THE SENSORY ORGANS
a. The eyes and insect vision
Fig. 259.—Different forms of compound eyes. A, a bug (Pyrrhocoris). B, worker bee. C, drone. D, male Bibio, a holoptic insect.—From Judeich and Nitsche.
Of the eyes of insects there are two kinds, the simple and the compound. Of the former there are usually three, arranged in a triangle near the top of the head, between the compound eyes (Fig. 259, B). The compound or facetted eyes, which are usually round and prominent, differ much in size and in the number of facets.
The number of facets varies from 12 in Lepisma,—though in a Brazilian beetle (Lathridius) there are only seven unequal facets,—to 50 in the ant, and up to 4000 in the house-fly, 12,000 in Acherontia atropos, 17,000 in Papilio, 20,000 in the dragon-fly (Æschna), 25,000 in a beetle (Mordella), while in Sphinx convolvuli, the number reaches 27,000. The size of the facets seems to bear some relation to that of the insect, but even in the smallest species none have been observed less than 1
2000 of an inch in diameter. Day-flying Lepidoptera have smaller facets than moths (Lubbock).
Fig. 260.—Section through the ocellus of a young Dyticus larva: ct, cuticula; l, corneal lens; gh, cells of the vitreous body, being modified hypodermal cells (hy); st, rods; re, retinal cells; no, optic nerve.—After Grenacher, from Lang.
The simple, or single-lensed eye (ocellus).—Morphologically the simple eye is a modified portion of the ectoderm, the pigment enclosing the retinal cells arising from specialized hypodermal cells, and covered by a specialized transparent portion of the cuticula, forming the corneal lens. The apparatus is supplied with a nerve, the fibres of which end in a rod or solid nerve-ending, as in other sensory organs.
As seen in the ocellus of Dyticus (Fig. 260), under the corneal lens the hypodermis forms a sort of pit, and the cells are modified to form the vitreous body (vitrella) and retina. Each retinal cell (re) is connected with a fibre from the optic nerve, contains pigment, and ends in a rod directed outwards towards the lens. The cells at the end of the pit or depression are, next to the lens, without pigment, and, growing in between the retina and the lens, fill it up, and thus form a sort of vitreous body.
The ocellus appears to be a direct heirloom from the eyes of worms, while the many-facetted compound eye of the crustaceans and of insects is peculiar to these classes. The compound eye of the myriopod Scutigera differs structurally in many respects from the compound eye of insects, and that of Limulus still more so.
It should be observed that in the young nymph of Ephemera, as well as in the semipupa of Bombus, each of the three ocelli are situated on separate sclerites. In Bombus the anterior ocellus has a double shape, being broad, transversely ovate, and not round like the two others, as if resulting from the fusion of what were originally two distinct ocelli.
The ocelli are not infrequently wanting, as in adult Dermaptera, in the Locustidæ, and in certain Hemiptera (Hydrocora). In Lepidoptera there are but two ocelli; in geometrid moths they are often atrophied, and they are absent in butterflies (except Pamphila).
The compound or facetted eye (ommateum).—The facetted arthropod eye is wonderfully complex and most delicately organized, being far more so than that of vertebrates or molluscs. The simplest or most primitive facetted eye appears to be that of Lepisma. As stated by Watase, the compound eye of arthropods is morphologically “a collection of ectodermic pits whose outer open ends face towards the sources of light, and whose inner ends are connected with the central nervous system by the optic nerve fibres.”
The facetted eye is composed of numerous simple eyes called ommatidia, each of which is complicated in structure. The elements which make up an ommatidium are the following: (1) The facet or cornea, which is a specialized portion of the cuticula; and (2), the crystalline lens or cone; (3), the nerve-ending or retinula, which is formed out of the retinula cells and the rhabdom or rod lying in its axis; and (4) of the pigment enclosing the lens and rod; the last three elements are derived from the hypodermis. The single eyes are separated from each other by pigment cells.
The facet or cornea.—This is biconvex, clear, transparent, usually hexagonal in outline, and refracts the light. The corneal lenses are cast in moulting.
