15.44 %). During 2012, the protein content varied from 15.53 to 17.60 % and 18.52 to 20.20 % in healthy and infested conditions, respectively. In healthy plants it was maximum in least susceptible JH 3459 (17.60 %), being on par with intermediate CM 143 (17.35 %), which were on par with highly susceptible Parkash (17.25 %); and minimum in other intermediate JH 31244 (15.53 %). In other genotypes it varied from 16.25 to 16.71 per cent. Under infested conditions, it was significantly higher in susceptible PMH 2 (20.20 %) with maximum increase of 24.31 per cent as compared to healthy plants. It was minimum in intermediate PMH 1 (18.52 %) which was on par with JH 3956 (18.78 %), JH 31244, CM 143 (18.87 % each), JH 3459 (18.96 %); and later was further on par with Parkash (19.29 %) and LM 16 (19.46 %). The per cent increase of protein content in infested conditions was minimum in other least susceptible JH 3459 (7.73 %) followed by intermediate CM 143 (8.76 %). However the susceptible hybrid Parkash (11.83 %) and other least susceptible hybrid JH 3956 (12.39 %) were intermediate in this regard. Thus the data again revealed that variations in protein contents did not have any relationship with relative susceptibility of test maize genotypes. The increase of protein content in infested plants indicated the synthesis of defense or structural protein to overcome insect injury (Sathya et al 2002). However, in present studies the variation in per cent increase in comparatively tolerant/ susceptible genotypes indicated that host plant might be involved in some kind of other defense mechanism to overcome shoot fly infestation. Similarly, non significant correlation of protein content with deadhearts (r = 0.41) and deadhearts proportion out of total incidence (r = 0.51) also indicated that the variation in protein content did not have any synchrony with susceptibility of test genotypes to A. naqvii (table 29). Similarly, Chamarthi et al (2011) reported non significant correlation between protein and deadhearts (r = 0.17) due to A. soccata in sorghum. However, a significant and positive correlation of nitrogen (r= 0.77) and crude protein content (r= 0.82) with deadhearts percentage was reported by Rao C N and Panwar (2001b). They concluded that resistant cultivars had significantly less nitrogen and crude protein, so are less favourable for insect growth and development. But such difference in biological parameters were not observed

108

in different test genotypes in the present studies as indicated by the non significant correlations of protein content with biological parameters of A. naqvii reared on different test genotypes (table 31). It was positive with adult longevity, larval survival, male and female pupal weight; and negative with larval period, pupal period and pupal survival.

Table 24. Effect of shoot fly, A. naqvii feeding on protein content (% dry weight) in the different test maize genotypes at 5th leaf stage during 2011 and 2012
Mean protein content (% dry weight) 2011 Genotypes JH 3459 PMH 2 JH 31244 H 3956 PMH 1 PARKASH LM 16 CM 143 CD (p=0.05) Healthy (H) 18.49 17.05 15.85 16.63 16.85 17.88 18.10 19.04 NS Infested (I) 22.98 25.56 22.59 22.03 22.13 23.55 23.01 21.98 NS % change (I-H/H x 100) 24.28 49.91 42.52 32.47 31.34 31.71 27.13 15.44 2012 Healthy (H) 17.60 16.25 15.53 16.71 16.54 17.25 16.54 17.35 0.55 Infested (I) 18.96 20.20 18.87 18.78 18.52 19.29 19.46 18.87 0.51 % change (I-H/H x 100) 7.73 24.31 21.51 12.39 11.97 11.83 17.65 8.76

Tannins: Table 25 depicts the data pertaining to tannins content on dry weight basis of shoot fly infested and non infested maize genotypes during spring 2011 and 2012. The data indicated that tannins content varied significantly among the genotypes under healthy and infested conditions during both the test years. In 2011, it varied from 2.97 to 3.85 mg/g and 2.95 to 3.88 mg/g under healthy and infested conditions, respectively. In healthy plants tannins content was maximum in CM 143 (3.85 mg/g) which was on par with LM 16 (3.71 mg/g), which was further on par with JH 3459 (3.54 mg/g) and Parkash (3.64 mg/g). It was minimum in JH 3956 (2.97 mg/g) being on par with JH 31244 (2.98 mg/g). PMH 2 (3.18 mg/g) and PMH 1 (3.22 mg/g) were intermediate in tannins content. Under infested conditions also, it was maximum in CM 143 (3.88 mg/g), being on par with LM 16 (3.81 mg/g) and PMH 1 (3.79 mg/g); and minimum in JH 3956 (2.95 mg/g). It was intermediate in JH 31244 (3.26 mg/g) which was on par with PMH 2 (3.32 mg/g) and it was further on par with Parkash (3.47 mg/g) and JH 3459 (3.52 mg/g). The tannins content increased or decreased in infested conditions in different genotypes irrespective of their susceptibility to A. naqvii. The maximum per cent increase and decrease was in an intermediate PMH 1 (17.70 %) and a susceptible hybrid Parkash (4.67 %), respectively among the test genotypes.

109

Table 25. Effect of shoot fly, A. naqvii feeding on tannins content (mg/g dry weight) in the different test maize genotypes at 5th leaf stage during 2011 and 2012 Mean tannins content (mg/g dry weight) 2011 Genotypes JH 3459 PMH 2 JH 31244 H 3956 PMH 1 PARKASH LM 16 CM 143 CD (p=0.05) Healthy (H) 3.54 3.18 2.98 2.97 3.22 3.64 3.71 3.85 0.18 Infested (I) 3.52 3.32 3.26 2.95 3.79 3.47 3.81 3.88 0.20 % change (I-H/H x 100) -0.56 4.40 9.40 -0.67 17.70 -4.67 2.70 0.78 2012 Healthy (H) 3.88 3.47 3.57 3.65 3.60 4.67 2.87 3.74 0.42 Infested (I) 3.50 3.37 3.59 3.63 4.02 3.63 3.36 3.84 0.30 % change (I-H/H x 100) -9.79 -2.88 0.56 -0.55 11.67 -22.27 17.07 2.67

In 2012, it varied from 2.87 to 4.67 mg/g and 3.36 to 4.02 mg/g under healthy and infested conditions, respectively. In healthy plants tannins content was maximum in susceptible hybrid Parkash (4.67 mg/g) and minimum in LM 16 (2.87 mg/g) while all the remaining genotypes being on par with each other (3.47 to 3.88 mg/g) formed intermediate group. However, in infested plants it was maximum in PMH 1 (4.02 mg/g) which was on par with CM 143 (3.84 mg/g) and minimum in LM 16 (3.36 mg/g) being on par with PMH 2 (3.37 mg/g) and JH 3459 (3.50 mg/g), which were further on par with JH 31244, JH 3956 and Parkash (3.59 -3.63 mg/g). Thus the variation in tannins content did not show any synchrony with the host plant reaction of different test genotypes to incidence of A. naqvii. The maximum per cent increase and decrease was in susceptible LM 16 (17.07 %) and another susceptible Parkash (22.27 %), respectively among different genotypes. The decrease in tannins content was recorded in susceptible hybrid Parkash and also in least susceptible hybrid JH 3459 and JH 3956 but its increase and decrease in another susceptible hybrid PMH 2 in 2011 and 2012 indicated that susceptibility to shoot fly in test genotypes is a complex phenomenon with interplay of more than one factors influencing the overall reaction of the genotypes to shoot fly. However, the increased tannins content in resistant genotypes was reported to be associated with defense mechanism of the plant in different crops. Bialczyk et al (1999) reported increase in the tannin content in tomato after attack of whitefly. Similarly, Shi ZhongLiang et al (2002) reported that tannins are important secondary metabolites

110

for induced resistance to blossom midge in wheat. In present study the tannins content was non significantly correlated with the biological parameters of shoot fly (table 31). The correlation with egg count and deadhearts was also non significant (table 29). However, in sorghum significant and negative correlation between tannin content with oviposition (r= -0.85) and shoot fly deadhearts (r= -0.86) was reported by Chamarthi et al (2011). Phenols: The data pertaining to the phenols content (mg/g) on dry weight basis of infested and healthy maize genotypes for both the test years have been presented in table 26. The data indicated that phenols content varied significantly among the genotypes under healthy in 2012 and infested conditions during both the test years. During 2011, the phenols content varied non significantly under healthy conditions being maximum in JH 3459 (31.55 mg/g) and minimum in JH 31244 (26.91 mg/g). In infested plants it was significantly higher in least susceptible hybrid JH 3459 (35.61 mg/g) and lower in susceptible inbred line LM 16 (28.00 mg/g). Phenols content was intermediate (30.97 to 32.57 mg/g) and on par with each other in the remaining genotypes irrespective of their susceptibility. There was increase or decrease of phenols content in infested and healthy conditions among different genotypes. The increase in phenols content under

infested conditions in test genotypes varied from 5.26 to 15.09 per cent being maximum in intermediate JH 31244 and minimum in susceptible Parkash, whereas, it decreased in another susceptible LM 16 (-7.44 %) only. During 2012, the phenols content in healthy plants was maximum in susceptible hybrid Parkash (26.33 mg/g) being on par with intermediate CM 143 (25.61 mg/g), least susceptible JH 3956 (25.10 mg/g) which were further on par with least susceptible JH 3459, intermediate PMH 1 (24.81 mg/g), an other intermediate JH 31244 (23.80 mg/g) and minimum in an other susceptible hybrid PMH 2 (20.68 mg/g). In infested conditions it was maximum in susceptible hybrid Parkash (29.74 mg/g) which was on par with a least susceptible hybrid JH 3459 (28.65 mg/g), a susceptible LM 16 (28.58 mg/g), an other susceptible PMH 2 (28.36 mg/g), intermediate JH 31244 (27.35 mg/g). It was minimum in an intermediate CM 143 (24.45 mg/g) which was on par with intermediate PMH 1 (25.25 mg/g) and both were on par with least susceptible JH 3956 (26.19 mg/g) and another intermediate JH 31244 (27.35 mg/g). As in 2011, phenol content increased in infested conditions in all the test genotypes except CM 143.The increase in phenols content varied from 1.77 to 37.14 per cent being maximum in susceptible PMH 2 and minimum in intermediate PMH 1, whereas the decrease in another susceptible CM 143 was 4.53 per cent. The results revealed that the phenols content was maximum in least susceptible hybrid JH 3459 and susceptible hybrid Parkash in different years under different conditions. Thus, phenols content did not reflect significance in providing resistance in test genotypes to shoot fly

111

infestation. Moreover, it was non significantly correlated with biological parameters of shoot fly in present studies, it was positively correlated with larval period, pupal period, adult longevity, larval survival and fecundity; and negatively with pupal survival, male and female pupal weight (table 31). The higher amount of phenolic compounds has been reported to be associated with shoot fly resistance in sorghum seedlings (Khurana and Verma 1983) and to stem borer, C. partellus in maize (Kabre and Ghorpade 1999). However, Chamarthi et al (2011) reported no significant differences in total phenols content of the test genotypes in sorghum and there was non significant correlation of phenols with oviposition (r= -0.14 ) and deadhearts incidence (r= 0.24). In the present studies too, it was non significantly correlated with egg counts and deadhearts (table 29). The non significant correlation of phenol content with different aspects of shoot fly infestation along with variations in genotypes belonging to different susceptibility group make phenols content as least putative marker in host plant resistance breeding programme against shoot fly. Table 26. Effect of shoot fly, A. naqvii feeding on Phenols content (mg/g dry weight) in the different test maize genotypes at 5th leaf stage during 2011 and 2012 Mean phenols content (mg/g dry weight) 2011 Genotypes JH 3459 PMH 2 JH 31244 H 3956 PMH 1 PARKASH LM 16 CM 143 CD (p=0.05) Healthy (H) 31.55 28.59 26.91 28.29 29.01 30.39 30.25 26.99 NS Infested (I) 35.61 32.13 30.97 31.26 32.57 31.99 28.00 30.97 2.65 % change (I-H/H x 100) 12.87 12.38 15.09 10.50 12.27 5.26 -7.44 14.75 2012 Healthy (H) 24.81 20.68 23.8 25.10 24.81 26.33 22.42 25.61 2.64 Infested (I) 28.65 28.36 27.35 26.19 25.25 29.74 28.58 24.45 3.34 % change (I-H/H x 100) 15.48 37.14 14.92 4.34 1.77 12.95 27.48 -4.53

Amino acids: The data pertaining to the amino acids content (mg/g) on dry weight basis of shoot fly infested and non infested maize test genotypes, varying significantly, during spring 2011 and 2012 have been presented in table 27. During 2011, the amino acids content varied from 5.03 to 9.69 mg/g and 5.42 to 9.96 mg/g under healthy and infested conditions, respectively. In healthy conditions it was maximum

