Insect morphology

Insects, like all arthropods, have no interior skeleton; instead, they have an exoskeleton, a hard outer layer made mostly of chitin that protects and supports the body. The insect body is divided into three parts: the head, thorax, and abdomen. The head is specialized for sensory input and food intake; the thorax, which is the anchor point for the legs and wings (if present), is specialized for locomotion; and the abdomen is for digestion, respiration, excretion, and reproduction. == External ==

Exoskeleton The insect's outer skeleton, the cuticle, consists of two layers; the epicuticle, which is a thin, waxy, water-resistant outer layer that lacks chitin, and the layer under it is called the procuticle. This is chitinous and much thicker than the epicuticle and has two layers, the outer is the exocuticle while the inner is the endocuticle. The tough and flexible endocuticle is built from numerous layers of fibrous chitin and proteins, crisscrossing each other in a sandwich pattern, while the exocuticle is rigid and sclerotized. The difference between the unmodified and modified forms is evident when comparing the body wall of a caterpillar (unmodified) to a beetle (modified). From the embryonic stages, a layer of columnar or cuboidal epithelial cells gives rise to the external cuticle and an internal basement membrane. The majority of insect material is inside of the endocuticle. The cuticle provides muscular support and acts as a protective shield as the insect develops. However, since it cannot grow, the external sclerotized part of the cuticle is periodically shed in a process called "molting". As the time for molting approaches, most of the exocuticle material is reabsorbed. In molting, the old cuticle separates from the epidermis (apolysis). Enzymatic molting fluid is then released between the old cuticle and epidermis, which separates the exocuticle by digesting the endocuticle and sequestering its material for the new cuticle. When the new cuticle has formed sufficiently, the epicuticle and reduced exocuticle are shed in ecdysis. The four principal regions of an insect body segment are the tergum or dorsal, sternum or ventral, and the two pleura or laterals. Hardened plates in the exoskeleton are called sclerites, which are subdivisions of the major regions – tergites, sternites, and pleurites, for respective regions tergum, sternum, and pleuron. Head The head in most insects is enclosed in a hard, heavily sclerotized, exoskeletal head capsule. This bears most of the sensory organs, including the antennae, ocelli, and compound eyes, along with the mouthparts. It consists of six segments that bear the paired head appendages, including the mouthparts, each pair on a specific segment. Thorax The insect thorax has three segments: the prothorax, mesothorax, and metathorax. The anterior segment, closest to the head, is the prothorax; its major features are the first pair of legs and the pronotum. The middle segment is the mesothorax; its major features are the second pair of legs and the anterior wings, if any. The third, the posterior, thoracic segment, abutting the abdomen, is the metathorax, which bears the third pair of legs and the posterior wings. Each segment is delineated by an intersegmental suture. Each segment has four basic regions. The dorsal surface is called the tergum (or notum, to distinguish it from the abdominal terga). The articular sclerites, or pteralia, of the wing base of the wing-flexing insects and their relations to the body and the wing veins, shown diagrammatically, are as follows: • Humeral plates • First Axillary • Second Axillary • Third Axillary • Fourth Axillary • '''Median plates (m, m')''' The humeral plate is usually a small sclerite on the anterior margin of the wing base, movable and articulated with the base of the costal vein. Odonata have their humeral plates greatly enlarged, The first axillary sclerite (lAx) is the anterior hinge plate of the wing base. Its anterior part is supported on the anterior notal wing process of the tergum (ANP); its posterior part articulates with the tergal margin. The anterior end of the sclerite is generally produced as a slender arm, the apex of which (e) is always associated with the base of the subcostal vein (Sc), though it is not united with the latter. The body of the sclerite articulates laterally with the second axillary. The second axillary sclerite (2Ax) is more variable in form than the first axillary, but its mechanical relations are no less definite. It is obliquely hinged to the outer margin of the body of the first axillary, and the radial vein (R) is always flexibly attached to its anterior end (d). The second axillary presents both a dorsal and ventral sclerotization in the wing base; its ventral surface rests upon the fulcral wing process of the pleuron. The second axillary, therefore, is the pivotal