Tuesday 20 October 2015

Achievements in Pheromones in India and its future scope in IPM:Pheromone Resourrces



Achievements in Pheromones in India and its  future scope in IPM

V.Nandagopal, Anand Prakash and J.Rao
Central Rice Research Institute (ICAR), Cuttack-753 006, Orissa

Introduction

Chemicals used for communication between the organisms are known as semio-chemicals or info-chemicals. These chemicals are also called “ assembling scents” by  Kettlewell (1942,1943,1946,1955). In Greek Pherein means  to carry  and hormone (to excite, stimulate) as described by Karlson and Butenandt (1959). Further Kirschenblatt (1957)  proposed the term “telegones”. Micklem (1959) termed it as “pherormone” instead of pheromone. Karlson and Luscher (1959) have given a more scientific clarification for the use of the term pheromone.

Pheromone has been in use in India over more than 30  years as a monitoring tool under All India Coordinating programme for some of the crops likes cotton, groundnut and pigeon pea.  The monitoring technology is well knitted in different centers in these crops.  The monitoring helps in developing forewarning systems and plan management strategies.  Unfortunately, as such efforts have not been made even at research station; the farmers are not convinced about practical utility of pheromone technology.  Some farmers use pheromone traps especially in cotton and pigeon pea eco system, as a mass-trapping tool. Surprisingly, in a recent study at the NRCG; a maximum of 1470 males/day/trap in case of Spodoptera litura was recorded showing its potentiality for mass trapping.

Even after 30 years of introduction of pheromone technology in Indian Agriculture, it remains still in its infancy. This may be attributed to, high cost of pheromone  chemicals, defects in manufacture of the lures and traps, insect related issues, difficulties in installation of traps at farmers level etc.  An analysis of these issues has been made and suggestions are given to overcome the same. Then only there be actual fillip to the novel, eco-friendly pheromone technology in Indian agriculture (Nandagopal, 2004).

In India, monitoring of the key insect pests such as H. armigera,  S. litura, P. gossypiella, were undertaken as early as 1985. Intensive work has been done on these insects spreading over Karnataka, Haryana, M.P, T. Nadu, A.P., Maharashtra, Himachal Pradesh, Punjab and Gujarat. Pheromone as a mass trapping tool have also been utilized in a few insect such as Pthorimoaea opercullela, S. litura, Chilo sacchariphagus indicus, H. armigera, A. modicella, Lymantria obfuscata Wlk (Porthetria dispar), Tortricid Cydia pomonella (L.), Codling moth, Cydia pomonella, and Pectinophora gossypiella (Saund.). The communication disruption as a  tool has been tried in S. litura, A. modicella, Peripleneta americana and Chilo auricilius in India with limited success. There are attempt made to design the trap for efficient trapping of the target insect, some of them were successful in the case of cotton, sugarcane and groundnut ecosystem. Interestingly, lot of work has been done on the basic aspects of pheromone since 1975.    


Basic studies
Intensive work has been done on the basic aspects of the sex pheromones in various insects and mammals in India since 1980.  Majority of the information are on the location of sex pheromone glands, occurrence of sex pheromones etc. These basic studies have given a fillip to the further studies on the pheromone work.  

Synthesis of Pheromone

The chemistry part of the pheromones is in its developing stage in India as sophisticated instruments are required for taking up of Isolation, Identification and synthesis of pheromones. Very few labs such as Pheromone groups of IICT, Hyderabad, Organic chemistry group of SPIC, Organic chemistry group of BARC have such facilities in India. Some  works  on the sex pheromone of the crop pests in  India has been taken up in  collaboration with foreign scientific organizations such as NRI, London;  TNO, Delft, Netherlands, NL and  CID, CSIC, Barcelona, Spain etc.

Monitoring:
Where monitoring can be applied ?
Yadav et al. (2004)  suggested that timely and efficient monitoring should be the foundation of sound IPM programs, without which no proper pest management decision can be made. A few of the examples of application of pheromones employed for monitoring purposes are as follows:
·         Early detection of pest
·         Mapping pest distribution
·         Quarantine inspection
·         Pest monitoring for action thresholds / decision support
·         Mapping insecticide resistance frequency
·         Monitoring parasites / predators
·         Estimation of population dynamics
·         Estimation of population densities
·         Timing of management procedures based on the threshold
·         Efficacy of  management procedures based on the threshold

The occurrence of the major insect pests is difficult to forecast unless it is noticed in a damaging proportion. Hence monitoring of the insect pests using various gadgets becomes imperative. There are various traps used in various situations and based on the behaviour of the target insects.  Light trap for adults of Lepidoptera, Coleoptera; sticky trap for  aphids, in some of the cases laser light traps are also used such as house fly;  micro, Lepidoptera, thrips etc. Now the emphasis is on the use of pheromones.  Pheromone traps are being used for all the major Lepidoptera insects, Coleoptera, Diptera etc. In the monitoring programme it is very important to recommend the required number of the traps to be used per unit area. Based on the trapping efficiency of the trap, the number of insects trapped, the management options have to be decided.  Careful consideration is to be given to the availability of the host plant, the mode of pupation, occurrence of natural enemies of the target insects etc.  Lot of research has to be done before recommending the number of traps to be used/unit area.

Pest monitoring using sex pheromones facilitates early detection of the pest occurrence, through moth population dynamics during crop season and this information helps in making decisions on the choice of pressing control tactics in other words precise timing of insecticide or bio-agents applications.  The table 1 gives the details of the pheromones used in Indian agriculture (Natarajan,2004).