The corneal lenses are circular in most cases where they are very convex, as in Lathridius and Batocera. The hexagonal ones are very irregular. When they are very convex the eye has a granular appearance, but when not greater than the convexity of the eye itself, the eye appears perfectly smooth (Bolbocerus, etc.). The facets in the lower part of the eye of Dineutes are a trifle larger than in the upper part (about nine to ten). In many insects the reverse is the case, the upper facets being larger than the lower, a notable instance being Anax. The intervening lines between the facets are often beset with hairs, sometimes very long and dense, as in the drone bee and Trichophthalmus; and the modifications of the hairs into scales which takes place on the body occurs on the eyes also, the scales on the eyes of some beetles of the family Colydiidæ being very large, arranged in lines over the eyes like tombstones (Trachypholis).[[45]]
Fig. 261.—Section through the eye of a fly (Musca vomitoria): c, cornea, or facet; pc, pseudocone; r, retinula; Rh, rhabdom; pg1, pg2, pg3, pigment cells; b.m, basilar membrane; T, Tt1, Tt2, trachea; tv, tracheal vesicle; t.a, terminal anastomosis; op, opticon; c.op, epiopticon; p.op, periopticon; n.c, nuclei; n.c.s, nerve-cell sheath; N.f, decussating nerve-fibres.—After Hickson, from Lubbock.
The crystalline lens or cone.—Behind or within the facets is a layer composed of the cones, behind which are the layers of retinulæ and rhabdoms, and which correspond to the layer of rods and cones, but not the retina as a whole, of vertebrate animals.
The crystalline lens is, when present, usually more or less conical, and consists of four or more hypodermis-cells.
The cones are of various shapes and sizes in insects of different groups, or are entirely wanting, and Grenacher has divided the eyes of insects into eucone, pseudocone, and acone. As the pseudocone seems, however, to be rather a modification of the eucone eye, the following division may be made:—
1. Eucone eyes, comprising those with a well-developed cone. They occur in Lepisma, Blatta (Fig. 262), and other Orthoptera, in Neuroptera, in Cicadidæ, in those Coleoptera with five tarsal joints, in the dipterous genus Corethra, and in the Lepidoptera and Hymenoptera (Fig. 263).
Fig. 262.—Ommatidium of cockroach (Periplaneta): lf, cornea; kk, crystalline cone; pg′ pigment cell; rl, retinula; rm, rhabdom.—After Grenacher, from Lubbock.
Fig. 263.—Two separate elements of the eucone eye of a bee; Lf, cornea; n, nucleus of Semper; Kk, crystalline cone; Pg, pigment cells; Rl, retinula; Rm, rhabdom.—After Grenacher, from Lubbock.
Fig. 264.—Three ommatidia of a pseudocone eye, diagrammatic: A, a separate ommatidium of Musca vomitoria, semi-diagrammatic: c, cornea; p.c, pseudocone; pg′, pigmented cells surrounding the pseudocone; p.g2, additional pigment cells; p.g3, basal pigment cells; n.p.c, nuclei of pseudocone; r, retinulæ; n.r, n.r′, nucleus of retinulæ; R, rhabdom; b.m, basal membrane; t.a, terminal anastomosis sending nerve-fibrils to the retinulæ. B, section through a retinula and rhabdom near the basal membrane, the six retinulæ (r) fused into a tube ensheathing the rhabdom (R).—After Hickson.
a. Pseudocone eyes; in which, instead of the crystalline lens or cone, there are four cells filled with a transparent fluid medium, and a smaller protoplasmic portion containing a nucleus (Muscidæ, Fig. 264, pc). Hickson states that the difference between the eucone and pseudocone eyes lies in the fact that in the pseudocone eye “the refracting body formed by the cone-cell lies behind the nuclei,” and in the eucone eye in front of it.
2. Acone eyes, where the cone or refracting body is wanting, but is represented by the four primitive cone-cells. Acone eyes occur in Forficulidæ, Hemiptera (except Cicadidæ), the nematocerous Diptera (Tipula, etc.), and those Coleoptera which have less than five tarsal joints.
The retinula and rod.—The retinula is morphologically a nerve-end cell, situated at the end of a nerve-fibril arising from the optic nerve. The elements of the retinula of Musca are six in number and surround the rhabdom (Fig. 264), which consists of a bundle of six long, delicate chitinous rods, more or less firmly united together (Fig. 264, R).
