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Short Views on Insect Biochemistry and Molecular Biology Vol.(1), October 2014

2014

Section I
Insect Biochemical approaches

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Short Views on Insect Biochemistry and Molecular Biology Vol.(1), 2014

IO

NAL B OO

SION
MIS

TERNA
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Short Views on Insect Biochemistry and Molecular Biology

Vol. (1), 00 00,
2009
Vol.
(1) 57 74, 2014

Review Article

Chapter 2

Modulation of Botanicals on Pests Biochemistry

K. Sahayaraj
Department of Advanced Zoology and Biotechnology,
St. Xavier's College (Autonomous), Palayamkottai 627 002, Tamil Nadu, India

Abstract

Insect lives in any agro-ecosystem both as beneficial and harmful organisms to crops. They
represent within insect orders particularly Lepidoptera, Hemiptera, Orthoptera, Coleoptera,
Thysanura, Blateria ect. The main consequence of insect pests includes infestation as well as acts
as vectors. These problems were apparently under control after the development of conventional
insecticides. However, side effects have been notices for microbes, plants, animals, and our
environments. One among the well-known alternative is botanicals. Some studies that have
searched for potential anti-insect products from plants have focused on repellent activity to
prevent insect pests infestation. Most studies, however, evaluated the toxicity of plant extracts and
plant-derived compounds, in order to control insect pests. To our knowledge, no attempts have
been made to discuss the biochemical impact of botanicals against pests. In this review we focus
on impacts of botanicals on carbohydrate (glycogen and trehalose), proteins (total proteins and
amino acids), lipids (total cholesterols and phospholipids), electrolytes (Na+, Cl, K+ and Ca2+
levels) and both digestive (Amylase, Glycosidases, Alkaline phosphatase, Lipases, Proteases) and
detoxication (esterase, oxidase, transoxidase hydrolases, glutothion transferase, cis-oxidase
hydrolases, adrin epoxidase, cytochrome P-450, alanine aminotransferase, asparate aminoteransferase, glutathione S-transferase, glutathione P-transferase) enzymes either directly or
indirectly to cause death or alter the behaviors which lead to death.

Key words: biochemical analysis, carbohydrates, lipid, enzymes, detoxification, mortality, plant defense
*For Correspondence (email: ksraj42@gmail.com)

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Overview
1. Introduction
2. Biological Traits
2.1. Feeding and Mortality
2.2. Reproduction and
Embryogenesis
3. Macromolecular profiles
3.1. Carbohydrates
3.1.1. Glycogen
3.1.2. Trehalose
3.2. Proteins
3.3. Lipids
3.3.1. Total Cholesterols
3.3.2. Phosopholipids
4. Electrolytics
5. Genomic DNA and RNA
6. Energy
7. Digestive enzymes
7.1. Digestive enzymes
7.1. Amylase
7.1.2. Glycosidases
7.1.3. Alkaline phosphatase
7.1.4. Lipases
7.1.5. Proteases
7.2. Detoxification enzymes
8. Conclusions and future direction
9. References

Review Article

1. Introduction
The environmental hazards by synthetic
chemical insecticides have necessitated the
search for the some alternative source of natural
origin for applying ecologically viable pest
control strategies. The usage of botanical,
aqueous extracts, solvent-based extracts, their
column fractions and also their bioactive
principles are one of the alternatives because of
the low environmental persistence and low
mammalian toxicity. The practice of using
botanical insecticides or plant extracts in
agriculture dates back at least to two millennia
in ancient Egypt, India, China, and Greece. In
Europe and North America, the documented use
of botanicals extends back to more than 150
years, dramatically predicting discoveries of the
major classes of synthetic chemical insecticides
beginning in the 1940s.

Taking into account the increasing utility of botanicals, it is imperative to know

about their impact on the biochemistry of insects. Aqueous extract or crude extracts
or column fractions or oil of a number of plants or their bioactive principles have
been used for the management of various economically important pests world-wide.
Large amount of works have been carried out world-wide and also voluminous texts
and general or specific reviews on pesticidal or insecticidal plants literature have been
published. Further, the secondary metabolic compounds synthesized by plants
(alkaloid, terpenoids, phenolics, non-protein amino acids, amines, cyanogenic
glycosides, Gossypol, Phytoestrogens, Carotenoids,) have an important role in protecting plants against insect pests and these compounds either alone or in combination
with other compound affect insects by being toxic causing death. However, some
insects and tolerate these toxicants depends upon their sensitivity.
Most of the literatures reveal that botanicals directly caused death or
morphological changes or behavioral changes and physiological changes which might
be indirectly due to the metabolic as well as catabolic biochemical changes induced
or modulated by botanicals in general and bioactive principle in particular. Further,
insects have developed a variety of strategies to thwart toxic molecules along with an
appropriate diverse mechanism to face varied encounters in nature/artificial
conditions (1-3). The series of these weapons depends only on the way in which in
built mechanisms have been devised for all sorts of molecules that are effective either
on a specific target or act as general shoot out signals. This kind of mechanism is
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essential to know the effectiveness of botanicals or resistance or susceptibility of

insects. Same concepts have been stressed by many people (4-9). However, very
limited studies have been initiated to evaluate the impact of botanicals on the
biochemistry of insects. This chapter highlights the issue and also provides a brief
account of the same.

2. Biological traits
2.1. Feeding and mortatlity
Feeding is a routine process for the growth and reproduction for maintaining all
live activities, which is inhibited by Vitex negundo (74%), Zelus efficinale (59%),
Azadirachta indica (Meliaceae) (53%). Citus sinensis (48%) treated groundnut shoot
fed Spodoptera litura resulting in loss of body weight by 74, 59, 53 and 48%,
respectively (Sahayaraj, 1998). This might be due to less/slow feeding of foliage and
diversion of energy from production of biomass (body flesh) to detoxification of plant
extract (4). Similarly Isman (10) reported azadirachtin reduced body weight, diet
consumption and utilization. Indirectly these might be due to starvation and slow
metabolic process due to chemical reaction (11) and physiological processes
including digestion and metabolism (1). Stop feeding subsequently to interfere with
all other biological and physiological activities.
The mechanism of action of most of the commercial botanical insecticides is
qualitatively similar to that of synthetic insecticides like DDT and other
organochlorine insecticides. One of the most important traditional as well as
commercial botanicals, pyrethrum products represent 80% of the total market of
global botanical insecticides (10) and are favored by organic growers because of their
low mammalian toxicity and environmental non-persistence making it chief among
the safest insecticides in use (12-14). Pyrethrums are mixed with piperonyl butoxide
(PBO) to increase insect mortality and to extend their shelf life. Further, in flying
insects, pyrethrins cause rapid knockdown effect, hyperactivity and convulsions
which result in neurotoxic action and block voltage-gated sodium channels in nerve
axons (2).
Essential oils are the integral part of integrative pest management (IPM)
components which consist of terpenes, benzene derivatives, hydrocarbons, and other
miscellaneous compounds. Essential oils are lipophilic in natures which generally
interfere with basic metabolic, biochemical, physiological, and behavioral functions
(15, 16) of insects leading to the death of the insect. Further, they interfere with the
neuromodulator octopamine (neurotransmitter, neurohormone, and circulating neurohormoneneuromodulator) (17) or GABA-gated chloride channels (18). Octopamine
disruption results in total breakdown of nervous system in insects, leading to paralysis
and subsequent death. Similarly, Octopamine mimicks like Eugenol increasing
intracellular calcium levels in cloned cells from the brain of Periplaneta americana
and Drosophila melanogaster (17).
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Acetylcholinesterase a key enzyme terminates nerve impulses by catalyzing the