Table 1. Pheromones used for monitoring

No.
Pest species
Pheromone components

CEREALS


Yellow stem borer, Scirphophaga incertulas
Hexadecenal
(Z)-9- Hexadecenal
(Z)-11- Hexadecenal
(Z)-11- Hexadecen-1-ol
(Z)-9- Octadecenal

 

Leaf folder,
Cnaphalocrocis medinalis
Hexadecyl acetate
(Z)-11-Hexadecenyl acetate
Octadecyl acetate
(Z)-13-Octadecenyl acetate

 

Leaf folder,
 Marasmia patnalis
(Z)-13-Octadecenyl acetate
(Z)-11-Hexadecenyl acetate

 

Maize stalk borer,
Chilo partellus
(Z)-11- Hexadecenal
(Z)-11- Hexadecen-1-ol

 

Striped stem borer,
Chilo suppressalis
Hexadecenal
(Z)-9- Hexadecenal
(Z)-11- Hexadecenal
(Z)-11- Hexadecen-1-ol
Octadecan-1-ol
(Z)-11- Octadecenal

 

Climbing cutworm, Mythimna separata
(Z)-11-Hexadecenal
(Z)-11- Hexadecen-1-ol
(Z)-11-Hexadecenyl acetate

 

Purple stem borer,
Sesamia inferens
(Z)-11-Hexadecenyl acetate
(Z)-11- Hexadecen-1-ol
(Z)-11-Hexadecenal

 

Rice green caterpillar, Naranga aenescens
(Z)-9-Tetradecenyl acetate
(Z)-9-Hexadecenyl acetate
(Z)-11-Hexadecenyl acetate

 

COTTON

 

Pink bollworm,
Pectinophora gossypiella
(Z,Z)-7,11-Hexadecadienyl acetate
(Z,E)-7,11-Hexadecadienyl acetate

 

Spotted bollworm,
Earias vittella
(E,E)-10,12-Hexadecadienal
(E,E)-10,12-Hexadecen-1-ol
Z11-Hexadecenal
Z11-Octadecenal

 

Spiny bollworm,
Earias insulana
(E,E)-10,12- Hexadecadienal

 

Gram pod borer,
Helicoverpa armigera
(Z)-9- Hexadecenal
(Z)-11- Hexadecenal
Hexadecenal
(Z)-11-Hexadecen-1-ol
Hexadecan-1-ol

 

Oriental tobacco budworm, Helicoverpa assulta
Hexadecenal
(Z)-9- Hexadecenal
(Z)-11- Hexadecenal
Hexadecencyl acetate
(Z)-9- Hexadecencyl acetate
(Z)-11- Hexadecencyl acetate
Hexadecen-1-ol
(Z)-9-Hexadecen-1-ol
(Z)-11-Hexadecen-1-ol

 

Tobacco caterpillar, Spodoptera litura
(Z)-9(E)-11-Tetradecadienyl acetate
(Z)-9(E)-12-Tetradecadienyl acetate

 

OILSEEDS

 

Castor semi looper,
Achaea janata
(Z,Z)-6,9-Heneicosadiene
(Z,Z,Z)-3,6,9-Heneicosatriene
(Z,Z)-9,12-Octadecadienal

 

Groundnut leaf miner, Aproaerema modicella
(Z)-7-Decenyl acetate
(E)-7-Decenyl acetate
(Z)-7,9-Decadienyl acetate

 

Tobacco caterpillar, Spodoptera litura
See under item 14

 

Red palm weevil, Rhynchophorus ferrugineus
(4S,5S)-4-methyl-5-nonanol
(4S)-4-methyl-5-nonanone

 

Rhinocerous beetle,
Orcytes rhinoceros
Ethyl-4-methyloctanoate
Ethyl-4-methylheptanoate
4-Methyloctanoic acid

 

VEGETABLES

 

Diamond-back moth,
Plutella xylostella
(Z)-11-Hexadecenal
(Z)- 11 -Hexadecenyl acetate
(Z)-11-Hexadecen-1-ol

 

Cabbage stem borer,
Hellula undalis
(E, E)-11, 13 - Hexadecadienal

 

Brinjal borer,
Leucinodes orbonalis
(E)- 11-Hexadecenyl acetate
(E)- 11-Hexadecen-1-ol

 

Cutworm,
Spodoptera exigua
Tetradecyl acetate
(Z)-9-Tetradecenyl acetate
(Z)-11-Tetradecenyl acetate
(Z,E)-9,12-Tetradecadienyl acetate
(Z,Z)-9,12-Tetradecadienyl acetate
(Z)-9-Tetradecen-1-ol

 

Tobacco caterpillar, Spodoptera litura
See under item 14

 

Gram pod borer,
Helicoverpa armigera
See under item 12

 

Earias sp.
See under item 10 and 11

 

Potato tuber moth, Phthorimaea operculella
(E, Z)-4,7-Tridecadienyl acetate
(E, Z, Z)-4,7,10-Tridecatrienyl acetate

 

Sweet potato weevil,
Cylas formicarius
(Z)-3-Dodecenol-(E)-2-butenoate

 

SUGARCANE

 

Sugarcane stalk borer,
Chilo auricilius
(Z)-7-Dodecenyl acetate
(Z)-8-Tridecenyl acetate
(Z)-9-Tetradecenyl acetate
(Z)-10-Pentadecenyl acetate

 

Early shoot borer,
Chilo infuscatellus
(Z)-11-Octadecen-1-ol

 

Internode borer,
Chilo sacchariphagus indicus
(Z)-13-Octadecenyl acetate
(Z)-13-Octadecen-1-ol

 

Armyworm,
Mythimna separata
See under item 6

 

Purple stem borer,
Sesamia inferens
See under item 7

 

TREE CROPS

 

Shoot and panicle borer of cardamom, turmeric and ginger,
Dichocrocis punctiferalis
(E)-10-Hexadecenal
(Z)-10-Hexadecenal


Krishnaiah (1995) reported several parameters that would enable efficient pheromone based monitoring for insect pests of rice  crop.  Krishnaiah et al, (1998) optimized three traps per acre as trap density for monitoring of rice yellow stem borer. Various parameters of an efficient pest monitoring system for rice pests are presented in Table 2.

Table 2: Pheromone based monitoring systems of rice insect pests
Insect
S. incertulas
C. suppressalis
S. inferens
C. medinalis
Components
(Z) 11-16: Al          (Z) 9-16: Al
( Z) 11-16: Al
(Z) 9-16: Al
(Z) 13-18: Al
(Z) 11-16: Ac
(Z) 11-16: OH
(Z) 11-16: Al
(Z) 13-18: Ac
(Z) 11-16: Ac
Best blend
3:1
10:1:1
40:10:1
10:1
Dose
2 to 5 mg
0.3 to 0.6 mg
2mg
1mg
Dispenser
rubber
rubber
rubber
polythene vial
Replacement period
3-4 weeks
4 weeks
3-4 weeks
4 weeks
Trap design
sleeve
plastic funnel
sleeve
delta sticky
Sleeve colour
white
yellow
-----
-----
Trap elevation
0.5-1 m
0.5 m
1.0m
canopy level

Monitoring system in cotton:

Several studies have been conducted on cotton pests to establish a quantitative relationship between pheromone trap catches of particular pest and the damage caused with an aim to develop relationship would help to forewarn. Utilization of pheromones of various cotton pests  as follows (Table 3).


