Insects, Their Ways and Means of Living/Chapter IV

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IN our human society each individual must obtain the things necessary for existence; the manner by which he acquires them, whether by one trade or another, by this means or by that, does not physically matter so long as he provides himself and his family with food, clothing, and shelter. Exactly so it is wit?a all forms of lire. The physical demands of living matter make certain things necessary for the maintenance of lire in that matter, but nature has no law specifying that any necessity shall be acquired in a certain manner. Lire itself is a circumscribed thing, but it bas complete freedom of choice in the ways and means of living. It is useless to attempt to make a definition of what living matter is, or of how it differs from non-living matter, for all definitions have failed to distinguish animate from non-animate substance. But we all know that living things are distinguishable from ordinary non-living things by the fact that they make some kind of response to changes in the contact between themselves and their environment. The "environment," of course, must be broadly interpreted. Biologically, it includes all things and forces that m any way touch upon living matter. Not only bas every plant and animal as a whole its environment, but every part of it has an environment. The cells of an animal's stomach, for example, have their environment in the blood and lymph on one side, the contents of the stomach on the other; in the energy of the nerves distributed to them; and in the effects of heat and cold that penetrate them.

The environmental conditions of the life of cells in a complex animal are too complicated for an elemental study; the elements of lire and its basic necessities are bet- ter understood in a simple organism, or in a one-celled animal; but for purposes of description, it is most con- venient to speak of the properties of mere protoplasm. AIl the vital needs of the most highly organized animal are .pr.esent in any part of the protoplasmic substance of which ?t ,s composed. Protoplasm is a chemical substance, or group of sub- stances, the structure of which is very comp]ex but is main- tained so long as there is no disturbance in the environ-

'??°- °?? BCIs ?BCIs ?BC]s A ]3 C FIG. 6?. Diagram show]ng the relation of the germ ce]ls (GCIs) and the body cells (BCIs) in successive generations A fertilized germ cell of generation A forms the germ cells and b?y cells of B, a fert]lized germ cell of B forms the germ cells and b?y cells of C, and so on. The offspring C of B derives nothing from the body cells of the parent B, but both offsp6ng C and parent B have a oemmon origin in a germ cell of A

metat. Let some least thing happen, however, such as a change in the temperature, in the strength of the light, in the weight of pressure, or in the chemical composition of the surrounding medium, and the protoplasmic molecules, in the presence ofoxygen, are likely to have the balance of their constituent particles upset, whereupon they partly decompose by the union of their less stable elements with oxygen to form simpler and more permanent compounds. The decomposition of the protoplasmic substances, like all processes of decomposition, liberates a certain amount of energy that had been stored in the making of the molecule, and this energy may manifest itselfin various ways. Ifit

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takes the form of a change of shape in the protoplasmic mass, or movement, we say the mass exhibits signs of life. The state of being alive, however, is more truly shown if the act can be repeated, for the essential property of living matter is its power of reverting to its former chemical composition, and its ability thus gained of again reacting to another change in the environment. In restoring its lost elements, it must get these elements anew from the environment, for it can not take them back from the sub- stances that have been lost. Here, expressed in its lowest terres, is the riddle of the physical basis of lire and of the incentive to evolution in the forms of lire. Not that these mvsteries are any more easily understood for being thus analyzed, but they are more nearly comprehended. Being alive is maintaining the power of repeating an action; it involves sensitivity to stimuli, the constant presence of free oxygen, elimination of waste, and a supply of substances from which carbon, hydrogen, nitrogen, and oxygen, or other necessary ele- ments, are readily available for replacement purposes. Evolution results from the continual effort of living matter to perform its lire processes in a more efficient manner, and the different groups of living things are the result of the different methods that lire has tried and found advan- tageous for accolnplishing its ends. Living organisms are machines that have become more and more complex in structure, but always for doing the same things. If animals may be compared with machines in their physical mechani?m, they are like them, too, in the fact that they wear out and are at last beyond repair. But here the simile ends, for when your car will no longer run, you must go to the dealer and order a new one. Nature provides continuous service by a much better scheme, for each organism is responsible for its own successor. This phase of lire, the replacement of individuals, opens another subject involving ways and means, and it, likewise, can be understood best in its simpler manifestations.

