Insects physiology. Lecture 1
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1 Insects physiology Lecture 1 1
2 Introduction The components that constitute the exoskeleton make an overwhelming contribution to the terrestrial success that arthropods can claim. Like the skin of vertebrates, the exoskeleton completely covers the insect and additionally provides an armor-like protective suit as a skeleton. Its most critical function is to serve as an interface between the insect and the environment, providing a barrier for the movement of water, ions, parasites, and environmental chemicals including insecticides. Introduction This barrier is especially significant for small animals like insects that have a high surface-to-volume ratio and therefore present a relatively large amount of surface area to the environment. The nature of the exoskeleton has thus had profound implications for growth, respiration, locomotion, and, from a human perspective, the design of chemicals that must penetrate the integument to be used as control agents. The exoskeleton also plays an important structural role in determining the form of the insect body and making possible the dramatic changes in form that accompany metamorphosis. 2
3 Introduction Insects exploit a variety of diverse habitats and diets,made possible by a developmental plasticity in body form and mouthpart structure. The rigidity that it provides allows for the insertion of muscles that can produce more precise locomotormovements than can the soft hydrostatic exoskeletons of the annelid worms. Introduction Although being surrounded by a rigid suit of 3
4 armor might limit the movement and environmental awareness of insects, the integument that makes up the exoskeleton is elastic in some areas to make flying and walking possible. Numerous sensory receptors that are concentrated in strategic areas provide windows to the outside world that allow the insect to respond appropriately to the environment. The terms integument and exoskeleton are often used interchangeably, depending on which functional component is under consideration. 4
5 The integument may comprise up to half the dry weight of some insects,representinga major investment of raw materials. However, because much of this is resorbedduring molting and even periods of starvation, the integument could also be viewed as a food reserve. The hydrocarbons that are deposited in the exoskeleton are responsible for releasing particular behavioral sequences that are involved in mating. Specific structures required for mate recognition, as well as chemicals such as pheromones and pigments that are deposited in the exoskeleton, are releasers for the stereotyped behaviors that are necessary for mating to occur. All of these features are provided by a single layer of epidermal cells and their secretions. 5
6 The exoskeleton provides a number of advantages, it does pose a major problem for growth. For insects with rigid exoskeletons to undergo significant amounts of growth, a new, larger exoskeleton must first be synthesized and the older one discarded. During this period of molting, the insect is relatively helpless against predators because flight or defense is difficult. Molting consumes time, energy, and metabolic resources. There is also a potential susceptibility for the loss of water because the insect can neither drink nor adjust its body to a changing environment. To reduce this susceptible period during the molting cycle, more advanced insects have evolved toward a reduced number of molts. Growth during the intermoltperiod is possible because the larvae of advanced holometabolousinsects generally have relatively unsclerotizedcuticles that can undergo a degree of stretching. 6
7 The growth and development of insects is largely a function of the growth and development of their integuments. The cuticularmolts that punctuate postembryonic growth are necessary if hard-bodied insects are to undergo any significant increases in size. Increases in body size do not always follow from molting Insects that are starved during the larval stage or molt to a diapauseform may actually molt to smaller individuals if they molt at all. 7
8 Apterygoteinsects continue to molt into the adult stage, but pterygote insects are incapable of molting as adults.. The inability of adult pterygote insects to molt is probably the result of the degeneration of the epidermal cells that produce the wing once it is formed. After the molt to the adult stage, the epidermal cells that make up the wing degenerate and the loss of the water contained within them makes it possible for the wing membranes to move rapidly for fl ight 8
9 The three types of metamorphosis in insects possible for the wing membranes to move rapidly for flighhowever, these dead cells can no longer initiate a molt nor synthesize a new cuticle. Withoutliving epidermal cells, another wing cuticle could not be formed if the insect molted again. Thus, the death of the cells that make flight possible also makes molting as an adult impossible. Only the winged mayfly subimagois capable of a molt to another winged form, but the subimagois a poor flier because living epidermal cells must remain as part of the wing 9
10 In many holometabolousinsects, considerable growth can occur during a single larval instarin the absence of a molt because, with the exception of the head capsule, the cuticle is extensible enough to accommodate some increases in size. The last instarof Manducasextacan grow from 1 g to over 9 g in weight without a molt because the pleated outer epicuticleof the exoskeleton is able to stretch to accommodate the growth that occurs within this instar. A molt may ultimately be necessary in order to acquire a larger head capsule and allow the sclerotized mouthparts to increase in size so the rate of food intake isincreasedto satisfy the demands of the larger body. The molting cycle and metamorphosis present some interesting developmental conditions. The particular developmental stage of an insect is referred to as an instar or stadium, 10
11 INSTARS, STADIA, AND HIDDEN PHASES One of the first steps in molting is apolysis, in which the old cuticle separates from the epidermis and new cuticle begins to be produced. With the old cuticle no longer directly attached to the epidermis, it has effectively been discarded, although it has not been shed, and the newly formed cuticle now represents the cuticle of the next instar. INSTARS, STADIA, AND HIDDEN PHASES For this reason, apolysisis said to mark the passage to the next instar, even though ecdysishas not yet occurred and the insect appears to still be in the skin of the earlier instar. An instaris therefore defined as the period between two apolysesand begins when the insect first becomes detached from its old skin. 11
12 INSTARS, STADIA, AND HIDDEN PHASES The instarthat is hidden under the old, unshed cuticle before ecdysisis referred to as the pharate instar. This distinction is more important for some instars than others. For example, some lepidopterans undergo diapause as pharate adults that are developmentally complete adults enclosed by the detached pupal cuticle. Although based on an external examination, it is easy to conclude that if the insect diapauses as a pupa, then the diagnosis would not be correct. 12
13 INSTARS, STADIA, AND HIDDEN PHASES The stadium that an insect is in is defined by its ecdyses; a stadium represents the interval between one ecdysisand the next. Therefore, at apolysis, an insect passes to another instarbut does not become the next stadium until after ecdysis. Apolysis, the separation of the epidermal cells from the cuticle, marks the beginning of the molt and the next instar. INSTARS, STADIA, AND HIDDEN PHASES The insect is an a pharate stage until ecdysisoccurs, the casting off the old cuticle. Ecdysismarks the beginning of the next stadium. At the apolysisfollowing the second instar, insect enters the third instarbut is still in the second until after ecdysis 13
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