Table 3. Utilization of pheromones of various cotton pests  as follows
Bollworm      trap ht              Predictability of larval damage       catch/night/trap                                                            
                                                                                     Expect egg laying in  the
Bollworm    (Plate plate 6 & 7)    for 3 nights               next 5-8  days
Pink boll                0.03m             8-10males/trap         Expect egg laying in  in the
worm                                            /night for 4 nights    next 10-15 days
Spotted                 0.03m              1 males/night           Expect egg laying in  in the
bollworm                                       for 3 nights              next 3-5  days
Spiny                    0.03m              1 males/night           Expect egg laying in  in the
bollworm                                      for 3 nights              next 3-5  days



Monitoring system in Groundnut:

The presence of sex pheromones of leaf miner has been identified for the first time (Nandagopal and Reddy, 1991) by the National Research Centre for Groundnut (NRCG) in collaboration with NRI, U.K. and field evaluation of the components of the sex pheromones have been also done by the NRCG. Experiments were also conducted by NRCG for standardizing the number of traps per unit area and use of the sex pheromone as one of the components of Integrated Pest Management (IPM) in groundnut where leaf miner damage is extensive (Plate 3) under Saurashtra conditions. Nandagopal and Soni (1993) have evaluated a number of traps   for      the    groundnut leaf   miner  (Aproaerema modiceela Deventer. Hall et al. (1993) have identified the total components of the pheromone compounds of leaf miner.

At Junagadh, a square glue trap developed at NRCG  (Nandagopal and Soni, 1993) was used, while in other centres water trap designed by ICRISAT was used. Different types of traps were designed and tried in the trapping of males of A. modicella (Plate 2a). The results indicated that a square glue trap designed with 22 cm side and with 4 entry holes (Nandagopal and Soni, 1993) was the most efficient in trapping the males.  The pheromone was also used for monitoring the activity of the moths in ground nut (Yadav et al., 2004).  A trap efficient to a tune of 60 to 90% has been developed for Spodoptera litura in groundnut.  Nandagopal et al. (1995) have reported on the effects  of certain  components  of  IPM  on  the  damage  and  yields  in  groundnut. Nandagopal (1998) explained about hove the eco-friendly management of Groundnut Leafminer in India can be achieved. Nandagopal et al. ( 1998) reported on the development of an efficient pheromone trap for Aproaerema modicella Dev. (Gelechiidae: Lepidoptera) population in groundnut crops. Further, Nandagopal  et al. ( 2006) have  developed a  viable efficient sex pheromone trap for Aproaerema modicell.  In  Spodoptera, Nandagopal et al.( 2003) reported on the ppulation dynamics of Spodoptera litura (F.) in relation to weather parameters in groundnut in India using pheromone trap catch. Nandagopal  et al.( 2004) have developed an efficient pheromone trap for field catch of Spodoptera litura Fabricius in groundnut and castor ecosystem.  Based on the pheromone trap catches the prediction of Spodoptera litura (F.) males in relation to key weather factors in groundnut have been reported. Holistic IPM was developed wherein the use of pheromone has been explained (Ghewande et al.,2002).

Monitoring system in red palm weevil Rhynchophorus ferrugineus Oliv.

Attraction of red palm weevil Rhynchophorus ferrugineus to ferrugineol based pheromone lures in coconut (Fig 4). The trapped weevils with different lures are given in table 4 (Faleiro, 2004)
.
Table 4. Weevils trapped with different lures
Sr. No.
Treatments
Strength of lure
Mean weevil catch per trap
Trial I
(23/12/02 to 24/1/03)
Trial II
(25/1/03 to 26/2/03)
Cumulative
Sex ratio
1
Pherobank RPW lure
400mg
3.52 (12.0)
2.90 (8.3)
3.26 (10.1)
1: 2.05
2
Pherobank RPW lure
700 mg
2.70 (7.0)
2.41 (5.6)
2.57 (6.3)
1: 2.8
3
Pherobank RPW lure
1000 mg
3.02 (8.7)
2.41 (5.6)
2.76 (7.1)
1: 1.39
4
Ferrolure+
800 mg
2.88 (8.0)
2.66 (7.6)
2.79 (7.8)
1: 1.35
5
ISCA Technology
900 mg
2.76 (7.3)
2.22 (4.6)
2.55 (6.0)
1: 1.25
6
CPCRI lure
0.157 g
1.56 (2.0)
1.26 (1.3)
1.47 (1.6)
1: 2.00
7
Ferrolure+ only
(no food)
800 mg
2.18 (4.3)
1.74 (2.6)
2.00 (3.5)
1: 1.63
8
Food only
(coconut petiole)
----
0.88 (0.3)
0.71 (0.0)
0.81(0.1)
1: 0.00
CD (P = 0.05)
0.49
NS
0.46
---


Monitoring of coffee white stem borer: Xylotrechus quadripes (Coleoptera: Cerambycidae):

Coffee  is one of the major plantation crops grown in India and occupies a place of pride among the plantation crops. Arabica and Robusta are the two types of coffee cultivated on commercial scale. Arabica is known for its quality and has high international demand. Coffee Arabica in India, China, Srilanka, Vietnam and East African Countries however, has been threatened by the presence of a dreaded pest called Coffee White Stem Borer, Xylotrechus quadripes, Chevrolat  (Coleoptera: cerambicidae) Fig 6)that has the potential to devastate the entire plantation if uncontrolled. Chemical methods developed to control the pest have their own disadvantages, as they are harmful to the environment. Biological methods of control proved to be a failure so far. Therefore, an alternate eco-friendly technique like use of sex pheromone has been developed after taking a clue that these beetles communicate each other through chemical medium. Studies showed that male borers attract the females by secreting sex pheromone. The sex pheromone components from the males have been successfully isolated, chemically characterized and synthesized in the laboratory. The components were identified to be (S)-2- hydroxy-3-decanone,             (R)-3-hydroxy-2-decanone and 2,3-dihydroxy octane. Laboratory bioassay conducted using the live beetles confirmed that (S)-2- hydroxy-3-decanone component is very active followed by (R)-3-hydroxy-2-decanone and the third component did not elicit any response from the females. Large-scale field trials were conducted for over 8 seasons (4 years) using the synthetic pheromone in sticky cross vane traps. Field trials showed that the male pheromone of coffee WSB could be effectively used to trap the female beetles in the field.