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The facts of reproduction in animais are not well ex- pressed by our name for them. Instead of "reproduc- tion," it would be-truer to say "repeated production," for individuals do not literally reproduce themselves. Genera- tions are serially related, not each to the preceding; they follow one another as do the buds along the twig of a tree,

Flç. 6 3. The external structure of an insect The body of a grasshopper dissected showing the head (H), the thorax and the abdomen (.4b). The head carries the eyes (E), the antennae (,llnt), and the mouth parts, which include the labrum (Lrn), the mandibles (Md), the maxillae {Mx), and the labium (Lb). The thorax consists of three segments {I, 2,?), the flrst separate and carrying the first legs (L,), the other two com- bined and carrying the wings (1¢'?, 11"?), and the second and third legs (L,, L.?). The abdomen consists of a series of segments; that of the grasshopper bas a large tympanal organ (Tre), probably an ear, on each side of its base. The end of the abdomen carries the external organs of reproduction and egg-laying

and buds on the same twig are identical or nearly so, not because one produces the next, but because all are the result of the same generative forces in the twig. If the spaces of the twig between the buds were shortened until

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one bud became contiguous with the one before, or became enveloped by it, a relation would be established between the two buds similar to that which exists between succes- sive generations of lire forms. The so-called parent gen-

eration, in other words, contains the germs of the succeeding generation, but it does not produce them. Each generation is simply the custodian of the germ cells entrust- ed to i t, and the "off- spring" resembles the parent, not because it is a chip off the parental block, but because both parent and offspring are developed from the same line of germ cells. Parents create the conditions under wl?i, ch the germ cells will de- velop; they nourish and protect them during the period of their develop- ment; and, when each generation has served the purpose of its ex- istence, it sooner or later dies. But the in- dividuals produced from

F,c. 64. The leg of a young grasshopper, showing the typical segmentation of an insect's leg The leg is supported on a pleural plate (Pi) in the lateral wall of its segment. The basal segment of the free part of the leg is the coxa (Cx), then comes a small trochanter (Tf), next a long femur (F) separated by the knee bend from the tibia (Tb), and lastly the foot, consisting of a sub-segmented tarsus (Tar), and a pair of terminal claws (CI) with an ad- hesive lobe hetween them

its germ cells do the same

for another set of germ cells produced simultaneously with themselves, and so on as long as the species persists. To express the facts of succession in each specific form of animal, then, we should analyze each generation into germ cells and an accompanying mass of protective cells which



forms a body, or soma, the so-called parent. Both the body, or somatic, cells and the germ cells are formed from a single primary cell, which, of course, is usually produced by the union of two incomplete germ cells, a spermatozoon and an egg. The primary germ cell divides, the daughter cells divide, the cells of this division again divide, and the division continues indefinitely until a mass of cells is pro- duced. At a very early stage of division, however, two groups of cells are set apart, one representing the germ cells, the other the somatic cells. The former refrain from further development at this time; the latter proceed to build up the body of the parent. The relation of the somatic cells to the germ cells may be represented diagram- matically as in Figure 62, except that the usual dual par- enrage and the union of germ cells is not expressed. The sexual form of reproduction is not necessary with all lower animals, nor with all generations of plants; in some insects the eggs can develop without fertilization. The fully-developed mass of somatic cells, whose real function is that of a servant to the germ cells, has assumed such an importance, as public servants are prone to do, that we ordinarily think of it, the body, the active sentient animal, as the essential thing. This attitude on our part is natural, for we, ourselves, are highly organized masses of somatic cells. From a cosmic standpoint, however, no creature is important. Species of animais and plants exist because they have found ways and means of living that have allowed them to survive, but the physical universe cares nothing about them--the sunshine is hOt made for them, the winds are not tempered to suit their conven- ience. Life must accept what it finds and make the best of it, and the question of how best to further its own wel- fare is the problem that conffonts every species. The sciences of anatomy and physiology are a study of the methods by which the soma, or body, bas contrived to meet the requirements imposed upon it by the unchanging laws of the physical universe. The methods adopted are as

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numerous as the species of plants and animais that have existed since lire began. A treatise on entomology, there- fore, is an account of the ways and means of living that insects have adopted and perfected in their somatic organ- ization. Before discussing insects in particular, however, we must understand a little more fully the principal con-

ditions of living that na- ture places on all forms of lire. As we have seen, lire is a series of chemical re- actions in a particular kind of matter that can carry on these reactions. A "reaction" is an action; and every act of living matter involves a break- ing down of some of the substances in the proto- plasm, the discharging of the waste materials, and the acquisition of new materia]s to replace those lost. The reaction is in- herent in the physical or chemical properties of protoplasmic compounds and depends upon the substances with which the protoplasm is sur- rounded. It is the func-


.'Fb ?:?!?ïllen

Fm. 6 5. Legs of a honeybee, showing special modifications A, outer surface of a hind leg, with a pollen basket on the tibia (Tb) loaded with pollen. B, a fore leg, showing the antenna cleaner (a) between the tibia and the tarsus, and the long, halry basal segment of the tarsus (t Tar), which is used as a brush for cleaning the body

tion of the creature's mechanism to see that the con- ditions surrounding its living cells are right for the con- tinuance of the cell reactlons. Each cell must be provided with the means of eliminating waste material and of restoring its lost material, since it can not utilize that which it has discarded.