hydrolysis of neurotransmitter, acetylcholine, in the nervous system of various
organisms including insects. This property led to the development of inhibitors of this
enzyme as insecticides, because insecticides covalently bind to the active site and
cause the death of the animal. But, many insects escapes from insecticide poisoning,
because they have an altered acetylcholinesterase which is less sensitive to the active
metabolite of the botanical insecticides and call them resistant insects. We can put
forth many examples: Drosophila strain (MH19), resistant to malathion. Zibaee and
Bandani (19) reported that A. annua extract inhibited the AChE activity in higher
doses. The alteration of AChE was observed in cockroach, Periplaneta americana L.,
at 4 ppm of azadirachtin (AZA) (20) and Senthil Nathan and his co-workers (21)
demonstrated that LC50 concentrations of AZA significantly inhibited the activity of
AChE compared with control.
2.2. Reproduction and embryogenesis
It was reported that methanoprere caused reproductive abnormalities such as
infertility and underdeveloped ovary in Dacus cucurbitae (22). Insects have molyph
serving as a repository for storing nutrients, besides those continuously derived from
digestion of ingested food, essentially comprising primary and secondary metabolism.
Zenobiotics including botanicals enter the diet, then into the haemolymph along with
nutrients. Subsequently they pass through all organs responsible for hormonal imbalance, regulatory and immunological activities along with those to govern all life
time vital functions. Hence, it is imperative to study the impact of plant chemicals on
the chemical composition of the haemolymph, as well as various organs causing
structural changes, as an aspect deserving greater attention.
Typical effects of various botanical applications on various types of insects are as follows:

1. Larvae that are marginally affected populous chimerical, with a mixture of larval
and pupal features (4).
2. Anterior part completely pupated and the posterior part did not moult popularly but
shrank and vice a versa (23).
3. Larvae are inhibition of chitin synergies.
4. Deformed head capsule and wide gap between abdominal segments (24), might be
due to necrotic changes in the CA and NS cells, and also due to retained food
consumption and digestive enzyme activity (25).
5. Deformed pupae with remnants of larval thoracic appendages (24) in Achea janata.
6. The molting phase results in mortality at ecdysis. The newly-formed cuticle lacks
chitin and is pale.
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In literature it was reported that Diflubenzaron (26); diamine furyl-s-triazin (27)

interfere with the deposition and biosynergistic of chitin inhibits molting and leads to
the death of insects. Deformed insects died during their cause of development. This is
a general observation by many authors. It was proposed by Sahayaraj (28) that this
may be due to the inhibitory action of plant toxins on chitin synthesis. Further, it was
explained that if synthesis of chitin is prevented the newly formed cuticle at the time
of pupation would divide chitin leading to the death of the insect (29). However,
Achaea janata Linn. fed with Christella parasitica extracts treated castor leaves,
caused more number of precocious pupa than Ipomea cornea extract treated leaf fed
larvae (24). Authors suggested that presence of phytoecdysones in the former plant
(Family Pterydopophyte) might be the reason. It was endorsed by Sahayaraj and his
co-workers (30) latter and was proposed that increased circulatory ecdysteroids titres
are generally accepted to invite pre-ecdysial events and moulting through hemocytes.
Two chromere derivatives namely, Precocene-I and Precocene- II identified
form the common bedding plant, Ageratum houstonianum. In adult insects, precocene
induces destruction of the corpora allata that results in the deficiency of
gonadotrophic hormone (JH) which, in turn, causes destruction of vitelogenesis and
other successive reproductive process (31-32). But corpora allate are selectively
destroyed by accumulation of metabolites of precocene formed in situ (33). Further,
decreasing amount of yolk protein in the oocytes cause deformed ovary with
disorganized oocytes in red cotton bug treated with Origanum vulgare oil (37).
Azadirachtin has two profound effects on insects (2):
i) at the physiological level; azadirachtin blocks the synthesis and release of molting
hormones (ecdysteroids) from the prothoracic gland, leading to incomplete ecdysis in
immature insects or in adult female insects, a similar mechanism of action leads to
sterility and
ii) azadirachtin is a potent antifeedant to many insects. Toxic principle of Vitex
negundo reduced pupation rate (4), indicating the presence of ecdysteroids (35) in the
plant.
3. Macromolecular profiles
All consumed foods are digested with anabolic, catabolic and metabolic
processes; fundamental and functional units are absorbed and utilized for growth,
development and reproduction via various metabolic pathways. Macromolecular
metabolism, carbohydrates (cellobiose, fructose, galactose, glucose, lactose, maltose,
melibiose, rassinose, sucrose and trehalose), proteins and lipids are generally
considered as macromolecule constituents of both vertebrate and invertebrate
including insects. In addition, macromolecules are synthesized through intermediary
metabolism, having specific storage place and storage form (Table 1). Macromolecule play important role in both catabolic and metabolic process subsequently in
development and reproduction. They are involved in multiple metabolic functions. As
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a result, consumption of food treated with botanicals in general particularly, the

bioactive principles alter the metabolism of carbohydrate, lipid and protein (36, 37).
For instance the results of Shoukry et al. (38) showed that treatment with Piper
cubeba and Salvia officinalis affects carbohydrate, lipid and protein levels in
haemolymph and protein fractions of Plodia interpunctella larvae.

Table 1. Macromolecular storage places, their storage and transportation forms: a general
concept.

Aspects

Macromolecules
Carbohydrate

Storage place
In what form it is
stored
In what form it is
transported
Metabolic form

Protein

Lipid

Fat body and

hemolymh
Glycogen or
trehalose
Trehalose

Hemolymh

Fat body

Imino acid
Imino acid

Fatty body or fat

body
Diacyl glycerol

Glucose

Imino acid

Fatty acid

3.1. Carbohydrates
Carbohydrate constitute an integral part of the exoskeleton of insects and their
role in physiological events such as moulting, metamorphosis, slight and diaphase has
been well documented . Both glycogen and trehalose are important carbohydrates of
insects and hence my researchers considered them for their studies. The concentration
of carbohydrates and other biochemical parameters mainly depend on the quality of
plants and animals in phytophagous and zoophagous insects respectively. Treatment
of second-instar larvae of Muscina stabulans with Cymbopogon citratus and
Rosmarinus officinalis oil induced a significant reduction in the carbohydrate content
during the whole pupal period (39). Abo El-Ghar et al. (37) also showed that
petroleum-ether extract of Ammi majus and Apium graveolens and acetone and
ethanol extracts of Melia azedarach fed with sixth instar larvae of Agrotis ipsilon
greatly reduced haemolymph carbohydrates. Further, similar impacts were also
caused by volatile oils of Lantana camara and Vitex aganus custus too as reported
(40). The authors recorded reduced total body carbohydrate in the larvae of Galleria
mellonella.