Mass Trapping

This is   a direct method of managing the insects.  Based on the number of insects trapped, a decision can be taken to continue the recommended number of traps/unit area for mass trapping or remove all the traps and use only for monitoring. Because, if the anticipated insects are going to occur only below ET levels, there is no need for continuing the traps meant for mass trapping which will lead to unnecessary waste of resources. If the population is of moderate level, mass trapping followed by application of bio-pesticides would be sufficient to contain the insect before reaching economically important level. If the insect is occurring in a endemic way, necessarily, we have to think of going for too many methods of available tactics to manage the insects.   The following are the insects  for which mass trapping was tried ( Table 5) (Nandagopal and Prasad, 2004).
Table 5. Various insects against which mass trapping was tried
Species
Location
Reference
Pthorimoaea opercullela
Karnataka and Tamil Nadu
Trivedi et al., 1994

Karnataka
Nandihalli et al., 1993

New Delhi
Gupta et al., 1990a,b

Uttar Pradesh
Siddiqi, 1989

Punjab
Anon,1981
Chilo auricilius
Uttar Pradesh,  Hariyana
David et al., 1986
S. litura  
Uttar Pradesh
Singh and Sachan,1991

Andra Pradesh
Krishnaiah,K.1986; Lalith kumari et al., 1992

Tamil Nadu
Dhandapani,1985

Gujarat
Nandagopal et al., 2004b
H. armigera
Andra Pradesh
Lalith kumari et al., 1992

Haryana
Pawar et al., 1984.

Gujarat
Patel et al 1985

Tamil Nadu
Balakrishnan, 2004a

Tamil Nadu
Balakrishnann et al., 2004b

Gujarat
Nandagopal et al., 2004b   

Gujarat
Patel et al., 2004f
Aproaerema modicella 
Gujarat
Nandagopal,1992;
Nandagopal et al., 2004a;
Nandagopal et al., 2004b

Andra Pradesh
Yadav et al., 2004
Lymantria americana (=Porthetria dispar)
Jammu Kashmir
Masoodi et al., 1990
Cydia pomonella
Jammu Kashmir
Pawar and Tuhan, 1985 

Ladakh
Pawar et al., 1982; Pawar et al., 1984
Pectinophora gossypiella
Tamil Nadu
Balasubramanian et al., 1979;
Satpute et al., 1985;
Balakrishnann et al., 2004a

Gujarat
Patel et al., 2004e
Periplaneta mericana
Karnataka
Urs et al., 1989
Orcytes rhinoceros
Andra Pradesh
Kalidas et al.,2004
Rhynchophorus ferrugineus
Goa
Faleiro et al., 2004

Karnataka
Kalleshwara swamy et al., 2004b
Scirpophaga incertulas
Andra Pradesh
Yadav et al., 2004 ;
Pasalu et al., 2004
 Xylotrechus quadripes
Karnataka
Jayarama et al., 2004
Chilo sacchariphagus indicus
Karnataka
Jayanth and Bhanu, 2004
Scirpophaga excerptalis
Karnataka
Jayanth and Bhanu, 2004
Chilo infuscatellus
Karnataka
Jayanth and Bhanu, 2004
Batocera dorsalis
Gujarat
Patel, 2004b

Gujarat
Patel, 2004c
Odoiporus longicollis
Tamil Nadu
Padmanaban et al., 2004b ;
Padmanaban et al., 2004c
Earias spp.
Tamil Nadu
Balakrishnan et al., 2004a,2004b

Gujarat
Patel et al., 2004f
Leucinodes orbonalis
Gujarat
Jhala, 2004
Cylas formicarius
Kerala
Pillai et al., 1996
Plutella xylostella
Karnataka
Reddy  and  Urs,1997

Mating Disruption

Male confusion is thought to be the result of ambient pheromone concentrations sufficient to hide the trails of the calling female to the large doses from diffuse sources such as microcapsules or larger doses of pheromone at the  point source dispensers such as tie-on polyethylene rope.  Added to the effect, is the adaptation of antennal receptor sites and for habituation of the insect’s central nervous system.  Specific receptor sites on the antennae respond to only the pheromone molecules.  When a receptor site is continually activated by high ambient concentrations of the pheromone, the resulting electrical signal diminishes (measured by an electroantennogram).  The receptor site becomes unresponsive and the insect becomes navigationally blink.  When the insect’s central nervous system is inundated with signals from the receptor sites, it becomes habituated and no longer remains cabable to provide the directed behaviour. The net result of confusion is that the male is unable to orient to any pheromone source and follow the upwind trail to a mate (Flint  and Doane, 1996).

Some possibilities of the mechanism of the confusion approach as put forth by Pedigo (2002) include:

1. Camouflage or covering up the natural pheromone scent of
    females
            2. Misdirection of males to scents from multiple point sources
                of synthetic pheromone, and
            3.Adaptation/habituation by desensitizing male antennal
               receptors through constant pheromone-like exposure

Bartell (1982) delineated five mechanisms by which mating disruption  could be successful as below:

1. Sensory adaptation of the pheromone receptors and central
    nerve  habitauation,
2. False trial following caused by a multiplicity of the trials
    produced by    synthetic odour source,
3. Inability of insects to discriminate odour trials from
    background,
4. Imbalance of sensory input caused by the release of an
    unnatural  ratio of pheromone components,
5. Peripheral sensilla blocked by pheromone analogues
           
The first preliminary field test demonstrating the potential of this approach was conducted in 1967 with the cabbage looper, Trichoplusia ni.  In this test, pheromone concentrations were shown to thwart males from being lured to female moths. 