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With the conditions of living granted, however, proto- plasm is still only potentially alive, for there is yet required a stimulus to set it into activity. The stimulus for lire activities comes from changes in the physical forms of energy that surround or infringe upon the potentially living substance; for, "live" matter, like all other matter, is subject to the law of inertia, which decrees that it must remain at rest until motion is imparted to it by other


FIO. 66. The head and mouth parts of a grasshopper A, facial view of the head, showing the positions of the antennae (dnt), the large cornpound eyes (E), the slrnple eyes, or ocelli (O), the broad front lip, or labrum (Lin) suspended from the cranium by the clypeus (Clp), and the bases of the mandibles (Md, Md) closed behind the labrum B, the rnouth parts separated from the head in relative positions, seen from in front: Hphy, hypopharynx, or tongue, attached to base of labiurn; Lb, labiuml Lin, labrum; Md, mandibles; Mx, maxillae

motion. A very small degree of stimulating energy, how- ever, may result in the release of a great quantity of stored energy. The food of all living matter must contain carbon, hydrogen, nitrogen, and oxygen. The mechanism of plants enables them to take these elements from com- pounds dissolved in the water of the soli. Animais must get them from other living things, or from the products of

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living things. Therefore, animals principally have de- veloped the power of movement; they have acquired grasp- ing organs of some sort, a mouth, and an alimentary canal for holding the food when once obtained. In the insects, the locomotory function is subserved by the legs and by the wings. Since all these organs, the three pairs of legs and the two pairs of wings, are carried by the thorax (Fig. 63, Th), this region of the body is distinctly the locomotor center of the insect. The legs (Fig. 64) are adapted, by modifications of structure in different species, for walking, running, leaping, digging, climbing, swim- ming, and for many varieties of each of these ways of pro- gression, fitting each species for its particular mode of living and of obtaining its food. The wings of insects are important accesslons to their locomotory equipment, since they greatly increase their means of getting about, and thereby extend their range of feeding. The legs, fur- thermore, are often modified in special ways to perform some function accessory to feeding. The honeybee, as is well known, has pollen-collecting brushes on its front legs (Fig. 65 B), and pollen-carrying baskets on its hind legs (A). The mantis, which captures other insects and eats them alive, bas its front legs made over into those efficient organs for grasping its prey and for holding the struggling victim which have already been described (Fig. 46). The principal organs by which insects obtain and ma- nipulate their food consist ?)f a set of appendages situated on the head in the neighborhood of the mouth, which, in their essential structure, are of the nature of the legs, for insects have no jaws comparable with those of vertebrate animais. The mouth appendages, or mouth parts as they are called, are very different in form in the various groups of insects that have different feeding habits, but in all cases they consist of the same fundamental pieces. Most important is a pair of jawlike appendages, known as the mandibles (Fig. 66 B, Md), placed at the sides of the mouth (A, Md), where they swing sidewise and close upon each other

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below the rnouth. Behind the rnandibles is a pair of maxdlae lB, Mx) of more cornplicated forrn, fitted rather for holding the food than for crushing it. Following the rnaxillae is a large under lip, or labium I Lb), having the

Ant -? C? Ht .-? /?I?] An ? Vent SoeGng ?C 6 7. Lençthw?se se«t{on of ? gr?shopper, show?ng the ?en?r?l ?he pnn«?p?/ mtoEn?l ornons, excep? the resp{?tory ?rache?l system and the organs of repr?uction anus; .g.t, anoenna; Br, brain; Cr, ?op: Ht, heart: l.t, intestine: Malpighian tubules: Mt?. mouth: Oe, oesophagus; SoeGn?, su?esoph?eal ganghon; ?ent, stomach (vent6culus); ?.VC, ventral nerve ?rd; ?, wings

structure of two rnaxi[lae united by their inner rnargins. A broad flap hangs downward befo're the rnouth to forrn an upper lip, or labrum (Lm). Between the rnouth ap- pendages and attached to the front of the Iabiurn there is a large rnedian lobe of the lower head wall behind the rnouth, known as the h.vpopharynx (Hphy). Insects feed, sorne on solid foods, others on [iquids, and their rnouth parts are rnodified accordingly. So it comes about that, according to their feeding habits, insects rnay be separated into two groups, which, like the fox and the stork, could not feed either at the table of the other. Those insects, such as the grasshoppers, the crickets, the beetles, and the caterpillars, that bite off pieces of food tissue and chew thern, have the rnandib[es and the other rnouth parts of the type described above. Insects that partake only of liquids, as do the plant lice, the cicadas, the rnoths, the butterflies, the rnosquitoes and other flies, have the