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3.1.1. Glycogen
Among the carbohydrates, cellobiose, fructose, galactose, glucose, lactose,
maltose, melibiose, rassinose, sucrose, strehaose; glycogen and trehalose play a very
crucial role in insects. Insects stored their carbohydrate as glycogen. This is utilized
during metamorphosis, as well as energy source. In order to maintain normal life
insects can produce more quantity of glycogen using trehalose. An increase in the
whole body glycogen in Spodoptera litura (41) as well as fat body glycogen in
Lipaphis erysimi (Kalt.) (42) and Achea janata (43) after treatment with plant extracts
(Calotropis gigantia, Vitex negundo and Pongamia glatora) and or botanical
bioactive compounds, Coumarin and methoprene ZR513, respectively has been
reported. This significant glycogen elevation is indicative of increased glycogenolysis
and/or increased body carbohydrate breakdown (41). They also suggested the inter
conversion between glycogen and trehalose in plant extracted Spodoptera litura
larvae.
In contrast, Capparis decidua aqueous fraction potentially reduced the body
glycogen content Sitophilus oryzae Linn (Coleoptera: Curculionidae) (44).This may
be due to i) depletion of glycogen indicating more and more utilization of food
reserves to cope up the botanical induced stress; ii) high release of glucagon,
corticosteroids and catecholamines which stimulates glucose production to combat
energy demand.
3.1.2. Trehalose
Trehalose is a non-reducing disaccharide comprising two glucose molecules;
present in high concentration as the main haemolymph sugar in insects. It is a
common, typical and major carbohydrate of insects, especially in the larval stage. Its
concentration depends on its rate of synthesis and utilization. Groundnut leaves
impregnated with plant extracts of A. indica, C, gigantia, P. pinnata and V. negundo
feeding topical application and contact toxicity against S. litura treahalose content
was investigated (45). Further, A. indica treatment reduced the trehalose content more
than V. negundo (65%), P. glatora (63%) and C. gigantia (47%) nearly 79%. This
might be due to azadiractin, ecdysteroids, pungamin and cardinalids respectively.
Perhaps such midway interfere in the nutrional ecology of the developing S. litura
larvae modulate the neuroendocrine system to inhibit appropriate hormone trehalogen
leading to hypotrehalogenic conditions (45). Similarly, Hayakawa et al. (1988)
observed decreased trehalose quantity in taurina treated category. However, after 96
hrs like prepulal stage, trehalose content increased to meet the energy request for
pupal formation of S. litura. The concept was also supported by Hiranoan Yamashita
(1984), who reported that the Trehalase was not only utilized for the development but
also essential for the transformation of larvae to pupae and also from pupae to adult.

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3.2. Proteins
Insect fat body is an organ analogous with mammalian liver, which performs
verity of different metabolic activities. Further, it is the place of intense biosynthetic
activities. Throughout the insect life it is the main source for the haemolymph protein.
Haemolymph proteins play an important role in insects for transport functions as well
as their enzyme action. Proteins are an integral part of the cuticle; haemolymph ant
plays an important role in metamorphosis and insect growth. Sahayaraj and Kalidas
(46) show that Padina pavonica (Linn.) (Phaeophyceae) extract significantly reduced
the total body protein of Dysdercus cingulatus (Fab.). Initially, it was found that
treatment of high doses of azadirachtin to the last larval instar of Epilachna varivestis
arrested the transformation of larvae to pupa (47). Later, it was also reported that seed
extract of Annona (48, 49), leaves extract of Lantana wightiana, Premna tomentosa
and Synedrella nodiflora (50) reduced the total body protein content of S. litura.
Similarly, Porterresia coarctata tackeeka leaf extract at different concentrations
showed significant reduction in protein and DNA content in the fat body and midgut
tissue (51). Larvicidal activity is mainly due to the reduction of protein content (52).
Observations revealed lower hemolymh protein, which varied with plant extracts.
Less protein in haemolymph could be attributed to reduction of protein synthesis in
plant extracts treated insects by deranging protein synthesis machinery. This could be
attributed to an adaptation of insect to overcome the phytochemical stress. Further,
changes in protein content probably reflect the balance between synthesis, storage,
transport and degradation of structural and functional nutrients during ontogeny as
well as response to particular physiological conditions. Incoming is due to either low
food intake or reduction in protein synthesis of higher mobilization of proteins.
Similar impact of azadirachtin and plumbagin was also reported by in Spodoptera
litura (53) and also in Helicoverpa armigera (54) and Corcyra (55) respectively. Both
topical application and oral application of Bryopsis plumose (Huds.) a marine algae
extract reduced the quality and quantity of the protein profile of hemolymh in
Hyblaea puera (Cramer) (Lepidoptera: Hyblaeidae) (56). Similar impact was also
caused by plant oils (57) in H. armigera.
In contrast, several reports show that botanicals increased the protein levels may
be due to the conversion of carbohydrates and lipids to proteins: Kinnear and
Thomson (58) suggested that an increased protein level was due to increased
synthesis of new proteins by the fat body, haemolymph and other tissues of the
larvae. Further, increased protein shows appearance of new peptide in hemolymh
upon botanical treatment, may liberate free radicals which affect nitrogenous
compounds directly; this in turn leads to breakdown of the peptide linkage, causing
fragmentation of protein molecules (Fig.1).
Amino acids are important constituents of insect body. These amino acids occur
as free amino acids having high concentration in insect hemolymh and derived to play
an important role in osmoregulation, energy production for fight, cocoon conjunction
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etc., act as units for protein synthesis etc. In Corcyra cephalonica azadiractin
decreased the total free aminoacids (TFA) level after 24 hrs of treatment the initial
decrease is due to increased neuromuscular activity of treated insect which results in
higher demands for energy. But at 48, 72 and 96 hrs TFA level increased (59).

Fig.1 Proposed hypothetical activities of botanical impacts on protein anabolic and catabolic
activities in insects

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3.3. Lipids
In addition to carbohydrate, lipids are of vital importance to many insects as
substrates for embryogenesis, metamorphosis and flight. Hazaricka and Baishya (22)
studied the impact of methoprene and azadirachtin on the lipid contents of insects.
Mostafa (36) reported that the total lipid content was significantly increased in
Trogoderma granarium when treated with plant extracts perhaps due to high
deposition of lipid together with low lipid utilization; they added that lipid
accumulation was more likely to be related directly to a lack of juvenile hormone.
Later, Abou El-Ela (60) also recorded similar result on Musca domestica after
treatment with water extracts of some plants. However, Shoukry and Hussein (40)
showed that treatment of third-instar larvae of Galleria mellonella with sublethal
concentrations of Lantana camara and Vitex agnus castus reduced the total lipids in
the last larval instar. Cholesterol and phospholipids are the important lipid
compounds of insects (61).
3.3.1. Total Cholesterols
The quality of total cholesterol was adversely affected by the water extracts of
Azadiracta indica, Pongamia globra, Vitex negundo, Calotropis gigantia (41),
probably suggesting the depletion of the reserve fats in Spodoptera litura under the
influence of bioactive substances present in the tested plants. Among the four plants,
V. negundo adversely (69%) affects cholesterol level, while it treated orally whereas
P. glabra affects when treated either topically (76%) or as contact (65%) toxicant
against S. litura. Further, the authors added, the diastic changes in the cholesterol
level in S. litura during the plant extracts treatment without being accompanied by
any changes in cholesterol level of haemolymph such as metabolic fluid or its
precensors, ecdysteroids. In general, the metabolism of tissue macromolecules is
known to be main physiological activity to cause haemolymph macromolecular level.
Interchange as well as confusion takes place between haemolymph and organs. Babu
et al. (62) reported azadirachtin reduced haemolymph lipid profile during gonadotropic period of Atractomorpha crenulata Fab. (Orthoptera: Acarididae). Similar
effects were also observed in methoprene and diflubenzeuon treated Dicladispa
armigera (63).
3.3.2. Phospholipids
Vitex negundo increased (54%) phospholipid content of S. litura more than
A. indica (51%) did (41). This increased level is due to decreased trehalose level and
the affected S. litura larvae took their energy from lipid sources for their activities. In
contrast, azadirachtin and ecdystone analogus (metheprene and diflubenzenation)
adversely affect subsequently reduce fat content of Altractomorpha creanulate (62)
and Diclodispa armigera (63).