Following this success, many studies were conducted to apply the approach but with little success.  These included work with insect pests of fruit crops, vegetables, field crops, forest and stored products.  In these studies, pheromone dispensing seemed the greatest obstacle to success. Subsequently, controlled-release dispensers were developed, and these have paved the way for successes in pest suppression.  One such type of dispenser is the Hercon® flake, produced by Hercon Environmental Company, Emigsville, Pan America.  Hercon flakes are multilayered plastic laminates, about 3 mm2 (1/8 in.2), that contain pheromone in the inner layer, or reservoir.  The outer layers of the flake serve as a protective barrier but allow the mixture to diffuse into the air.  As the material is dissipated into the air, a replacement quantity automatically moves outward from the reservoir to maintain the desired surface concentration and to give long-lasting effectiveness.  The flakes are applied somewhat like a conventional insecticide with the use of a special device, the Hercon dispenser pod, attached to an airplane. 

This unit automatically combines the flakes with a special sticker that causes them to adhere to foliage as they land.

 

ii) Mating disruption  technique in the key insect pests in India


In a very few cases, mating disruption technique was successful in India.  Based on the information available, communication disruption may be successful in crops where dense foliage results in reduced wind velocity and thus pheromone persists for a few hours. Some examples are Chilo auricilius in sugarcane in Uttar Pradesh (Nesbitt et al., 1980),  Spodoptera litura  in groundnut in Andhra Pradesh  (Rao et al., 1989), Hall et al. (1994) used the pheromones for mating disruption of cotton bollworms and rice stemborer in developing countries. Slow-release pheromone formulations have been used for control by mating disruption of cotton bollworms (Pectinophora gossypiella, Earias vittella and E. insulana) in Pakistan and yellow rice stem borer (Scirpophaga incertulas) in India. Single applications of pheromone formulations gave season-long control and final yields at least as good as those achieved with conventional insecticides, and savings of up to five applications of insecticides against cotton bollworms and two against rice stem borer.

Mating disruptions in  Aproaerema modicella  in pigeon pea intercropped with groundnut in Tamil Nadu ( our unpublished work) reported to be successful.  Similarly, Pasalu et al. (2004) and  Cork et al. (1996) have used unnatural blend of pheromones for the successful mating disruption of  rice yellow stem borer, Scirpophaga incertulas. The first trial compared the efficacy of 2 formulations containing 1:10:1 and 1:10:0 ratios of (Z)-9-hexadecenal, (Z)-11-hexadecenal and (Z)-9-octadecenal, components of the sex pheromone of yellow stem borer and a commercially-available formulation, Selibate CS, containing the related pheromone of Chilo suppressalis, a 1:10:1 blend of (Z)-9-hexadecenal, (Z)-11-hexadecenal and (Z)-13-octadecenal, with farmers’ practice plots treated with insecticides. Pheromone trap catches indicated that in each of the pheromone-treated plots the catches of adult males were reduced by up to 98% compared with catches in the insecticide-treated plots, suggesting that pheromone-mediated communication was disrupted. Larval damage ranged from 5.7 to 8.1% white heads (WH) in the insecticide-treated plots compared to a significantly reduced range of 2.1 to 2.4% WH in the pheromone-treated plots.

Cork and Basu (1996) also attempted to manage  rice yellow stem borer by mating disruption with a PVC resin formulation of the sex pheromone of C.suppressalis  in West Bengal.

Dhawan and  Sidhu (1978) have also reported on the effect of location of gossyplure traps on catches of the pink bollworm, Pectinophora gossypiella males. They conducted field studies in 1979-80 in India to observe the effect of position of gossyplure-baited traps on the capture of males of this gelechiid pest on the cotton. Traps placed at 25 cm above the crop canopy or near to stacked cotton stalks had the highest catch. Before the start of flowering, catches in traps inside and outside the cotton field were similar, but during flowering catches were lower in traps placed outside the cotton fields.  Taneja and Jayaswal (1986) worked  out  the population dynamics of P. gossypiella on upland cotton in Haryana. The number of adult males caught in traps baited with the sex pheromone gossyplure increased from mid-July to a peak in September-November, and declined thereafter. The incidence of larvae on cotton flowers peaked in mid- to late August, whereas the incidence of larvae in bolls increased during the cropping season. Using regression techniques, a linear relationship was established  between the per cent incidence of larvae in flowers and bolls and the number of males caught/ trap/night. Damage caused by the larvae was positively correlated with trap catches during the consequative crop seasons. 

Rup and  Sharma (1978) studied the role of the sex pheromone in the mating behaviour of pulse beetle, Callosobruchus maculates (F.), with a simple olfactometer in the laboratory. Observations confirmed an earlier finding that only the females of this species produce an attractant. The insignificant attraction of the females to the males, males to males and females to females confirmed the absence of an aggregation pheromone in this bruchid. Kanaujia and  Sidhu (1979) were   the pioneers who worked on pheromones of storage insects in India. The effects of age, mating, time of day and larval diet on the production of sex pheromone by adult females of the Angoumois grain moth Sitotroga cerealella were investigated in the populations reared on maize, wheat and  rice grains in the laboratory at Ludhiana. Pheromone production commenced in female pupae 1.5 days before adult emergence, reached a peak intensity 1-2 days after emergence and declined very slightly thereafter, persisting even in dead females. After mating, it declined to a level below that of virgin females but did not cease altogether. Pheromone production occurred throughout the diel but was highest at 24.00-04.00 h. There was no significant difference between the pheromone content of females that developed from larvae reared on the 3 different kinds of grain. The same authors have also studied on the factors affecting the responsiveness of males to female sex pheromone in S. cerealella (Kanaujia and  Sidhu, 1980). The laboratory studies were carried out in India on the response of males to the female sex pheromone, with regard to age, mating, pheromone concentration and time of day. Mating  response of males increased with increase pheromone concentration. Males of all the age-groups tested (0.25-9 days) showed similar responses to the pheromone. Mating induced a very high reversible inhibition of response that was independent of pheromone concentration and persisted for 14 h after mating. Although males responded to the pheromone at all hours of the day and night, a peak was observed from  24.00 to 05.00 h with a gradual increase during the dark hours and an abrupt decrease during the light hours.