112

in least susceptible genotype JH 3459 (9.69 mg/g) being on par with susceptible genotypes PMH 2 and CM 143 (9.45 mg/g each); and minimum in the remaining least susceptible hybrid JH 3956 (5.03 mg/g). It was intermediate and on par in intermediate JH 31244, susceptible LM 16 and intermediate PMH 1 (7.10 to 7.91 mg/g) and later was also on par with another susceptible Parkash (7.99 mg/g). Under infested conditions it was maximum in intermediate PMH 1 (9.96 mg/g), being on par with susceptible PMH 2 (9.76 mg/g) which was further on par with least susceptible JH 3459 (9.06 mg/g), susceptible CM 143 (9.02 mg/g), another susceptible LM 16 (8.94 mg/g) and intermediate JH 31244 (8.68 mg/g). The amino acids content was minimum in least susceptible JH 3956 (5.03 and 5.42 mg/g) both under healthy and infested conditions, whereas it was intermediate in susceptible Parkash (7.63 mg/g). It varied irrespective of infested and healthy conditions among different genotypes. The increase in amino acids content was lower in susceptible PMH 2 (3.28 %) and higher in intermediate PMH 1 and susceptible LM 16 (25.92 %); and the decrease was lower in susceptible Parkash (-4.51 %) and higher in least susceptible JH 3459 (-6.50 %) under infested conditions in the test genotypes. During 2012, the amino acids content ranged from 5.50 to 8.48 mg/g and 5.67 to 9.16 mg/g in healthy and infested conditions, respectively. In healthy condition it was maximum in susceptible Parkash (8.48 mg/g) which was on par with least susceptible JH 3459 (7.82 mg/g) and intermediate CM 143 (7.75 mg/g). As in 2011, the amino acids content was minimum in least susceptible JH 3956 (5.50 mg/g) which was on par with intermediate JH 31244 (5.90 mg/g) and susceptible LM 16 (6.50 mg/g). It was intermediate in susceptible PMH 2 (7.00 mg/g) and intermediate PMH 1 (7.16 mg/g). In infested conditions it was again maximum in susceptible Parkash (9.16 mg/g) and minimum in least susceptible JH 3956 (5.67 mg/g). After Parkash, the amino acids content in PMH 1 (7.70 mg/g) also being on par with PMH 2, JH 3459, CM 143 , JH 31244 and LM 16 (6.76-7.56 mg/g) formed the intermediate group. As in 2011, the amino acids content increase or decrease in infested conditions in the test genotypes. In 2012, increase in amino acids varied from 3.09 to 16.78 per cent in intermediate JH 31244 and minimum in least susceptible JH 3956, whereas it decreased -5.42 and -5.75 per cent in intermediate CM 143 and least susceptible JH 3459, respectively. The amino acids was more in susceptible hybrids PMH 2 and intermediate PMH 1; and Parkash in 2011 and 2012, respectively and lower content in least susceptible hybrid JH 3956 during both the tears, indicated roll of amino acids in contributing towards susceptibility. But its effect on biological parameters of shoot fly as indicated by non significant correlation coefficients is not evident (table 31). The correlation was positive with adult longevity, larval survival, male and female pupal weight; and negative with larval period, pupal period, pupal survival and fecundity. Kabre and Ghorpade (1999) also reported that

113

relationship between free amino acids content and stem borer susceptibility in maize to C. partellus was not evident. Singh and Jotwani (1980c) also reported very little differences in amino acids among the susceptible and resistant varieties in sorghum. This might be due to the fact very little work has been done to determine specific amino acids contributing in the resistance to shoot fly. Table 27. Effect of shoot fly, A. naqvii feeding on free amino acids content (mg/g dry weight) in different test maize genotypes at 5th leaf stage during 2011 and 2012 Mean free amino acids content (mg/g dry weight) 2011 Genotypes JH 3459 PMH 2 JH 31244 H 3956 PMH 1 PARKASH LM 16 CM 143 CD (p=0.05) Healthy (H) 9.69 9.45 7.10 5.03 7.91 7.99 7.10 9.45 0.86 Infested (I) 9.06 9.76 8.68 5.42 9.96 7.63 8.94 9.02 1.43 % change (I-H/H x 100) -6.50 3.28 22.25 7.75 25.92 -4.51 25.92 -4.55 2012 Healthy (H) 7.82 7.00 5.90 5.50 7.16 8.48 6.50 7.75 1.18 Infested (I) 7.37 7.56 6.89 5.67 7.70 9.16 6.76 7.33 0.86 % change (I-H/H x 100) -5.75 8.00 16.78 3.09 7.54 8.02 4.00 -5.42

Reducing sugars: The data pertaining to the reducing sugars content (mg/g) on dry weight basis of different maize genotypes during spring 2011 and 2012 have been presented in Table 28. The data indicated that reducing sugars varied significantly among the genotypes under healthy and infested conditions during both the years. During 2011, the reducing sugars varied from 11.98 to 16.12 mg/g and 8.97 to 14.31 mg/g under healthy and infested conditions, respectively. It was maximum in susceptible genotype LM 16 (16.12 mg/g), being on par with susceptible Parkash (15.96 mg/g) which were further on par with intermediate JH 31244 (15.15 mg/g); and minimum in intermediate CM 143 (11.98 mg/g) being on par with intermediate PMH 1 (12.42 mg/g) which were further on par with least susceptible JH 3956 (12.76 mg/g). The contents were intermediate and on par in least susceptible JH 3459, susceptible PMH 2 and intermediate JH 31244 (13.87 to 15.15 mg/g). Under infested conditions, the content was maximum in susceptible hybrid Parkash (14.31 mg/g) which was on par with another susceptible PMH 2 (12.89 mg/g), which were further on par with

114

remaining susceptible LM 16 (12.07 mg/g), intermediate PMH 1 (11.90 mg/g), least susceptible JH 3956 (11.68 mg/g) and intermediate JH 31244 (11.37 mg/g). It was minimum in least susceptible hybrid JH 3459 (8.97 mg/g) which was on par with comparatively less susceptible inbred CM 143 (10.75 mg/g). The reducing content decreased in all the tested genotypes under infested conditions. It varied from 4.19 to 35.33 per cent, being maximum in JH 3459 and minimum in PMH 1. During 2012, the reducing sugar varied from 15.76 to 20.47 mg/g and 11.45 to 18.65 mg/g under healthy and infested conditions, respectively. As in 2011, in healthy condition it was maximum in susceptible genotype LM 16 (20.47 mg/g), being on par with another susceptible Parkash (20.30 mg/g ) which were further on par with the remaining susceptible PMH 2 (18.26 mg/g). It was minimum in intermediate PMH 1(15.76 mg/g), being on par with least susceptible JH 3956 (15.80 mg/g) which were further on par with remaining susceptible JH 3459 (16.21 mg/g) and intermediate CM 143 (16.32 mg/g), another intermediate JH 31244 (18.14 mg/g) was on par with both the groups. Under infested conditions it was maximum in susceptible hybrid Parkash (18.65 mg/g) and minimum in least susceptible hybrid JH 3459 (11.45 mg/g), which was on par with intermediate CM 143 (13.23 mg/g). The reducing sugar content was intermediate in least susceptible JH 3956 (13.43 mg/g), being on par with intermediate JH 31244 (13.49 mg/g) which were on par with susceptible PMH 2 (14.53 mg/g) and susceptible LM 16 (14.55 mg/g) and these were on par with intermediate PMH 1 (15.35 mg/g). As in 2011, the reducing sugar content decreased in infested conditions and it varied from 2.60 to 29.36 per cent, being maximum in least susceptible JH 3459 and minimum in intermediate PMH 1. The data revealed that under infested conditions, the highly susceptible hybrid Parkash had higher sugar content whereas it was lower in least susceptible hybrid JH 3459 during both the test years. Moreover, the decrease in sugar content was also higher in least susceptible JH 3459. Thus, reducing sugars were related to susceptibility to shoot fly, A. naqvii in the test genotypes. Similarly, higher sugar content was reported to be associated with the susceptibility to shoot fly in sorghum (Chamarthi et al 2011). In the present studies, reducing sugars had positive and significant correlation (table 31) with larval survival and fecundity (r = 0.87* and 0.71*, respectively). Accordingly, the larval survival was lower in JH 3459 and maximum in Parkash; and fecundity was also minimum and maximum in these genotypes (table 20). Sekhon and Kanta (1994) reported that maize plants resistant to spotted stem borer, C. partellus had lower amounts of sugar. Similarly, Kabre and

115

Ghorpade (1999) reported that total (r= 0.89) and reducing sugars (r= 0.92) were significantly correlated with stem borer susceptibility in maize.

Table 28. Effect of shoot fly, A. naqvii feeding on reducing sugars content (mg/g dry weight) in the different test maize genotypes at 5th leaf stage during 2011 and 2012 Mean reducing sugars content (mg/g dry weight) 2011 Genotypes JH 3459 PMH 2 JH 31244 H 3956 PMH 1 PARKASH LM 16 CM 143 CD (p=0.05) Healthy (H) 13.87 14.17 15.15 12.76 12.42 15.96 16.12 11.98 2.48 Infested (I) 8.97 12.89 11.37 11.68 11.90 14.31 12.07 10.75 1.78 % change (I-H/H x 100) -35.33 -9.03 -24.95 -8.46 -4.19 -10.34 -25.12 -10.27 2012 Healthy (H) 16.21 18.26 18.14 15.80 15.76 20.30 20.47 16.32 2.45 Infested (I) 11.45 14.53 13.49 13.43 15.35 18.65 14.55 13.23 1.81 % change (I-H/H x 100) -29.36 -20.43 -25.63 -15.00 -2.60 -8.13 -28.92 -18.93

The correlations (table 29) of reducing sugars content was positive and significant with per cent leaf injury (r = 0.77*), deadhearts (r= 0.88*) and oviposition (r= 0.88*) by shoot fly further indicating the role of reducing sugars in imparting susceptibility in different test maize genotypes. Similarly positive and significant correlation between deadhearts percentage (r= 0.91*) and oviposition (r= 0.61*) by shoot fly with reducing sugars was observed in sorghum by Singh and Jotwani (1980c). According to Singh et al (2004) total sugars in sorghum seedlings showed significant and positive correlations with deadhearts due to shoot fly infestation and they further suggested that reducing and total sugars play a phagostimulatory role. Alternatively, low levels of reducing sugars appear to play a direct role in antixenosis to feeding and/or antibiosis. Hence, to conclude among biochemical constituents reducing sugar content only being positively associated with oviposition, leaf injury and deadhearts incidence could act as a reliable biochemical marker for selection of genotypes in shoot fly resistance breeding programmes.

116

Table 29. Correlation coefficients of shoot fly, A. naqvii damage parameters under field conditions with morphological and biochemical parameters of the test genotypes Leaf Deadhearts injury (%) (%) Morphological parameters -0.05 0.06 -0.14 0.20 0.54 0.53 0.45 0.48 0.01 -0.06 -0.36 0.14 -0.45 0.02 -0.05 0.79* 0.89* 0.85* 0.76* 0.08 -0.28 -0.33 0.32 0.24 0.06 0.15 0.28 0.34 0.26 0.36 0.21 -0.41 -0.55 Mean egg count Proportion dead hearts out of total incidence 0.48 0.84* 0.09 0.25 -0.43 -0.40 -0.48 -0.21 0.11 -0.35 -0.41

Parameters

Leaf surface wetness score Seedling viguor score Leaf glossiness score Leaf sheath pigmentation score Leaf length (cm) Leaf width (cm) Leaf area (cm ) Stem girth (cm) Trichome length (m) Trichome angle No. of trichomes
2

Biochemical parameters Chlorophyll ( CCI) Moisture (%) Protein (%) Tannins (mg/g) Phenols (mg/ g) Amino acids (mg/ g) Reducing sugars (mg/ g) -0.10 -0.05 0.17 -0.37 -0.26 -0.01 0.81* 0.39 -0.24 0.01 -0.06 0.13 0.16 0.77* -0.32 -0.23 0.41 0.23 -0.14 0.34 0.88* -0.81* 0.13 0.51 0.26 -0.42 0.38 0.41

* Significant at 5 % level of significance

117

Table 30. Correlation coefficients of shoot fly, A. naqvii damage parameters under field conditions and biological parameters of A. naqvii reared on the different test genotypes Mean egg count Leaf injury (%) Deadhearts (%) Proportion dead hearts out of total incidence

Parameters

Damage parameters Leaf injury (%) Deadhearts (%) Proportion dead hearts out of total incidence 0.75* 0.86* 0.43 0.70* 0.01 0.70*

Biological parameters Larval period (days) Pupal period (days) Adult longevity (days) Larval survival (%) Pupal survival (%) Male pupal wt (mg) Female pupal wt (mg) Fecundity -0.24 0.01 -0.35 0.59 0.05 0.37 0.66 0.70* 0.01 0.33 - 0. 07 0.48 0.04 -0.07 0.13 0.59 -0.61 -0.21 -0.54 0.86* -0.08 0.27 0.66 0.58 -0.84* -0.60 -0.71* 0.68 -0.15 0.51 0.83* 0.15

* Significant at 5 % level of significance

118

Table 31. Correlation coefficients of shoot fly, A. naqvii biological parameters reared on the different test genotypes with their morphological and biochemical parameters Larval period (days) Pupal period (days) Adult Larval Pupal longevity survival survival (days) (%) (%) Morphological parameters 0.59 -0.51 -0.28 -0.78* 0.45 0.32 0.39 0.28 -0.40 0.27 0.76* -0.73* 0.19 0.14 0.29 0.07 0.10 0.05 0.04 0.60 -0.47 -0.73* -0.14 -0.20 0.63 0.63 -0.15 -0.03 -0.05 -0.38 -0.14 -0.39 -0.21 Pupal weight (mg) Male Female Fecundity

Parameters

Leaf surface wetness score Seedling viguor score Leaf glossiness score Leaf sheath pigmentation score Leaf length (cm) Leaf width (cm) Leaf area (cm2) Stem girth (cm) Trichome length (m) Trichome angle No. of trichomes Chlorophyll ( CCI) Moisture (%) Protein (%) Tannins (mg/g) Phenols (mg/g) Amino acids (mg/g) Reducing sugars (mg/g)

0.87* -0.57 -0.23 -0.48 0.48 0.42 0.48 0.38 -0.52 0.45 0.67

0.65 -0.54 -0.07 -0.20 0.55 0.56 0.60 0.54 0.09 0.40 -0.05

0.21 0.71* -0.29 -0.05 -0.14 -0.16 -0.23 0.19 -0.49 0.31 0.17

-0.27 0.76* -0.05 0.17 -0.15 -0.17 -0.26 0.03 -0.22 -0.06 -0.20

-0.06 -0.25 -0.41 0.12 0.57 0.51 0.46 0.37 0.08 -0.01 -0.07

Biochemical parameters 0.73* -0.42 -0.37 -0.49 0.26 -0.18 -0.51 0.74* 0.14 -0.58 -0.16 0.16 -0.05 -0.09 0.46 -0.54 0.09 -0.18 0.66 0.11 -0.44 -0.47 0.29 0.45 0.42 0.03 0.23 0.87* 0.36 -0.02 -0.58 -0.32 -0.29 -0.64 0.12 -0.37 -0.14 0.13 -0.36 -0.63 0.26 -0.13 -0.58 -0.03 0.33 -0.23 -0.57 0.23 0.35 -0.17 -0.13 0.41 -0.30 0.07 -0.19 0.71*

* Significant at 5 % level of significance

119

4.4.

