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Insect biology The science that deals with insect’s individual developmental history. It is divided into the study of reproduction, embryology, post embryology and adult Insect Reproduction Most species of insects have males and females that mate and reproduce sexually Some insects reproduce asexually, without the joining of male and female gametes. Types of reproduction Sexual reproduction Parthenogenesis Polyembryony Viviparity Paedogenesis Facultative parthenogenesis Obligate parthenogenesis Ovoviviparity Adenotrophic viviparity Pseudoplacental viviparity Haemocoelous viviparity Sexual reproduction Most insects reproduce sexually, by the joining of male and female gametes or sex cells. Gametes are produced by a special kind of cellular reproduction. parthenogenesis A form of reproduction in which the ovum develops into a new individual without fertilization. Natural parthenogenesis has been observed in the aphid. Facultative parthenogenesis sporadic parthenogenesis : Silkworm/phasmatodea Obligate parthenogenesis Cyclical parthenogenesis: aphids heterogeny Obligate parthenogenesis Constant parthenogenesis: In many social insects, such as the honeybee and the ant, the unfertilized eggs give rise to the male drones and the fertilized eggs to the female workers and queens. Polyembryony Parasitic hymenopteran Viviparity Ovoviviparity Nutrition was supplied by egg. Embryo finishes development in the mother’s body. Egg hatches in the mother’s body. Larva leaves mother’s body after hatching. Scale, aphid, thrips, house fly, parasitic fly Viviparity Adenotrophic viviparity Pupiparity Nutrition of embryo was supplied by egg. Embryo finishes development in the mother’s body. Egg hatches in the mother’s body. Larvae leave mother’s body until they are access to pupae. Some species in the house fly family. Viviparity Pseudoplacental viviparity No or little yolk in egg Nutrition was absorbed by pseudoplacenta in the mother’ body. Some species of aphids, Dermaptera and cockroaches Viviparity Haemocoelous viviparity Ovary breaks when embryo finishes development. Eggs are released in the mother’s haemocoele Larvae feed on mother’s organs until they come out from mother’ body before puparity. Paedogenesis Paedogenesis is a form of neoteny in insects in which the larval stage reproduces without maturing first. It occurs in the females of certain beetles, Strepsiptera, bagworms, and gall midges. Paedogenesis is the precocious development of sexual maturity in a larva. Insect reproduction Short life cycles-most go through generation in 1-6 weeks Large number of offspring / female-100-2,000 eggs Fruit flies-2 week life cycle26 generations/year100 eggs / female In 1 year from 1 male and 1 female if all offspring survive to breed would produce 1041 flies Ontogenesis Preembryonic development: sperm/egg Embryonic development Postembryonic development: after finishing embryonic development to adult Egg In most insects, life begins as an independent egg. This type of reproduction is known as ovipary. Manufactured within the female's genital system Released from her body through an ovipositor Production of eggs by the female is called öogenesis The egg-laying process is known as oviposition. Each insect species produces eggs that are genetically unique and often physically distinctive as well -- spherical, ovate, conical, sausage-shaped, barrel-shaped, or torpedo-shaped. Each egg is composed of only a single living cell -the female gamete. Some types of eggs Mode of ovipositing on the object surface:insects secret gland An ovipositing butterfly Eggs of a tent caterpillar Eggs of a true bug (Hemiptera) On the plant surface In the host organization In the concealment locus In the water In the earth Insects protect their eggs Egg structure Egg shell-chorion Vitelline membrane Periplasm Nucleus Micropyle Yolk Cytoplasm Cytoplasm reticulum chorion Periplasm Vitelline membrane An egg's cell membrane is known as the vitelline membrane . It is a phospholipid bilayer similar in structure to most other animal membranes. It surrounds the entire contents of the egg cell, most of which consists of yolk (food for the soonto-develop embryo). The cell's cytoplasm is usually distributed in a thin band just inside the vitelline membrane (where it is commonly called periplasm ) and in diffuse strands that run throughout the yolk ( cytoplasmic reticulum ). The egg cell's nucleus (haploid) lies within the yolk, usually close to one end of the egg. Near the opposite end, the öosome (a region of higher optical density) may be visible as a dark region in the more translucent yolk. The egg's anterior/posterior polarity is determined by the relative positions of the nucleus and the öosome. The egg is covered by a protective "shell" of protein secreted before oviposition by accessory glands in the female's reproductive system. This egg shell, called the chorion , is sculptured with microscopic grooves or ridges that may be visible only under an electron microscope. chorion The chorion is perforated by microscopic pores (called aeropyles ) that allow respiratory exchange of oxygen and carbon dioxide with relatively little loss of water. The micropyle , a special opening near the anterior end of the chorion, serves as a gateway for entry of sperm during