Sharma and  Deora (1980) reported that females of rice weevil, Sitophilus oryzae  produced and released a sex pheromone on the day after adult emergence but the optimum level of response by males occurred on the fifth day. Pheromone responsiveness of males varied with the time of day showing the maximum response between 10.00 to 14.00 h and the minimum at 02.00 h. A normal photoperiod (LD 14:10) was required for the full male response to the pheromone. Mated and unmated males responded similarly. Starvation of males and females enhanced male responsiveness.
Mating disruption:

Nesbitt et al. (1986) have reported the first ever work on the sugarcane internode borer in India in collaboration with Sugarcane Breeding Institute, Coimbatore. Four pheromone components were detected in ovipositor washings and volatiles from females of the pyralid Chilo auricilius using combined gas chromatography and electroantennography. The components were identified as (I) (Z)-7-dodecenyl acetate [looplure], (II) (Z)-8-tridecenyl acetate, (III) (Z)-9-tetradecenyl acetate and (IV) (Z)-10-pentadecenyl acetate by comparison of their gas chromatographic behaviour with that of synthetic standards. In field tests carried out in northern India during 1982-84, a combination of II, III and IV components in their naturally occurring ratio (8:4:1) had shown a highly attractive synthetic source of pheromones. Looplure was found to reduce catches of males of C. auricilius, both when dispensed with the other 3 components and when released from dispensers surrounding a trap baited with the other 3 components.

Studies on mate seeking behaviour and anemotactic response of tropical warehouse moth, Ephestia cautella to (Z,E)-9,12-tetradecadienyl acetate were made by Singh and  Majumder (1983). Probably, this is a first  attempt on the synthesis of  a   pheromone in the India by Majee et al. (1983), who have synthesed Queenbee pheromone. Chattopadhyay et al. (1983) also synthesized the pheromones  of the queen bee and cabbage looper from aleuritic acid. The effects of various concentrations of (9Z,12E)-9,12-tetradecadienyl acetate on the mate-seeking and upwind anemotactic behaviour of  E. cautella were studied in the laboratory in India. The mating rate was inversely related to anemotactic flight at all pheromone concentrations tested. Extensive mating was observed during periods of low flight activity. The pheromone had no effect on calling females. Much anemotactic zigzagging flight activity was recorded 5-15 min after exposure to the pheromone at a rate of 10-3 ng, while there was little straight flight activity in comparison with zigzagging flight at all pheromone dosages. The highest upwind anemotactic response was observed 5-15 min after exposure to 10-4 ng of the pheromone. Olfactory studies indicated decreased response to increasing dosages of the pheromone. 
Siddiqi and  Khan (1983) have established the presence of a female sex pheromone which was detected for the first  time in the grass hopper, Hieroglyphus nigrorepletus (or indeed in any acridid) in the laboratory at Aligarh Muslim University, India, by means of a glass olfactometer. The pheromone was secreted in the digestive tract and expelled with the excreta, and it was perceived by the male through the antennae. No pheromone was secreted by newly emerged females, but secretion increased from 72 h to 8-9 days after emergence, when the first mating usually occurred; older females continued to release the pheromone, although in reduced quantities up to the age of 29 days.
Chari et al. (1985) have studied the populations of the noctuid H. armigera and  monitored with pheromone and light traps in field studies in bidi tobacco in Gujarat, India, during 1981-83. The pheromone traps were baited with 1 mg of a mixture of the synthetic pheromones (Z)-11-hexadecenal and (Z)-9-hexadecenal in the ratio 97:3. Catches in the pheromone traps peaked in December-April while catches in the light traps fluctuated. Light-trap catches were found to be significantly influenced by weather parameters and the moon phase. The study indicated that pheromone traps were more reliable for monitoring populations of H. armigera than light traps.

Sain and  Kalode (1985) have used the virgin female of the rice gall midge, Orseolia oryzae to attract males of the same species using various types of traps was investigated in field studies in India in 1982. Of three types of traps tested, only the delta trap was found suitable. When traps containing 3 and 6 virgin females placed in the middle of rice fields, 24 and 125 males were caught/night, respectively. No males were trapped in the absence of virgin females, indicating the presence of strong female pheromone. The population of O. oryzae was monitored weekly with the delta traps. The average catches varied from 2.3 males/trap/night in December to 120 in October, while there were no catches in February and May-July. The trap had several advantages over the light trap presently used for monitoring. A technique for catching live males of O. oryzae for laboratory cultures using an inverted funnel trap was described. 

A good amount of information has been brought out by  Sahu and  Hameed  (1983) on the pheromones in insect control. After five years of  the earlier report on the Induction of oestrous cycle irregularities in mice: presence of pheromonal stimulus in urine and excreta of females was reported by Gangrade and  Dominic (1983). They  have reported that Oestrous cycle irregularities and prolongation of the duration of vaginal cornification were induced in individually housed female by exposure to the urine of female housed in single-sex groups, or to bedding soiled by the excreta of such female. The results suggest the involvement of a non-volatile urinary contact pheromone in the incidence of oestrous cycle irregularities in unisexually grouped mice. Chandla et al. (1987) worked on monitoring of adult potato tuber moth, Phthorimaea operculella, with sex pheromone by conducting in potato fields at Shimla, India, during 1983-84. Sex-pheromone-baited water traps were found to be effective as a means of detecting and monitoring the occurrence of adults of   P. operculella at an early stage, so that control measures could be applied at the most appropriate time. The traps were baited with (4E,7Z)-4,7-tridecadienyl acetate and (4E,7Z,10Z)-4,7,10-tridecatrien-1-ol acetate. A total of 575 and 631 adult males were caught during May to August in 1983 and 1984, respectively.