Cultural and chemical control of shoot fly, Atherigona spp. on spring maize: The experiment on cultural and chemical control of shoot fly, Atherigona spp. on spring

maize was conducted during spring 2011 and 2012 and the data recorded on various aspects have been presented in Table 32-37. 4.4.1 Oviposition by shoot fly, Atherigona spp.: The incidence of egg counts of shoot fly recorded in different treatments during different observations did not differ significantly during 2011 and 2012 (table 32 and 33) except at 6 DAG during 2012. The overall mean egg counts in all the treatments were comparatively low in different treatments during 2011 (ranging from 0.11-0.17 per plant) as compared to 2012 (ranging from 0.21 -0.47 per plant). The mean egg laying per plant was on par in different treatments in all the observations recorded during 2011 (upto 18 DAG) and 2012 (upto 15 DAG) except at 6 DAG during 2012 where it differed significantly. In 2012 at 6 DAG it was maximum in T 11 (untreated control) having 1.07 eggs per plant being on par with T 4 (with two rows of unprotected sorghum sown as a trap crop), T6 (two rows of protected sorghum sown as a trap crop) and T 7 (where first irrigation was given at emergence) each having 1.00 eggs per plant (table 33). It was minimum in T8 (where first irrigation was given after one week of emergence) (0.27 eggs/plant) and followed by T2 i.e. with higher dose of thiamethoxam (0.60 eggs/ plant), T 5 (with single row of protected sorghum sown as a trap crop) with 0.63 eggs per plant, T1 ( with lower dose of thiamethoxam) with 0.80 eggs per plant, T3 (with single row of unprotected sorghum sown as trap crop) with 0.90 eggs per plant. T9 (granular application of carbofuran) and T10 (seed treatment with imidacloprid 600 FS ) each having 0.93 eggs per plant formed intermediate group where egg laying was either more or on par with the above treatments. Marwaha et al (1984) reported that high relative humidity (75-80 %) and rainfall in the last week of February with sporadic drizzling in March proved to be conducive to shoot fly incidence in terms of higher oviposition and deadhearts formation. In the present studies during 2011 too (table 32), the egg laying by Atherigona spp was maximum at 9 DAG in T8 where first irrigation was given one week after emergence however, it was minimum in same treatment during 2012 when observed at 6 DAG i.e. one day before the application of irrigation. These differences in egg laying in these treatments during 2011 and 2012 indicate the importance of increased moisture level for egg laying of the pest. Thus, both the findings indicate the role of changed microclimate of crop to influence the egg laying behavior of shoot fly, Atherigona spp. The variation in egg counts in T8 in both years might also be due to variations in abundance of shoot fly population. In the present studies the trend in egg laying in all the treatments remained positive upto 9 and 6 DAG, however, after that it remained negative upto 18 and 15 DAG (last observations in both cases)

120

Table 32. The oviposition by shoot fly, Atherigona spp. in different treatments during spring 2011 Mean egg count per plant 12 15 6 DAG 9 DAG DAG DAG 0.07 (1.03) 0.00 (1.00) 0.07 (1.03) 0.00 (1.00) 0.00 (1.00) 0.00 (1.00) 0.00 (1.00) 0.00 (1.00) 0.00 (1.00) 0.03 (1.02) 0.00 (1.00) (NS) 0.27 (1.12) 0.27 (1.12) 0.30 (1.14) 0.37 (1.17) 0.33 (1.15) 0.30 (1.14) 0.30 (1.14) 0.40 (1.18) 0.27 (1.12) 0.27 (1.12) 0.30 (1.14) (NS) 0.23 (1.11) 0.20 (1.09) 0.27 (1.12) 0.20 (1.09) 0.23 (1.11) 0.20 (1.09) 0.23 (1.11) 0.23 (1.11) 0.20 (1.09) 0.20 (1.09) 0.23 (1.11) (NS) 0.20 (1.09) 0.23 (1.11) 0.23 (1.11) 0.23 (1.11) 0.23 (1.11) 0.13 (1.06) 0.13 (1.06) 0.07 (1.03) 0.17 (1.08) 0.13 (1.06) 0.23 (1.11) (NS)

Treatments T1 - Thiamethoxam 30 FS @ 4 g per kg seed one day before sowing T2 - Thiamethoxam 30 FS @ 6 g per kg seed one day before sowing T3 - Single row of unprotected sorghum sown as a trap crop T4 - Two rows of unprotected sorghum sown as a trap crop T5 - Single row of protected sorghum sown as a trap crop T6- Two rows of protected sorghum sown as a trap crop T7- First irrigation at crop emergence T8- First irrigation after one week of crop emergence T9- Carbofuran 3 G @ 5 kg per acre at sowing (standard) T10 - Imidacloprid 600 FS @ 6 ml per kg seed (standard) T11 - Untreated control CD (p=0.05)

3 DAG 0.03 (1.02) 0.00 (1.00) 0.00 (1.00) 0.03 (1.02) 0.03 (1.02) 0.00 (1.00) 0.00 (1.00) 0.03 (1.02) 0.07 (1.03) 0.03 (1.02) 0.00 (1.00) (NS)

18 DAG 0.10 (1.05) 0.07 (1.03) 0.13 (1.06) 0.20 (1.09) 1.00 (1.05) 0.13 (1.06) 0.07 (1.03) 0.17 (1.08) 0.07 (1.03) 0.13 (1.06) 0.07 (1.03) (NS)

Mean 0.15

0.13

0.15

0.17

0.16

0.11

0.12 0.15 0.13 0.12 0.14

Figures in parentheses are n+1 transformed values DAG: days after germination

121

Table 33. The oviposition by shoot fly, Atherigona spp. in different treatments during spring 2012 Mean egg count per plant Treatments T1 - Thiamethoxam 30 FS @ 4 g per kg seed one day before sowing T2 - Thiamethoxam 30 FS @ 6 g per kg seed one day before sowing T3 - Single row of unprotected sorghum sown as a trap crop T4 - Two rows of unprotected sorghum sown as a trap crop T5 - Single row of protected sorghum sown as a trap crop T6- Two rows of protected sorghum sown as a trap crop T7- First irrigation at crop emergence T8- First irrigation after one week of crop emergence T9- Carbofuran 3 G @ 5 kg per acre at sowing (standard) T10 - Imidacloprid 600 FS @ 6 ml per kg seed (standard) T11 - Untreated control CD (p=0.05) 3 DAG 0.20 (1.09) 0.17 (1.08) 0.23 (1.11) 0.23 (1.11) 0.20 (1.09) 0.33 (1.15) 0.27 (1.12) 0.10 (1.05) 0.23 (1.11) 0.23 (1.11) 0.30 (1.14) (NS) 6 DAG 0.80 (1.34) 0.60 (1.26) 0.90 (1.37) 1.00 (1.41) 0.63 (1.28) 1.00 (1.41) 1.00 (1.41) 0.27 (1.12) 0.93 (1.39) 0.93 (1.39) 1.07 (1.44) (0.12) 9 DAG 0.53 (1.24) 0.43 (1.19) 0.53 (1.24) 0.60 (1.26) 0.43 (1.19) 0.43 (1.19) 0.37 (1.17) 0.30 (1.14) 0.70 (1.30) 0.63 (1.28) 0.47 (1.21) (NS) 12 DAG 0.30 (1.14) 0.20 (1.09) 0.27 (1.13) 0.30 (1.14) 0.30 (1.14) 0.23 (1.11) 0.20 (1.09) 0.23 (1.11) 0.33 (1.15) 0.27 (1.12) 0.27 (1.12) (NS) 15 DAG 0.13 (1.06) 0.10 (1.05) 0.13 (1.06) 0.13 (1.06) 0.17 (1.08) 0.10 (1.05) 0.10 (1.05) 0.13 (1.06) 0.17 (1.08) 0.10 (1.05) 0.13 (1.06) (NS) Mean 0.39

0.30

0.41

0.45

0.35

0.42

0.39

0.21

0.47

0.43

0.45

Figures in parentheses are n+1 transformed values DAG: days after germination

122

during both the test years, respectively. Which is in line with the findings of Delobel (1981), Rao and Panwar (1992, 1996b) and Hari et al (2013) who also reported that the shoot fly spp. i.e. A. soccata and A. naqvii showed non discriminate behavior towards different cultivars receiving different chemical treatments and the flies laid eggs at random on the crop. In the present studies, there was no significant reduction in the egg laying on crop in seed treatment before sowing or where chemicals was applied to soil at sowing. However, Katole et al (2003) reported that shoot fly, A. soccata on sorghum preferred healthy seedlings i.e. treated with carbosulfan (32.66 eggs) followed by imidacloprid (25.33-33.33 eggs) and thiamethoxam (26.33 - 30.66 eggs) per 5 seedlings in comparison to weak seedlings i.e. treated with acetamiprid seed treatment (6.33 to 8.00 eggs). The overall mean number of egg laid per plant in treatments irrespective of different observations with sorghum sown as a trap crop in different combinations (T3- T6) varied from 0.11 to 0.17 and 0.35 to 0.45 in 2011 and 2012, respectively which was comparable to egg counts in untreated control (0.14 and 0.45) in the respective years and thus indicated that sorghum is not a successful trap crop in reducing the egg laying in the spring sown maize as a main crop. This may be due to the fact that in sown spring maize A. naqvii was more dominating rather than A. soccata species of shoot fly having more preference towards sorghum and the maize seedlings are comparatively more vigorous at early stage than that of sorghum. Similarly, Sarup et al (1986) had also reported that the oviposition soon after the emergence of maize seedlings at 2-3 leaf stage by A. naqvii restricted the success of sorghum as a trap crop. During 2012, another experiment was conducted with four selected treatments i.e. spring maize sown with two rows of protected sorghum as a trap crop (T6), delayed irrigation up to one week of emergence (T 8) and recommended seed treatment with imidacloprid 600 FS @ 6 ml/ kg seed before sowing along with untreated control, with a view that sorghum may be more effective as trap crop due to its better growth with rise in temperature. However, the egg laying did not differ significantly among different treatments when observed at 5 and 10 DAG in this experiment also (table 34). It was minimum in T8 (0.17and 0.20/ plant) and maximum in T6 (0.33/ plant) and T11 (0.40/ plant) at both these observations, respectively. The higher egg count of 0.73 per sorghum seedlings at 10 DAG though indicated oviposition preference of shoot fly species towards sorghum, but it could not reduced oviposition on maize crop as apparent from the different treatments. non significant differences in

123

Table 34. The shoot fly, Atherigona spp. incidence and grain yield in 2nd experiment during spring 2012. Mean egg count per plant Treatments 5 DAG T6Two rows of protected sorghum sown as trap crop T8- First irrigation after one week of crop emergence T10 - Imidacloprid 600 FS @ 6 ml per kg seed (standard) T11 - Untreated control CD (p=0.05) T6- Egg count per sorghum seedling 0.33 (1.15) 0.17 (1.08) 0.20 (1.09) 0.27 (1.12) (NS) 10 DAG 0.23 (1.11) 0.20 (1.09) 0.23 (1.11) 0.40 (1.18) (NS) 0.73 Per cent shoot fly incidence 10 DAG LI DH (%) (%) 26.34 10.83 15 DAG LI DH (%) (%) 19.46 16.65 Yield (q/ha) 45.63

26.89

8.95

16.57

11.54

50.22

1.79

0.37

0.35

0.72

69.60

30.83 3.62

15.66 2.39

28.17 5.81

22.53 6.52

38.81 8.91

Figures in parentheses are n+1 transformed values DAG: days after germination; LI: Leaf injury, DH: deadhearts 4.4.2. Incidence of shoot fly, Atherigona spp.: During 2011 and 2012 the leaf injury and deadhearts incidence due to shoot fly differed significantly in different treatments except leaf injury during 2011 and deadhearts during 2012 at 7 DAG, where all the treatments were on par with each other (Table 35 and 36) and in general, it increased progressively from 7 to 21 DAG during both the years and consequently the

differences become more pronounced at the last observation i.e. 21 DAG.The leaf injury incidence at 7 DAG varied from 0.32 2.02 and 3.36 16.01 per cent during 2011 and 2012, respectively. Though the differences were non significant during 2011, yet the minimum leaf injury was observed in T10 (seed treatment with imidacloprid) and maximum in T6 (with two rows of protected sorghum sown as a trap crop). During 2012, it was minimum in T10 (3.36 %), being on par with T9 (carbofuran 3 G @ 5 kg per acre at sowing) with incidence of 3.71 per cent, which were also on par with T1 and T2 (seed treatment with thiamethoxam 30 FS @ 4 g/ kg seed with incidence of 7.73% and at 6 g/kg seed with incidence of 6.67 %). It was maximum in untreated control T11 (16.01 %), being on par T3 and T5 (single of unprotected and protected sorghum sown as a trap crop with incidence of 14.38 -15.10 % ), which were further on par with T8 (first irrigation after one week of emergence), T6 (two rows of protected sorghum sown as a trap crop),