sclerite of the wing base, and it specifically manipulates the radial vein. although, most Neoptera have a jugal fold just behind vein 3A on the forewings. It is sometimes also present on the hindwings. Where the anal area of the hindwing is large, as in Orthoptera and Blattodea, the whole of this part may be folded under the anterior part of the wing along a vannal fold a little posterior to the claval furrow. In addition, in Orthoptera and Blattodea, the anal area is folded like a fan along the veins, the anal veins being convex, at the crests of the folds, and the accessory veins concave. Whereas the claval furrow and jugal fold are probably homologous in different species, the vannal fold varies in position in different taxa. Folding is produced by a muscle arising on the pleuron and inserted into the third axillary sclerite in such a way that when it contracts, the sclerite pivots about its points of articulation with the posterior notal process and the second axillary sclerite. Similarly, protibia, mesotibia, and metatibia refer to the tibiae of the front, middle and hind legs. The tarsus of insects corresponds to the penultimate segment of a generalized arthropod limb, which is the segment called the propodite in Crustacea. In adult insects, it is commonly subdivided into two to five subsegments, or tarsomeres, but in the Protura, some Collembola, and most holometabolous insect larvae it preserves the primitive form of a simple segment. The subsegments of the adult insect tarsus are usually freely movable on one another by inflected connecting membranes, but the tarsus never has intrinsic muscles. The tarsus of adult pterygote insects having fewer than five subsegments is probably specialized by the loss of one or more subsegments or by a fusion of adjoining subsegments. In the tarsi of Acrididae, the long basal piece is composed of three united tarsomeres, leaving the fourth and the fifth. The basal tarsomere is sometimes conspicuously enlarged and is distinguished as the basitarsus. On the under surfaces of the tarsal subsegments in certain Orthoptera, there are small pads, the tarsal pulvilli, or euplantulae. The tarsus is occasionally fused with the tibia in larval insects, forming a tibiotarsal segment; in some cases, it appears to be eliminated or reduced to a rudiment between the tibia and the pretarsus. A similar structure in nymphal stoneflies (Plecoptera) is of uncertain homology. == Internal ==

Nervous system The nervous system of an insect can be divided into a brain and a ventral nerve cord. The head capsule is made up of six fused segments, each with a pair of ganglia, or a cluster of nerve cells outside of the brain. The first three pairs of ganglia are fused into the brain, while the three following pairs are fused into a structure of three pairs of ganglia under the insect's esophagus, called the subesophageal ganglion. Some cockroaches have just six ganglia in the abdomen, whereas the wasp Vespa crabro has only two in the thorax and three in the abdomen. Some insects, like the house fly Musca domestica, have all the body ganglia fused into a single large thoracic ganglion. At least a few insects have nociceptors, cells that detect and transmit sensations of pain. This was discovered in 2003 by studying the variation in reactions of larvae of the common fruitfly Drosophila to the touch of a heated probe and an unheated one. The larvae reacted to the touch of the heated probe with a stereotypical rolling behavior that was not exhibited when the larvae were touched by the unheated probe. Although nociception has been demonstrated in insects, there is not a consensus that insects feel pain consciously. Digestive system An insect uses its digestive system for all steps in food processing: digestion, absorption, and feces delivery and elimination. Most of this food is ingested in the form of macromolecules and other complex substances like proteins, polysaccharides, fats, and nucleic acids. These macromolecules must be broken down by catabolic reactions into smaller molecules like amino acids and simple sugars before being used by cells of the body for energy, growth, or reproduction. This break-down process is known as digestion. The main structure of an insect's digestive system is a long-enclosed tube called the alimentary canal (or gut), which runs lengthwise through the body. The alimentary canal directs food in one direction: from the mouth to the anus. The gut is where almost all of insects' digestion takes place. It can be divided into three sections – the foregut, midgut and hindgut – each of which performs a different process of digestion. In addition to the alimentary canal, insects also have paired salivary glands and salivary reservoirs. These structures usually reside in the thorax, adjacent to the foregut. Some insects, like flies, have extra-oral digestion. Insects using extra-oral