Trap design
Limited information is available on the development of traps for various insects in Indian agriculture.  There has been efficient trap design for groundnut leaf miner (Nandagopal and Soni,1993) in groundnut ecosystem, for H armigera (Pawar et al.,1988 and Nandagopal et al.,2003),  For sugarcane internode borer, Chilo sachariphagus indicus  Kapur  (David et al., 1986), for S. litura, (Prasad et al., 1985;Krishnananda and Satyanarayana, 1985).  The efficiency of these traps were between 60 to 90 %.  In the case of Pectinophora gossypiella (Saunders)  in cotton ecosystem an interesting observation was reported (Nandagopal et al.,2003).  The trap and the lures used for H. armigera attracted the Pectinophora males.  Reed et al (1975)   and Satpute et al (1985) reported a efficient trap for P. gossypiella.

ii) Trap design developed


Information  available on the development of traps for various insects in Indian agriculture is limited. Efficient traps were designed for H armigera (Tamhanker et al., 1993; Pawar et al.,1988 and Nandagopal et al.,2003); groundnut leaf miner (Nandagopal and Soni,1993; Nandagopal et al., 2006); Sugarcane internode borer, Chilo sachariphagus indicus  (David et al., 1986); and  Spodoptera litura, (Prasad et al., 1985; Krishnananda and Satyanarayana, 1985; Ranga Rao et al., 1993).  The efficiency of these traps were between 60 to 90%. The trap and the lures used for H. armigera attracted the Pectinophora males in pigeon pea  ecosystem (Nandagopal et al.,2003). Reed et al (1975) and Satpute et al (1985) reported an efficient trap for P. gossypiella. The efficacy of 4 types of pheromone traps for catching adult males of Spodoptera litura in groundnut fields was evaluated in studies in India in 1988 (Table 6). There was no significant difference in the performance of single- and double-funnel traps, and the single-funnel trap (20 cm in diameter) captured more moths than any other type of trap. Septa of 4 weeks or less exposure attracted most moths. One and 2 traps per ha caught significantly fewer moths than 4 and more traps per ha; however, there was no significant improvement in capture when 4 or more traps per ha were installed. Night observations indicated that many moths escaped from sleeve traps. Single plastic funnel trap found suitable when being used in monitoring S. litura in the national monitoring network in India (Roa et al., 1991) (Nandagopal and Prasad, 2004).
Table 6 : Trap designs developed in India
Species

1.1          Reference

Pectinophora gossypiella (Saunders) (Gelechiidae:Lepidoptera)
Reed et al., 1975

Nandagopal et al.,2003
Helicoverpa (Hub.)
(Noctuidae:Lepidoptera)
Pawar et al.,1988;
Tamhankar et al., 1993;
Nandagopal et al.,2003
Sitotroga cerealella (Olivier)
(Gelechiidae: Lepidoptera).
Kanaujia and Sidhu, 1981.
S. litura
(Noctuidae:Lepidoptera)
Prasad et al., 1985;
Krishnananda and Satyanarayana, 1985
Chilo sachariphagus indicus  Kapur
(Pyralidae:Lepidoptera)
David et al., 1986 ;
PCI (pvt) Ltd,Bngalore

Aproaerema modicella  (Dev.)
 (Gelechiidae:Lepidoptera)
Nandagopal and Soni,1993,
Nandagopal et al., 2004; 2006
Ranga Rao et al., 1993
Spodoptera litura
Ranga Rao et al., 1991
Bactrocera dorasalis walker
Patel et al., 2004


iii) Traps available and improved

There are many kinds of traps available for various important insect and mite pests. These include Sleeve trap, Water trap made of Galvonized Iron trays or Plastic Watta –Trap, Sticky trap such as Delta trap, Diomond trap, Panel trap, Unit trap, Rhegalatis trap (Plate 2.).  The plate 3. is the stika trap named David’s trap has been developed for groundnut leaf miner in early 1990s.

There are particular trap for each of the species of insects belonging to a family/order to trap a specific pest. For example water traps are mostly used for Pyralid moths of the order Lepidoptera. When it was modified using a plastic, the trap is used for  catching males of  brinjal fruit and shoot borer  and  Groundnut leaf miner a Gelechiidae of order Lepidopera (Plate 4.).

A typiocal example of how a available trap can be improved  has been demonstrated by Nandagopal et al., (2006) which has been presented below:

Groundnut leaf miner (GLM), Aproaerema modicella Dev. (Gelechiidae: Lepidoptera) is a serious oligophagus insect pest of groundnut, soybean and  other few  legume crops in south and south East Asia (Mohammad, 1981). GLM populations fluctuate widely between seasons.  Population peaks have been reported in July and August in Thailand (Campbell, 1983), at the end of the rainy season (September and October) specially in drought or low rain fall years (Amin, 1983), in November and December (Mohammad, 1981) as well as at the end of the post-rainy season i.e. March and April in Bangladesh and India (Islam et al., 1983).

The identification of sex pheromone of GLM  (Nandagopal and Reddy, 1990; Hall et al., 1994) has enabled to monitor this pest with pheromone trap fabricated based on the behavior of this species (Nandagopal and Soni, 1993). Male moth catches of pheromone traps at ICRISAT, Hyderabad showed that two highest peak populations of this pest appear in  30-33 standard weeks (July-August) with 4-5 generations during rainy season and 6-11 standard weeks (February-March) with 6 generations during post-rainy season.  Several years of pheromone trap data is required to indicate population fluctuations of GLM over seasons. 

To mitigate the hazardous consequences due to indiscriminate use of chemical pesticides and reduce the cost of cultivation, use of sex pheromones for monitoring, mass trapping and mating disruption has become essential tool in IPM programme. Experiments were carried out on developing the efficient pheromone trap based on the behavior of this pest. Among various designs of traps such as delta, funnel, sticky and water trap evaluated against GLM, water trap was found effective (Ranga Rao et al., 1993). Based on the trapping efficiency of the trap, number of insects trapped, the management options have to be decided. Hence the present investigation was taken up to evaluate different types of traps available in market and make necessary modifications so as to increase the trapping efficiency. 

Randomized field experiment was conducted at  National Research Centre for Groundnut (NRCG),  Junagadh, Gujarat during rainy season of 2004. Pheromone lures of polythene vials impregnated with the blend of   (Z,Z) 7-9- Decadienyl acetate (1) + (E) 7-Decenyl acetate (II) + (Z) 7-Decenyl acetate (III) in the ratio of 10:2:1.4 manufactured by Pheromone Research Group, Indian Institute of Chemical Technology (IICT), Hyderabad were used in the experiment.  Each trap was fitted with lures loaded with 3 mg pheromone. The traps were placed at 0.5 m above the ground in the groundnut crop using the inverted ‘ L’ shaped bamboo sticks (Ranga Rao et al., 1993) and with a distance of 10 m apart on all the sides.