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mouth parts fitted for sucking, or for piercing and sucking. Some of the sucking types of mouth parts will be described in other chapters (Figs. ???, ?63, ?83) , but it will be seen that all are merely adaptations of form based on the ordi- nary biting type of mouth appendages. The fossil records of t'he history of insects show that the sucking insects are the more recent products of evolution, since all the earlier kinds of insects, the cockroaches and their kin, have typical biting mouth parts. The principal thing to observe concerning the organs of feeding, in a study of the physiological aspect of anatomy, is that they serve in all cases to pass the natural food materials from the outside of the animal into the alimen- tary canal, and to l?ive them whatever crushing or masti- canon is necessarv. It is within the alimentary canal, therefore, that the next steps toward the final nutrition of the animal take place. The a]imentary cana] of most insects is a simple tube (Fig. 68), extend'ing either straight through the body, or

FI{I. 68. The alimentary canal of a grasshopper ?IInt, anterior intestine; dn, anus; Cr, crop; GC, gastric caeca, pouches of the stomach; Hphy, hypopharynx (tongue); Lb, base of labium; Mal, Malpighian tubules; Mlnt, raid-intestine; l?lth, mouth; Oe, oesophagus; Rect, hind intestine (rectum); SIGI, salivary glands opening by their united ducts at base of hypo- pharynx; I/ent, ventriculus (stomach)

making only a few turns or loops in its course. It con- sists of three principal parts, of which the middle part is the true stomach, or ventriculus (Fent) as it is called by insect anatomists. The first part of the tube includes a

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pharynx irnrnediately behind the rnouth, followed by a narrower, tubular oesophagus (Oe), after which comes a sac- like enlargernent, or crop (Cr), in which the food is tern- porarily stored, and finally an antecharnber to the stornach, narned the proventriculus. The third part of the alimen- tary canal, connecting the stomach with the anal opening, is the intestine, usually cornposed of a narrow anterior part, and a wide posterior part, or rectum (Rect). Muscle layers surrounding the entire alirnentary tube cause the food to be swallowed and to be passed along from one section to the next toward the rear exit. With the taking of the food into the alirnentary canal, the/natter of nutrition is by no rneans accornplished, for the animal is still confronted with the problern of getting the nutrient rnaterials into the inside of its body, where alone they can be used. The alirnentary tube bas no openings anywhere along its course into the body cavity. Whatever food substances the tissues of the animal receive, therefore, rnust be taken through the walls of the tube in which they are inclosed, alad this transposition is accorn- plished by dissolving thern in a liquid. Most of the nutri- ent rnaterials in the raw food rnatter, however, are not soluble in ordinary liquids; they rnusr be changed chern- ically into a forrn that will dissolve. The process of get- ring the nutrient parts of the raw foodstuff into solution constitutes digestion. The digestive liquids in insects are furnished mostly by the stornach walls or the walls of tubular glands that open into the stornach, but the secretion of a pair of large glands, called the salivary glands (Fig. 68, SIGI), which open between the rnouth parts, perhaps bas m sorne cases a digestive action on the food as it is taken into the mouth. Digestion is a purely chemical process, but it must be a rapid one. Consequently the digestive juices contain not only substances that will transforrn the food rnaterials into soluble cornpounds, but other substances that will

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speed up these reactions, for otherwise the animal would starve on a full stomach by reason of the slowness of its gastric service. The quickening substances of the diges- tive fluids are called enzymes, and each kind of enzyme acts on only one class of food material. An animal's prac- tical digestive powers, therefore, depend entirely upon the specific enzymes its digestive liquids contain. Lacking this or that enzyme it can not digest the things that depend upon it, and usually its instincts are correlated with its enzymes so that it does not fill its stomach with food it can not digest. A few analyses of the digestive liquids of in- sects have been made, enough to show that their digestive processes depend upon the presence of the same enzymes as those of other animais, including man. The grosser digestive substances, in cooperation with the'enzymes, soon change all the parts of the food ma- terials in the stomach that the animal needs for its suste- nance into soluble compounds which are dissolved in the liquid part of the digestive secretions. Thus is produced a rich, nutrient juice within the alimentary canal which can be absorbed through the walls of the stomach and intes- tine and can so enter the closed cavity of the body. The next problem is that of distribution, for still the food ma- terials must reach the individual cells of the tissues that compose the animal. The insect's way of feeding, of digesting its food, and of absorbing it is not essentially different from that of the higher animais, including ourselves, for alimentation is a very old and fundamental function of all animais. Its means of distributing the digested food within its body, however, is quite different from that of vertebrates. The absorbed pabulum, instead of being received into a set of lymphatic vessels and from these sent into blood-filled tubes to be pumped to all parts of the organism, goes directly from the alimentary walls into the general body cavity, which is filled with a liquid that bathes the inner surfaces of all the body tissues. This body liquid is called

