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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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(1), 00 – 00, 2009 Vol. (1) 57 – 74,

2014

Modulation of Botanicals on Pest’s 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. analysis, carbohydrates, carbohydrates, lipid, enzymes, enzymes, detoxification, detoxification, mortality, mortality, plant defense  Key words: biochemical analysis, *For Correspondence (email: [email protected] [email protected])

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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 58  Printed in the Unitated States of of America, 2014  ISBN: 978-1-63315-205-2

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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. compounds. Essential oils are lipophilic in natures which generally 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 neurohormone–neuromodulator) (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). 59  Printed in the Unitated States of America, 2014  ISBN: 978-1-63315-205-2

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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 im balance, 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. 60  Printed in the Unitated States of of America, 2014  ISBN: 978-1-63315-205-2

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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). (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 61  Printed in the Unitated States of America, 2014  ISBN: 978-1-63315-205-2

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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 synt hesis machinery. This could b e 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 64  Printed in the Unitated States of of America, 2014  ISBN: 978-1-63315-205-2

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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  lipids  are of vital importance to many insects as substrates for embryogenesis, metamorphosis and flight. 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) (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 l eaves 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 level was + 2+ higher while  D. cingulatus was fed with cotton seed. Similarly K   and Ca  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) (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 68  Printed in the Unitated States of of America, 2014  ISBN: 978-1-63315-205-2

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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) 69  Printed in the Unitated States of America, 2014  ISBN: 978-1-63315-205-2

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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 gly cogens 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 -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 70  Printed in the Unitated States of of America, 2014  ISBN: 978-1-63315-205-2

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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. It’s (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 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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Babu, R., Murugan, K. and Vanithakumari, G. (1996) Interference of Azadirachtin on the food utilization efficiency and midgut enzymatic profiles of Helicoverpa of Helicoverpa armigera, armigera, Indian J. Environ. Toxicol. 6: 81–84.

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Zibaee, A., Bandani, A.R. and Ramzi, S. (2008) Lipase and invertase activities in midgut and salivary glands ofChilo Chilo suppressalis (Walker) (Lepidoptera, Pyralidae), rice striped stem borer. Invertebrate borer. Invertebrate Survival J. 5: J. 5: 180-189.

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Zibaee, A., Bandani, A.R. and Ramzi, S. (2009) Characterization of alpha and beta glucosidases in midgut and salivary glands of Chilo suppressalis Walker suppressalis Walker (Lepidoptera: Pyralidae), rice striped stem borer.Comptes borer.Comptes Rendus Biologies, 332: Biologies, 332: 633–641.

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Hemmingi, J.D.C. and Lindroth, R.L. (1999) Effects of light and nutrient availability on aspen: growth, phytochemistry and insect  performance. J.  performance. J. Chemical Ecology, 26: Ecology, 26: 1687–1714.

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Hemmingi, J.D.C. and Lindroth, R.L. (2000) Effects of phenolic glycosides and protein on gypsy moth (Lepidoptera: Lymantriidae) and forest tent caterpillar (Lepidoptera: Lasiocampidae) performance and detoxication activities. Environmental activities. Environmental Entomol.  Entomol.  29: 1108–1115.

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Sakharov, I.Y., Makarova, I.E., and Ermolin, G.A. (1989) Chemical modification and composition of tetrameric isozyme K of alkaline phosphatase from harp seal intestinal mucosa.Comparative mucosa.Comparative Biochem. Physiol. Part B: Comparative Biochem. 92(1): Biochem. 92(1): 119-122.

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Senthil Nathan, S. and Sehoon, K. (2006). Effects of Melia of Melia azedarach  azedarach  L. extract on the teak defoliafor Hyblaea defoliafor  Hyblaea puera   Cramer

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Terra, W.R. and Ferriera, C. (2005) Biochemistry of digestion. In: Comprehensive molecular insect science by Lawrence I. Gilbert,

(Lepidoptera: Hyblaeidae), Crop Prot. 25: Prot. 25: 287–291. Kostas Iatrou, and Sarjeet S. Gill, Elsevier, USA, Vol.(3): 171-224.