Four types of traps namely Acrylic Sticka trap (small), Acrylic Sticka trap (big), plastic tray trap, Wota-T trap were used in the study. Wota – T trap from Pest Control India (PCI), Bangalore was used with castor oil and water+ kerosene, while castor oil was used in the remaining traps as trapping material. Castor oil was smeared on the inside and sides of the trap as trapping material. All the traps with three replications were installed in groundnut fields where GLM infestation was more than 60 percent. The positions of pheromone traps were changed clockwise  on alternate day to eliminate locational/positional effect, if any on the trap catches. The traps were arranged following the procedure of Nandagopal (1992).  Each trap was checked everyday for male moths trapped and observations were recorded for five consecutive days. The data obtained  was statistically analyzed using two way analysis of variance.
The results of the present study showed that the difference in the male moths trapped in various traps were highly significant. Out of the five different types of traps evaluated against A. modicella, catches of male moths were significantly higher in Wota –T trap compared to other traps tested (Table 1). Wota –T trap with castor oil was found superior (196.8 male moths /trap/day) which was on par with  Wota –T trap having water and kerosene as trapping material (194.8 males/trap/ day).

The traps with two and four openings differed significantly, where  4 entry opening trap caught more male moths compared to two opening trap. Even the bigger acrylic trap which was found to be superior in our previous trials found to trap only 66 male moths /day however, the number of moths trapped in bigger acrylic trap with four openings were more than the smaller acrylic trap with two openings. The probable reason could be four opening of the bigger acrylic trap facilitate more number of male moths to enter into the trap compared to  trap with two openings.

Since GLM is a passive flier with limited distance by itself and mostly carried by the downward wind whose direction may change. It is this which makes the four opening trap ideal as there is an opening at each of the four corners of the compass to  facilitate the catching of the pest. The low population of males trapped in small acrylic sticka trap clearly indicates the difficulty that the male pest have when trying to negotiate entry to the trap when the wind is blowing in the wrong direction.

It is interesting to note that though the trapping area is more in Plastic tray trap (2040 cm2) compared to Wota-T trap (1584 cm2) the number of male moths trapped are less in plastic tray trap (108.5 male moths /trap/day)  than in Wota –T trap (196.8 male moths /trap/day) indicating that the efficiency of the trap depends upon the design of the trap rather than the total trapping area. It was observed that  the males when visited the lures  run all around the lure, probably in search of the females, when found otherwise, they tried to move out. This behaviour of moving out is very important in designing the trap. When we measured the angles of movement and the site of the catch, it was below 45º angle and those which were caught on the sides were of more than 45º angle (Fig. 4). However, there may be certain % of males moths might have escaped by flying above 45 ° angle (ACD) up to just below 180° angle BDE   (Fig. 5).  Probably the increased angle might have facilitated for  catching those males which may even move above 45 ° angle. 

Ranga Rao et al. (1993) reported open water trap having 28cm x 22cm with a gap of 11 cm clearance was effective against GLM. The water trap used by the Ranga Rao et al., (1993) was made up GI sheets fitted with a rod. The cost of the water trap was estimated to be Rs. 200 per trap as per  the local manufactures. Whereas the Wota –T trap used in this trial is about Rs. 39 per trap. There was no significant difference between castor oil and water + kerosene as trapping material in Wota –T trap indicating that these can be efficiently used for trapping of GLM however, we recommend to use castor oil in the Wota –T trap for easy and economic considerations (Nandagopal et al., 2006) Table 7 .


Table 7: GLM moth catches in different types of traps (five nights)
Type of trap
Mean moth catches/ day
Acrylic stika trap (small)
18.89
Acrylic stika trap (bigger)
66.33
Plastic tray trap
107.55
Wota –T  trap (Castor oil)
196.89
Wota–T trap (water+ kerosene)
194.78
SEm
26.57
CD (5%)
86.67
CV(%)
39.4


For further readings:

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Balakrishnan,N., R.K.Murali Baskaran and N.R. Mahadevan.2004. Pheromone trap catches of bollworms in different IPM modules in rainfed cotton ecosystem P16. In  National Seminar on Trends in pheromone research and technology, 6-7 February,2004, National Research Centre for Groundnut, Junagadh, Gujarat.
Balasubramanian M; Murugesan S; Parameswaran S. 1979. Trapping of the cotton pink bollworm Pectinophora gossypiella (Saunders) Gelechiidae, Lepidoptera moths using synthetic gossyplure. Pesticides. 1979, 13: 6, 49,51.
Bartell,R.J.1982. Mechanism of communication disruption by pheromone in the control of Lepidoptera: a review. Physiological Entomology, 7:353-354.
Campbell, W.V. 1983. Management of arthropods on peanut in southeast Asia. Ann. Rep. Peanut Collab. Res. Support Prog. (CRSP), University of  Georgia, Georgia Experiment Station, USA.
Chandla,V.K.,  Bhalla,O.P.,  and Dogra,G.S. 1987. Monitoring of adult potato tubermoth, Phthorimaea operculella Zeller, with sex pheromone. National-Academy-Science-Letters. 1987, vol 10: 11, 397-399.
Chari,M.S., Patel,A.R., Rao,B.S., Bharpoda,T.M., and Patel,N.M. 1985.Population studies on tobacco capsule borer Heliothis armigera Hubner. Tobacco-Research. 1985, vol 11: 2, 98-104.
Chari,MS; Patel,AR; Rao,BS; Bharpoda,TM; Patel,NM.1985. Population studies on tobacco capsule borer Heliothis armigera Huber Tobacco, Research. 1985, 11: 2, 98,104.
Chattopadhyay,A.,  Mamdapur,V.R., and  Chadha,M.S.1983. Synthesis of  the queen bee and cabbage looper the pheromones  of from aleuritic acid. Indian Journal of Chemistry,B. 1983, 22: 2, 158-159.
Cork, A. and Basu, S.K. 1996.Control of the yellow stem borer, Scirpophaga incertulas by mating disruption with a PVC resin formulation of the sex pheromone of Chilo suppressalis (Lepidoptera: Pyralidae) in India. Bull. Entomol. Res. 86: 1, 1-9.
Cork,A,, Souza-Kde, Krishnaiah,K. Kumar,D.V.S.S.R., Reddy,A.A., and  Casagrande,E. 1996.  Control of yellow stem borer, Scirpophaga incertulas (Lepidoptera: Pyralidae) by mating disruption on rice in India: effect of unnatural pheromone blends and application time on efficacy. Bulletin-of-Entomological-Research. 1996, vol 86: 5, 515-524.
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