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the "blood" of the insect, but it is a colorless or slightly yellow-tinted lymph. It is kept in motion, however, by a pulsating vessel, or heart, lying

Fro. 69. Diagram of the typical structure of an insect's heart and supporting dia- phragm, with the course of the circulating blood marked by ,?o, aorta, or anterior tubular part of the heart without lateral openings; Dpk, mem- branous diaphragm; Ht, ante- rior three chambers of the heart, which usually extends to the posterior end of the body; Md, muscles of dia- phragm, the fibers spreading from the body wall to the heart; Ost, ostium, or oneofthe lateral openings into the heart chambers

in the dorsal part of the body; and by this means the food, now dissolved ira the body liquid, is carried into the spaces between the various organs, where the cells of the latter can have access to it. The heart of the insect is a slender tube suspended along the midline of the back close to the dorsal wall of the body (Fig. 67, 1/t). It has intake apertures along its sides I Vig. 69, Ost), and its anterior end opens into the body cavitv. It pulsates for- ward, by means of muscle fibers in its walls, thereby sucking the blood in through the lateral openings and discharging it by way of the front exit. An im- perfect circulation of the blood is thus established through the spaces between the organs of the body cavity, sufficient for the purposes of so small an animal as ara insect. The final act of nutrition comes now when the blood, charged with the nutrient materials ab- sorbed from the digested food in the alimentary canal, brings these materials into contact with the inner tissues. The tissue cells, by the inherent power of

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all living rnatter (which depends on the laws of osrnosis and on chernical affinity), take for thernselves whatever they need from the menu offered by the blood, and with this rnatter they build up their own substance. It is evident, therefore, that the blood rnust contain a suffi- cient quantity and variety of dietary elernents to satisfy all possible cell appetites; that the stornach's walls and their associated glands must furnish the enzyrnes appro- priate for rnaking the necessary elernents available from the raw food rnatter in the stornach; and, finally, that it rnust be a part of the instincts of each animal species to consume such native foodstuffs of its environment as will supply every variety of nourishing elernents that the cells dernand. As we have seen, the dernand for food comes from the loss of rnaterials that are decornposed in the tissues during cell activitv. Better stated, perhaps, the chernical break- down with[n the cell is the cause of the cell activity, or is the cell activity itself. The way in which the activity is expressed does not rnatter; whether by the contraction of a muscle cell, the secretion of a gland cell, the generation of nerve energy by a nerve cell, or just the rninirnurn activity that rnaintains'life, the result is the same always--the loss of certain substances. But, as with most chernical reduc- tion processes, the protoplasrnic activity depends upon the presence of available oxygen; for the decornposition of the unstable substances of the protoplasrn is the result of the affinity of sorne of their elernents for oxygen. Conse- quently, when the stimulus for action comes over a nerve from a nerve center, a sudden reorganization takes place between these protoplasrnic elernents and the oxygen atorns which results in the formation of water, carbon dioxide, and various stable nitrogenous cornpounds. The substances discarded as a result of the cell activi- ties are waste products, and nlust be elirninated from the organisrn for their presence would clog the further activity of the cells or would be poisonous to thern. The anirnai,



therefore, must have, in addition toits mechanisms for bringing food and oxygen to the cells, a means for the re- moral of wastes. The supplying of oxygen and the removing of carbon dioxide and some of the excess water are accomplished by respiration. Respiration is primarily the exchange of gases between the cells of the body and the outside air. If an animal is suflîciently small and soft-skinned, the gas ex- change can be made directly by diffusion through the skin. Larger animais, however, must have a device for conveying air into the body where the tissues wil| have closer access toit. It will be evident, then, that there is not neces- sarily only one way of accomplishing the purposes of respiration. Vertebrate animais inhale air into a sac or pair of sacs, called the lungs, through the very thin walls of which the oxygen and carbon dioxide can go into and out of the blood respectively. The blood contains a special oxygen carrier in the red matter, hemog|obin, of its red corpuscles, by means of which the oxygen taken in from the air is transported to the tissues. The carbon dioxide is carried from the tissues partly by the hemoglobin, and partly dissolved in the blood liquid. Insects have no lungs, nor have they hemoglobin in their blood, which, as we have seen, is merely the liquid that fills the spaces of the body cavity between the organs, lnsects have adopted and perfected a method of getting air dis- tributed through their bodies quite different from that of the vertebrates. They have a system of air tubes, called tracheae (Fig. 7o), opening from the exterior by small breathing pores, or spiracles (Sp), along the sides of the body, and branching minutely within the body to all parts of the tissues. By this means the air is conveyed directly to the parts where respiration takes place. There are usually in insects ten pairs of spiracles, two on the sides of the thorax, and eight on the abdomen. The spiracles communicate with a pair of large tracheal trunks lying