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Johnson, D.E., Brookhart, G.L., Kramer, K.J., Barnett, B.D. and McGaughey, W.H. (1990) Resistance to Bacillus thuringiensis by thuringiensis by the Indian meal moth Plodia moth Plodia interpunctella: interpunctella: Comparison of midgut proteinase from susceptible and resistant larvae. J.  J. Invertebrate  Pathology, 55: 235–244.

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Zibaee, A. and Bandani, A.R. (2010) Effects of Artemisia of Artemisia annua L. annua L. (Asteracea) on digestive enzymes profiles and cellular immune reactions of sunn pest, Eurygaster pest, Eurygaster integriceps  integriceps  (Heteroptera: Scutellaridae), against Beauvaria against Beauvaria bassiana. bassiana .  Bulletin of Entomological  Res. 100: 185-196.

80. 81.

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) mori ) by biochemical markers. Insect markers. Insect Sci. 12: 87-94.

82.

Dugravot, S., Thibout, E., Abo‐Ghalia, A., and Huignard, J. (2004) How a specialist and a non‐specialist insect cope with dimethyl disulfide produced by Allium by Allium porrum. porrum. Entomologia Experimentalis et Applicata, Applicata, 113(3): 173-179.

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Yerasi, B., and Chitra, K.C. (2000) Response of aspartate and alanine aminotransferase to sublethal concentration of annonain by Spodoptera litura (Fab.). litura  (Fab.). Indian  Indian J. Entomol. 62(4): Entomol. 62(4): 367-370.

84.

Wilkinson, C.F. (1976). Insecticide synergism, pp. 195-218. In R.L. Metcalf and J.J. McKelvey, Jr. [Eds.], Insecticides  Insecticides for the  Future: Needs and Prospects. John Prospects. John Wiley and Sons, New York.

85.

 Naqvi, S.H. (1986) Biological evaluation of fresh neem e xtracts and some est erase a ctivity in insects. Proc, rd3  Int. Neem Conf.  Nairobi. 315-330.

86.

Grant, D.F., and Matsumura, F. (1989) Glutathione-S-transferase 1 and 2 in susceptible and insecticide resistant Aedes resistant Aedes aegypti.. Pesticide Biochem. and Physiol. 33(2): 132-143. aegypti

87.

Francis, F., Vanhaelen, N. and Haubruge, E. (2005) Glutathione S-transferases in the adaptation to plant secondary metabolites in the Myzus the Myzus persicae aphid. persicae aphid. Arch.  Arch. Insect Biochem. Physiol. 58: Physiol. 58: 166–174.

Article History:

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

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Table Contents

Page No.

Preface Forward message Contributor’s Reviewers  Acknolwedgement

i ii iii iv v

Volume 1 Section I:

Insect Biochemical approaches

1. Introduction to Insect Molecular Biology. Biology.

3

 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, Patnaik, and P. Michailova

2.

Modulation of Botanicals on pests’ biochemistry.

57

Sahayaraj, K.

3.

Detoxication, stress and immune responses i n insect antenna:

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new insights from transcriptomics.  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 of Dendroctonus armandi Tsai 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 Sclerotization–Hardening 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 planthopper,  Nilaparvata lugens. lugens.

233

Yanyuan Bao, Raman Chandrasekar, Chuan-Xi Zhang

10. Immune Pathways in Anopheles in Anopheles gambiae.

253

 Maria L. Simões 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 silkworm, Bombyx  Hiroaki Abe, Tsuguru Fujii and Raman Chandrasekar

13. Molecular characterization and DNA barcoding for identification of

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agriculturally important insects’.  Rakshit Ojha, Jalali, S.K., and Venkatesan, T.

