semiochemicals for sustainable pest management · 2020. 6. 29. · ijrar1bjp202 international...
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Semiochemicals for sustainable pest management
Sonawane Bhushan Laxman1, Kushal Thakur1, Bhade Khemraj Shankar1, Jyoti B. Patil1, Pathma J*1
Abstract
Modifying the chemical ecology of insects which could change insect behaviour is an area under exploration and
could provide ample opportunities to manage noxious herbivore pests damaging food crops leading to loss in
agricultural production and causing human health hazards by inflicting poisonous bites and vectoring dreadful
pathogens. Semiochemicals are organic signalling molecules which triggers intra and inter specific
communication among living organisms. Insect pheromones are an important group of intra specific
semiochemicals which have gained commercial importance with respect to pest surveillance and management
programs. They assist in pest monitoring, and mass trapping and cause mating disturbance in insect pests aiding
in pest management. This review highlights the classification of semiochemicals, their mode of action,
techniques involved in pheromone isolation and characterization, commercial formulations available and their
utilization in eco-friendly pest management.
Keywords: Semiochemicals, pheromones, pest management
1. Introduction
Semiochemical originates from a Greek word “semeon” meaning signal or sign. These are biologically active
organic substances which induces behavioural changes in living organisms. Insects use these semiochemicals to:
locate their hosts for food and shelter; avoid competition; organise themselves; evade natural enemies; reach their
mates for courtship etc., Molecular weight of the semiochemicals decides its volatile nature thereby enabling
insect communication and its success in pest management programs. Compounds with molecular weight ranging
between 80-300 with 5 to 20 carbon atoms are volatile in nature and naturally allow long distance
communication in insects (Wilson and Bossert 1963). They include small hydrophobic molecules as well as
water soluble peptides. History of use of pheromones for pest management dates back to 19th century when the
sex pheromone from female Bombyx mori was identified and isolated by Butenandt and his team in 1959
(Butenandt et al. 1959). Olfactory receptors were later identified in drosophila and silk moth (Clyne et al. 1999;
Vosshall et al. 1999; Krieger et al. 2005) which intensified the interest in insect chemical ecology and pheromone
research. The research also entered molecular level and the olfactory receptor of sex pheromone bombykol in
Bombyx mori viz., BmOR-1 and the role of CD36-related receptor in pheromone detection in Drosophila were
documented (Sakurai et al. 2004; Benton et al. 2007). Till date pheromones have been isolated from nearly 1500
insect species. Parallel concepts on insect chemical ecology and development of analytical techniques such as
1Department of Entomology, School of Agriculture, Lovely Professional University, Punjab
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gas chromatography opened up new avenues in exploring the possibilities of using these semiochemicals as an
alternative to hazardous chemical insecticides for pest management (Brossut 1997; Cork 2004). Understanding
the chemical ecology of insects, their behavioural responses to the signalling molecules apart from identifying
the key signalling compound and studying their chemical nature will provide us better insights on their utility in
insect pest management. Extraction and structural elucidation of these natural molecules will enable us to
characterise them and artificially synthesis them without loss in their properties and with improved
environmental stability.
2. Classification
Semiochemicals are molecules of biological origin eliciting a behavioural response in the organisms of the same
species (intraspecific) or different species (interspecific). Based on the origin and the nature of response they
elicit they can be classified broadly as Pheromones (intra specific semiochemicals) and allelochemicals
(interspecific semiochemicals). Pheromones can be further classified as sex pheromones (produced by one sex
which attracts the opposite sex of the same species), aggregation pheromones (attracts both male and females
irrespective of the sex producing the pheromone and in most cases are produced by males), Trail marking
pheromones (chemical communication in ants), Alarm pheromones (chemical communication in aphids and
honey bees) and host marking pheromones (chemical communication in parasiotoids). Allelochemicals can be
divided as Allomones (benefitting the emitter), Kairomones (benefitting the receiver), Synamones (benefitting
both emitter and receiver) and apneumone (substances originating from non-living material benefitting certain
organisms and detrimental to other). Few semiochemicals can also elicit both inter and intraspecific
communication.
3. Mode of action
Use of semiochemicals has become an integral part of eco-friendly integrated pest management and
semiochemicals are considered as ‘new generation pest control agents’(Suckling 2000; Tinzaara et al. 2002).