along the sides of the body (Fig. 7o), and from these trunks are ?iven off branches into each body segment and into the head, which go to the alimentary canal, the heart, the nervous system, the muscles, and to all the other organs, where they break up into finer branches that

terminate in minute end tubes going practically to every cell of the body. Many insects breathe by regular movements of expansion and contrac- tion of the under surface of the abdo- men, but experimenters have not yet agreed as to whether the air goes in and out of the same spiracles or whether it enters one set and is ex- pelled through another. It is probable that the fresh air goes into the smaller tracheal branches principally by gas diffusion, for some insects make no perceptible respiratory movements. The actual exchange of oxygen from the air and carbon dioxide from the tissues takes place through the rhin walls of the minute end tubes of the tracheae. Since these tubes lie in im- mediate contact with the cell surfaces the gases do not have to go far in order to reach their destinations, and the insect has little need of an oxygen carrier in its blood--its whole body, practically, is a lung. And yet some investigations have made it appear likely that the insect blood does con- tain an oxygen carrier that functions in a manner similar to that of the hemoglobin of vertebrate blood, though the importance of oxygen transportation in insect physiology has


F?e. 7 ?. Respirator l' system of a eaterpillar. The external breathing apertures, or spiracles (S/?, S/?), along the sides of the body open into lateral tracheal trunks (a, a), which are connected crosswise by transverse tubes (b, b) and give off mi- nutely branching tra- cheae into all parts of the head (H) and body


hOt been determined. In any case, the tracheal method of respiration must be a very efficient one; for, consider- ing the activity of insects, especially the rate at which the wing muscles act during flight, the consumption of oxygen must at times be pretty high. The activity of insects depends very much, as every one knows, upon the temperature. We have all observed how the bouse flies disappear upon the first cold snap in the fall and then surprise us by showing up again when the weather turns warm, just after we have taken down the screens. AIl insects depend largely upon external warmth for the heat necessary to maintain cellu[ar activity. While their movements produce heat, they have no means of con- serving this heat in their bodies, as have "warm-blooded" animais. That insects radiate heat, however, is very evident from the high temperature that bees can maintain in their hives during winter by motion of the wings. All insects exhale much water vapor from their spiracles, an- other evidence of the production of heat in their bodies. The solid matter thrown off from the cells in activity is discharged into the blood. These waste materials, which are mostly compounds of nitrogen in the form of salts, must then be removed from the b]ood, for their accumula- tion in the body would be injurious to the tissues. In vertebrate animais, the nitrogenous wastes are eliminated by the kidneys. Insects have a set of tubes, comparable with the kidneys in function, which open into the intestine at the junction of the latter with the stomach (Fig. 68, Mal), and which are named, after their discoverer, the Malpighian tubule«. These tubes extend through the prin- cipal spaces of the body cavity, where they are looped and tangled like threads about the other organs and are continually bathed in the blood. The cells of the tube walls pick out the nitrogenous wastes from the blood and discharge them into the intestine, whence they are passed to the exterior with the undigested food refuse. We thus see that the inside of an insect is hOt an unor-



ganized mass of pulp, as believed by those people whose education in such naatters comes principally from under- foot. The physical unity of all fornas of lire makes it neces- sary that every creature naust perforna the sanae vital functions. The insects have, in naany respects, adopted their own wavs of accomplishing these functions, but, as already pointC out, the naeans of doing a thing does not count with nature so long as the end results are attained. The essential conditions are the supply of necessities and the removal of wastes. The body of a conaplex aninaal naay be likened to a great factory, in which the individual workers are represented by the cells, and groups of workers by the organs. That the factory naay accornplish its purpose, the activities of each worker naust be coordinated with those of all the other workers by orders fro?n a directing office. Just so, the ac- tivities of the cells and organs of the animal must be con- trolled and coordinated; and the directing office of the aninaal organization is the central nervous systena. The work of almost every cell in the body is ordered and con- trolled by a "nerve inapulse" sent toit over a nerve fiber frona a nerve center. The inner structure of the nervous tissues and the work- ing mechanisna of the nerve centers are essentially alike in all animals, but the form and arrangenaent of the nerve tissue masses and the distribution of the nerve fibers naay differ nhuch according to the plan of the general body or- ganization. The insects, instead of following the verte- brate plan of having the central nerve cord along the back inclosed in a bony sheath, have round it just as well for their purposes to l?ave the principal nerve cord lying free in the lower part of the body (Fig. 67, I'.VC). In the head there is a brain (Figs. ?7, 7 -?, Br) situated above the oesophagus (Fig. 67, Oe), but it is connected by a pair of cords with another nerve mass below the pharynx in the lower part of the head (SoeG?lg). Frona this nerve naass another pair of nerve cords goes to a third nerve mass