digestion expel digestive enzymes onto their food to break it down. This strategy allows insects to extract a significant proportion of the available nutrients from the food source. Midgut Once food leaves the crop, it passes to the midgut (element 13 in numbered diagram), also known as the mesenteron, where the majority of digestion takes place. Microscopic projections from the midgut wall, called microvilli, increase the surface area of the wall and allow more nutrients to be absorbed; they tend to be close to the origin of the midgut. In some insects, the role of the microvilli and where they are located may vary. For example, specialized microvilli producing digestive enzymes may more likely be near the end of the midgut, and absorption near the origin or beginning of the midgut. Hindgut In the hindgut (element 16 in numbered diagram), or proctodaeum, undigested food particles are joined by uric acid to form fecal pellets. The rectum absorbs 90% of the water in these fecal pellets, and the dry pellet is then eliminated through the anus (element 17), completing the process of digestion. The uric acid is formed using hemolymph waste products diffused from the Malpighian tubules (element 20). It is then emptied directly into the alimentary canal, at the junction between the midgut and hindgut. The number of Malpighian tubules possessed by a given insect varies between species, ranging from only two tubules in some insects to over 100 tubules in others. Air is taken in through openings on the sides of the abdomen called spiracles. There are many different patterns of gas exchange demonstrated by different groups of insects. Gas exchange patterns in insects can range from continuous and diffusive ventilation, to discontinuous gas exchange. Diffusive ventilation is simply a form of continuous gas exchange that occurs by diffusion rather than physically taking in the oxygen. Some species of insect that are submerged also have adaptations to aid in respiration. As larvae, many insects have gills that can extract oxygen dissolved in water, while others need to rise to the water surface to replenish air supplies, which may be held or trapped in special structures. Circulatory system Insect blood or haemolymph's main function is that of transport and it bathes the insect's body organs. Making up usually less than 25% of an insect's body weight, it transports hormones, nutrients and wastes and has a role in, osmoregulation, temperature control, immunity, storage (water, carbohydrates and fats) and skeletal function. It also plays an essential part in the moulting process. An additional role of the haemolymph in some orders, can be that of predatory defence. It can contain unpalatable and malodourous chemicals that will act as a deterrent to predators. Body fluids enter through one-way valved ostia, which are openings situated along the length of the combined aorta and heart organ. Pumping of the haemolymph occurs by waves of peristaltic contraction, originating at the body's posterior end, pumping forwards into the dorsal vessel, out via the aorta and then into the head where it flows out into the haemocoel. Reproductive system Female Female insects are able make eggs, receive and store sperm, manipulate sperm from different males, and lay eggs. Their reproductive systems are made up of a pair of ovaries, accessory glands, one or more spermathecae, and ducts connecting these parts. The ovaries make eggs and accessory glands produce the substances to help package and lay the eggs. Spermathecae store sperm for varying periods of time and, along with portions of the oviducts, can control sperm use. The ducts and spermathecae are lined with a cuticle. == Internal morphology of different taxa ==

Blattodea Cockroaches are most common in tropical and subtropical climates. Some species are in close association with human dwellings and widely found around garbage or in the kitchen. Cockroaches are generally omnivorous with the exception of the wood-eating species such as Cryptocercus; these roaches are incapable of digesting cellulose themselves but have symbiotic relationships with various protozoans and bacteria that digest the cellulose, allowing them to extract the nutrients. The similarity of these symbionts in the genus Cryptocercus to those in termites are such that it has been suggested that they are more closely related to termites than to other cockroaches, and current research strongly supports this hypothesis of relationships. All species studied so far carry the obligate mutualistic endosymbiont bacterium Blattabacterium, with the exception of Nocticola australiensis, an Australian cave dwelling species without eyes, pigment or wings, and which recent genetic studies indicates are very primitive cockroaches. Cockroaches, like all insects, breathe through a system of tubes called tracheae. The tracheae of insects are