fertilization. A female receives sperm from her male partner during the act of mating---insemination She can store that sperm for long periods of time in a special part of her reproductive system, the spermatheca. As a developing egg moves past the opening to the spermatheca, a few sperm are released onto its surface. Fertilization The sperm swim toward the micropyle -- the first one to reach its destination enters and injects its nucleus into the egg. The sperm nucleus quickly fuses with the egg nucleus to form a diploid zygote -- a one-celled embryo. This event is known as fertilization. After the egg is fertilized, it undergoes a period of rapid growth and development known as embryogenesis. EMBRYOGENESIS A developmental process that usually begins once the egg has been fertilized. It involves multiplication of cells (by mitosis) and their subsequent growth, movement, and differentiation into all the tissues and organs of a living insect. EMBRYOGENESIS Cleavage and formation of blastoderm Formation of germ band, embryonic layer and embryonic envelop Segmentation of embryo and formation of appendages Formation of organizations and systems Blastokinesis and disappearance of embryonic envelop Cleavage and formation of blastoderm This process of nuclear division is known as superficial cleavage As they form, the cleavage nuclei migrate through the yolk toward the perimeter of the egg. They settle in the band of periplasm where they engineer the construction of membranes to form individual cells. The end result of "cleavage" is the blastoderm -a one-cell-thick layer of cells surrounding the yolk. Formation of blastoderm Formation of blastoderm The first cleavage nuclei to reach the vicinity of the öosome are "reserved" for future reproductive purposes -- they do not travel to the periplasm and do not form any part of the blastoderm. Instead, they stop dividing and form germ cells that remain segregated throughout much of embryogenesis. These cells will eventually migrate into the developing gonads (ovaries or testes) to become primary öocytes or spermatocytes. Only when the adult insect finally reaches sexual maturity will these cells begin dividing (by meiosis) to form gametes of the next generation (eggs or sperm). Germ cells never grow or divide during embryogenesis, so DNA for the next generation is "conserved" from the very beginning of development. This strategy has a clear selective advantage: it minimizes the risk that an error in replication will accidently be passed on to the next generation. Formation of germ band Blastoderm cells on one side of the egg begin to enlarge and multiply. This region, known as the germ band (or ventral plate), is where the embryo's body will develop. The rest of the cells in the blastoderm become part of a membrane (the serosa) that forms the yolk sac. Cells from the serosa grow around the germ band, enclosing the embryo in an amniotic membrane Formation of germ band At this stage of development, when the embryo is not much more than a single layer of cells, a group of control genes (called homeotic selector genes) become active. These genes encode for proteins that contain a special active site (the homeobox) for binding with DNA. They interact with specific locations in the genome where they function as switches for activating (or inhibiting) the expression of other genes Basically, each selector gene controls the expression of certain other genes within a restricted domain of cells based on their location in the germ band. By regulating activity within a suite of genes that produce hormone-like "organizer" chemicals, cellsurface receptors, and structural elements, the selector genes guide the development of individual cells and channel them into different "career paths". This process, called differentiation, continues until the fundamental body plan is mapped out -- first into general regions along an anterio-posterior axis, then into individual segments, and finally into specialized structures or appendages. As the germ band enlarges, it begins to lengthen and fold into a sausage shape with one layer of cells on the outside (the ectoderm) and another layer of cells on the inside (the mesoderm). An important developmental milestone, called dorsal closure, occurs when the lateral edges of the germ band meet and fuse along the dorsal midline of the embryo's body. Ectoderm cells grow and differentiate to form the epidermis, the brain and nervous system, and most of the insect's respiratory (tracheal) system. In addition, the ectoderm invaginates (folds inward) at the front and rear of the embryo's body to create front and rear portions of the digestive system (foregut and hindgut). Mesoderm cells differentiate to form other internal structures such as muscles, glands, heart, blood, fat body, and reproductive organs. The midgut develops from a third germ layer (the endoderm) that arises near the fore- and hindgut invaginations and eventually fuses with them to complete the alimentary canal Developmental Fate of Insect Germ Layers Ectoderm: Epidermis, exocrine glands, brain and nervous system, sense organs, foregut and hindgut, respiratory system, external genitalia. Mesoderm: Heart, blood, circulatory system, muscles, endocrine glands, fat body, gonads (ovaries and testes). Endoderm: Midgut. During its early development, the