The six elements of the retinula of Musca are in their outer or distal portion free from one another, but towards their base are fused into a sheath (Fig. 264, r). They are true nerve-end cells, as shown by Müller and by Max Schultze, their views having been confirmed by Grenacher and by Hickson. The relations of the nerves to the rods after passing through the basal membrane is seen in Fig. 266.
The pigment.—The cones or pseudocones are mostly buried in pigment, as well as the rods; and the pigment forms two layers. The outer of the two layers is called the iris pigment (Fig. 265, e, iris tapetum), and the inner (f) the retinal pigment.
Between the ommatidia internally there occur, according to Hickson, pigment cells (Fig. 264, p.g3), each of which stands on the basilar membrane and sends a fine process outwards towards the internal process of the external pigment-cell (p.g2). A long, slender tracheal vesicle also passes in between the retinulæ.
Fig. 265.—Two ommatidia from the eye of Colymbetes fuscus, × 160: a, cornea; b, cone; c, rhabdom; d, basal membrane, with nerve filaments below it: e, iris pigment; f, retina pigment.—After Exner, from Sharp.
The basilar membrane.—This is a thin fenestrate membrane (Fig. 261) separating the cones and rods from the optic tract (Fig. 264, b.m). It is perforated for the passage of tracheal diverticula and of the optic nerve fibrils. It separates the dioptric or instrumental portion of the eye from the percipient portion, i.e. the optic tract.
The optic tract.—This is the optic ganglion of earlier writers, and appears to be the percipient portion of the eye, as opposed to the dioptric portion. If the reader will examine Figs. 249 and 261, he will see that it consists of three distinct ganglionic swellings, i.e. the opticon, epiopticon, and periopticon, whose structure is very complicated. In Musca (Fig. 261) the first ganglionic swelling (opticon) is separated from the brain by a slight constriction, which Berger regards as the homologue of the optic nerve of the other arthropods. It consists of a very fine granular matrix traversed throughout by a fine meshwork of minute fibrillæ, the neurospongium of Hickson. In the young cockroach (Periplaneta) the optic nerve separating the cerebral ganglion from the opticon is much longer in proportion than it is in the adult blow-fly.
Fig. 266.—Periopticon and terminal anastomosis of Agrion, showing the character of the elements of the periopticon (p.op) and the structure of the terminal anastomosis (t.a). 1. The first layer of the terminal anastomosis, consisting of a plexus of fibrils and nerve-cells (n.c). 2. The second layer, in which the fibrils are collected together in bundles. 3. The final optic plexus and nerve-cells. 4. The layer in which the optic fibrils are collected in bundles to be distributed to the retinulæ (r); b.m, basal membrane.—After Hickson.
The second ganglionic swelling (epiopticon, Fig. 261, c.op) is separated from the opticon by a tract of fine nerve-fibrils, which partially decussate; at the decussation two or three larger nerve-cells may be seen. It also contains a few scattered nerve-cells (n.c). The third ganglionic swelling (periopticon, p.op) is separated from the others by a bundle of long optic nerve-fibrils, which cross one another. It is composed of a number of cylindrical masses of neurospongium arranged side by side (Fig. 261, p.op). Between these elements of the periopticon, which do not seem to bear any relation to the number of ommatidia, a single nerve-cell is very frequently seen. The periopticon does not occur in Periplaneta and Nepa (Hickson). The three optic ganglia thus described, together with the cerebral ganglia, are surrounded by a sheath of densely packed nerve-cells.
Bearing in mind the fact that the retinulæ are the nerve-end cells of the fibres passing through the periopticon, it will be well to read the following account, by Hickson, of the terminal anastomosis of the optic fibrils in the periopticon of Agrion bifurcatum, and to examine his sketch (Fig. 266):
“The terminal anastomosis of Agrion may be conveniently divided into four regions. First the region (1) lying nearest to the periopticon in which the nerve-cells are numerous, and the fibrils leaving the periopticon form a complicated plexus; the region (2) next to this, in which the fibrils have collected into bundles separated by spaces occupied by very thin-walled tracheæ in which there are no spiral markings, and lymph-spaces; next, the region (3) in which the fibrils form a final plexus, and in which there are again a considerable number of nerve-cells; and, lastly, the region (4) in which the fibrils are again collected into bundles, separated by spaces containing tracheæ, which perforate the basement membrane to supply the retinulæ.”