‘Semiochemicals’ are ‘infochemicals’ which provides chemical information that mediates behavioural and
ecological interactions between the organisms such as plants and insects sharing a common ecosystem. These
infochemicals when released into the environment by the emitter are detected by the chemoreceptors present on
different body parts and in many cases concentrated on the antennae of the receiver insect in which a behavioural
change is inflicted. Molecular pathway of olfactory perception involves multi-gene families comprising of
odorant-binding proteins (OBPs), chemosensory proteins (CSPs) and odorant receptors (ORs) with helps in
odour recognition and response (Siciliano et al. 2014). Certain signals are short-lived providing an quick message
such as forewarning danger (alarm pheromone) or reproductive readiness while certain signals reach longer
distances and persist for long time and are involved in attracting the opposite sex, or mark the geographical
boundary etc., Pheromones are target specific and highly safe to other non-target organisms occupying the
ecosystem, highly active in extremely small quantities in range of nanograms, thereby pose no pollution hazard
to the environment. Additionally, insects with protected lifestyle living within plant tissues with lesser
possibilities to be exposed to insecticides can be easily managed by use of pheromones. Thus these
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semiochemicals could be exploited for pest surveillance and management programmes with high safety and
feasibility (Witzgall et al. 2010).
4. Use of semiochemicals in pest management programmes
4.1 Pest monitoring
Pest monitoring is an integral part of pest management program and provides timely information on the
occurrence of regular pests and migration of invasive pests both qualitatively and quantitatively which will
enable us to assess risk and take up timely pest control measures based on the economic threshold levels and
integrate all pest management tactics appropriately and thereby avoid potential damage to the crops and
environment (Suckling 2000). Pheromones helps us in monitoring programmes involving pests of agro-
ecosystems, horticulture ecosystems, forest ecosystems, storage products and house-hold pests (Wall 1989;
Burkholder 1990; Borden 1993). Wall et al. (1987) devised an action threshold for Cydia nigricana (pea moth)
based on trap catches and this was the first widely adopted use of infochemical for pest monitoring system. The
sensitivity of pheromones traps to efficiently detect insects in low population density enables effective tracking
of invasive species in the early introduction phase prior to establishment (El-Sayed et al. 2006; Liebhold and
Tobin 2008). However the success of the use of pheromones in pest monitoring programmes is decided by
numerous factors including formulation of pheromones, chemical purity, dispenser quality, release rates, trap
design, trap location, data analysis and interpretation, environmental factors (eg. strong winds), insect species
(strong flyers) and expert decision systems. In few cases the trap catches did not correlate with the damage levels
in the field (Srivastava et al. 1992; David and Birch 1986; Tadas et al. 1994). A strict quality control system is
essential to ensure the success of the pheromones in pest monitoring as well as trapping (Arn et al. 1997).
4.2 Pest management
4.2.1 Attract and affect: In this technique the semiochemical is used as a lure to attract insect towards a
hazardous agent viz., insecticide or pathogen or sterilant which can attract and kill or attract and infect or attract
and sterilize the insect accordingly thereby reducing their population and ecological fitness (El-Shafie and
Faleiro 2017).
4.2.2. Mating disruption: Mating disruption refers to meddling with the process of mating naturally in sexually
reproducing insects by miss-orienting the males to the synthetic pheromones placed in dispensers (micro
capsules/ hollow fibers) resulting in reduced oviposotion (Carde and Minks 1995; Harari et al. 2015). In many
cases female produces sex attractants to attract males and use of these attractants will lure the males and trap
them making them unavailable for mating. Conditions including early eclosion of males and protandry adds
strength to this annihilation technique (Witzgall et al. 2010). Installation of traps containing sex pheromones in
the natural environment creates an artificial signalling thereby reducing the probability of a male reaching the
female resulting in mating disturbance, increasing the probability of females laying unfertilised, inviable eggs
thereby reducing the off-spring population. This technology has found a great success in controlling many
lepidopteran crop pest species including Pectinophora gossypiella (pink boll worm of cotton), Helicoverpa
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armigera (highly polyhagous American boll worm), Spodoptera litura (highly polyhagous tobacco caterpillar),
Chilo suppressalis and Scirpophaga incertulas (stem borers of cereals), Cydia pomonella (apple codling moth),
Epiphyas postvittana (light-brown apple moth) and Lobesia botrana (European berry moth in vineyards) etc.,
and coleopteran pests such as Cylas formicarius, C. puncticollis, C. brunneus (sweet potato weevils) etc (Carde
and Minks 1995; Cork et al. 1996; Tamhankar et al. 2000; Downham et al. 2001; Witzgall et al. 2008; Reddy and
Guerrero 2010). However this technique also has some limitations which includes potential risk of mated females
intruding the treated area unless this technique is employed over a large area. Thus success of this technique
requires a large scale community participation rather than adaptation by individual farmers (Carde and Minks
1995; Staten et al. 1997). Additionally, very high population of the targeted insect which will reduce the
competition between natural and synthetic lures also hinders the success of this technique (Carde and Minks
1995). Furthermore the lures should be designed to release the attractant at sufficient strength to disrupt natural
mating. However, this technique will fail to control long lived species with ability to undergo multiple mating’s
as the effectiveness of mating disturbance lies in a large proportion of males failing to mate the females
(Suckling 2000). Mechanisms of mating disruption includes competitive attraction or false trail following
(density-dependant), Camouflage (density-independent), desensitization and sensory imbalance (El-Shafie and
Faleiro 2017).