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4. Electrolytics
Calcium ion plays an important role in the regulation of muscle contraction, cell
motility and activity of the nervous system. Tephrosia purpurea (Linn.) (Fabaceae)
and Acalypha indica (Linn.) (Euphorbiaceae) crude extract impacts on the
reproductive organs electrolytes level in alimentary canal and detoxication enzyme
level in the fat body and intestine of Dysdercus cingulatus (Fab.) (Pyrrhocoridae) was
studied by Sahayaraj and Shoba (64). The authors study was the first in a series
aiming to record the mineral elements level in the red cotton bug in relation to crude
plant extracts ingestion through seed or leaves of the cotton plant. Mineral elements
are found in most plant-feeding insects because they are constituents of plants.
Sodium, potassium and calcium level was high in cotton seed and leaves respectively
fed pest reveals that part of the plant determines the level of mineral elements in
insects. In T. purpurea extracts treated cotton seed fed insect, the sodium level
significantly blacked, at the same time while the extract incorporated in to the cotton
leaves, sodium level slightly increased, because the bioactive of this plant selectively
block the sodium channels in the slow inactivated state. An opposite trend was
observed for A. indica extracts. Type of feed such as cotton seed and leaves also
influence the electrolyte level of D. cingulatus. For instance, both Na+, Cl level was
higher while D. cingulatus was fed with cotton seed. Similarly K+ and Ca2+ level was
higher when D. cingulatus was fed with cotton leaves. But incorporation of A. indica
and T. purpurea bioactive principles further reduced these electrolytes. It shows that
tested plants interfere with the physiology of D. cingulatus.
5. Genomic DNA and RNA
Chloroform extracts of Padina pavonica reduce whole body DNA content (30%)
of Dysdercus cingulatus (46). Similarly, it was observed that the seaweed Sargassum
tenerrimum extracts, and chromatographic fractions reduce genomic DNA content of
D. cingulatus along with the reduction of total body protein (65). In another study,
Capparis decidua hexane extract 55% reduced the DNA content and acetone extract
reduced 70% RNA content in Sitophilus oryzae (44) which is due to inhibition of
nucleic acid synthesis at cellular level and catabolism get increased which results in
low availability of nucleic acids.
6. Energy
A flavanoids rotenone / rotenoids extracted plants belonging to the roots of
Derris, Lonchocarpus, Tephrosia is a mitochondrial poison, which blocks electron
transport chain and prevents energy production (66).This blocking phosphorylation of
ADP to ATP stand inhibiting insect metabolism.

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7. Digestive Enzymes
7.1. Digestive enzymes
Insect midgut exhibited the activity of amylase, invertase, maltase and proteinase
in its ventricular region and thus possessed the capacity to digest starch, sucrose,
maltose and protein. The utilization of nutrients viz., carbohydrates, proteins and
lipids from available food plants depends on the digestive enzymes. They play a
major role in the body of insects by converting complex food materials into
micromolecules necessary to provide energy and metabolites for growth,
development and other vital functions. Botanicals in general, inhibit digestive
enzymes of insects (67).
7.1.1. Amylase
Amylase is one of the key enzymes involved in digestion of carbohydrate
metabolism in insects. In 1987, Saleem and Shakoori reported that sub-lethal
concentrations of pyrethroids decreased the gut a-amylase activity of Tribolium
castaneum Herbst (Coleoptera: Tenebrionidae) (68). Similar impact was also reported
(8) in elm leaf beetle treated by A. annua extract and also in Eurygaster integriceps
Puton (Hemiptera: Scutelleridae) due to Artemisia annua extract treatment (67).
Hemionitis arifolia (Brun) Tmore reduced the amylase activity more than A. indica in
S. litura (69) did. Similar reduction in alkaline phosphate activity was also observed
in S. litura (53). Babu et al. (70) also recorded similar results in Helicoverpa armigera
(Hubner) treated with azadirachtin. Sahayaraj and Antony (69) concluded that the
reduction in amylase as well as Alkaline phosphatase (AP) activities as a result of P.
glabora and Hemionitis arifolia treatment might be due to the rupture of epithelial
cells, which are the sites for enzyme secretion. Further, they clearly reported that A.
indica does not cause such a kind of alimentary canal disruption. Indigestion leads to
starvation and subsequent death.
7.1.2. Glycosidases
Generally, after amylase, glycosidases digest carbohydrate oligomers into
monosaccharides (71, 72). On the other hand, glycosidases catalyze the hydrolysis of
terminal, non-reducing 1, 4-linked alpha-D-glucose residues with releasing of alphaD-glucose. Treating the E. integriceps adults (67) and gypsy moth (Lymantriidae) and
forest tent caterpillar (Lasiocampidae) (73, 74) with different concentrations of A.
annua extract and phenolic components respectively howed the reduction in the
activity of glucosidases.
7.1.3. Alkaline phosphatase
ALP is primarily found in the intestinal epithelium of animals and its major
function is to provide phosphate ions from mononucleotide and ribonucleoproteins
for a variety of metabolic processes. Alkaline phosphatase (ALP) play a role in the
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maintenance of energy supply of the living cells as they catalyze the phosphate esters
and release the high energy phosphate bonds which are utilized for reveal metabolic
process. Pongamia globra aerial part extracts highly reduced AP activity of S. litura
than Crystella parasitica (69). It is suggested by the authors that the reduction in
nutritional measures might be the influential factor for the reduction of enzyme
activity in the midgut. Along with alkaline phosphatase and acid phosphatase are the
hydrolytic enzymes, which hydrolyze phosphomonoesters under alkaline or acid
conditions, respectively.
ALP is involved in the transphosphorylation reaction and the midgut has the
highest ALP and ACP activity as compared to other tissues (75). The overall activity
of ALP and ACP decreased due to increasing of plant extract concentrations so that
there were significant differences among control and three treatments. These findings
coincided with other reports of plant extract treatments of insects. For example,
Senthil Nathan (76) showed that treatment of rice plants with Melia azedarach Juss
(Meliaceae) extracts decreased the activity level of ALP in Cnaphalcrocis medinalis
(Guenee). In another study, it was reported that feeding Spodoptera litura Fabricius
(Lepidoptera: Noctuidae) on Ricinus communis L. treated with azadirachtin decreases
the amount of this enzyme in the midgut (9).
7.1.4. Lipases
Very limited information is available about activity of digestive lipases in
relation to botanicals. Senthil Nathan et al. (76) reported that treating Cnaphalocrocis
medinalis (Guenee) (Lepidoptera: Pyralidae) with azadirachtin and other neem
components sharply decreased the activity level of lipase in the midgut. Similar
observation was reported in the midgut of Chilo suppressalis Walker (Lepidoptera:
Pyralidae) (71) and E. integriceps (67) due to A. annua extract treatment.
7.1.5. Proteases
Proteases sub-classes like serine, cysteine, and aspartic proteinases have a
crucial role in food digestion by insects (77). Studies by Johnson et al. (78),
Senthil-Nathan et al. (76) and Zibaee and Bandani (67) inferred that botanical
insecticides interfere with the production of certain types of proteases and disable
them to digest ingested proteins. It was investigated that either alone or in
combination A. annua and Lavandula stoechas decreased the digestive enzyme
activity except for protease and lipase of Hyphantria cunea Drury (Lepidoptera:
Arctiidae) (79). The Km is the measurement of the stability of the enzyme-substrate
complex and a high Km would indicate weak binding while a low Km would indicate
strong binding (80). Zibaee and Bandani (67) argued that A. annua extract increased
the value of Km in E. integriceps. Further, it was observed that plant extracts can bind
to the enzyme at the same time as the enzyme binds to the substrate, and this binding
affects the binding of the substrate and vice versa (79, 80). Zibaee and Bandani (79)
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also recorded negative effect of A. annua extract on nodule formation and