124

T7 (first irrigation at emergence) and T4 (two rows of unprotected sorghum sown as a trap crop), with incidence ranging from 11.31-15.10 per cent. The deadhearts incidence was observed only in T8 (0.65 %) during 2011. In 2012, there was no deadhearts formation in T 9 and T10 and it was maximum in T3 (3.66 %); and it varied from 0.34-2.43 per cent in remaining treatments, At 14 DAG, the leaf injury incidence ranged from 2.21 - 23.79 and 4.04 42.83 per cent during 2011 and 2012, respectively (table 35). In 2011, it was cooperatively low in plot sown with seed treatment (T1, T2 and T10) being minimum in T10 (2.21 %) which was on par with T2 (10.14 %). The maximum leaf injury was observed in T 7 (23.79 %), being on par with T11 (22.92 %). The remaining treatments i.e. T1, T3, T4, T5, T6, T8 and T9 were also on par with each other with leaf injury ranging from 13.46-18.73 per cent. Similarly in 2012 (table 36), leaf injury was minimum in T10 (4.04 %) being on par with T9 (6.10 %) and maximum in T4 (42.83 %) which was also on par with T6, T8, T11and T3 (33.35-40.26 %). The incidence was intermediate in T2 (seed treatment with higher dose of thiamethoxam with incidence of 25.27 %) which was also on par with treatments having higher incidence. The deadhearts incidence varied from 1.26 16.18 and 0.68 25.43 per cent during 2011 and 2012, respectively. In 2011 it was minimum in T 10 (1.26 %) which was also on par with T2 (3.73 %) which was further on par with T 9 (6.64 %). It was maximum in T11(untreated control with incidence of 16.18 %) which was also on par with T 7 (15.21 %) followed by T3 (10.94 %). T3 was on par with T1, T2, T6 and T8, all on par with each other (8.01-9.53 %). This group was on par with T9 also. The lower dose of thiamethoxam as seed treatment (T1), application of carbofuran at sowing, restricted irrigation at emergence (T 8) ; and T3, T4, T5 and T6 i.e. different treatments with sorghum sown as a trap crop being on par with each other (8.01 10.94 %) were better than untreated control in reducing deadhearts formation. Similarly in 2012, deadhearts incidence was significantly lower in T 10 (0.68 %) being on par with T9 (1.33 %) was followed by T2 (10.58 %) and T8 (restricted irrigation with incidence of 13.06 %), both being on par with each other were further on par with T 1 and T7 (16.10 -16.34 %). (table 21). It was maximum in T4 (25.43 %), which being on par with treatments of sorghum sown as a trap crop treatments (20.05 - 22.39 %) were also on par with untreated control (22.27 %). At 21 DAG the treatment differences were more prominent as indicated by non overlapping of different treatment groups for shoot fly incidence. The per cent leaf injury ranged from 13.25 40.67 and 4.33 - 28.63 per cent during 2011 and 2012, respectively. In 2011, it was significantly lower (13.25 %) in T10 (seed treatment with imidacloprid) and higher in T 11 (untreated control with incidence of 40.67 %) which was on par with single row of protected sorghum sown as a trap-T5 (40.61 %) and both were also on par with other treatments of sorghum sown as a trap crop T6,T4 and T3 (33.00 - 34.94 %) and T7 (first irrigation at emergence with

125

incidence of 37.42 %). The leaf injury (27.32 and 30.85 %) in thiamethoxam seed treatments (T 1 and T2), T8 (first irrigation after one week of emergence with incidence of 26.43 %) and recommended application of carbofuran at sowing T9 (29.58 %) being on par with each other was lower than that in untreated control. In 2012 also, the leaf injury being minimum in T 10 (4.33 %) and on par with T9 (4.34 %) was significantly lower than the remaining treatments Table 35. The shoot fly, Atherigona spp. incidence in different treatments during spring 2011. Mean shoot fly incidence at 7 DAG Treatments T1 - Thiamethoxam 30 FS @ 4 g per kg seed one day before sowing T2 - Thiamethoxam 30 FS @ 6 g per kg seed one day before sowing T3- Single row of unprotected sorghum sown as a trap crop T4- Two rows of unprotected sorghum sown as a trap crop T5- Single row of protected sorghum sown as a trap crop T6- Two rows of protected sorghum sown as a trap crop T7 - First irrigation at crop emergence T8 - First irrigation after one week of crop emergence T9- Carbofuran 3 G @ 5 kg per acre at sowing (standard) T10 - Imidacloprid 600 FS @ 6 ml per kg seed (standard) T11 - Untreated control CD (p=0.05) LI (%) 0.71 DH (%) 0.00 14 DAG LI (%) 13.46 DH (%) 8.01 21 DAG LI (%) 30.85 DH (%) 17.95

1.01 0.33 0.34 0.67 2.02 1.03 1.65 0.67 0.32 1.35 NS

0.00 0.00 0.00 0.00 0.00 0.00 0.65 0.00 0.00 0.00 0.27

10.14 17.24 13.69 15.38 16.17 23.79 18.73 17.28 2.21 22.92 8.71

3.73 10.94 8.90 8.68 8.77 15.21 9.53 6.64 1.26 16.18 3.59

27.32 34.94 33.53 40.61 33.00 37.42 26.43 29.58 13.25 40.67 8.17

15.20 19.89 21.92 21.01 20.58 22.60 17.22 11.95 3.47 23.83 4.11

DAG: days after germination; LI: Leaf injury, DH: deadhearts

126

Table 36.

The shoot fly, Atherigona spp. incidence in different treatments during spring 2012. Mean shoot fly incidence at 7 DAG 14 DAG LI (%) 31.24 DH (%) 16.10 21 DAG LI (%) 27.50 DH (%) 25.33 Mean of 2011 & 2012 at 21 DAG LI DH (%) (%) 29.17 21.64

Treatments T1 - Thiamethoxam 30 FS @ 4 g per kg seed one day before sowing T2 - Thiamethoxam 30 FS @ 6 g per kg seed one day before sowing T3- Single row of unprotected sorghum sown as a trap crop T4- Two rows of unprotected sorghum sown as a trap crop T5- Single row of protected sorghum sown as a trap crop T6- Two rows of protected sorghum sown as a trap crop T7 - First irrigation at crop emergence T8 - First irrigation after one week of crop emergence T9- Carbofuran 3 G @ 5 kg per acre at sowing (standard) T10 - Imidacloprid 600 FS @ 6 ml per kg seed (standard) T11 - Untreated control CD (p=0.05)

LI (%) 7.73

DH (%) 0.67

6.67

1.05

25.27

10.58

28.63

16.22

27.97

15.71

15.10

3.66

33.35

20.43

27.06

28.10

31.00

23.99

11.31

2.43

42.83

25.43

25.89

31.52

29.72

26.72

14.38

1.35

30.34

20.05

21.96

31.49

31.28

26.25

13.18 12.66 13.46

0.34 1.68 1.92

40.26 28.36 36.88

22.39 16.34 13.06

22.71 20.73 16.81

30.83 30.92 19.10

27.85 29.07 21.62

25.71 26.76 18.16

3.71

0.00

6.10

1.33

4.34

3.02

16.96

7.49

3.36 16.01 6.23

0.00 1.98 NS

4.04 36.08 10.89

0.68 22.27 6.72

4.33 23.64 12.22

1.09 32.02 4.88

8.79 32.15 7.14

2.28 27.92 3.10

DAG: days after germination; LI: Leaf injury, DH: deadhearts

127

including untreated control (16.81-27.50 %) being maximum in T2 (28.63 %). The deadhearts incidence ranged from 3.47 22.60 and 1.09 32.02 per cent during 2011 and 2012, respectively. In 2011, it was significantly low in T10 (recommended imidacloprid seed treatment with incidence of 3.47 %) followed by T9 (with recommended application of granular application of carbofuran) which was on par with T2 (higher doses of thiamethoxam for seed treatment). T 9 and T2 with 11.95 and 15.20 per cent deadhearts formed intermediate group. The incidence was significantly more in sorghum sown as trap a crop treatments T3, T6, T5 and T4 (19.89-21.92 %) and T7 (first irrigation at emergence with incidence of 22.60 %), which were on par with each other as well as with untreated control (23.83 %) . Although, incidence of deadhearts in T8 (17.22 %) and T1 (17.95 %) was significantly lower than untreated control but was more than that in recommended treatments (T9 and T10). During 2012 also, deadhearts incidence being on par with each other was significantly lower in T10 and T 9 (1.09 and 3.02 %). It was maximum in T11 (untreated control with incidence of 32.02 %), which was on par with treatments T 3 T6 having sorghum sown as a trap crop (28.10 - 31.52 %) and T7 (30.92 %) again proving that sowing of sorghum as intercrop in spring maize was not effective in reducing the shoot fly incidence. The higher dose of thiamethoxam as seed treatment T2 (16.22 %), and irrigation after one week of emergence T8 (19.10 %) forming intermediate group, showed some potential in controlling shoot fly infestation in comparison to untreated control (32.02 %). In 2012 also, T 1 (thiamethoxam seed treatments at lower dose) was not found effective in reducing deadhearts incidence (25.33 %). During 2011 and 2012 there was an increase in deadhearts incidence at higher dose of thiamethoxam from 3.73 15.20 per cent and 10.58 16.22 in 7 days i.e. from 14 DAG to 21 DAG, respectively which implies that insecticide may not be persisted at a level required to provide protection to shoot fly in spring sown maize. The delayed irrigation i.e. up to one week after emergence (T 8) to some extent was found effective in reducing shoot fly incidence in 2012. However, increase in incidence from 14 to 21 DAG in this treatment during 2011 may be attributed to the corresponding more egg laying during susceptible crop growth period at that site. The pooled data for shoot fly incidence recorded at 21 DAG for 2011 and 2012 (table 36) also showed that the leaf injury was significantly lower in recommended imidacloprid seed treatment-T10 (8.79 %) followed by granular application of carbofuran-T9 (16.96 %), which was also on par with delayed irrigation-T8 (21.62 %). It was maximum in untreated control (32.15 %), which was on par with all the remaining treatments (27.97 31.28 %). The differences were more clear by considering the observations of deadhearts incidence that again revealed that it was minimum in recommended T10 (imidacloprid seed treatment with incidence of 2.28 %) followed by T9 ( granular application of carbofuran with incidence of 7.49 %). The incidence was

128

intermediate and being on par with each other in T2 (higher dose of thiamethoxam with incidence of 15.71 %) and T8 (delayed irrigation with incidence of 18.16 %) but significantly less than that in untreated control (27.92 %). The sorghum sown as a trap crop in treatments T 3 T6 and first irrigation at emergence T7 (23.99 - 26.76 %) being on par with each other as well as untreated control. Similar results were obtained in March sown experiment conducted during 2012, the observation recorded on 10 and 15 DAG only revealed that as in scheduled experiments imidacloprid seed treatment plot (T10) had significantly lower leaf injury (1.79 to 0.35 %) incidence (table 34) followed by two rows of protected sorghum sown as a trap crop - T6 (26.34 19.46 %) and delayed irrigation - T8 (26.89 16.57 %), both being on par with each other but significantly better than untreated control (30.83 28.17 %) at both the observations. The deadhearts incidence was also minimum in T10 (imidacloprid seed treatment plot with incidence of 0.37 to 0.72 %) at these observations. It was lower in T 8 (8.95 and 11.54 %) than untreated control (15.56 and 22.53 %) at 10 and 15 DAG, however, only at 10 DAG in T 6 (10.83 %) ; and deadhearts was on par with untreated control (22.53 %) at 15 DAG in this treatment (16.65 %). This showed that sorghum as a trap crop is comparatively less effective than the other treatments in reducing shoot fly incidence in early march sown crop also. The grain yield (q/ ha) differed significantly among different treatments during both the test years (table 37). In 2011, it was maximum in T10 (86.37 q/ ha) which was on par with T9 (77.73 q/ ha) which was further on par with higher dose of thiamethoxam seed treatment-T2 (70.94 q/ ha). It was minimum in untreated control (52.74 q/ ha) which was on par with T7 (53.10 q/ ha) and all the treatments of sorghum as trap crop (54.54 57.95 q/ha) except T4 i.e. two rows of protected sorghum as trap crop. The yield levels were intermediate in lower dose of thiamethoxam T1 (65.02 q/ ha) and delayed irrigation- T8 (66.39 q/ha). Similarly in 2012, the grain yield, being on par with T9 (79.55 q/ha), was maximum in T10 (85.78 q/ ha). The higher dose of thiamethoxam T2 (67.11 q/ ha) and delayed irrigation T8 (62.78 q/ha) being on par with each other and also with T1 (lower dose of thiamethoxam) had significantly more yield than the remaining treatments. It was minimum in untreated control which was on par with T 5, T6, T7 (47.56- 50.67 q/ha), which were further on par with T3 and T4 (51.56-53.78 q/ha). The pooled data of both the test years revealed similar results with significantly more yield in T 10 (86.08 q/ha) followed by T9 (78.48 q/ha) and it was intermediate in T2 (69.01 q/ha) and T8 (64.58 q/ha). It was minimum in untreated control (48.16 q/ha) which was on par with T 5, T6, T7 (50.56 52.95 q/ha). The sorghum trap crop treatments (T3- T6) being on par with each other have significantly lower grain yield than recommended insecticide treatments (T9 and T10 ). The present study revealed that the sorghum as a trap crop proved comparatively less effective in management of shoot fly,