4.2.3. Mass trapping: This is a method of attracting and trapping adult insects in large numbers by installing
more number of pheromone traps which will reduce the insect population in the subsequent generation (Birch
and Haynes 1982). The success of mass trapping using sex pheromones again depends on various factors and is
decided by the number of traps deployed, trapping frequency, nature of insects involved, integration of this with
other crop management tactics including crop sanitation etc. High insect population density, high reproductive
rate of females, polygamous nature of males and availability of a number of natural attractant sites are factors
responsible for diminishing the success rate of mass trapping using sex pheromones (Cuthbert et al. 1977; Birch
and Haynes 1982; Weislein 1992; James et al. 1996; Tamhankar et al. 2000). Mass trapping technology could be
effective in insects with limited dispersal ability and slow population build up with longer life span and lower
fecundity (k-stratergies) (Giblin-Davis et al. 1996a). ‘Trap spillover’ is another mechanism which might cause
increased damage to the host crop surrounding the traps making the traps less effective in certain cases as
reported with Japanese beetle, Popillia japonica (Switzer et al. 2009).
4.2.4. Push-pull strategy: This strategy involves the use of two different semiochemicals (an attractant and
repellent) stimulating a contradictory behaviour in target pests and their natural enemy complex. In depth
knowledge on arthropods chemical ecology is essential to utilize this strategy for successful pest management.
This stratergy involves use of semiochemicals that make the protected resource unattractive to the pests (push)
and attracting them towards a lure (pull) where they can be destroyed (Pyke 1987; Cook et al. 2007).
5. Factors affecting the effectiveness of use of pheromones for pest management
Numerous factors that decide the success of utilization of pheromones for pest management programmes
includes trap color, trap design, pheromone dose, pheromone release rate, spatial and temporal patterns of trap
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arrangement (De Groot and De Barr 1998; Giblin-Davis et al. 1996b); environmental factors including wind
velocity, direction, temperature, rainfall, humidity (Jansson et al. 1989; Sappington and Spurgeon 2000), pest
density, pest biology and physiology (age, sexual maturity, response to pheromones, mating status) etc., (Jansson
et al., 1991; Tinzaara et al. 2002). Another important edaphic factor deciding the success of utilization of
semiochemical in pest management is the large scale adaptation of the strategy over a wide area (Witzgall et al.
2010). Additionally, synergistic effect of certain host plant volatiles with the insect pheromones have been
reported and reviewed (Light et al. 1993; Reddy and Guerrero 2010).
6. Isolation, identification and synthesis of semiochemicals
Isolation and structural elucidation of semiochemical is a tedious process involving numerous steps and
sophisticated instrumentation (Silverstein et al. 1967). Basic steps involved includes compound extraction,
bioactivity evaluation, bioactive compound identification, chemical synthesis and bioactivity evaluation of the
synthesised compound and mass production etc., Compound extraction can be done by various methods such as
solvent-assisted extraction followed by distillation, Static/ Dynamic Headspace Sampling, Solid-Phase
Extraction, Solid-Phase Micro-Extraction etc., Confirmation of bioactivity with suitable bioassays,
Electroantennography, Single Sensillum Recording, Audiovisual Analysis, Sensor based detection and
behavioral studies (Linear/ Two-Path/ four or more path Olfactometry, Wind-Tunnel Assay/ no-choice tests etc).
