The Anatomy and Physiology of Insects in Relation to the Dissemination of Plant Diseases

Because the insects play a vital role in the dissemination of plant pathogens, So it is very important to understand the anatomy and physiology of different organs of insects and how they aids in the dissemination of the plant pathogens.

Exoskeleton
Exoskeleton of an insect provides protection to the internal delicate structures of an insects body, avoids desiccation and provides site of attachment to the different body organs. It can be divided into three main parts; 1. Epidermis: Cellular, living layer. 2. Cuticle: Non cellular, secreted by epidermis made up of chitin. Can further be divided into; Exocuticle: Darker in color. Endocuticle: Lighter in color.

3. Basement membrane: Thin layer below epidermis. In some cases there is an other layer above cuticle called epicuticle is present. Again can be divided into; Cement layer Wax layer Upper epicuticle Lower epicuticle

As far as the dissemination of pathogens with exoskeleton is concerned the dissemination is totally mechanical, which is greatly inhanced by numerous cuticular processes found on an insect body. These cuticular processes can be of two main types; Setae: Contain membranous articulation at their base and are moveable. Microtrichogen: Does not contain membranous articulation at their base and are immoveable. These cuticular processes along with body appendages greatly enhance the mechanical transmission of pathogens to the healthy plants.

Mouth parts
Because the wounds produced by the feeding of insects provide an effective way of transmission of pathogens, So it is very important to understand the anatomy and physiology of different types of mouthparts and they help in the spread of pathogens. There are six main types of mouthparts; 1. Chewing 2. Rasping sucking 3. Piercing sucking 4. Sponging 5. Siphoning 6. Chewing lapping These mouthparts are made up of eight essential structures which may be highly modified or completely absent in these different type of mouthparts. These are; Labrum Epipharynx Hypopharynx A pair of mandibles A pair of maxillae Labium

1. Chewing mouthparts:
Represented by the mouthparts of grasshoppers, crickets, termites, caterpillars of Lepidoptera and grubs of Coleoptera , which are important vectors of several destructive diseases. These type of mouthparts are used to pinch off or eat out the part of plant tissue into the mouth of insect, provides the site of entry to the pathogens into the plants body. Different parts of an insect mouthparts has its own role; 1. Labrum - plate (sclerite) that serves as upper lip in insects with chewing mouthparts. Helps to pull food into the mouth. 2. Mandible - analogous to jaw. Used to chew, cut, and tear food, to carry things, to fight, and to mold wax. Move from side to side rather than up and down. 3. Maxillae - 2nd pair of feeding appendages, used for food handling and sensing. More complicated than the mandibles but working in the same manner. 4. Labium - analogous to lower lip. They function to close the mouth below or behind. Evolved from paired maxillae-like structures that are fused along the center line. 5. Epipharynx- found on the ventral side of labrum and has the sensors of taste. 6. Hypopharynx- functions as tongue.

2. Rasping Sucking:
These type of mouthparts found in thrips, responsible of several destructive diseases in plants. The

mouthparts are cone shaped, below head and bent backwards. Cone shaped portion is made up of clypeus, labrum, maxillae and labium. While there is a stylet between this cone which is a main sucking organ is composed of 1 hypopharynx, 1 mandible and 2 maxillae.

3. Piercing sucking:
Represented by the mouthparts of plant bugs, sucking lice, biting lice, members of Homoptera and Hemiptera. We will only discuss about the mouthparts of Homoptera and Hemiptera because they are involved in the inoculation and ingression of the plant pathogens. The mouthparts are modified into beaklike proboscis. Maxillae are doubly grooved and form the salivary canal and food canal, while the mandibles are used to pierce the plant surface to enter the proboscis into the plant surface.

4. Sponging mouthparts:
Found in the adults of

specialized flies. The labium is modified into elbow shaped proboscis and the labrum into sponge like sucking organ labella. There are several transverse channels in labella which are filled in by capillary action when labella is placed over liquid food and sucked up by the

flies. During the feeding of solid food by the flies the crop contents of the fore-gut of flies are regurgitated by the flies on to the plant surface which contains bacteria that helps in the digestion or dissolving of plant surface. Now these crop contents are again sucked up along with the dissolved food. It has been found that these crop contents of the fore-gut also contain plant pathogenic bacteria, So in this way plant pathogens can transfer into the plant body. Some Dipterous contains prestomal teeth which are used to lacerate the plant surface.

5. Siphoning mouthparts:
These type of mouthparts are found in moths and butterflies of Lepidoptera. Labrum and mandibles are completely absent. Feeding proboscis is formed by the galea of maxillae. These type of mouthparts cannot penetrate the plant body. The transfer of pathogens only occurs mechanically.

6. Chewing lapping:
These type of mouthparts are found in honeybees and many Hymenoptera. The transfer of pathogens having these type of mouthparts also occurs mechanically.

Salivary glands
Every insect is provided with a pair of salivary glands located on each side of fore-gut in thorax. They produce saliva which act as a solvent, converts sugar into starch and reduces the viscosity of the food so it can pass the alimentary canal which is 1-2 in diameter. Saliva of some species of insect is toxicogenic for the plants. It has been found that during feeding by the insects on the virus infected plants takes virus particles into their body which some how makes their way into the salivary glands and when these insects feeds on healthy

plants these viruses transfer into them along with saliva. Spores of Ergot of fungi and soft rot of bacteria spreads in this way.