129

whereas T2 and T8 showed some potential in integrated strategies for management of shoot fly in future. Similar results were obtained in 2nd experiment conducted with selected treatments during 2012 (table 34). The recommended seed treatment with imidacloprid recorded significantly higher grain yield (69.60 q/ha), while delayed irrigation treatment T8 (50.22 q/ha) was also significantly better than untreated control (38.81 q/ha). However, sorghum as trap crop sown even in March T6 (45.63 q/ha) was also on par with untreated control as well as T 8. Thus again it revealed the effectiveness of recommended insecticide treatments and potential of delayed irrigation up to one week of emergence in management of shoot fly in spring sown maize. Similar to present studies, Table 37. The grain yield (q/ ha) in different treatments during spring 2011 and 2012 Mean grain yield during Treatments T1 - Thiamethoxam 30 FS @ 4 g per kg seed one day before sowing T2 - Thiamethoxam 30 FS @ 6 g per kg seed one day before sowing T3- Single row of unprotected sorghum sown as a trap crop T4- Two rows of unprotected sorghum sown as a trap crop T5- Single row of protected sorghum sown as a trap crop T6- Two rows of protected sorghum sown as a trap crop T7 - First irrigation at crop emergence T8 - First irrigation after one week of crop emergence T9- Carbofuran 3 G @ 5 kg per acre at sowing (standard) T10 - Imidacloprid 600 FS @ 6 ml per kg seed (standard) T11 - Untreated control CD (p=0.05) 2011 65.02 70.94 57.95 59.30 54.44 55.26 53.10 66.39 77.73 86.37 52.74 9.05 2012 60.00 67.11 51.56 53.78 47.56 50.67 48.14 62.78 79.55 85.78 43.56 7.78 Mean of 2011 & 2012 62.54 69.01 54.77 56.55 50.99 52.95 50.56 64.58 78.48 86.08 48.16 5.79

According to Sajjan and Sekhon (1985), Rao and Panwar (1992, 1995c), Kanta et al (2006), Jindal and Hari (2008a, 2011) as in the present studies application of granular insecticides and imidacloprid as seed treatment was effective in managing shoot fly infestation on spring sown maize crop in north India. But thiamethoxam appeared to be effective only at higher doses

130

of 6 ml/ kg seed in the present studies, however, the effectiveness of thiamethoxam 70 WS @ 2 g/ kg seed of sorghum in reducing dead hearts incidence (7.94 %) was also reported by Kumar and Prabhuraj (2007) but comparatively less effectiveness of thiamethoxam 70 WS @ 3.0 and 5.0 g/kg seed (17.02 and 15.10 % deadhearts at respective doses) in spring sown maize was reported by Jindal and Hari (2011). These differences might be due to different sowing time and prevalent predominant species of shoot fly in different studies conducted on spring maize and sorghum. The higher shoot fly incidence in treatments with sorghum as trap crops, in the present studies indicated that this cultural control method was less effective in management of shoot fly in spring sown maize. Sarup et al (1986) similarly reported the ineffectiveness of sorghum as trap crop, where deadhearts incidence of 24.8-37.9 and 56.4-71.2 per cent was recorded on maize plants having varied proportion of sorghum plants as compared to incidence of 27.2 and 64.7 per cent on sole maize plots, during 1985 and 1986, respectively. The optimization of irrigation frequency in relation to shoot fly population dynamics seems to be an important strategy in its management. The overall perusal of the data indicated that these finding may be integrated with time of sowing, use of tolerant cultivars and population dynamics of shoot fly vis-a-vis irrigation frequency may be integrated to strengthen the IPM for shoot fly in spring sown maize.

131

Chapter V SUMMARY Maize, Zea mays (L.) (Poaceae), ranking third in area after wheat and rice is one of the important cereal crops in the world. The generation of new agricultural technology in India has led to round the year cropping of maize and offers a bright prospect in crop diversification in the Punjab. The cultivation of spring maize i.e. February sown crop is becoming popular because of its higher yield potential. The continuous planting of maize has led to the invasion of polyphagous shoot flies species as the key pests on spring sown maize in the northern India (Sarup et al 1984). Moreover, the incidence and extent of losses by insect pests complex to the newly recommended cultivation of maize during spring season in the Punjab is not available. Shoot fly, Atherigona naqvii Steyskal is a serious pest of spring sown maize in the Punjab (Sandhu and Kaushal 1976, Singh and Kanta 2004) causing upto 85.8 per cent deadhearts incidence in the Punjab (Sajjan and Sekhon 1985). As the adult flies oviposit on the emerging seedlings and in cracks and crevices around the seedlings and shoot fly maggot feeding inside the leaf whorls is not easily accessible to insecticides sprayed that is why, the seed treatment and soil application of insecticides are recommended for its control. The exploitation of host plant resistant also holds a good potential for the management of shoot fly. Resistance to shoot fly in sorghum is expressed in terms of oviposition non-preference, antibiosis and tolerance (Dhillon et al 2005a). However, the mechanism of host plant resistance to shoot fly and factors influencing these mechanisms have not been identified in the promising maize cultivars for spring season crop in the Punjab. These factors can be quantified or monitored easily in plant populations and such characters can be used as "marker traits" in screening and selection for resistance to insect pests (Chamarthi 2008). Moreover, there is a need to develop integrated pest management (IPM) strategies involving the cultural methods of management of shoot fly spp. in spring season. Keeping in view the above facts, the present studies on incidence of insect pests and management of shoot fly, Atherigona spp. in spring sown maize were undertaken with the following objectives:

a) To study the populations build up of insect pests and their extent of losses in spring maize. b) To establish the morphological and biochemical bases of resistance against shoot fly in different promising cultivars.
c) To formulate integrated approaches for management of shoot fly in spring sown maize. To meet these objectives, studies were conducted at the Research Farm and the Maize Entomology Laboratory, Department of Plant Breeding and Genetics, Punjab Agricultural University (PAU), Ludhiana, during spring 2011 and 2012. One of the field experiments was also

132

carried out at the farmers field in village Chagra of district Hoshiarpur (Punjab). The biochemical analysis was carried out in the Quality Laboratory, Department of Plant Breeding and Genetics, PAU, Ludhiana. Population build up of insect pests of spring maize at Hoshiarpur: The experiment on population build up of insect pests of spring maize on two maize hybrids JH 3459 and PMH 2 at two nitrogen levels of 120 and 90 kg ha -1 under unprotected conditions was conducted at Hoshiarpur during spring 2011 and 2012. The number of eggs laid by shoot fly, Atherigona spp. per plant did not differ significantly during different observations i.e. 7, 14 and 21 days after germination (DAG) in both the test hybrids and at different nitrogen level during 2011 and 2012. The interaction of hybrids and nitrogen levels was also non significant during both the test years. The leaf injury incidence irrespective of hybrids was numerically less at higher nitrogen level (23.14 and 31.97 %) than that at lower one (23.19 and 33.04 %) in the respective test years. It differed significantly irrespective of nitrogen levels, among hybrids during 2012, being more in PMH 2 (39.09 %) than that in JH 3459 (25.91%). Irrespective of hybrids, significantly more deadhearts were formed at nitrogen level of 120 kg ha -1 (14.65 %) than that in 90 kg ha-1 (11.99 %) during 2011 only. However, irrespective of nitrogen level, the hybrid PMH 2 (16.40 and 25.58 %) had more deadhearts incidence than JH 3459 (10.24 and 16.40 %) in both the test years. The non significant interaction of hybrids and nitrogen level for deadhearts incidence again implies that hybrid PMH 2 had higher incidence than JH 3459 at both the nitrogen levels. This infestation level (in terms of deadhearts incidence) confirmed the key pest status of shoot fly in spring sown maize in the Punjab. In the present studies, the incidence of maize stem borer, Chilo partellus; corn earworm, Helicoverpa armigera; armyworm, Mythimna separata; pyrilla, Pyrilla perpusilla; small brown planthopper, Laodelphax striatellus and grasshopper spp. and beneficial arthropods i.e. coccinellids and spiders at Hoshiarpur was sporadic in spring sown maize, but shoot fly was observed to be the major yield reducing factor during both the test years. The differences in the grain yield (q/ ha) between the genotypes as well as under different nitrogen levels in yield, even at varied infestation levels of insect pests, might be due to the inherent yield potential of the test genotypes. Estimation of losses caused by insect pests in spring sown maize The experiment on insect pests incidence and estimation of losses caused by them in spring sown maize was conducted at PAU, Ludhiana on hybrid JH 3459 and PMH 2 sown under three sowing time i.e. on end January, mid and end February under protected/ unprotected conditions against shoot fly. The maximum egg counts of Atherigona spp. were observed at 10 DAG, irrespective of sowing date, in 2011 and at 17, 7 and 12 DAG in end January, mid February and end February sown crops, respectively during 2012. However, the egg counts differed non significantly in the hybrids and crop protection conditions except for few observations in 2012.

133

The egg counts had positive but non significant correlation with maximum and mean air temperature, but minimum temperature had a more (+ve) influence on egg laying by shoot fly. The relative humidity (R.H.) of air had non significant correlation with egg laying by shoot fly during both the test years. The mean leaf injury and deadhearts incidence due to shoot fly was higher in end February followed by mid February sown crop and being the least in end January sown crop during both the test years. The mean leaf injury incidence, irrespective of hybrids, was lower in protected conditions (5.97 and 5.56 %) than that in unprotected conditions (30.60 and 15.02 %) during 2011 and 2012, respectively. Similarly, deadhearts incidence was also lower in protected conditions (1.52 and 4.03 %) than that in unprotected conditions (16.55 and 25.01 %) during 2011 and 2012, respectively. The mean deadhearts incidence, irrespective of crop protection conditions, was significantly lower in JH 3459 than that in PMH 2 at all the sowing times during both the test years. The increase in deadhearts incidence was 4.21 to 9.99 and 3.91 to 15.40 per cent in 2011 and 2012, respectively from 10 to 15 DAG in the mid February sown crop, supporting the fact that the shoot fly damage can also be minimized with suitable control measures other than recommended seed treatment and soil application of insecticides at the first appearance of infestation also. The incidence (leaf injury and deadhearts) increased with rise of minimum air temperature in maize crops sown at different times, however, R.H. was not found correlated with shoot fly damage. Apart from incidence of shoot fly, the high incidence of small brown planthopper, Laodelphax striatellus (Falln) was also observed for the first time in spring sown maize in the Punjab during spring 2011, where as the incidence of other insect pests of maize was low in the crops sown at 3 different times during both the test years. The highest population counts of L. striatellus were observed in 3rd week of April (96.06 186.08/ sweep) and 4th week of April (0.16 1.49/ plant) on crop sown at different times during 2011 and 2012, respectively.The occurrence of small brown planthopper could be a limiting factor in future, being a vector of number of viruses on cereal crops including maize. The correlation of its population counts was negative with maximum (r= - 0.14 to - 0.52) and mean air temperature (r= - 0.13 to - 0.57) in crops sown at different times. The higher population counts of coccinellids were also observed in correspondence with the higher population of small brown planthopper during 2011, highlights the need of detailed studies on the insect biodiversity and their interaction with environmental factors, in future. The grain yield losses due to shoot fly increased with delay in sowing of spring maize as it caused 27.93 - 37.97, 38.21 - 38.66; and 38.99 - 45.04 % losses in end January, mid and end February sown crop, respectively during 2011 and 2012. Relative susceptibility of different genotypes to shoot fly, Atherigona naqvii and role of various morphological and biochemical plant characteristics in resistance to A. naqvii: The screening of 8 maize genotypes i.e. six hybrids (JH 3459, PMH 2, JH 31244, JH 3956, PMH 1 and Parkash) and 2 inbred lines (LM 16 and CM 143) against Atherigona naqvii

134

was conducted to determine the role of various morphological and biochemical plant traits in resistance to shoot fly, A. naqvii during spring 2011 and 2012. The mean number of eggs per plant in the test genotypes differed non significantly in all the observations during both the test years except at 3 DAG in 2011 indicated that the antixenosis for oviposition in maize to A. naqvii was not observed in the test genotypes. Only reducing sugars content showed positive and significant correlation (r = 0.81*) with oviposition of A. naqvii in the test genotypes. The differential reaction in test genotypes to A. naqvii infestation became evident at 20 DAG and that too on the basis of deadhearts incidence only. The genotypes i.e. JH 3459 (10.49 %) & JH 3956 (10.95 %); CM 143 (15.16 %), PMH 1 (15.35 %) & JH 31244 (16.70 %); and LM 16 (21.48 %), PMH 2 (21.52 %) & Parkash (23.57 %) on the basis of mean per cent deadhearts incidence were categorized as least, intermediate and highly susceptible to shoot fly, A. naqvii, respectively. The relative tolerance to A. naqvii infestation in terms of proportion of deadhearts out of shoot fly incidence (leaf injury + deadhearts) among different genotypes was almost similar to above grouping. This proportion was maximum and minimum in LM 16 (47.43 %) and JH 3956 (32.45 %), respectively. Moreover, the correlation between this proportion and leaf injury was non significant but the correlation was positive and significant with deadhearts (r=0.70*) again proved that tolerance is key mechanism of resistance in different genotypes and these had differential capacity to withstand the infestation of A. naqvii. The genotypes LM 16, PMH 2 and Parkash (susceptible genotypes) having lower larval and pupal periods; and higher larval survival, male & female pupal weight and fecundity also suggested the role of antibiosis mechanism in addition to tolerance in least susceptible cultivars. The higher larval survival (51.04 and 63.87 %) in susceptible group i.e. LM 16, PMH 2 and Parkash might have resulted in comparative higher deadhearts formation and subsequently in higher proportion of deadhearts out of total incidence. The various morphological and biochemical traits of the seedlings were studied at 5th leaf stage to ascertain their role in host plant resistance to A. naqvii in different genotypes. Among morphological traits leaf glossiness, leaf sheath pigmentation, leaf surface wetness, leaf trichome traits did not influence the oviposition and damage by A. naqvii in test genotypes. Only the seedling vigour was negatively correlated (r = 0.84*) with proportion of deadhearts out of total incidence and low seedling vigour in susceptible hybrid PMH 2 ( with high seedling vigour score of 2.83) indicated that seedling vigour is related to tolerance in maize genotypes against shoot fly. The leaf length, width and the stem girth was more in susceptible genotypes can serve as reliable markers in host plant resistance breeding programmes against A. naqvii. Among biochemical factors only reducing sugars content being phagostimulatory to feeding maggots had positive and significant correlation with larval survival (r = 0.87*), fecundity (0.71*), leaf injury (r = 0.77), deadhearts (r= 0.88) and oviposition (r= 0.88) by shoot fly indicates its role in imparting susceptibility to A. naqvii in test maize genotypes.