phenoloxidase activity of E. integriceps. High activity of midgut protease indicates a
great utilization of exogenous proteins, more physiological activity to perform and
development.
7.2. Detoxification enzymes
Insect herbivores can increase their detoxification activities against a particular
plant poison/toxin in response to prolonged or short ingestion of the same
compounds. Studies reveal that esterase, oxidase, transoxidase hydrolases, glutothion
transferase, cis-oxidase hydrolases, adrin epoxidase, cytochrome P-450, alanine
aminotransferase (ALAT), asparate aminoteransferase (ALAT), glutathione
S-transferase (GST); glutathione P- transferase (GPT) role in phase II of enzyme
detoxification. Further, for the first time Upadhyay (44) reported that acid and
alkaline phosphatase have been studied as enzymes significant in detoxification.
Zibaee et al. (71) proposed that the amino transferases are important components of
amino acid catabolism; which is involved in transferring an amino group from one
amino acid to a keto acid. Further, Etebari et al. (81) reported that both the AST and
the ALT serve as a strategic link between the carbohydrates and protein metabolism
and are known to be altered during various physiological and pathological conditions.
It is well established that the GSTs play a key role in detoxication in insects. Dugrano
et al. (82) reported that Allium porrum highly induced the activity of GST in
Callosobruchus maculatus Fab. adults and Acrolepiopsis assectella Zellee larvae.
However, Sahayaraj and Antony (69) observed an opposite trend in S. litura
indicating that the insect is susceptible to the plant extracts they studied. Recent
reports says, Tephrosia purpurea (Linn.) seed extract induce the production of AAT
and ALAT, both in fat body and intestine of Dysdercus cingulatus (64), indicating the
turnover of amino acids and glutamate formation during metamorphosis in red cotton
bug. Further, they proposed the inter conversion between glycogens and trehalose in
plant extracts treatment. Previously it was studied the impact of ammonia, an
important chemical constituent of Annona squemosa, on AAT and ALAT levels of
S. litura (83). Proteins also play an important role in detoxication process by
synthesizing microsomal detoxifying enzyme which assists in detoxification (84).
Esterase activity was altered by neem extract (85). The responses of EST to
botanical insecticides were significantly due to using different concentrations of
extract and long exposure. In the early stage, plant extract stimulated the expression
of EST body to increase the detoxification ability (19). In the late stage, because of a
toxic effect and time EST activity was suppressed. Grant and Matsumura (86)
proposed that Glutathione S-transferases (GST) are the mainly cytosolic enzymes
which catalyze the conjugation of electrophile molecules with reduced glutathione
(GSH), potentially toxic substances become more water soluble and generally less
toxic. Further, it plays an important role in insecticide resistance and is involved in
the metabolism of organophosphorus and organochlorine compounds (72). In
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addition to organophosphorus and organochlorine compounds, other xenobiotics such

as plant defense allelochemicals against phytophagous insects induce GST activity
(86, 87). By treating Artemisia annua L. (Asteracea) extracts on E. integriceps adults
(67) the activity level of GST in 24 h post treatment increased significantly for both
substrates (CDNB, DCNB) of the enzyme. Its (two or one) activity was dosedependent and increased by exposuring higher concentration of plant extract. The
influence of plant allelochemicals on GST activity was not limited to the herbivores
and was observed in several predators too (87).
8. Conclusions and future direction
Plant insect and phytopathogens protection is an important issue for the
agricultural community. Pesticides have been used so far which leads to biotic and
abiotic impacts, hence an alternative method of biointensive integrated pest
management (BIPM) where botanical insecticides play an important role. In order to
avoid multitude impacts of botanicals crude extracts or their column chromatographic
factions or bioactive principles, insects have developed a wide variety of defensive
mechanisms which are not studied deeply so far. Botanical induced or altered
macromolecules, micromoleucles including electrolytes, genomic DNA, energy giving
molecules, digestive and detoxication enzymes analyses would be essential for a better
understanding of the potential molecular mechanism of the response to various
phytophagous insects. Hence this review provides an insight into molecular
interference of the botanicals. In future more quantity of work is necessary to utilize
many screened plants to bring them in to formulation which can be made available at a
low cost to our farming community.
Acknowledgements
I acknowledge Rev. Dr. V. Gilbert Camillus, S.J. for the support and encouragements. I also express my sincere thanks to DST, Delhi (HR/OY/Z-13/96; SP/SO/C51/99) and MoEs, New Delhi (F.No. 14/23/2008-ERS/RE) for the grant support.

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Zibaee, A. and Bandani, A.R. (2010) Effects of Artemisia annua L. (Asteracea) on digestive enzymes profiles and cellular immune
reactions of sunn pest, Eurygaster integriceps (Heteroptera: Scutellaridae), against Beauvaria bassiana. Bulletin of Entomological
Res. 100: 185-196.
Stryer L. (1995) Biochemistry. W.H. Freeman and Company, New York, pp. 415.
Etebari, K., Mirhodeini, S.Z. and Matindoost, L. (2005) A study on intraspecific biodiversity of eight groups of silkworm (Bombyx
mori) by biochemical markers. Insect Sci. 12: 87-94.
Dugravot, S., Thibout, E., AboGhalia, A., and Huignard, J. (2004) How a specialist and a nonspecialist insect cope with dimethyl
disulfide produced by Allium porrum. Entomologia Experimentalis et Applicata, 113(3): 173-179.
Yerasi, B., and Chitra, K.C. (2000) Response of aspartate and alanine aminotransferase to sublethal concentration of annonain by
Spodoptera litura (Fab.). Indian J. Entomol. 62(4): 367-370.
Wilkinson, C.F. (1976). Insecticide synergism, pp. 195-218. In R.L. Metcalf and J.J. McKelvey, Jr. [Eds.], Insecticides for the
Future: Needs and Prospects. John Wiley and Sons, New York.
Naqvi, S.H. (1986) Biological evaluation of fresh neem extracts and some esterase activity in insects. Proc, 3rd Int. Neem Conf.
Nairobi. 315-330.
Grant, D.F., and Matsumura, F. (1989) Glutathione-S-transferase 1 and 2 in susceptible and insecticide resistant Aedes
aegypti. Pesticide Biochem. and Physiol. 33(2): 132-143.
Francis, F., Vanhaelen, N. and Haubruge, E. (2005) Glutathione S-transferases in the adaptation to plant secondary metabolites in
the Myzus persicae aphid. Arch. Insect Biochem. Physiol. 58: 166174.

Article History:

Received on 10th July 2013; Revised on 15th May 2014 ; Accepted on 10th June 2014 and
Published on 30th October2014.

Printed in the Unitated States of America, 2014

ISBN: 978-1-63315-205-2

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Table Contents

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Page No.