14. Polytene chromosomes and their si gnificance for Taxonomy,

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Speciation and Genotoxicology  Paraskeva V. Michailova

15. Insect exuvium extracted DNA marker: a good complementary molecular taxonomic characteristics with special reference to mosquitoes.

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

Index

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Volume 2

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 Montagné, Olivier Mirabeau, Emmanuelle Jacquin-Joly

18. Molecular architecture, phylogeny and biogeography of pheromone

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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 :

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Its Current Status.  Atanu Bhattacharyya , Raman Chandrasekar, 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.

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 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 i mportant component of

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novel approaches to insect pest management. Usha Rani, P.

26. Mosquito control using biological larvicides: Current Scenario.

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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.

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 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, Chandrasekar, Yeon Soo Han

30. Computer-aided pesticide design: A short view

685

 Jitrayut Jitonnom

Index

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

First Edition: Volume 1, 2 October 2014 Total No. Pages: 398 + 372 = 770

Edited by by Raman Chandrasekar Chandrasekar B.K. Tyagi Zhong Zheng Gui Gerald R. Reeck © Copyright Reserved Published by International Book Mission©, Academic Publisher, South India.

Printed in the K-State Union, Copy and Printing Printi ng services, Kansas State University, Manhattan 66506, KS, USA. This publication is considered to provide accurate and authoritative information with regards to the subject matter has been obtained by its authors. The publisher has taken reasonable care in the preparation of this book volume. However, the publisher and its authors shall in no event be liable for any errors or omission arising out of use of this information and specifically disclaim any implied warranties or merchantability or fitness for any particular use. No part of these books may be reproduced, stored in retrieval system or transmitted in any form or by any means, electronic, mechanical, photocopying or otherwide, without the prior permission of the Copyright owner. Application for such permission, with a statement of the purpose and extend for the reproduction, should be addressed to the publisher (IBM, Academic Publisher).

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Tel. +91-431-2311187 Tel. +1-859-608-7694 (USA) 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” 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 designing.  In short this book provides comprehensive reviews of recent research from various geographic areas around the world and contributing author’s 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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Short Views on

Insect Biochemistry and Molecular Biology Edited by Raman Chandrasekar, 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, 4–Sarojini 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 Palayamkottai 627 002, Tamil Nadu, India.

Prof. Chuan-Xi Zhang Zhang,,

Prof. David Siaussat

Dr. Maria L. Simões

Université Pierre et Marie Curie (Paris 6/UPMC), UMR 1272A Physiologie de l'Insecte: Signalisation et Communication (PISC), 7 Quai Saint Bernard, Batiment A - 4ème étage  bureau 410, 75252 Paris Paris Cedex 05, France. France.

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

Prof. Shou-An Xie College of Forestry,  Northwest A & F University University Yangling, Shaanxi 712100, China ，

Dr. Raman Chandrasekar 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.

Institute of Insect Science, Zhejiang University, China.

UEI Parasitologia Médica, Centro de Malária e Outras Doenças 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.

Dr. Hiroaki Abe Tokyo University of Agriculture and Technology, Japan.

Dr. S.K. Jalali  National Bureau of of Agriculturally 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.

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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, 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 School of Science University of Phayao, Thailand.

Prof. K. Murugan Department of Zoology, School of Life Sciences, Bharathiar University, Coimbatore - 641 046, India.

Prof. Immo A. Hansen Department of Biology,  New Mexico State University, University, Las Cruces, NM, USA.

Dr. Ronald J. Nachman USDA-ARS, Food Animal Protection Research Laboratory, USA.

Dr. Hari C Sharma International 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.

Prof. B.R. Pittendrigh Department of Entomology, University of Illinois, Urbana-Champaign, IL, 61801, USA .

Dr. Subbiah Poopathi 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, Bioinformatics, Faculty of Engineering & Technology, SRM University, Kattankulathur, 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 l’Insecte, 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 São 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 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 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 receiving new proposals for the new edition 2015 - 2017. International Book Mission  Academic Publisher Manager

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