135

Cultural and chemical control of shoot fly, Atherigona spp. on spring maize: The experiments to develop integrated management strategies for shoot fly, Atherigona spp. in spring sown maize were conducted by treating the seed with thiamethoxam 30 FS @ 4 & 6 ml/ kg seed, using sorghum as a trap crop in 4 different combinations, altering of irrigation frequency; and seed treatment & soil application of imidacloprid 600 FS @ 6 ml per kg seed & carbofuran 3G @ 5 kg/ acre (as standard checks), respectively along with untreated control at PAU, Ludhiana on susceptible hybrid PMH 2 during 2011 and 2012. The mean egg laying per plant was on par in different treatments in all the observations recorded during both the test years, except at 6 DAG during 2012 where it was maximum in untreated control (1.07 eggs/ plant) and minimum where first irrigation was given after one week of emergence (0.27 eggs/ plant) thus indicating the role of moisture level in the field for egg laying of the pest. Non significant reduction in egg laying was observed in the standard checks. The egg counts in untreated control (0.14 and 0.45/ plant) were comparable in sorghum sown as a trap crop treatments (0.11 to 0.17 and 0.35 to 0.45/ plant) in the respective years. This indicated that, sorghum as a trap crop is not successful in reducing the egg laying on the main spring sown maize crop. The mean deadhearts incidence in both the years was minimum in standard checks i.e. imidacloprid seed treatment (2.28 %) followed by granular application of carbofuran (7.49 %). The incidence was intermediate, being on par with each other in the treatments with higher dose of thiamethoxam (15.71 %) and delayed irrigation (18.16 %) was significantly less than that in untreated control (27.92 %). The sorghum sown as a trap crop (in all combinations) and first irrigation given at emergence (23.99 - 26.76 %) were on par with each other and also with untreated control. This showed that sorghum as a trap crop was not effective in reducing shoot fly incidence but seed treatment with higher dose of thiamethoxam and delayed irrigation showed potential in integrated strategies for management of shoot fly in future. The pooled data of grain yield of both the test years revealed that it was also significantly more in seed treatment with imidacloprid (86.08 q/ ha) followed by soil application of carbofuran 3G (78.48 q/ ha) and it was intermediate in seed treatment with higher dose of thiamethoxam (69.01 q/ ha) and delayed irrigation (64.58 q/ ha). However, it was minimum in untreated control (48.16 q/ ha) and treatments with sorghum sown as a trap crop had significantly lower grain yield than the standard checks. The present studies revealed that the overall strategy for shoot fly management in spring sown maize should involve integration of sowing time, use of comparatively tolerant cultivars, enhanced seedling vigour, careful monitoring of the pest at susceptible stage i.e. 3-10 days after germination vis-a-vis alteration in irrigation and adoption of plant protection measures at sowing or at the first appearance of the shoot fly infestation.

136

REFERENCES

Anonymous (2006) Package of Practices for Kharif Crops. Pp. 21. Punjab Agricultural University, Ludhiana. Anonymous (2010) Package of Practices for Rabi Crops. Pp. 81. Punjab Agricultural University, Ludhiana. Anonymous (2011) 55th Annual Progress Report. Pp. 13. All Indian Coordinated Research Project. Directorate of Maize Research, New Delhi. Anonymous (2012) Package of Practices for Kharif Crops. Pp. 23. Punjab Agricultural University, Ludhiana. AOAC (2000) Official methods of analysis: Association of official analytic chemists, 17th edn. Gaithersburg, MD, USA. Arabjafari K H and Jalali S K (2007) Identification and analysis of host plant resistance in leading maize genotypes against spotted stem borer, Chilo partellus (Swinhoe) (Lepidoptera: Pyralidae). Pak J Biol Sci 10: 188595. Beck S D (1965) Resistance of plants to insects. Annu Rev Entomol 10: 20732. Bhise H T, Desai B B and Chavan U D (1997) Assessments of some biochemical parameters responsible for shoot fly resistance in sorghum. J Maharashtra agric Univ 21: 12729. Bialczyk J, Lechowski Z and Libik A (1999) The protective action of tannins against glasshouse whitefly in tomato seedlings. J agric Sci 133:197201. Bozzola J J and Russell L D (1999) Electron Microscopy: Principles and techniques for biologists. 2nd edn. Pp. 670. Jones and Bartlett Publishers, Inc. Sudbury, MA, USA Caciagli P (1991) Effect of anaesthesia with carbon dioxide on vectoring ability and survival of the planthopper Laodelphax striatellus. Ann Appl Biol 119: 25764. Chamarthi S K (2008) Biochemical mechanisms of resistance to shoot fly, Atherigona soccata (Rondani) in sorghum, Sorghum bicolor (L.) Moench. Ph.D. dissertation. Jawaharlal Nehru Technological University Hyderabad, Andhra Pradesh, India. Chamarthi S K, Sharma H C, Sahrawat K L, Narasu L M and Dhillon M K (2011) Physicochemical mechanisms of resistance to shoot fly, Atherigona soccata in sorghum, Sorghum bicolor. J Appl Entomol 135: 44655. Chatterji S M, Bhamburkar M W, Marwaha K K, Panwar V P S and Siddiqui K H (1973) Evaluation of some systemic insecticides for the control of shoot fly infesting spring maize. Indian J Ent 35: 16163. Chavan M H, Phadnawis B N, Hudge V S and Salunkhe M R (1990) Biochemical basis of shoot fly tolerant sorghum genotypes. Ann Plant Physiol 4: 21520.

137

Classen D, Arnason J T, Serratos J A, Lambert J D H, Nozzolillo C and Philogne B J R (1990) Correlation of phenolic acid content of maize to resistance to Sitophilus zeamais, the maize weevil, in CIMMYT's collections. J Chem Ecol 16: 30115. Davies J C and Seshu Reddy K V (1981) Shoot fly species and their graminaceous hosts in Andhra Pradesh, India. Insect Sci Applic 2: 3338. Deeming J C (1971) Some species of Atherigona Rondani (Diptera: Muscidae) from northern Nigeria, with special reference to those injurious to cereal crops . Bull Ent Res 61: 133 90. Delobel A G L (1981) The distribution of the eggs of the sorghum shoot fly, Atherigona soccata Rondani (Diptera: Muscidae). Insect Sci Applic 2: 6366. Delobel A G L (1983) Humidity effects on Atherigona soccata: Egg development and hatch. Ent exp appl 33: 26975. Dhillon M K (2004) Effects of cytoplasmic male-sterility on expression of resistance to sorghum shoot fly, Atherigona soccata (Rondani). Ph.D. dissertation. Chaudhary Charan Singh Haryana Agricultural University, Hisar, Haryana, India. Dhillon M K, Sharma H C, Reddy B V S, Singh R, Naresh J S and Kai Z (2005b) Relative susceptibility of different male-sterile cytoplasms in sorghum to shoot fly, Atherigona soccata. Euphytica 144: 27583. Dhillon M K, Sharma H C, Singh R and Naresh J S (2005a) Mechanisms of resistance to shoot fly, Atherigona soccata in sorghum. Euphytica 144: 30112. Dhillon M K, Sharma H C, Singh R and Naresh J S (2006) Influence of cytoplasmic male-sterility on expression of physico-chemical traits associated with resistance to sorghum shoot fly, Atherigona soccata Rondani. SABRAO J 38: 10522. Doggett H, Starks K J and Eberhart S A (1970) Breeding for resistance to the sorghum shoot fly. Crop Sci 10: 52831. Doharey K L, Srivastava B G, Jotwani M G and Dang K (1977) Effect of temperature and humidity on the development of Atherigona soccata Rondani. Indian J Ent 39: 21115. Elahi M and Khan N (1973) Free amino acids of Pakistani wheat varieties. J agric Food Chem 21: 74344. Endo S and Tsurumachi M (2000) Insecticide resistance and insensitive acetycholinesterase in small brown planthopper, Laodelphax striatellus. J Pestic Sci 25: 39597. Frazer B D (1988) Coccinellidae.In: Minks A K and Haaewijin B (eds.) Aphids-their biology, natural enemies and control. Vol. B. Pp. 231-47. Elsevier, New York, USA. Gagandeep and Kanta U (2007a) Estimation of losses due to shoot fly Atherigona naqvii Steyskal in spring maize. J Insect Sci 20: 15255.

138

Gagandeep and Kanta U (2007b) Influence of date of sowing on incidence of Atherigona naqvii Steyskal in spring maize. J Insect Sci 20: 25456. Gao B L, Wu J, Huang S J, Mu L F and Han Z J (2008) Insecticide resistance in field populations of Laodelphax striatellus Falln (Homoptera: Delphacidae) in China and its possible mechanisms. Int J Pest Manag 54:1319. Gomez K A and Gomez A A (1984) Statistical Procedures for Agricultural Research, 2nd edn. Pp. 653. Wiley-interscience, New York. Hari N S and Jindal J (2008) Chemical control and damage potential of shoot fly, Atherigona naqvii Steyskal on spring sown maize. Indian J Plant Prot 36: 26365. Hari N S and Jindal J (2009) Assessment of Napier millet (Pennisetum purpureum x P. glaucum) and sorghum (Sorghum bicolor) trap crops for the management of Chilo partellus on maize. Bull Ent Res 99: 13137. Hari N S, Jindal J, Aggarwal N and Hari J K (2013) Egg distribution of Atherigona naqvii Steyskal (Diptera: Anthomyiidae) infesting spring maize. Indian J Plant Prot (accepted). Hewett EW (1977) Some effects of infestation on plants: A physiological viewpoint. New Zealand Entomol 6: 23543. Jalali S K and Singh S P (2003) Biological control of C. partellus (Swinhoe) in fodder maize by inundative releases of parasitoids. Indian J Plant Prot 31: 9395. Jayanthi P D K, Reddy B V S, Gour T B and Reddy D D R (2002) Early seedling vigour in sorghum and its relationship with resistance to shoot fly, Atherigona soccata Rondani. J ent Res 26: 93100. Jindal J, Aggarwal N, Singh G, Kumar M and Grewal M S (2012) Effect of different methods of sowing on the incidence of shoot fly, Atherigona naqvii Steyskal in spring sown maize. Pp. 92930.In: Crop Improv, Special issue on International conference on Sustainable Agriculture for Food and Livelihood Security, PAU, Ludhiana, India. Jindal J and Hari N S (2008a) Evaluation of seed treatment and foliar sprays against shoot fly, Atherigona naqvii Steyskal on spring sown maize. J Insect Sci 21: 24952. Jindal J and Hari N S (2008b) Studies on components of resistance in maize genotypes to Chilo partellus (Swinhoe). Indian J Ent 70: 31418. Jindal J and Hari N S (2011) Efficacy of seed treatments against shoot fly, Atherigona naqvii Steyskal on spring maize in Punjab. J Insect Sci 24: 8690. Jindal J, Hari N S, Grewal M S and Chawla J S (2007) Evaluation of different genotypes against shoot fly, Atherigona naqvii Steyskal in spring sown maize. Crop Improv 34: 16062.

139

Jotwani M G, Marwaha K K, Srivastava K M and Young W R(1970) Seasonal incidence of shoot fly (Atherigona varia soccata Rond.) in jowar hybrids at Delhi. Indian J Ent 32: 715. Kabre G B and Ghorpade S A (1999) Susceptibility to maize stem borer, Chilo partellus (Swinhoe) in relation to sugars, proteins and free amino acids content of maize germplasm and F1 hybrids. J Insect Sci 12: 3740. Kamatar M Y and Salimath P M (2003) Morphological traits of sorghum associated with resistance to shoot fly, Atherigona soccata Rondani. Indian J Plant Prot 31: 7377. Kamatar M Y, Salimath P M, Ravi Kumar R L and Swamy Rao T (2003) Heterosis for biochemical traits governing resistance to shoot fly in sorghum [Sorghum bicolor (L.) Moench]. Indian J Genet 63: 12427. Kanta U, Singh D P and Sekhon S S (2006) Chemical control of shoot fly, Atherigona naqvii Steyskal on spring maize in Punjab. J Insect Sci 19: 10506. Karanjkar R R, Chundurwar R D and Borikar S T (1992) Correlations and path analysis of shoot fly resistance in sorghum. J Maharashtra agric Univ 17: 38991. Katole S R, Kudale N V, Thakare S M and Sadawarte A K (2003) Management of sorghum tissue borers with imidacloprid and thiamethoxam, the new systemic insecticides. Pestology 27: 2022. Khan R M and Singh A P (1980) effect of temperature on the seasonal incidence of shoot fly, Atherigona naqvii in wheat at Jaipur, India. Trop Pest Manag 26: 25457. Khurana A D and Verma A N (1983) Some biochemical plant characters in relation to susceptibility of sorghum to stem borer and shoot fly. Indian J Ent 45: 2937. Kishore P (1991) Distinguishing characters of eggs of two major shoot fly species, Atherigona soccata Rondani and Atherigona naqvii Steyskal. J ent Res 15: 14143. Klun J A, Guthrie W D, Hallauer A R and Russell W A (1970) Genetic nature of the concentration of 2,4-dihydroxy-7-methoxy-(2H)-1,4-benzoxazin-3(4H)-one and resistance to the European corn borer in a diallel set of eleven maize inbreds. Crop Sci 10 : 8790. Kumar J and Chander R (1987) Occurrences of shoot fly Atherigona naqvii Steyskal (Diptera: Muscidae) on maize. Himachal J agric Res 10: 64. Kumar P, Sekhar J C and Chaudhary R (2005) Management of insect-pests of maize in tropics. In: Zaidi P H and Singh N N (eds) Stresses on Maize in Tropics. Pp. 298323. Directorate of Maize Research, New Delhi. Kumar S and Singh R (1998) Inheritance of tannin in relation to shoot fly resistance in sorghum. Cereal Res Commun 26: 27173. Kumar V and Kanta U (2011) Management of insect pest complex in maize In: Arora R, Singh B and Dhawan A K (eds) Theory and Practice of Integrated Pest Management. Pp. 338 369. Scientific publishers, India.