D o

Fro. 7 ?. The nervous system of the head of a grass- hopper, as seen by removal of the facial wall .4ntNv, antennal nerve; 1Br, gBr,3Br, the three parts of the brain; CoeCon, circumoesophageal connectives; 3Corn, suboesophageal commissure of the third Iobes of the brain; FrGng, frontal ganglion? FrCon, frontal ganglion connective with the brain; LbNv, labial nerve; LraNv, labral nerve; MdNv mandibular nerve; MxNv, maxillary nerve; O, simple eye; OpL, optic lobe connected with the brain; RNv, recurrent nerve; 8oeGng, suboesophageal ganglion

lying against the lower wall of the first body seg- ment (Fig. 7?, Gng ?), which is likewise connect- ed with a fourth mass in the sec- ondsegment,and so on. The cen- tral nervous sys- tem of the insect thus consists of a series of small nerve masses united by double nerve cords. The nerve masses are known as gan- glia (Gng), and the uniting cords are called the connectives (Fig. 7?, Con). Typi- cally there is a ganglion for each of the first eleven body seg- ments, besides the brain and the lower gan- glioia of the head.

The brain of an insect (Fig. 7 ?) has a highly complex internal structure, but it is a less important controlling center than is the brain of a vertebrate animal. The other ganglia have much independence of function, each giving the stimuli for movements of its own segment. For this


reason, the head of an insect may be cut off and the rest of the creature may still be able to walk and to do various other things until it dies of starvation. Similarly, with some species, the abdomen may be severed and the insect will still eat, though the food runs out of the cut end of the alimentary canal. The detached abdomen may lay eggs, if properly stimulated. Though the insect thus appears to be largely a creature of automatic regulations, acts are hOt initiated without the brain, and full coordination of the functions is possible only when the entire nervous system is intact. The active elements of the nerve centers are nerve cells; the nerve fibers are merely conducting threads extended from the cells. If the nerve force that stimulates the other kinds of cells into activity comes from nerve cells, the question then arises as to whence comes the primary stimulus that activates the nerve cells. We must discard the old idea that nerve cells act automatically; being mat- ter, they are subject to the laws of matter--they are inert until compelled fo act. The stimulus of the nerve cells comes from something outside of them, either from the environmental forces of the external world or from sub- stances formed by other cells within the body. Nothing is known definitely of the internal stimuli of insects, but there can be no doubt that substances are formed by the physiological activiries of the insect tissues, similar to the horrnones, or secretions of the ductless glands of other animais, that control action in other organs either directly or through the nervous system. Thus, some in- ternal condition must prompt the insect to feed when its stomach is empty, and the entrance of food into its pharynx must stimulate the alimentary glands to prepare the diges- tive juices. Probably a secretion from the reproductive organs of the female, when the eggs are ripe in the ovaries, gives the stimulus for mating, and later sets into motion the reflexes that govern the laying of the eggs. The cater- pillar spins its cocoon at the proper rime for doing so; the




Gris'2 l+Ii+lll


GnçVl - - ..... VI

FIG. 7 2. The general nervous system of a grass- hopper, as seen from above /lnt, antenna; ?/o, aorta;/i'r, brain; Cer, cercus; E, compound eye; Gngl, ganglion of prothorax; Gng2, ganglion of mesothorax; Gng3+I+II+III, compound ganglion of metathorax, comprising the ganglia belonging to the metathorax and the first three abdominal segments; Gngll/oEngl/llI, ganglia of the fourth to eighth abdominal seg- ments; O, ocelli; Proc, proctodeum, or posterior part of alimentary canal; .ça, suranal plate; .çeglI--X, second to tenth segments of abdomen; .çoeGng, suboesophageal ganglion; .çtom, stomo- deum, or anterior part of alimcntary canal

stimulus, most like- ly, comes from the products of physio- logical changes be- ginning to take place in the body that will soon result in the transformation of the caterpillar into a chrysalis, a stage when the insect needs the protection of a cocoon. These activities of insects we call instincts, but the term is simply a cover for our ignor- ance of the processes that cause them. External stimuli are things of the outer environment that affect the living organism. They in- clude matter, elec- tromagnetic energy, and gravity; but the known stimuli do not comprise all the activities of matter or of the "ether." The common stimuli are: pressure of solids, liquids, and gases; humidity; chemical qualities (odors and tastes);