Alimentary canal

Practically, in all cases of biological transmission of plant diseases alimentary canal is directly involved. So, it is very important to understand the anatomy and physiology of alimentary canal and they harbor plant pathogens. Alimentary canal can be broadly divided into three main parts; Stomodaeum or Fore-gut Mesenteron or Mid-gut Proctodaeum or Hind-gut

Stomodaeum:
It may be divided into; Buccal cavity Pharynx Oesophagus Crop Proventriculus 6

Buccal cavity: It is a just mouth opening. In ants it is modified into infra buccal cavity. During the feeding of liquid food solid contamination gather here and thrown out in the form of solid pellets it has been found that they pellets contain pathogenic spores of fungi. Fungus cultivating ants also start new colony of fungus in this way. Pharynx: Composed of strong dilateral muscles, which act as a pump during sucking in sucking insects. Oesophagus: Just act as a connecting tube between pharynx and crop. Crop: It is a main storage organ of food. Some digestion may also take place in crop. In some insects it is modified into lateral diverticulum. It has been found that the crop contents of Diptera are liquid and contain plant pathogenic bacteria (Detail has been mentioned earlier). Proventriculus: Its walls are highly seclarotized and contain spines to convert the food into smaller particles. Also called as Chewing stomach or Gizzard.

Mesenteron:
It is also called as ventriculus. Food enters into ventriculus through cardiac valve. It is the main digestion portion and also the digested food absorbed here. Epithelium of mid-gut is modified into columnary cells, which secrete digestive juices. The food is covered into a peritrophic membrane. Gastric cecae: There are blind pouches in mesenteron called gastric cecae. Which no. and location in mesenteron varies in different insects. It mainly works as secretary organ but has different functions in different insects. It has been found that congenital transmission of pathogens can take place of here. Because it contain many plant pathogens. A striking modification of the alimentary canal occurs in Homoptera who contain filter chamber. In order to get sufficient amount of nitrogenous food they had to suck large amount of cell sap, which contains water and soluble carbohydrates. So, these filter chambers filter the nitrogenous food and fatty materials from the cell sap, the rest sugar solution is pass out the body in the form of honey dew. On this honey dew sooty mould develops and the normal functioning of the leaf retarded.

Proctodaeum:
Digested food enters into the hind-gut through pyloric valve. Proctodaeum can be divided into;

Fore-intestine Hind-intestine

Fore-intestine: This can be divided into Ileum and Colon. Some absorption may also take place in ileum. Malpighian tubules open into the ileum which immerse into the haemolymph, gather the metabolic waste and transfer it to the ileum. In some species of insects it has been found that they also produce a substance which is used to woven cocoons for pupation. After that the waste material transfers into colon and subsequently into hind-intestine. Hind-intestine: It is composed of Rectum and Rectal pads. Waste material enters into the rectum and from there it is ejected from the anus with the help of rectal pads.

Organs of Reproduction

Organs of reproduction plays vital role in the congenital transfer of pathogens, So it is very important to study them that how they can involve in the transfer of pathogens to the next generation or to the healthy plant.

Because the female insects are mostly involved in the transfer of pathogens, So we will only discuss the organs of reproduction of only female insect. Female reproductive system can be divided into two main parts; Internal Genitalia External Genitalia

Internal genitalia: It can be divided into several parts; Ovary: There are two pairs of ovaries in insects. Each of them are composed of two or more Ovarioles (depends upon the species). The germ cells are produced in ovarioles in the portion of Germarium just below the terminal filament. After that these germ cells transfers into site of Vitallarium where they are nourished by the nurse or nutritive cells and transforms into oogonium and then oocyte. The epithelium of ovarioles is composed of follicle cells which may also provide nutrition to these cells. They also provide the protective covering Chorion to the oocytes. On the basis of presence or absence of nurse cells ovarioles can be divided into two types; Planoistic: Nurse cells absent. Meroistic: Nurse cells present. During feeding nurse cells gather around the germ cell and there protoplasm is diffused into the germ cells. At this time some pathogens may also enter into the oocytes and congenital transfer may occur. After formation of egg they are transferred into the Lateral oviducts. In some insects they may be modified into Calyx for temporarily storage of the eggs. Then these enters into Median oviduct and from there they enters into Vagina through Gonopore (regulates the flow of eggs). Vagina: It acts as a pouch for copulation and also called as Bursa Copulatrix. It has been found that many pathogens develop into the vagina and when eggs enters into vagina they enters into it and transfers congenitally. Spermatheca: They are also called seminal vesicles. In this the sperms stores after mating. Many plant pathogens are also found in spermatheca and they enter into eggs during fertilization. Spermathical glands: They provide the fluid which transfers with the sperms.

Accessory glands: They provide the gluing or adhesive substance for the eggs so that they could remain attached to each other or to the substrate. They also called as Colletrial glands. In some stinging Hymenoptera they also provide a toxic substance. In some insects the vagina and anus are close together to ensure the congenital transfer of pathogens from the alimentary canal. Rectum contains pouches in which pathogens are present. So, when eggs pass through vagina they pushes the walls of these pouches, results in the squeeze out of these pathogens at the same time as eggs pass through vagina, and contaminates eggs. External Genitalia: It is composed of copulatory organs and organs of oviposition. In some insects the organ of oviposition is modified into a long ovipositor along with 8 and 9th abdominal segment. It helps insects to lay eggs into the plant tissues. So, ovipositional punctures made by these ovipositors provide an entry site to the pathogens into the plant.
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