140

Kumar V J and Prabhuraj A (2007) Bioeffficacy of new seed treatment chemicals against shoot fly and shoot bug on rabi sorghum in north Karnatka. Indian J Crop Sci 2: 4445. Kumarasinghe N C and Wratten S D (1996) The sugarcane lophopid planthopper, Pyrilla perpusilla (Homoptera: Lophopidae): A review of its biology, pest status and control. Bull Ent Res 86: 48598. Kundu G G and Kishore P (1971) Increasing range of distribution of Atherigona naqvii Steyskal as a pest of wheat. Entomologists Newsl. 1: 38. Kundu G G and Kishore P (1972) New host record of Atherigona naqvii Steyskal (Anthomyiidae: Diptera) from India, together with new record of its three hymenopterous parasites. Indian J Ent 34: 8081. Lawani S M (1982) A review of the effects of various agronomic practices on cereal stem borer populations. Trop Pest Manag 28: 26676. Levin D A (1973) The role of plant trichomes in plant defense. Q Rev Biol 48: 315. Li W , Guo H F, Wang R F, Liu B S, Zhong W F and Fang J C (2009) Comparison of the life tables of Laodelphax striatellus (Homoptera: Delphacidae) on different host plants. Acta Entomol Sin 52: 53136. Mahadeva A and Nagaveni V (2011) Alterations in the biochemical components and photosynthetic pigments of mulberry (Morus Spp.) attacked by leaf roller (Diaphania pulverulentalis) pest. Afr J Biochem Res 5: 36572. Maiti R K and Gibson R W (1983) Trichomes in segregating generations of sorghum matings, II. Associations with shoot fly resistance. Crop Sci 23: 7679. Manuwoto S and Scriber M (1985) Consumption and utilization of experimentally altered corn by Southern armyworm: iron, nitrogen, and cyclic hydroxamates. J Chem Ecol 11: 1469 83. Marwaha K K, Siddiqui K H, Panwar V P S and Sarup P (1984) Peak period of activity of shoot fly species (Atherigona soccata Rondani and A. naqvii Steyskal) in spring sown maize germplasm. J ent Res 8: 199208. Marwaha K K, Siddiqui K H and Sarup P (1985) Evaluation of various insecticides against the shoot fly complex (Atherigona soccata Rondani and A. naqvii Steyskal) in spring sown maize. J ent Res 9: 100103. Marwaha K K, Siddiqui K H and Sarup P (1986) Response of indigenous maize germplasm to heavy natural infestation of shoot fly species, Atherigona soccata Rondani and A. naqvii Steyskal in spring season. J ent Res 10: 10911. Marwaha K K, Siddiqui K H and Sarup P and Singh J P (1987) Minimal application of insecticidal formulation affects the dead-heart formation caused by shoot fly species, Atherigona soccata Rondani and A. naqvii Steyskal in spring sown maize. J ent Res 11: 17577.

141

Mathur L M L (1987) Bibliography of Maize Pests in India. Pp. 4047. All India Coordinated Maize Improvement Project, IARI. New Delhi. Mathur L M L (1988) Symptomatology of pest infestation in Indian maize and its control - Thrips and shoot fly. Pesticide 22: 2730. McMullen M D, Frey M and Degenhardt J (2009) Genetics and Biochemistry of Insect Resistance in Maize. In: Bennetzen J L and Hake S C (eds) The Maize Handbook. Pp. 27189. Springer, New York, USA. Meksongsee B, Chawanapong M, Sangkasuwan U and Poonyathaworn P (1981) The biology and control of the sorghum shoot fly, Atherigona soccata Rondani, in Thailand. Insect Sci Applic 2: 11116. Mote U N, Phadnis B N and Taley Y M (1979) Studies on some physiological factors of shoot fly resistance in sorghum. Sorghum Newsl 22: 6667. Mote U N and Ramshe D G (1987) Effect of fertilizers and plant population on the incidence of shoot fly (Atherigona soccata Rondani) on some sorghum cultivars. Indian J Ent 49: 49095. Mote U N and Shahane A K (1994) Biophysical and biochemical characters of sorghum varieties contributing resistance to delphacid, aphid and leaf sugary exudations. Indian J Ent 56: 11322. Nair S K, Khandge S V, Varshneya M C and Naidu T V R (1995) Effects of weather parameters on the incidence and development of shoot fly. J Maharashtra agric Univ 20: 8384. Naitam N R and Sukhani T R (1989) Studies on effect of soil, weather and other crop factors on insecticidal efficacy against sorghum shoot fly. Indian J Ent 51: 35657. Nelson N (1944) A photometric adoption of somogyi method for determination of glucose. J Biol Chem 15: 37580. Noda H and Saito T (1979) Effects of high temperature on the development of Laodelphax striatellus (Homoptera: Delphacidae) and on its intracellular yeastlike symbiotes. Appl Ent Zool 14: 6475. Nwanze K F (1997) Screening for resistance to sorghum shoot fly. In: Sharma H C, Singh F and Nwanze K F (eds) Plant Resistance to Insects in Sorghum. Pp. 35-37. International Crops Research Institute for the Semi-Arid Tropics, Patancheru, Andhra Pradesh, India. Nwanze K F, Pring R J, Sree P S, Butler D R, Reddy Y V R and Soman P (1992) Resistance in sorghum to the shoot fly, Atherigona soccata: Epicuticular wax and wetness of the central whorl leaf of young seedlings. Ann Appl Biol 120: 37382. Nwanze K F, Reddy Y V R and Soman P (1990) The roles of leaf surface wetness in larval behavior of the sorghum shoot fly, Atherigona soccata. Ent exp appl 56: 18795. Ogwaro K (1978) Ovipositional behavior and host plant preference of the sorghum shoot fly, Atherigona soccata (Diptera: Anthomyiidae). Ent exp appl 23: 18999.

142

Painter R H (1958) Resistance of plants to insects. Annu Rev Entomol 3: 26790. Pandey K C and Sharma V K (1980) Comparative susceptibility of some released and elite maize germplasm against shoot fly, Atherigona spp. in spring sown maize. Indian J Ent 42: 2123. Pant N C and Kalode M B (1964) Pests of wheat, maize and millet. Pp.28184. In: Entomology in India 1938-1963. Indian J Ent, Silver Jubilee No., Entomological Society of India, New Delhi. Panwar V P S (1995) Reaction of promising maize germplasm to shoot fly species ( Atherigona soccata Rondani and A. naqvii Steyskal) in spring season. Indian J Ent 57: 7072. Panwar V P S (1998) Insect pests and their management in maize. Indian Farming 48: 9799. Panwar V P S (2005) Management of shoot fly (Atherigona spp) in springsown maize in tropics, In: Zaidi P H and Singh N N (eds) Stresses on Maize in Tropics. Pp. 37695. Directorate of Maize Research, New Delhi. Panwar V P S, Kumar J and Parmar B S (1999a) Field appraisal of controlled release formulations of phorate against the shoot fly infesting maize. Ann Pl Prot Sci 7: 80 86. Panwar V P S, Mukherjee B K and Ahuja V P (2000) Maize inbreds tolerant to tissue borers, Chilo partellus and Atherigona spp. Indian J Genet 60: 7175. Panwar V P S and Sarup P (1985) Distribution and host-plant of shoot fly species attacking maize in different parts of the world. J ent Res 9: 20717. Panwar V P S and Sarup P (1986) Introduction nursery contributes resistant maize germplasm to shoot fly complex (Atherigona soccata Rondani and A. naqvii Steyskal) in spring sown maize. J ent Res 10: 14446. Panwar V P S and Sarup P (1988) Differential reaction of promising maize germplasm to shoot fly species (Atherigona soccata Rondani and A. naqvii Steyskal). J ent Res 12: 4144. Panwar V P S and Sharma R K (1998) Evaluation of maize germplasm for locating multiple resistance sources to Chilo partellus and Atherigona spp. Indian J Ent 60: 17176. Panwar V P S, Siddiqui K H and Singh N N (1997) Insect pest scenario in maize based cropping system. Indian J Ent 59: 17. Panwar V P S, Singh N N and Sharma R K (1999 b) Impact of previous-season farming practices and fertilization regimes on Chilo partellus (Swinhoe) infestation in two high yielding maize cultivars. Ann agric Res 20: 22026. Patel G M and Sukhani T R (1990) Screening of sorghum genotypes for resistance to shoot fly, Atherigona soccata Rondani. Indian J Ent 52: 18.

143

Patil S S, Narkhede B N and Barbate K K (2006) Effects of biochemical constituents with shoot fly resistance in sorghum. Agric Sci Dig 26: 7982. Pont A C (1972) A review of the oriental species of Atherigona Rondani (Diptera: Muscidae) of economic importance. In: Jotwani M G and Young W R (eds) Control of Sorghum Shoot Fly. Pp. 27104. Oxford & IBH, New Delhi, India. Raga N, Ito K, Matsui M and Okada M (1988) Effects of temperature on adult longevity, fertility and rate of transovarial passage of rice stripe virus in the small brown planthopper, Laodelphax striatellus Falln (Homoptera: Delphacidae). Appl Ent Zool 23: 6775. Raina A K (1981) Movement, feeding behaviour and growth of larvae of the sorghum shoot fly, Atherigona soccata. Insect Sci Applic 2: 7181. Raina A K (1982) Fecundity and oviposition behavior of the sorghum shoot fly, Atherigona soccata. Ent exp appl 31: 38185. Raina A K (1985) Mechanisms of resistance to shoot fly in sorghum: A review.In: Kumble V (ed) Proc. International Sorghum Entomology Workshop, Pp. 131136, Texas A & M University, College Station, Texas, U.S.A. & International Crops Research Institute for the Semi-Arid Tropics, Patancheru 502324, Andhra Pradesh, India. Raina A K and Kibuka J G (1983) Oviposition and survival of the sorghum shoot fly on intercropping maize and sorghum. Ent exp appl 34: 10710. Raina A K, Thindwa H Z, Othieno S M and Corkhill R T (1981) Resistance in sorghum to sorghum shoot fly: Larval development and adult longevity and fecundity on selected cultivars. Insect Sci Applic 2: 99103. Raina A K, Thindwa H Z, Othieno S M and Douglass L W (1984) Resistance in sorghum to shoot fly (Diptera: Muscidae) and oviposition on selected cultivars. J econ Ent 77: 64851. Ramachandra Rao Y (1924) Notes on pests investigated in Madras during the year 1921 22. Rep Proc 5th ent. Meet, Pp. 1922. Pusa Rao C N and Panwar V P S (2001a) Morphological plant factors affecting resistance to Atherigona spp. in maize. Indian J Genet 61: 31417. Rao C N and Panwar V P S (2001b) Biochemical plant factors affecting resistance to Atherigona spp. in maize. Ann Pl Prot Sci 9: 3742. Rao C N and Panwar V P S (2001c) Morphological plant factors affecting resistance to Atherigona spp. in maize. Ann agric Res New Series 22: 27375. Rao C N and Panwar V P S (2002) Biochemical plant factors affecting resistance to Chilo partellus (Swinhoe) in maize. Ann Pl Prot Sci 10: 2830. Rao K R and Panwar V P S (1992) Efficacy of insecticides/plant products against shoot fly species (Atherigona soccata and A. naqvii) in spring sown maize. Pestic Res J 4: 5557.

144

Rao K R and Panwar V P S (1994) Exploring possibility of varied plant population levels of maize for the control of shoot fly species (Atherigona soccata Rondani and A. naqvii Steyskal) in spring season. Indian J Ent 56: 25659. Rao K R and Panwar V P S (1995 a) Effect of sowing dates on the incidence of shoot fly (Atherigona soccata and A. naqvii) in spring sown maize. Ann Pl Prot Sci 3: 3840. Rao K R and Panwar V P S (1995 b) Factors affecting resistance in selected maize varieties to shoot fly species (Atherigona soccata Rondani and A. naqvii Steyskal) in spring season. Shaspa 2: 13540. Rao K R and Panwar V P S (1995 c) Efficacy of seed treatment with carbosulfan against shoot fly species (Atherigona soccata and A. naqvii) in spring sown maize. Pestic Res J 7: 4248. Rao K R and Panwar V P S (1995 d) Efficacy of mixture of carbosulfan treated and untreated maize seeds for the control of shoot fly species (Atherigona soccata and A. naqvii) in spring season. Pestic Res J 7: 14950. Rao K R and Panwar V P S (1995 e) Intercropping of maize with the millets and pulses for the management of shoot fly (Atherigona soccata Rondani and A. naqvii Steyskal) in the spring season. Ann Pl Prot Sci 3: 11517. Rao K R and Panwar V P S (1996a) Location of sources of resistance among maize germplasm against shoot fly species (Atherigona soccata and A. naqvii) during spring. Ann Pl Prot Sci 4: 4749. Rao K R and Panwar V P S (1996 b) Relative efficacy of some insecticides and neem products for the control of shoot flies (Atherigona spp.) in spring sown maize. Pestic Res J 8: 12431. Rathore Y S, Chatterji S M and Asnani V L (1969) A note on the screening of maize varieties for resistance to shoot fly, Atherigona spp. Indian J Ent 31: 27778. Rice E L (1984) Allelopathy.2nd edn. Pp. 422.Academic Press, Orlando, Florida, USA. Sadasivam S and Manickam A (1992) Phenolics. In: Biochemical methods for Agricultural Sciences. Pp. 187-88. Wiley Eastern Ltd., New Delhi, India. Sajjan S S, Sekhon S S and Kanta U (1982) Control of jassid , Zygnidia manaliensis (Singh) and thrips, Anaphothrips sudanensis Trybom infesting maize. J ent Res 6: 1821. Sajjan S S and Sekhon S S (1985) Control of shoot fly, Atherigona naqvii Steyskal (Diptera: Anthomyiidae) on maize through soil and seed treatment. Indian J Ent 47: 37175. Sandhu G S, Dhaliwal G S and Sidhu B S (1986) Resistance to forage sorghum to shoot fly, Atherigona soccata (Rond). Indian J agric Sci 56: 75356. Sandhu G S and Kaushal K K (1972) Incidence of yellow hairy caterpillar, Euproctis virguncula Walker in Punjab. Entomol Newsl 2: 4344.