embryo's body is rather worm-like in appearance. Individual segments first become visible near the anterior end (the protocephalon) where ectodermal tissue differentiates into the brain and compound eyes. Bud-like swellings develop in front of the mouth opening. They will eventually grow to form the labrum and the antennae. Segments behind the mouth also develop bud-like swellings. Each of the first three post-oral segments form paired appendages that become mouthparts: mandibles, maxillae, and labium. The next three post-oral segments develop into the thorax -- they form appendages that become walking legs. Segments of the abdomen also develop limb buds but these soon shrink and disappear -- perhaps they are vestigal remnants of abdominal appendages found in more primitive arthropods (like millipedes and centipedes). In general, the rate of embryonic development depends on temperature (insects are poikilothermic) and on species-specific characteristics of development. Embryogenesis ends when the yolk's contents have been consumed: the immature insect is fully formed and ready to hatch from the egg. During the hatching process (often called eclosion) the young insect may chew its way through the egg's chorion or it may swell in size by imbibing air until the egg shell "cracks" along a predetermined line of weakness. Formations of inner layer Three stages of embryonic development A protopod B polypod C oligopod Embryonic development of tobacco hornworm .Manduca sexta eggs. M. sexta embryo 19 M. sexta egg showing micropyle hours after fertilization M. sexta embryo 57 hours after fertilization M. sexta embryo 115 hours after fertilization. M. sexta embryo 37 hours afterfertilization Newly emerged larva showing the head Egg hatching Insect eggs close to hatching showing embrios with red "eye-spots". Once the hatchling emerges, it is called a first instar nymph (or larva). As it grows, it will continue to develop and mature. These post-embryonic changes are known as morphogenesis. MORPHOGENESIS Once an insect hatches from the egg it is usually able to survive on its own, but it is small, wingless, and sexually immature. Its primary role in life is to eat and grow. If it survives, it will periodically outgrow and replace its exoskeleton (a process known as molting). In many species, there are other physical changes that also occur as the insect gets older (growth of wings and development of external genitalia, for example). Collectively, all changes that involve growth, molting, and maturation are known as morphogenesis. Instar Timeline of MORPHOGENESIS Molting The molting process is triggered by hormones released when an insect's growth reaches the physical limits of its exoskeleton. Each molt represents the end of one growth stage (instar) and the beginning of another In some insect species the number of instars is constant (typically from 3 to 15), but in others it may vary in response to temperature, food availability, or other environmental factors. Molting stops when the insect becomes an adult -energy for growth is then channeled into production of eggs and sperm. An insect cannot survive without the support and protection of its exoskeleton, so a new, larger replacement must be constructed inside the old one -- much like putting an overcoat under a sweater! The molting process begins when epidermal cells respond to hormonal changes by increasing their rate of protein synthesis. This quickly leads to apolysis -- physical separation of the epidermis from the old endocuticle. Epidermal cells fill the resulting gap with an inactive molting fluid and then secrete a special lipoprotein (the cuticulin layer) that insulates and protects them from the molting fluid's digestive action. This cuticulin layer becomes part of the new exoskeleton's epicuticle After formation of the cuticulin layer, molting fluid becomes activated and chemically "digests" the endocuticle of the old exoskeleton. Break-down products (amino acids and chitin microfibrils) pass through the cuticulin layer where they are recycled by the epidermal cells and secreted under the cuticulin layer as new procuticle (soft and wrinkled). Pore canals within the procuticle allow movement of lipids and proteins toward the new epicuticle where wax and cement layers form. When the new exoskeleton is ready, muscular contractions and intake of air cause the insect's body to swell until the old exoskeleton splits open along lines of weakness (ecdysial sutures). The insect sheds its old exoskeleton (ecdysis) and continues to fully expand the new one. Over the next few hours, sclerites will harden and darken as quinone cross-linkages form within the exocuticle. This process (called sclerotization or tanning) gives the exoskeleton its final texture and appearance An insect that is actively constructing new exoskeleton is said to be in a pharate condition. During the days or weeks of this process there may be very little evidence of change. Ecdysis, however, occurs quickly (in minutes to hours). A newly molted insect is soft and largely unpigmented (white or ivory). It is said to be in a teneral condition until the process of tanning is completed (usually a day or two). Summary of Molting Step Step Step Step Step Step Step Step Step 1: 2: 3: 4: 5: 6: 7: 8: 9: Apolysis -- separation of old exoskeleton from epidermis