Preface
Forward message
Contributors
Reviewers
Acknolwedgement

i
ii
iii
iv
v

Volume1
Section I: Insect Biochemical approaches

1. Introduction to Insect Molecular Biology.

Raman Chandrasekar, P.G., Brintha, Enoch Y.Park, Paolo Pelsoi, Fei Liu,
Marian Goldsmith, Anthony Ejiofor, B.R., Pittendrigh, Y.S., Han,
Fernando G. Noriega, Manickam Sugumaran, B.K., Tyagi, Zhong Zheng Gui,
Fang Zhu, Bharath Bhusan Patnaik, and P. Michailova

2.

Modulation of Botanicals on pests biochemistry.

57

Sahayaraj, K.

3.

Detoxication, stress and immune responses in insect antenna:

new insights from transcriptomics.

75

David Siaussat, Thomas Chertemps and Martine Maibeche

4.

Application of isotopically labeled compounds and tandem mass

spectrometry for studying metabolic pathways in mosquitoes.

99

Stacy Mazzalupo and PatriciaY.Scaraffia

5.

Field Response of Dendroctonus armandi Tsai & Li (Coleoptera:

Scolytinae) to Synthetic Semiochemicals in Shaanxi, China.

127

Shou-An Xie, Shu-Jie L.V., Hui-Chen, Raman Chandrasekar

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Section II: Insect Growth

6. Insect Cuticular SclerotizationHardening Mechanisms and Enzymes.

149

Manickam Sugumaran

7. New Approaches to Study Juvenile Hormone Biosynthesis in Insects.

185

Crisalejandra Rivera-Perez, Marcela Nouzova and Fernando G. Noriega

8. The regulatory biosynthetic pathway of juvenile hormone.

217

Zhentao Sheng and Raman Chandrasekar

Section III:

Insect Immunity

9. The innate immune network in a hemimetabolous insect, the brown

planthopper, Nilaparvata lugens.

233

Yanyuan Bao, Raman Chandrasekar, Chuan-Xi Zhang

10. Immune Pathways in Anopheles gambiae.

253

Maria L. Simes and Raman Chandrasekar

11. Key biochemical markers in silkworms challenged with immuno-

271

elicitors and their association in genetic resistance for survival.

Somasundaram, P., Chandraskear, R., Kumar,K.A., and Manjula, A.

Section IV:

Insect Molecular Genetics

12. The recent progress of the W and Z chromosome studies of the

291

silkworm, Bombyx mori

Hiroaki Abe, Tsuguru Fujii and Raman Chandrasekar

13. Molecular characterization and DNA barcoding for identification of

317

agriculturally important insects.

Rakshit Ojha, Jalali, S.K., and Venkatesan, T.

14. Polytene chromosomes and their significance for Taxonomy,

331

Speciation and Genotoxicology

Paraskeva V. Michailova

15. Insect exuvium extracted DNA marker: a good complementary

molecular taxonomic characteristics with special reference
to mosquitoes.

355

Dhanenjeyan, K. J., Paramasivam, R., Thanmozhi, V., Chandrasekar,R., and Tyagi, B.K.
Index

363

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Volume2
Section V:

Molecular Biology of Insect Pheromones

16. Understanding the functions of sex-peptide receptors?

373

Orly Hanin, Ada Rafaeli

17. Current views on the function and evolution of olfactory receptors

385

in Lepidoptera.

Arthur de Fouchier, Nicolas Montagn, Olivier Mirabeau, Emmanuelle Jacquin-Joly

18. Molecular architecture, phylogeny and biogeography of pheromone

409

biosynthesis and reception genes / proteins in Lepidoptera.

Jian-Cheng Chang, P. Malini, R. Srinivasan

Section VI:

Insect Molecular Biology

19. Application of Nanoparticles in sustainable Agriculture :

429

Its Current Status.

Atanu Bhattacharyya , Raman Chandrasekar, Asit Kumar Chandra,

Timothy T. Epidi and Prakasham, R.S.

20. Mosquito Ribonucleotide Reductase: A Site for Control.

449

Daphne Q.-D. Pham, Victor H. Perez, Lissette Velasquez, Dharty Bhakta,

Erica L. Berzin, Guoli Zhou, and Joy. J. Winzerling.

21. Green protocol for synthesis of metal nanoparticles

to control insect pests.

473

Murugan, K., Chandrasekar, R., Panneerselvam, C., Naresh Kumar, A.,

Madhiyazhagan, P., Mahesh Kumar, P., Jiang-Shiou Hwang, Jiang Wei

22. Aquaporins in Blood-Feeding Arthropods.

497

Lisa L. Drake, Hitoshi Tsujimoto, Immo A. Hansen

23. Mimetic analogs of three insect neuropeptide classes

509

for pest management.

Ronald J. Nachman

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Section VII:

Insect Pest Management through

Biochemical and Molecular approaches

24. Induced resistance in plants against insect pests and

533

counter-adaptation by insect pests.

Abdul Rashid War and Hari C Sharma

25. Insect Chemical communication - an important component of

549

novel approaches to insect pest management.

Usha Rani, P.

26. Mosquito control using biological larvicides: Current Scenario.

575

Subbiah Poopathi, C. Mani and R. Chandrasekar

27. Application of RNAi toward insecticide resistance management.

595

Fang Zhu, Yingjun Cui, Douglas B. Walsh, Laura C. Lavine

Section VIII:

Insect Bioinformatics

28. Entomo-informatics: A prelude to the concepts in Bioinformatics.

621

Habeeb, S.K.M. and Raman Chandrasekar

29. Molecular expression and structure-function relationships of

633

apolipophorin III in insects with special reference to innate immunity.

Bharat Bhusan Patnaik, Raman Chandrasekar, Yeon Soo Han

30. Computer-aided pesticide design: A short view

685

Jitrayut Jitonnom

Index

709

xx

ISBN No. 978-1-63315-205-2 (USA)

First Edition: Volume 1, 2 October 2014

Total No. Pages: 398 + 372 = 770

Edited by Raman Chandrasekar

B.K. Tyagi
Zhong Zheng Gui
Gerald R. Reeck
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Book Mission Project # 2: Initiated on June 2010; Completed on March 2014 and Published on Oct. 2014.

Volume 1 & 2, October 2014

Short Views on Insect Biochemistry

and Molecular Biology

PREFACE
Entomology as a science of inter-depended branches like biochemistry, molecular entomology, insect
biotechnology; has made rapid progress in its attributes in the light of modern discoveries. This also
implies that there is an urgent need to manage the available resources scientifically for the good of man.
In the past five decades, entomology in the world/country has taken giant steps ahead. Continued
research has evolved better pest management through molecular approaches. The aim of the Short
Views on Insect Biochemistry and Molecular Biology book is to integrate perspectives across
biochemistry and molecular biology, physiology, immunology, molecular evolution, genetics,
developmental biology and reproduction of insects. This century is proclaimed as the Era of
Biotechnology and its consists of all types of Mol-Bio applications, which is an essential component for
a through understanding of the Insect Biology. This volume 1 & 2 (8 section with 30 chapters)
establishes a thorough understanding of physiological and biochemical functions of proteins, genes in
insects life processes; the topics dealt with in the individual chapters include chemistry of the insect
cuticle, hormone and growth regulators; biochemical defenses of insects; the biochemistry of the toxic
and detoxification action; modern molecular genetics and evolution; inter- and intra-specific chemical
communication and behavior; insect pheromone and molecular architecture, phylogeny and chemical
control of insect by using insect pheromones biotechnology; insect modern biology and novel plant
chemical and microbial insecticides for insect control, followed by a discussion of the various
mechanisms of resistance (both behavioral and physiological) and resistance management; modern insect
pest management through biochemical and molecular approaches; Mimetic analogs of insect
neuropeptide for pest management; entomo-informatics and computer-aided pesticide designing. In short
this book provides comprehensive reviews of recent research from various geographic areas around the
world and contributing authors area recognized experts (leading entomologist/scientist) in their
respective filed of molecular entomology. We will miss this collaboration now it has ended, but will feel
rewarded if this book is appreciated by our team/colleagues and remarkable mile stone in entomology
field.
This book emphasizes upon the need for and relevance of studying molecular aspects of entomology in
Universities, Agricultural Universities and other centers of molecular research. To encompass this
knowledge and, particularly disseminate it to the scientific community free of cost, was the major
inspiring force behind the launch of Short Views on Insect Biochemistry and Molecular Biology.