It would seem as if the decussation of the optic nerve-fibrils were a matter of primary importance, as it so generally occurs, but in the young of that most generalized of all pterygote insects, the cockroach (Periplaneta), Hickson states that the optic nerve-fibrils which leave the periopticon pass without decussating to the ommateum, and in the adult there is only a partial decussation. In Nepa there is no decussation, but the anastomosis is complicated by the presence of looped and transverse anastomoses.
Looking at the eye as a whole, Hickson regards all the nerve structure of the eye lying between the crystalline cone-layer and the true optic nerve to be analogous with the retina of other animals. With Ciaccio, Berger, and others, he does not regard the layer composed of the retinulæ and rhabdoms as the equivalent of the retina of vertebrates, etc.
Origin of the facetted eye.—The two kinds of eye, the simple and the compound, are supposed to have been derived from a primitive type, resembling the single eye (ommatidium) of the acone eye of Tipula. As stated by Lang, “an increase of the elements of this primitive eye led to the formation of the ocellus; an increase in number of the primitive eyes, and their approximation, led to the formation of the compound facet eye.” This view is suggested, he says, by the groups of closely contiguous single eyes of the myriopods, considered in connection with the compound eye of Scutigera. Grenacher looks upon simple (ocelli) and compound eyes as “sisters,” not derived from one another, but from a common parentage.
Immature insects rarely possess compound eyes; they are only known to occur in the nymphs of Odonata and Ephemeridæ, and in the larvæ and pupa of Corethra.
Mode of vision by single eyes or ocelli.—In their simplest condition, the eyes of worms and other of the lower invertebrates, probably only enable those animals to distinguish light from darkness. The ocelli of spiders and of many insects, however, probably enable them, as Lubbock remarks, to see as our eyes do. The simple lens throws on the retina an image, which is perceived by the fine terminations of the optic nerve. The ocelli of different arthropods differ, however, very much in degree of complexity.
Müller considered that the power of vision of ocelli “is probably confined to the perception of very near objects.”
“This may be inferred,” Müller states, “partly from their existing principally in larvæ and apterous insects, and partly from several observations which I have made relative to the position of these simple eyes. In the genus Empusa the head is so prolonged over the middle inferior eye that, in the locomotion of the animal, the nearest objects can only come within the range. In Locusta cornuta, also, the same eye lies beneath the prolongation of the head.... In the Orthoptera generally, also, the simple eyes are, in consequence of the depressed position of the head, directed downwards towards the surface upon which the insects are moving.”[[46]] Lowne considers that in the ocellus of Eristalis, the great convexity of the lens must give it a very short focus, and the comparatively small number of rods render the picture of even very near objects quite imperfect and practically useless for purposes of vision, and that the function of the ocelli is “the perception of the intensity and the direction of light, rather than of vision, in the ordinary acceptation of the term.”
Réaumur, Marcel de Serres, Dugès, and Forel have shown by experiment, that in insects which possess both ocelli and compound eyes, the former may be covered over without materially affecting the movements of the animals, while if the facetted eyes are covered, they act as if in the dark (Lubbock).
While Plateau regards the ocelli as of scarcely any use to the insect, and Forel claims that wasps, humble bees, ants, etc., walk or fly almost equally well without as with the aid of their ocelli, Lubbock demurs to this view, and says the same experiments of Forel’s might almost be quoted to prove the same with reference to the compound eyes. Indeed, the writer has observed that in caves, eyeless beetles apparently run about as freely and with as much purpose, as their eyed relatives in the open air.
Plateau has recently shown that caterpillars which have ocelli alone are very short-sighted, not seeing objects at a distance beyond one or two centimetres, and it has been fully proved by Plateau and others, that spiders, with their well-formed ocelli, are myopic, and have little power of making out distinctly the shape of the objects they see.
On the whole, we are rather inclined to agree with Lubbock and Forel, that the ocelli are useful in dark places and for near vision. They are, as Lubbock states, especially developed in insects, such as ants, bees, and wasps, which live partly in the open light and partly in the dark recesses of nests. Moreover, the night-flying moths nearly all possess ocelli, while with one known exception (Pamphila) they are wanting in butterflies.
