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:
Bollworm trap ht Predictability of larval damage catch/night/trap
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
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:
Anonymous.1981. An operational
field trial project in India for suppression of the cotton pink bollworm, Pectinophora
gossypiella (Saunders) (Gelechiidae: Lepidoptera) employing 'gossyplure
hollow fiber' controlled release sex pheromone formulation.Technical,Report,,Plant,Protection,Adviser,to,the,Government,of,India.
1981, recd. 1983, No. 1, [7+] 44 pp.
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.
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