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sound, heat, light, and gravity. Most of these things stim- ulate the nerve centers indirectly through nerves connected with the skin or with specialized parts of the skin called sense organs. An animal can respond, therefore, only to those stimuli, or to the degrees of a particular stimulus, to which it is sensitive. If, for example, an animal has no re- ceptive apparatus for sound waves, it will not be affected by sound; if it is not sensitized to certain wave lengths of light, the corresponding colors will not stimulate it. There are few ki?Ms of natural activities in the environment that animais do not perceive; but even our own perceptive powers fMI far short of registering all the degrees of any activity that are known to exist and which the physicist tan measure. |nsects respond to most of the kinds of stimuli that we perceive by out senses; but if we say that they see, hear, smell, taste, or touch we make the implication that insects have COllSCiOUSlleSS. | t is most likely that their reactions to external stimuli are for the most part performed un- consciously, and that their behavior under the effect of a stimulus is an automatic action entirely comparable to out reflex actions. Behavioristic acts that result from reflexes the biologist calls tropi?m?. Coordinated groups of tropisms constitute an instinct, though, as we have seen, an instinct may depend also on internal stimuli. It can not be said that consciousness does hOt play a small part in determining the activities of some insects, es- pecially of those species in which memory, i.e., stored impressions, appears to give a power of choice between different conditions presented. The subject of insect psychology, however, ls too intricate to be discussed here. The phases of lire thus far described, the complexity of physical organization, the response to stimuli, the phenomena of consciousness from their lowest to their highest manifestations, all pertain to the soma. Yet» somehow, the plan of the edifice is carried along in the



germ cells, and by them the whole somatic structure is rebuilt with but little change of detail from generation to generation. This phase of lire activity is still a mystery to us, for no attempted explanation seems adequate to account for the organizing power resident in the germ cells that accomplishes the familiar facts of repeated

k-X DO? A B F?ç. 73- Di?rams ?f the internal organ? of reproduction in insects A, the female otgans, ?mprising a pair ?f ??aries (Or), ea«h «omp?sed ?f ? group ?f ?g tubul?s (oe), a pair ?f ovidu«ts (DOv),and a median ourlet tube,?r v?ina (g?), ?ith usually a pair of «olleterial glands (CIÇI) dis«ha?ing into the ?agina, and a sperm roEeptade» ?r spermath?a (?p?), opening from the up?r surface of the latter B, the male o?ans, comprising a pair of restes (Tes) com?s? of spermatic tubules, a pair of s?rm ducts, or vasa deferentia (?D), a pair of s?rm vesicles (gS), and an outlet tube, or ductus ejaculatorius (DES, with usually a pair of mucous glands (MGI) discharging into the ducts of the sperm vesicles development which we call repr?uction._ When we can explain the repetition of buds along the twig, we may have a key to the secret of the germ cells?and possibly to that of organic evolution. The organs that house the germ cells in the mature insect consist of a pair of ovaries in the female (Fig. 73 A, Or) in which the eggs mature, and of a pair of testes in the [???l


male (B, Tes) in which the spermatozoa reach their com- plete growth. Appropriate ducts connect the ovaries or the testes with the exterior near the rear end of the body. The female usually has a sac connected with the egg duct (A, Spm) in which the sperm, received at mating, are stored until the eggs are ready to be laid, when they are

VIII IX X Cet" I I / .-A t I

Fxc. 74- The oviposltor of a long-horned grasshopper, a member of the katydid family, showing the typical structure of the egg-laying organ of female insects A, the ovipos'itor (Ovp) in natural condition, projecting from near the posterior end of the body B, the parts of the ovlpositor separated, showing the six component pieces, two arising from theeighth abdominal segment (FIII), and four from the nlnth (IX). /ln, anus; Cet, cerci; IX, ninth abdominal segment; Ovp, ovipositor; FgO, vaginal opening; FIH, eighth abdominal segment; X» tenth abdominal segment extruded upon the latter and bring about fertilization. The egg cells ordinarily are all alike, but the spermatozoa are of two kinds; and according to the kind of sperm re- ceived by any particular egg, the future individual will be male or female.



The germ cells accompanying each new soma undergo a series of transformations within the parent body before they themselves are capable of accomplishing their pur- pose. They multiply enornaously. With some animais, only a few of them ever produce new members of the race; but with insects, whose motto is "safety in numbers," each species produces every season a great abundance of new individuals, to the end that the many forces arrayed against them may not bring about their extermination. The world seems full of forces oppos.ed to organized lire. But the truth is, all organization ?s an opposition to established forces. The reason that the forms of life now existing have held their places in nature is that they have found and perfected wavs and means of opposing, for a time, the forces that tend to the dissipation of energy. i.ife is a revolt against inertia. Those species that have died out are extinct, either because they came to the end of their resources, or because they became so inflexibly adapted to a certain kind-of lire that they were unable to meet the emergency of a change ira the conditions that made this lire possible. Eflïciency ira the ordinary means of living, rather than specialization for a particular way of living, appears to be the best guarantee of continued existence.

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