attached to the spiracles, excluding the head. Thus cockroaches, like all insects, are not dependent on the mouth and windpipe to breathe. The valves open when the CO2 level in the insect rises to a high level; then the CO2 diffuses out of the tracheae to the outside and fresh O2 diffuses in. Unlike in vertebrates that depend on blood for transporting O2 and CO2, the tracheal system brings the air directly to cells, the tracheal tubes branching continually like a tree until their finest divisions, tracheoles, are associated with each cell, allowing gaseous oxygen to dissolve in the cytoplasm lying across the fine cuticle lining of the tracheole. CO2 diffuses out of the cell into the tracheole. While cockroaches do not have lungs and thus do not actively breathe in the vertebrate lung manner, in some very large species the body musculature may contract rhythmically to forcibly move air out and in the spiracles; this may be considered a form of breathing. Coleoptera The digestive system of beetles is primarily based on plants, which they for the most part feed upon, with mostly the anterior midgut performing digestion. However, in predatory species (e.g., Carabidae) most digestion occurs in the crop by means of midgut enzymes. In Elateridae species, the predatory larvae defecate enzymes on their prey, with digestion being extraorally. Like other insect species, beetles have hemolymph instead of blood. The open circulatory system of the beetle is driven by a tube-like heart attached to the top inside of the thorax. Different glands specialize for different pheromones produced for finding mates. Pheromones from species of Rutelinae are produced from epithelial cells lining the inner surface of the apical abdominal segments or amino acid-based pheromones of Melolonthinae from eversible glands on the abdominal apex. Other species produce different types of pheromones. Dermestids produce esters, and species of Elateridae produce fatty-acid-derived aldehydes and acetates. While Bombardier beetles have well-developed, like other carabid beetles, pygidial glands that empty from the lateral edges of the intersegment membranes between the seventh and eighth abdominal segments. The gland is made of two containing chambers. The first holds hydroquinones and hydrogen peroxide, with the second holding just hydrogen peroxide plus catalases. These chemicals mix and result in an explosive ejection, forming temperatures of around 100 C, with the breakdown of hydroquinone to H2 + O2 + quinone, with the O2 propelling the excretion. Several species of the genus Cicindela in the family Cicindelidae have ears on the dorsal surface of the first abdominal segment beneath the wing; two tribes in the family Dynastinae (Scarabaeidae) have ears just beneath the pronotal shield or neck membrane. The ears of both families are to ultrasonic frequencies, with strong evidence that they function to detect the presence of bats via their ultrasonic echolocation. Even though beetles constitute a large order and live in a variety of niches, examples of hearing is surprisingly lacking in species, though it is likely that most are just undiscovered. The female lays her eggs as close to the food source as possible, and development is very rapid, allowing the larva to consume as much food as possible in a short period of time before transforming into the adult. The eggs hatch soon after being laid, or the flies are ovoviviparous, with the larva hatching inside the mother. Hormones and the glands that produce them run the development of butterflies and moths as they go through their life cycle, called the endocrine system. The first insect hormone PTTH (Prothoracicotropic hormone) operates the species life cycle and diapause (see the relates section). This hormone is produced by corpora allata and corpora cardiaca, where it is also stored. Some glands are specialized to perform certain task such as producing silk or producing saliva in the palpi. In the circulatory system, hemolymph, or insect blood, is used to circulate heat in a form of thermoregulation, where muscles contraction produces heat, which is transferred to the rest of the body when conditions are unfavorable. In lepidopteran species, hemolymph is circulated through the veins in the wings by some form of pulsating organ, either by the heart or by the intake of air into the trachea. Air is taken in through spiracles along the sides of the abdomen and thorax supplying the trachea with oxygen as it goes through the lepidopteran's respiratory system. There are three different tracheae supplying oxygen diffusing oxygen throughout the species body: The dorsal, ventral, and visceral. The dorsal tracheae supply oxygen to the dorsal musculature and vessels, while the ventral tracheae supply the ventral musculature and nerve cord, and the visceral tracheae supply the guts, fat bodies, and gonads. == See also ==