Finally, remarks Lubbock, “Whatever the special function of ocelli may be, it seems clear that they must see in the same manner as our eyes do—that is to say, the image must be reversed. On the other hand, in the case of compound eyes, it seems probable that the vision is direct, and the difficulty of accounting for the existence in the same animal of two such different kinds of eyes is certainly enhanced by the fact that, as it would seem, the image given by the medial eyes is reversed, while that of the lateral ones is direct” (p. 181).
Mode of vision by facetted eyes.—The complexity of the facetted eyes of insects is amazing, and difficult to account for unless we accept the mosaic theory of Müller, who maintained that the distinctness of the image formed by such an eye will be greater in proportion to the number of separate cones. His famous theory is thus stated: “An image formed by several thousand separate points, of which each corresponds to a distinct field of vision in the external world, will resemble a piece of mosaic work, and a better idea cannot be conceived of the image of external objects which will be depicted on the retina of beings endowed with such organs of vision, than by comparing it with perfect work of that kind.”
Fig. 267.—From Lubbock.
How vision is effected by a many-facetted eye is thus explained by Lubbock: “Let a number of transparent tubes, or cones with opaque walls, be ranged side by side in front of the retina, and separated from one another by black pigment. In this case the only light which can reach the optic nerve will be that which falls on any given tube in the direction of its axis.” For instance, in Fig. 267, the light from a will pass to a′, that from b to b′, that from c to c′, and so on. The light from c, which falls on the other tubes, will not reach the nerve, but will impinge on the sides and be absorbed by the pigment. Thus, though the light from c will illuminate the whole surface of the eye, it will only affect the nerve at c′.
According to this view those rays of light only which pass directly through the crystalline cones, or are reflected from their sides, can reach the corresponding nerve-fibres. The others fall on, and are absorbed by, the pigment which separates the different facets. Hence each cone receives light only from a very small portion of the field of vision, and the rays so received are collected into one spot of light.
It follows from this theory that the larger and more convex the eye, the wider will be its field of vision, while the smaller and more numerous are the facets, the more distinct will be the vision (Lubbock).
The theory is certainly supported by the shape and size and the immense number of facets of the eye of the dragon-fly, which all concede to see better, and at a longer range, than probably any other insect.
Müller’s mosaic theory was generally received, until doubted and criticised by Gottsche (1852), Dor (1861), Plateau, and others. As Lubbock in his excellent summary states, Gottsche’s observation (previously made by Leeuwenhoek) that each separate cornea gives a separate and distinct image, was made on the eye of the blow-fly, which does not possess a true crystalline cone. Plateau’s objection loses its force, since he seems to have had in his mind, as Lubbock states, Gottsche’s, rather than Müller’s, theory.
Müller’s theory is supported by Boll, Grenacher, Lubbock, Watase, and especially by Exner, who has given much attention to the subject of the vision of insects, and is the weightiest authority on the subject.
Gottsche’s view that each of the facetted eyes makes a distinct image which partially overlaps and is combined with all the images made by the other facets, was shown by Grenacher to be untenable, after repeating Gottsche’s experiments with the eyes of moths, in which the crystalline cones are firm and attached to the cornea. He was thus able to remove the soft parts, and to look through the cones and the cornea. When the microscope was focussed at the inner end of the cone, a spot of light was visible, but no image. As the object-glass was moved forward, the image gradually came into view, and then disappeared again. Here, then, the image is formed in the interior of the cone itself.
Exner attempted to make this experiment with the eye of Hydrophilus, but in that insect the crystalline cones always came away from the cornea. “He, however, calculated the focal length, refraction, etc., of the cornea, and concluded that, even if, in spite of the crystalline cone, an image could be formed, it would fall much behind the retinula.”
“In these cases, then,” adds Lubbock, “an image is out of the question. Moreover, as the cone tapers to a point, there would, in fact, be no room for an image, which must be received on an appropriate surface. In many insect eyes, indeed, as in those of the cockchafer, the crystalline cone is drawn out into a thread, which expands again before reaching the retinula. Such an arrangement seems fatal to any idea of an image.”