145

Sandhu G S and Kaushal K K (1976) Occurrence and biology of maize shoot fly in India. Entomol Rec 88: 27. Sandhu G S, Singh B and Kanta U (1974a) Control of cereal thrips, Anaphothrips sudanensis Trybom on maize with systemic insecticides. J Res Punjab agric Univ 11: 40304. Sandhu G S, Singh J, Kaushal K K and Sajjan S S (1974b) The biology of Euproctis virguncula Walker (Lymantridae: Lepidoptera) on cereals in the Punjab. J Res Punjab agric Univ 11: 18790. Santiago F S and Mendoza E M T (1983) Changes in some chemical constituents of maize during infestation by the Asiatic corn borer, Ostrinia furnacalis Guene. Philipp J Crop Sci 8:13339. Sarup P, Marwaha K K, Panwar V P S and Siddiqui K H (1978) Identification of sources of resistance to maize stalk borer, Chilo partellus (Swinhoe) amongst world maize germplasm comprising International Nursery. J ent Res 2 : 15459. Sarup P, Marwaha K K, Siddiqui K H and Panwar V P S (1979) Investigations on major insectspests of maize with special reference to insect-plant relationship: Research Bulletin, New Series, No. 2. Pp. 183 Division of Entomology, IARI, New Delhi. Sarup P and Panwar V P S (1986) Location of sources of resistance to the shoot fly species complex (Atherigona naqvii Steyskal and A. soccata Rondani) in spring sown maize germplasm comprising different maturity groups. J ent Res 10 : 2733. Sarup P and Panwar V P S (1987) Preferential plant age and site for oviposition by Atherigona naqvii Steyskal and Atherigona soccata Rondani in spring sown maize. J ent Res 11: 15054. Sarup P, Siddiqui K H, Marwaha K K and Panwar V P S (1984) Changing pest complex of maize as exemplified by the shoot fly (Atherigona spp.) preference for hosts in spring season. J ent Res 8: 11519. Sarup P, Siddiqui K H and Marwaha K K (1986) Possible limitations in exploring the trap of preferred host sorghum for the control of shoot fly complex (Atherigona soccata Rondani and A. naqvii Steyskal) in spring sown maize. J ent Res 10: 11924. Sathya P K, Sreedhar S, Singhvi N R, Kodandaramaiah J and Sen A K (2002) Post thrips infestation biochemical changes in leaves of mulberry (Morus spp.). Plant Arch 2: 85 88. Schoonhoven L M (1968) Chemosensory basis of host plant selection. Annu Rev Entomol 13: 11536. Sekhon S S, Dhillon B S and Kanta U (1993) Host plant resistance in maize to insect pests. In: Dhaliwal G S and Dilawari V K (eds) Advances in Host Plant Resistance to Insects. Pp. 79-109. Kalyani Publishers, Ludhiana, India.

146

Sekhon S S and Kanta U (1994) Mechanisms and bases of resistance in maize to spotted stem borer. Proc Insect resistant maize: recent advances and utilization. Pp. 10611. International Maize and Wheat Improvement Center, Mexico. Sekhon S S, Sajjan S S and Bhinder S S (1986) Influence of cultural practices on the incidence of shoot fly, Atherigona naqvii Steyskal on the winter maize in the Punjab. Indian J Ecol 13: 10508. Seshu Reddy K V and Davies J C (1977) Species of Atherigona in Andhra Pradesh. PANS 23: 37983. Shahzad M A, Shaheen M S, Khan M T H and Iqbal B (2006) Field screening of promising cultivars of maize against shoot fly (Atherigona soccata Rond.) and maize borer, Chilo partellus (Swin.) during spring season. Pak Entomol 28: 1517. Sharma G C, Jotwani M G, Rana B S and Rao N G (1977) Resistance to the sorghum shoot fly, Atherigona soccata (Rond.) and its genetic analysis. J ent Res 1: 112. Sharma H C (1985) Strategies for pest control in sorghum in India. Trop Pest Manag 31: 16785. Sharma H C, Nwanze K F and Subramanian V (1997) Mechanisms of resistance to insects and their usefulness in sorghum improvement. In: Sharma H C, Singh F and Nwanze K F (eds.) Plant Resistance to Insects in Sorghum. Pp. 81100. International Crops Research Institute for the Semi-Arid Tropics, Patancheru, Andhra Pradesh, India. Sharma H C, Taneja S L, Leuschner K and Nwanze K F (1992) Techniques to screen sorghum for resistance to insect pests: Information Bull No. 32. Pp. 48. International Crops Research Institute for the Semi-Arid Tropics, Patancheru , Andhra Pradesh, India. Sharma R C, Kumar P and Sharma R K (2006) Status and Prospects of Integrated Pest Management Strategies in Selected Crops: Maize. In: Singh Amerika, Sharma O P and Garg D K (eds) Integrated Pest Management-Principles and Applications, Vol.2: Applications, Pp.10226. CBS Publishers, New Delhi, India. Sharma R K and Panwar V P S (1998) Search for source for resistance amongst maize germplasm against Atherigona soccata and A. naqvii during spring season. Ann agric Res 19: 359 60. Sharma V K and Singh J M (1975) Screening for resistance to Atherigona spp. in spring sown maize. Indian J Ent 37: 3943. Sharma V K, Kanaujia K R and Choudhary R N (1979) Relative efficacy of foliar spray of insecticides for the control of shoot fly, Atherigona spp. in spring maize. Pestology 3: 2022. Shi ZhongLiang, Qiu SongYing , Ma AiPing, Xu GangYuan, Wu JiPing and Lu LaHu (2002) Studies on mechanisms of biochemical resistance to blossom midge ( Sitodiplosis mosellana Gelin) in wheat. J Triticeae Crops 22: 6365. Shukla K K (1979) Occurrence of a new insect, small brown plant hopper Laodelphax striatellus (Falln), in India. Curr Sci 48: 548.

147

Siddiqui K H, Marwaha K K and Sarup P (1988) Differential response of maize germplasm to shoot fly species, Atherigona soccata Rondani and A. naqvii Steyskal) under natural conditions. J ent Res 12: 16065. Siddiqui K H and Marwaha K K (1994) The Vistas of Maize Entomology in India. Pp.136. Kalyani Publishers, Ludhiana, India. Siddiqui K H, Panwar V P S, Marwaha K K and Singh N N (1995) Reaction of parental inbreds, single crosses involved in experimental maize hybrids for resistance to stalk borer, Chilo partellus (Swinhoe) under artificial infestation. Shaspa 2: 14143. Singh A P and Khan R M (1980) A new pest of wheat and a new host of wheat pests in India. Entomologists mon Mag 115: 20708. Singh B U, Padmaja P G and Seetharama N (2004) Stability of biochemical constituents and their relationships with resistance to shoot fly, Atherigona soccata (Rondani) in seedling sorghum. Euphytica 136: 27989. Singh H and Kanta U (2004) Influence of methods and dates of sowing of summer maize on damage caused by shoot fly, Atherigona naqvii Steyskal. J Insect Sci 17: 3840. Singh H and Kanta U (2006) Evaluation of some granular insecticides as spot treatment for the management of shoot fly, Atherigona naqvii Steyskal on summer maize. J Insect Sci 19: 11011. Singh R (1991) Studies on the damage and losses caused by jassid, Zygnidia manaliensis (Singh) to winter and spring maize and its control. M.Sc. Thesis, Punjab Agricultural University, Ludhiana, India. Singh R and Narayana K L (1978) Influence of different varieties of sorghum on the biology of sorghum shoot fly. Indian J agric Sci 48: 812. Singh S P (1998) Field efficacy of some bio-pesticides against shoot fly and stem borer in forage sorghum. Forage Res 24: 17778. Singh S P and Jotwani M G (1980a) Mechanisms of resistance in sorghum to shoot fly I. Ovipositional non-preference. Indian J Ent 42: 35360. Singh S P and Jotwani M G (1980b) Mechanisms of resistance in sorghum to shoot fly IV. Role of morphological characters of seedlings. Indian J Ent 42: 80608. Singh S P and Jotwani M G (1980c)Mechanism of resistance in sorghum shoot fly III. Biochemical basis of resistance. Indian J Ent 42: 55166. Singh S P and Jotwani M G (1980d) Mechanisms of resistance in sorghum to shoot fly II. Antibiosis. Indian J Ent 42: 24047. Singh S P, Lodhi G P and Naresh J S (1995) Ovipositional preference of shoot fly, Atherigona soccata (Rondani) in forage sorghum. Crop Res 10:22325.

148

Singh S P and Verma A N (1988) Monitoring of shoot fly, Atherigona soccata (Rondani) in traps and their periodic incidence in sorghum. Crop Res 1: 7683. Singh U C, Misra U S, Dhamdhere S V and Dwivedi U S (1985) Carry-over of the sorghum stalk borer, Chilo partellus (Swinhoe) in off-season in different crops. J ent Res 9: 17073. Sinha R B P (1989) Studies on residues and residual toxicity of some promising systemic insecticides against some maize pests. Ph.D. dissertation. Indian Agricultural Research Institute, New Delhi, India. Smith C M (2004) Plant Resistance Against Pests: Issues and Strategies. In: Koul O, Dhaliwal G S and Cuperus G (eds) Integrated Pest Management: Potential, Constraints and Challenges. Pp. 14767. CABI Publication, Oxon, UK. Smith R F (1970) Pesticides: their use and limitations in pest management. In : Rabb R L and Guthrie F E (eds) Concepts of pest management. Pp. 629. North Carolina Sate University, Raleigh, USA. Sodhi and Sekhon (1997) Incidence of maize jassid, Zygnidia manaliensis (Singh) on maize in Ludhiana, Punjab. Pest mgmt eco Zoo 5: 9194. Solomon M E (1973) Ecology in relation to the management of insects. In: Geier P W, Clark L R, Anderson D J and Nix H A (eds) Insects: studies in population management. Pp. 153 67. Ecological society of Australia, Canberra. Soto P E (1972) Mass rearing of sorghum shoot fly and screening for host plant resistance under greenhouse conditions. In: Jotwani M G and Young W R (eds) Control of sorghum shoot fly. Pp. 137-38. Oxford & IBH, New Delhi, India. Soto P E (1974) Ovipositional preference and antibiosis in relation to resistance to sorghum shoot fly. J econ Ent 67: 26567. Srinivas P S and Panwar V P S (2003) Combined effects of intercropping maize with pulses and potash fertilizer on stem borer, Chilo partellus (Swinhoe) infestation in maize. Ann agric Res 24: 46165. Srivastava B K and Khan R M (1961) Field tests with newer insecticides for control of the maize armyworms. Indian agric 5: 11314. Srivastava K P, Young W R and Bhatia S K (1969) A new record of stem fly, Atherigona naqvii Steyskal, causing damage to barley. Indian J Ent 31: 29091. Srivastava R P and Pandey Y D (1968) Biology of Atherigona oryzae Malloch (Diptera: Anthomyiidae) a new pest of wheat in Rajasthan. Ann Arid Zone 7: 13238. Swain T and Hills W E (1959) The phenolic constituents of Prunus domestica: The quantitative analysis of phenolic constituents. J Sci Food Agric 10: 6368.

149

Taneja S L and Leuschner K (1985) Resistance screening and mechanisms of resistance in sorghum to shoot fly. In: Kumble V (ed) : Proc. International Sorghum Entomology Workshop. Pp.115129, Texas A&M University, College Station, Texas, USA & International Crops Research Institute for the Semi- Arid Tropics, Patancheru 502324, Andhra Pradesh, India. Taneja S L, Seshu Reddy K and Leuschner K (1986) Monitoring of shoot fly population in sorghum. Indian J Plant Prot 14: 2936. Verma G D, Singh J and Singh I P (1979) Field evaluation of some insecticidal treatments for control of Rhyacia herculea (Corti and Draudt) on rabi maize in Bihar. Entomon 4:129 31. Verma T and Singh S P (2000) Morpho-physio plant characters associated with shoot fly resistance in sorghum. Haryana agric Uni J Res 30: 4143. Vidhyasekaran P (1988) Physiology of disease resistance in plants. CRC press, Boca raton, Florida, USA. Wang Y, Cai Q N, Zhang Q W and Han Y (2006) Effect of the secondary substances from wheat on the growth and digestive physiology of cotton bollworm. Eur J Ent 103: 25558. Zhang A M, Liu X D, Zhai B P and Gu X Y (2008) Influences of temperature on biological characteristics of the small brown planthopper, Laodelphax striatellus (Falln) (Hemiptera: Delphacidae). Acta Entomol Sin 51: 64045.

150