Editors

Raman Chandrasekar
Brij Kishore Tyagi

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ShortViewson

InsectBiochemistryand
MolecularBiology
Editedby
Raman Chandrasekar, Ph.D.,
Kansas State University, USA.
B.K.Tyagi, Ph.D.,
Centre for Research in Medical Entomology (ICMR), India.
Zhong Zheng Gui, Ph.D.,
Jiangsu University of Science and Technology,
Sericultural Research Institute, Chinese Academy of
Agricultural Sciences, China.
Gerald R. Reeck, Ph.D.,
Kansas State University, USA.

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Contributing Authors

Dr. B.K.Tyagi

Prof.Fernando G. Noriega

Centre for Research in Medical Entomology,

4Sarojini Street, Chinna Chokkikulam,
Madurai 625002 (TN), India

Department of Biological Sciences

HLS 227, Florida International University
11200 SW 8th St, Miami, FL 33199, USA.

Prof. Gui Zhongzheng

Dr. Zhentao Sheng

Sericultural Research Institute,

Chinese Academy of Agricultural
Sciences, Zhenjiang, 212018,
Jiangsu, P. R. China.

Chicogo University, Chicogo, USA.

Prof. K. Sahayaraj

Prof.Yanyuan Bao
Institute of Insect Science,
Zhejiang University, China.

Dept. of Advanced Zoology and Biotechnology,

St. Xavier's College
Palayamkottai 627 002, Tamil Nadu, India.

Prof. Chuan-Xi Zhang,

Prof. David Siaussat

Dr. Maria L. Simes

Universit Pierre et Marie Curie (Paris 6/UPMC),

UMR 1272A Physiologie de l'Insecte:
Signalisation et Communication (PISC),
7 Quai Saint Bernard, Batiment A - 4me tage bureau 410, 75252 Paris Cedex 05, France.

Prof. PatriciaY.Scaraffia
Department of Tropical Medicine,
Tulane University, New Orleans,
LA 70112, USA.

Prof. Shou-An Xie

Institute of Insect Science,

Zhejiang University, China.

UEI Parasitologia Mdica,

Centro de Malria e Outras Doenas Tropicais,
Instituto de Higiene e Medicina Tropical,
Rua da Junqueira 96, 1300 Lisboa,
Portugal.

Dr. P. Somasundaram
Central Sericultural Germplasm Resources Centre,
P.B.No.44, Thally Road,
Hosur-635109,
Tamilnadu, India.

College of Forestry,
Northwest A & F University
Yangling, Shaanxi 712100, China

Dr. Hiroaki Abe

Dr. Raman Chandrasekar

Dr. S.K. Jalali

Department of Biochemistry and Molecular

Biophysics, Kanas State University,
Manhattan, 66506, KS, USA.

Prof. Gerald R. Reeck

Department of Biochem. and Molecular
Biophyscis, Kansas State University, KS, USA.

Prof. Manickam Sugumaran

Department of Biology
University of Massachusetts Boston
100 Morrissey Blvd,
Boston, MA 02125, USA.

Tokyo University of Agriculture and Technology,

Japan.

National Bureau of Agriculturally Important

Insects, ICAR, India.

Prof. Paraskeva V. Michailova

Institute of Biodiversity and
Ecosystem Research,
1 Tzar Osvoboditel boulv
Bulgarian Academy of Sciences
Sofia 1000, Bulgaria.

Prof. Ada Rafaeli

Associate Director for Academic Affairs &
International Cooperation
Agricultural Research Organization,
The Volcani Center, P. O. Box 6,
Bet Dagan 50250, Iseral.

ix

Prof. Emmanuelle Jacquin-Joly

Dr. Fei Liu

UMR PISC Physiologie de l'insecte

INRA, Route de Saint-Cyr
78026 Versailles cedex, France..

Department of Biological Science & Technol.,

Shaanxi Xueqian Normal University,
Shaanxi, China.

Dr. R. Srinivasan

Prof. Marian Goldsmith

Entomologist and Head of Entomology Group

AVRDC-The World Vegetable Center
60 Yi Ming Liao, Shanhua
Tainan 74151, Taiwan.

Biological Sciences Department,

University of Rhode Island,
Kingston, RI 02881, USA

Prof. Atanu Bhattacharyya

Prof. Anthony Ejiofor

Vidyasagar College for Women,

Post Graduate Department of Environmental
Science,
University of Kolkata, India.

Department of Biological Sciences,

College of Agriculture, Human & Natural
Sciences, Tennessee State University,
3500 John A Merritt Blvd., Nashville,
Tennessee 37209, USA.

Prof. Daphne Q.-D. Pham

Dr. Bharath Bhusan Patnaik

Dept of Biological Sciences,

University of Wisconsin-Parkside,
900 Wood Road, Kensoha,
WI 53144, USA.

School of Biotechnology,
Trident Academy of Creative Technology
(TACT), Bhubaneswar 751013 Odisha, India.

Prof. Jitrayut Jitonnom

Prof. B.R. Pittendrigh

School of Science
University of Phayao, Thailand.

Department of Entomology,
University of Illinois, Urbana-Champaign, IL,
61801, USA
.

Prof. K. Murugan

Dr. Subbiah Poopathi

Department of Zoology, School of Life Sciences,

Bharathiar University,
Coimbatore - 641 046, India.

Prof. Immo A. Hansen

Department of Biology,
New Mexico State University,
Las Cruces, NM, USA.

Dr. Ronald J. Nachman

USDA-ARS,
Food Animal Protection Research Laboratory,
USA.

Dr. Hari C Sharma

International Crops Research Institute for the
Semi-Arid Tropics (ICRISAT), Patancheru502324,
Andhra Pradesh, India.

Prof. Paolo Pelsoi

State Key Laboratory for Biology Plant Diseases
and Insect Pests, Institute of Plant Protection,
Chinease Academy of Agricultural Sciences,
Bejing, China.

Unit of Microbiology and Immunology,

Vector Control Research Centre
(Indian Council of Medical Research),
Medical complex, Indira Nagar,
Puducherry 60 5006, India.

Dr. P.Usha Rani

Biology and Biotechnology Division
Indian Institute of Chemical Technology
(CSIR)Taranaka,
Hyderabad - 500 007 (AP), India.

Dr. Fang Zhu

Irrigated Agriculture Research and Extension
Center, Dept.of Entomology,
Washington State University,
Prosser, WA, USA.