Lubbock thus sums up the reasons which seem to favor Müller’s theory of mosaic vision, and to oppose Gottsche’s view: “(1) In certain cases, as in Hyperia, there are no lenses, and consequently there can be no image; (2) the image would generally be destroyed by the crystalline cone; (3) in some cases it would seem that the image would be formed completely behind the eye, while in others, again, it would be too near the cornea; (4) a pointed retina seems incompatible with a clear image; (5) any true projection of an image would in certain species be precluded by the presence of impenetrable pigment, which only leaves a minute central passage for the light-rays; (6) even the clearest image would be useless, from the absence of a suitable receptive surface, since both the small number and mode of combination of the elements composing that surface seem to preclude it from receiving more than a single impression; (7) no system of accommodation has yet been discovered; finally (8), a combination of many thousand relatively complete eyes seems quite useless and incomprehensible.”
In his most recent work (1890) on the eyes of crustacea and insects, Exner states that the numerous simple eyes which make up the compound eye have each a cornea, but it is more or less flat, and the crystalline part of the eye has not the shape of a lens, but of a “lens cylinder,” that is, of a cylinder which is composed of sheets of transparent tissue, the refracting powers of which decrease toward the periphery of the cylinder. If an eye of this kind is removed and freed of the pigment which surrounds it, objects may be looked at through it from behind; but its field of vision is very small, and the direct images received from each separate eye are either produced close to one another on the retina (or rather the retinulæ of all the eyes) or superposed. In this last case no less than thirty separate images may be superposed, which is supposed to be of great use to night-flying insects. Exner claims that many other advantages result from the compound nature of an insect’s eye. Thus the mobile pigment, which corresponds to our iris, can take different positions, either between the separate eyes or behind the lens cylinders, in which case it acts as so many screens to intercept the over-abundance of light. Exner finds that with its compound eyes the common glow-worm (Lampyris) is capable of distinguishing large signboard letters at a distance of ten or more feet, as well as extremely fine lines engraved one-hundredth of an inch apart, if they are at a distance of less than half an inch from the eye. Exner substantiates the truth of the results of Plateau’s experiments, and claims that while the compound eye is inferior to the vertebrate eye for making out the forms of objects, it is superior to the latter in distinguishing the smallest movements of objects in the total field of vision.
More recently Mallock has given some optical reasons to show that Müller’s view is the true one. He concludes, and thus agrees with Plateau, that insects do not see well, at any rate as regards their power of defining distant objects, and their behavior certainly favors this view. It might be asked, What advantage, then, have insects with compound eyes over those with simple eyes? Mallock answers, that the advantage over simple-eyed animals lies in the fact that there is hardly any practical limit to the nearness of the objects they can examine. “With the composite eye, indeed, the closer the object the better the sight, for the greater will be the number of lenses employed to produce the impression; whereas, in the simple eye the focal length of the lens limits the distance at which a distinct view can be obtained.” He gives a table containing measures of the diameters and angles between the axes of the lenses of various insect eyes, and states that the best of the eyes would give a picture about as good as if executed in rather coarse woodwork and viewed at a distance of a foot, “and although a distant landscape could only be indifferently represented on such a coarse-grained structure, it would do very well for things near enough to occupy a considerable part of the field of view.”
The principal use of the facetted eye to perceive the movements of animals.—Plateau adopts Exner’s views as to the use of the facetted eye in perceiving the movements of other animals. He therefore concludes that insects and other arthropods with compound eyes do not distinguish the form of objects; but with Exner he believes that their vision consists mainly in the perception of moving bodies.
Most animals seem but little impressed by the form of their enemies or of their victims, though their attention is immediately excited by the slightest displacement. Hunters, fishermen, and entomologists have made in confirmation of this view numerous and demonstrative observations.
Though the production of an image in the facetted eye of the insect seems impossible, we can easily conceive, says Plateau, how it can ascertain the existence of a movement. Indeed, if a luminous object is placed before a compound eye, it will illuminate a whole group of simple eyes or facets; moreover, the centre of this group will be clearer than the rest. Every movement of the luminous body will displace the centre of clearness; some of the facets not illuminated will first receive the light, and others will reënter into the shade; some nervous terminations will be excited anew, while those which were so formerly will cease to be. Hence the facetted eyes are not complete visual organs, but mainly organs of orientation.