Bioactive compound identification by various analytical techniques viz., gas chromatography coupled with:
Flame Ionization Detectors (GC-FID) or electroantennography (GC-EAG)/ Mass Spectrometry (GC-MS) or
Nuclear Magnetic Resonance (NMR). Additionally semiochemical identification can also be done through
‘reverse chemical ecology’ approach which utilises molecular biology and bioinformatics tools (Leal et al. 2008).
If the pheromones are of glandular origin as in case of lepidopterans the gland can be detached for
semiochemical extraction. However this method has many disadvantages including the inclusion of non-targeted
compounds with targeted semiochemical as emitted by the active organ and hence in situ volatile collection
through head space collection is more advantageous. Other methods include passing pure air onto the insect
trapped in an aeration chamber and trapping the exhaust air on columns with suitable absorbent (Super Q), or
subjecting the air from outlet to GC-MS analysis using or solid phase micro-extraction (SPME) technique where
the volatiles from the test sample carrier by the air passed are trapped in needle like fiber coated with silica
positioned in a cylinder resembling a syringe. The fiber is inserted into an inlet maintained at high temperature to
vaporise and elute the trapped compounds for analysis by GC or GCMS (Pawliszyn 2000). Mass spectroscopic
analysis (MS) and proton nuclear magnetic resonance (NMR) helps in identification of active compounds.
Differential analysis by 2D NMR spectroscopy (DANS) can help in identification of small bioactive molecules at
very low concentrations from a mixture of compounds (Robinette et al. 2011; Soroker et al. 2015). Advanced
molecular tools enable us to decipher the chemical signals involved. An odarant binding protein was used to
identify an ovipositional attractant compound in mosquito by use of ‘Reverse chemical ecology approach’ (Leal
et al. 2008; Pickett et al. 2010). Further research and use of electrophysiological imaging will enable odorant
receptors ligand identification and their expression in heterologous cell system for high-throughput screening
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(Wetzel et al. 2001; Hallem et al. 2006; Kiely et al. 2007) or the combined use of structural chemistry, bio
informatics and statistics software could calculate physicochemical odor metrics which inturn predicts the
neuronal responses (Haddad et al. 2008). Few extensively studied and commercially available semiochemicals
for pest management is tabulated (Table 1).
7. Conclusion
Growing population with shrinking land and other resources is a challenge to agriculturists to produce quality
food to assure demand supply balance. The situation is in turn aggravated by climate change posing threat to crop
production. Climate change has a tremendous impact in the crop pests and had enhanced their pest status,
intensified their vector and migration potential adding to their noxious and invasive nature. Continuous use of
chemical insecticides to tackle the pests has led to insecticide resistance, pest resurgence and replacement. Use of
eco-friendly pest control alternatives is the need of the hour. Use of semiochemicals could be an effective
alternative but requires interdisciplinary research involving entomologists, molecular biologists, chemists and
ecologists for understanding the pest biology, chemical ecology, molecular mechanisms involved and successful
identification of bioactive molecules eliciting a behavioural change in insects so as to synthesise them and utilize
them to manipulate insect behaviour in the agro-ecosystems. In addition, the success of integration of
semiochemicals in pest management is decided by a large scale adaptation of this technique, cost effectiveness
and efficacy of the method utilized for field application. Intense focussed research and combination of classical
techniques with advanced technology could work out marvels and identify better, eco-friendly, cost effective
solutions for sustainable pest management.
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Table 1 Pheromones for pest management
Infochemical class Associated Insect Order Key compound References
Sex pheromone Helicoverpa armigera Lepidoptera Helilure ((Z)-11-hexadecenal (I) and (Z)-11-
hexadecen-1-ol(II),
Nesbitt et al. (1979)
Sex pheromone Spodoptera litura Lepidoptera Spodolure: (Z,E)-9,11- and (Z,E)-9,12-
tetradecadienyl acetate (Z9,E11-14:Ac;
Z9,E12-14:Ac). / cis-9, trans-11- and cis-9,
trans-12-tetra-decdien-1-ol acetates
Tamaki et al. (1973)
Sex pheromone Pectinophora
gossypiella
Lepidoptera Gossyplure: blend of cis and trans isomers of
7,11-hexadecadienyl acetate
Hummel et al. (1973)
Sex pheromone Tuta absoluta Lepidoptera triene, (E, Z, Z)-3,8,11- tetradecatrien-1-yl
acetate
Svatos et al. (1996)
Sex pheromone Leucinodes orbonalis Lepidoptera (E)-11-hexadecenyl acetate (E11-16:Ac) Cork et al. (2001)
Sex pheromone Cydia (=Laspeyresia)
pomonella
Lepidoptera Codlemone: trans-8, trans-10-Dodecadien-1-ol Roelofs et al. (1971)
Synthetic attractants Noctuid moths Lepidoptera Phenylacetaldehyde, β-myrcene, 3-methyl-1-
butanol and acetic acid
Haynes et al. (1991); Heath et al.