Secretion of inactive molting fluid by epidermis Production of cuticulin layer for new exoskeleton Activation of molting fluid Digestion and absorption of old endocuticle Epidermis secretes new procuticle Ecdysis -- shedding the old exo- and epicuticle Expansion of new integument Tanning -- sclerotization of new exocuticle Exoskeleton traits fixed in size new exoskeleton Incorporates the changes that are part of metamorphosis. Initially soft and is larger than the old exoskeleton. Therefore, change in body form comes about as a series of steps. Stages between each molt are called instars. first stage which emerged from the egg is the first instar or nymph. -- to grow must shed its skin or molt -- may be four or five instars before the adult stage is reached Cicada Ecdysis An adult cicada (Homoptera) just after molting Metamorphosis In nearly all insects growth involves a metamorphosis, that is, a transformation in form and in way of life, especially from larva to adult Types of Metamorphosis anamorphosis epimorphosis prometamorphosis Incomplete metamorphosis hemimetamorphosis paurometamorphosis hyperpaurometamorphosis complete metamorphosis hypermetamorphosis anamorphosis Abdomen segments added with each molt: 9-12 Little or no change between the immature and adult form except in size and development of the sexual organs. Protura epimorphosis In a few very primitive, wingless insects (such as the silverfish) there is no metamorphosis. The insect emerges from the egg as a miniature adult and the only futher changes are in size and in maturation of the reproductive organs. Collembola, diplura, thysanura Prometamorphosis Nymphs of mayflies have 1245 aquatic instars, and wings are visible in older nymphs. Between the nymph stage and adult stage there is a subimago The subimago is fully winged and flying Mayfly nymphs Incomplete metamorphosis Incomplete, or gradual, metamorphosis is seen in members of less advanced orders (such as locusts and their relatives and the true bugs). The larva, often called a nymph (or, if aquatic, a naiad) is usually similar in form to the adult, but lacks wings. The wings begin as external bumps on the larva, and the adult emerges from the last molt without having undergone a pupal stage. Incomplete Metamorphosis 3 Insect Stages Eggs Larvae • Body form resembles adult • No wings Adults • No increase in size • Reproduction • Wings fully grown if present Example: Squash Bug HOMOPTERA / HETEROPTERA (THE TRUE BUGS: SPITTLE BUGS, APHIDS, ETC.) paurometamorphosis hemimetamorphosis Of an insect with aquatic young undergoing incomplete metamorphosis in which the young does not resemble the adult The young stage is called naiads hyperpaurometamorphosis Thysanoptera are hemimetabolous Their final immature stage is quiescent, non-feeding, and enclosed in a silken cocoon. This developmental stage, called a "pupa’’ Thrips represent an "intermediate" stage between hemi- and holometabolous development. Male scale Complete metamorphosis Complete metamorphosis is characteristic of over 80% of all insect species The wingless, wormlike larva is completely unlike the adult, and its chief activities are eating and growing. After several molts the larva enters a quiescent stage called the pupa; the pupa does not eat and usually does not move, but within the exoskeleton a major transformation occurs that involves the reorganization of organ systems as well as the development of such adult external structures as wings and compound eyes. In some insects the pupa is enclosed in a protective case, called the cocoon, built by the larva just before pupation. When the transformation is complete the final molt occurs: the adult emerges, its wings fill with blood and expand, and the new exoskeleton hardens. The chief function of the adult is propagation; in some species it does not eat. Do small butterflies grow up to be big butterflies? Do small butterflies grow up to be big butterflies? Complete Metamorphosis 4 Insect Stages – Eggs – Larvae – Pupae • Transformation from larva to adult • True legs, wings, antennae are formed – Adults • No increase in size • Reproduction • Short Life span LEPIDOPTERA (BUTTERFLIES AND MOTHS) COLEOPTERA (BEETLES) DIPTERA (FLIES) Adult and immature insects with complete metamorphosis feed on the different food egg 1st 2nd instar larva 3rd pupa adult The Emergence Of A Monarch The Monarch chrysalis is one of nature's most beautiful creations. This butterfly wears a crown of gold on jade green. About 24 hours before the emergence of the adult butterfly, the chrysalis becomes completely transparent, revealing the new butterfly inside. Breaking free of the chrysalis, a Monarch greets the world. After struggling free of the chrysalis, the Monarch immediately begins to inflate its wings with a reservoir of blood contained in its swollen abdomen. As the wings inflate, the body of the butterfly attains its normal proportions. When the wings are fully inflated, the insect expels any excess fluid and rests. In a few hours, with its wings dried and hardened, the Monarch will take wing on its first flight. Hypermetamorphosis Meloidae or blister beetles, undergo what is referred to as hypermetamorphosis. In this situation, growth is by complete metamorphosis, but there are distinct changes in external form and habits at each successive larval molt