Prof. S.K.M. Habeeb

Department of Bioinformatics,
Faculty of Engineering & Technology,
SRM University, Kattankulathur,
Chennai 603203, Tamilnadu, India.

Prof. Yeon Soo Han

Division of Plant Biotechnology,
College of Agriculture & Life Science,
Chonnam National University,
Gwangju 500-757, South Korea

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Reviewer & External supportive members

Prof. Michael Riehle, Department of Entomology, University of Arizona, USA.

Dr. Dawn L.Geiser, College of Agriculture and Life Sciences, University of Arizona, USA.
Prof. Young Jung Kwon, School of Applied Biosci., Kyungpook National University, South Korea.
Dr. Kaliappandar Nellaiappan, CuriRx Inc. USA.
Prof. Patricia Y. Scaraffia, Department of Tropical Medicine, Tulane University, USA.
Prof. Richard Newcomb, Plant & Food Research, University of Auckland, New Zealand.
Dr. S. Krishnaswamy, School of Biotechnology, Madurai Kamaraj University, South India.
Dr. Mary-Anne Hartley, University of Lausanne, Switzerland.
Dr. Igor F. Zhimulev, Institute of Molecular and Cellular Biology, Novosibirsk, Russia.
Dr. S. Subramanin, Indian Agricultural Research Institute. India.
Prof. Gustavo F. Martins, Departament de Biologia Geral, Universidade Federal de Vicosa, Brazil.
Prof. Helena Janols, Infektionsklinien, Skanes Universitetsisjukhus, Sweden.
Prof. Donald R.Barnard, USDA, Agricultural Research Service, CMAVE, USA.
Dr. Keith White, Faculty of Life Science, University of Manchester, UK.
Prof. Marten J.Edwards, Biology Department, Muhlenberg College, USA.
Prof. E. Warchalowska-Sliwa, Polish Academy of Sciences, Poland.
Dr. K. Balakrishnan, Department of Immunology, Madurai Kamaraj University, India.
Dr. J.Joe Hull, USAD-ARS, Arid Land Agricultural Research Centre, USA.
Dr. Neil Audsley, The Food & Environment Research Agency, UK.
Dr. Raman Chandrasekar, Kansas State University, USA.
Dr. B.K. Tyagi, Centre for Research in Medical Entomology (ICMR), Madurai, TN, India.
Prof. Zhongzheng Gui, Sericulture Research Institute, Chinese Academy of Agricultural Sci., China.
Dr. Fang Zhu, Irrigated Agril. Research and Extension Center, Washington State University, USA.
Prof. K. Murugan, Department of Zoology, Bharathiar University, Coimbatore, India.
Dr. Xiao-Wei Wang, Institute of Insect Science, Zhejiang University, China.
Dr. Haijun Xu, Institute of Insect Science, Zhejiang University, China.
Dr. Alisha Anderson, CSIRO Ecosystem Sciences, Australia.
Prof. Eric D.Dodds, Department of Chemistry, University of Nebraska-Lincoln, USA.
Prof. P. Mosae Selvakumar, Department of Chemistry, Karnaya University, Coimbatore, India.
Prof. A.K.Dikshit, Indian Agriculture Research Institute, New Delhi.
Prof. K.R.S. Sambasiva Rao, Dept. of Biotech. & Zoology, Acharya Nagarjuna University, India
Dr. R. Rangeshwaran, National Bureau of Agriculturally Important Insects, Banglore, India.
Dr. V. Selvanarayanan, Faculty of Agriculture, Annamalai University, Tamil Nadu, India.
Prof. Fernando G. Noriega, Florida International University, Miami, USA.
Prof. Ada Rafaeli, Department of Food Quality and Safety, A.R.O., Israel.
Prof. Daphne Q.-D. Pham, Dept. of Biological Sciences, University of Wisconsin-Parkside, USA.
Prof. Emmanuelle Jacquin-Joly, INRA, UMR 1272 Physiologie de lInsecte, Versailles, France.
Prof. Manickam Sugumaran, University of Massachusetts Boston, USA.
Prof. Nannan Liu, Auburn University, USA.
Prof. Michihiro Kobyashi, Nagoya University, Japan.
Prof. Enoch Y.Park, Innovative Joint Research Center, Shizuoka University, Japan.
Prof. Luiz Paulo Moura ANDRIOLI, Universidade de So Paulo, SP - Brazil
Prof. SHIMADA Toru, The University of Tokyo, Japan.
Prof. Erjun Ling, Institute of Plant Physiology and Ecology, China.

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Acknowledgements
Writing and publishing a book requires the assistance of individuals who are
creative, talented, and hard-working. All of these qualities were present in the
individuals assembled to produce this book volume. I would like to express my
heartfelt gratitude to my former teacher Prof. Seo Sook Jae, (GSNU, South Korea),
Prof. Subba Reddy Palli (University of Kentucky, USA), and other external mentors
Prof. Marian R. Goldsmith (University of Rhode Island, USA), Prof. Enoch Y. Park
(Shizuoka University, Japan), Prof. M. Kobayashi (Nagoya University, Japan), Prof.
CHU Jang Hann (National University of Singapore, Singapore), Prof. Thomas W.
Sappington (USDA-ARS, USA), Prof. Fernando G. Noriega (Florida International
University, USA), Dr. Srinivasan Ramasamy, AVRDC, The World Vegetable
Center, Taiwan), Dr. H.C. Sharam (ICRISAT, India), who inspiration and
supported me at many ways for the commencement of this International Book
Mission Program. The book mission program was initiated on May 2010,
completed on March 2014 and published on October 2014. I have no words to
express my feeling for all those who provided valuable contributions from USA,
South Korea, Japan, China, India, Thailand, Taiwan, Bulgaria, France, Iseral, and
Portugal (Contributors name list, see page no. v) and made the completion of this
book possible. We express our appreciation to the following people (Reviewer
name list, see page no. vii) who reviewed various part of the manuscript as it was
being developed and improved quality of each chapter. I thank the ICMR, New
Delhi, and Chinese Academy of Agricultural, China, and Kansas State University for
support from several aspects. Many others (scientists and publishers) have also
allowed us to use their materials in the various chapters, their color image have then
been converted to gray color/BW. Iam especially indebted to International Book
Mission Organization, Academic Publishing Services for the production of book. I
thank my Co-Editors for their continuous vigilance over the book project and for
always giving advance notice of the editing and proofreading schedules. I thank also
my Brintha, P.G., (my wife), who in all possible way, encouragement helped
transform our original efforts into an acceptable final form. I apologize to those
whose work could not be cited owing to space considerations limitation. Further, I
wish to recognize the moral support extended by colleagues and friends. I hope that
this volume will inspire interest on the diverse aspects of insect biochemistry and
molecular biology in aspiring and established scientists.
Raman Chandrasekar

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A Note from the Publisher

Dear Readers,
This edition represents the first number of the Short Views on Insect
Biochemistry and Molecular Biology book series published by International
Book Mission. It serves to show the public how important entomology field in
expanding basic knowledge or in the development of new technologies nowadays,
in virtually all fields of knowledge. We called for piece of work falling into two
volumes (Basic and Advance aspects).
Far from being complete, the 30 chapters clearly structured and simply explained
experts contributions may provide an overview about current and prominent
advances in insect biochemistry and molecular biology which will help students and
researchers to broaden their knowledge and to gain an understanding of both the
challenges and the opportunities behind each approach.
We look forward to receiving new proposals for the new edition 2015 - 2017.
International Book Mission
Academic Publisher
Manager

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