(1992); Landolt and Alfaro (2001);
Landolt et al. (2006, 2007)
Sex pheromone Agriotes sp. Coleoptera Geranyl octanoate Toth et al. (2008)
Sex pheromone Xylotrechus quadripes Coleoptera (S)-2-hydroxy-3-decanone Hall et al. (2006)
Aggregation pheromone Oryctes rhinoceros Coleoptera Ethyl 4-methyloctanoate, Hallett et al. (1995)
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Aggregation pheromone Ips duplicatus Coleoptera Ipsdienol and E-myrcenol, Schlyter et al. (2003)
Aggregation pheromone Rhynchophorus
palmarum
Coleoptera Rhynchophorol: (4S)-2-methyl-(5E)-hepten-4-
ol
Rochat et al. (1991); Oehlschlager
et al. (1992)
Aggregation pheromone Rhynchophorus
ferrugineus
Coleoptera Ferrugineol: (4S,5S)-4-methyl-5-nonanol and
4-methyl-5-nonanone
Giblin-Davis et al. (1996); Perez
et al. (1996)
Aggregation pheromone Cosmopolites
sordidus
Coleoptera Sordidin Reddy et al. (2009)
Aggregation pheromone Anthonomus grandis Coleoptera Grandlure Tumlinson et al. (1969)
Synthetic pheromone
blend
Popillia japonica Coleoptera Japonilure with floral compounds, eugenol,
geraniol and phenethyl propionate
Switzer et al. (2009)
Anti-aggregation
pheromones
Dendroctonus
ponderosae
Coleoptera verbenone Gillette et al. (2009a)
Anti-aggregation
pheromones
D. pseudotsugae, Coleoptera methylcyclohexenone (MCH) Gillette et al. (2009b)
Feeding Attractants Diabrotica virgifera
virgifera
Coleoptera Cucurbitacin (plant metabolite) Toth et al. (2007)
Lure Agrilus planipennis Coleoptera Volatile Sesquiterpenes from Fraxinus
pennsylvanica
Crook et al. (2008)
Alarm pheromones Various aphids
species
Hemiptera (E)-β-farnesene Bowers et al. (1972)
Synthetic Bactrocera cucurbitae Diptera Cue-lure Witzgall et al. (2010)
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Parapheromones and
attractants
Synthetic
Parapheromones and
attractants
Bactrocera dorsalis Diptera Methyl eugenol Witzgall et al. (2010)
Synthetic Sex
pheromone and food
attractant
Bactrocera oleae Diptera (1,7)-dioxaspiro-[5,5,]-undane (olean), n-
nonanal, α-pinene, ethyl dodecanoate and
ammonium bicarbonate
Mazomenos and Haniotakis
(1985); Broumas et al. (2002)
Synthetic
Parapheromones and
attractants
Ceratitis capitata Diptera Trimedlure Witzgall et al. (2010)
Feeding Attractants Musca domestica Diptera Muscalure: (Z)-9- tricosene Butler et al. (2007); Geden et al.
(2009).
Synthetic lure Glossina spp., Diptera Mixture of 1-octen-3-ol, 3-n-propyl phenol,
and 4-methylphenol (p-cresol) plus acetone or
methyl ethyl ketone in a separate dispenser
Bursell et al. 1988; Vale et al.
(1988)
Synthetic lure Lucilia cuprina Diptera Blend of 2-mercaptoethanol, indole, butanoic
acid, and sodium sulfide
Ward and Farrell (2003); Urech et
al. (2004)
Attractants (food baits) Social wasps Hymenoptera Mixture of acetic acid, with isobutanol or
heptyl butyrate or butyl butyrate
Landolt et al. (2000)
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