Plateau experimented in the following way: In a darkened room, with two differently shaped but nearly equal light-openings, one square and open, the other subdivided into a number of small holes, and therefore of more difficult egress, he observed the choices of opening made by insects flying from the other end of the room. Careful practical provisions were made to eliminate error; the light-intensity of the two openings was as far as possible equalized or else noted, and no trees or other external objects were in view. The room was not darkened beyond the limit at which ordinary type ceases to be readable, otherwise the insects refused to fly (it is well known that during the passage of a thick cloud insects usually cease to fly). These observations were made on insects both with or without ocelli, in addition to the compound eyes, and with the same results.
From repeated experiments on flies, bees, etc., butterflies and moths, dragon-flies and beetles, Plateau concludes that insects with compound eyes do not notice differences in form of openings in a half-darkened room, but fly with equal readiness to the apparently easy and apparently difficult way of escape; that they are attracted to the more intensely lighted opening, or to one with apparently greater surface; hence he concludes that they cannot distinguish the form of objects, at least only to a very slight extent, though they readily perceive objects in motion.
One result of his experiments is that insects only utilize their eyes to choose between a white luminous orifice in a dark chamber, or another orifice, or group of orifices, equally white. They are guided neither by odorous emanations nor by differences of color. He thinks that bees have as bad sight and act almost exactly as flies.
From numerous experiments on Odonata, Coleoptera, Lepidoptera, Diptera, and Hymenoptera Plateau arrives provisionally at the following conclusions:
1. Diurnal insects have need of a quick strong light, and cannot direct their movements in partial obscurity.
2. Insects with compound eyes do not notice differences of form existing between two light orifices, and are deceived by an excess of luminous intensity as well as by the apparent excess of surface. In short, they do not distinguish the form of objects, or if they do, distinguish them very badly.
Lubbock, however, does not fully accept Plateau’s experiments with the windows, and thinks they discern the form of bodies better than Plateau supposes.
How far can insects see?—It is now supposed that no insects can perceive objects at a greater distance than about six feet. On an average Lepidoptera can see the movements of rather large bodies 1.50 meters, but Hymenoptera only 58 cm., and Diptera 68 cm.; while the firefly (Lampyris) can see tolerably well the form of large objects at a distance of over two meters.
Until further experiments are made, it seems probable, then, that few if any insects have acute sight, that they see objects best when moving, and on the whole—except dragon-flies and other predaceous, swiftly flying insects, such as certain flies, wasps, and bees, which have very large rounded eyes—insects are guided mainly rather by the sense of smell than of sight.
Relation of sight to the color of eyes.—It appears from the observations of Girschner that those Diptera with eyes of a uniform color see better than those with brightly banded or spotted eyes. Thus those flies (Asilidæ, Empidæ, Leptidæ, Dolichopidæ) whose predaceous habits requires good or quick sight have uniformly dark eyes, as have also such flies as live constantly on the wing, i.e., the holoptic Bombyliidæ, Syrphidæ, Pipunculidæ, etc., whose eyes are also very large.
Those flies whose larvæ are parasitic on other animals have eyes of a uniform color that they may readily detect the most suitable host for their young; such are the Bombyliidæ, Conopidæ, Pipunculidæ, and Tachinidæ.
Certain flies which live in the clear sunlight, as many Dolichopidæ, some Bombyliidæ, and certain Tabanidæ (Tabanus, Chrysops, Hæmatopota), and which are often easily caught with the hand, have eyes spotted or banded with bright or metallic colors. This is also a sexual trait, as the males of some horse-flies visiting flowers have eyes of a single color, the spots and bands surviving only on the lower and hinder parts of the eye, while their voracious blood-sucking females have the entire eye spotted or banded (Kolbe).
The color-sense of insects.—Insects, as Spengel first suggested, appear to be able to distinguish the color of objects. Lubbock has experimentally proved that bees, wasps, and ants have this power, blue being the favorite color of the honey-bee, and violet of ants, which are sensitive to ultra-violet rays.
It is well known that butterflies will descend from a position high in the air, mistaking white bits of paper for white flowers; while, as we have observed, white butterflies (Pieris) prefer white flowers, and yellow butterflies (Colias) appear to alight on yellow flowers in preference to white ones.
The late Mr. S. L. Elliott once informed us that on a red barn with white trimmings he observed that white moths (Spilosoma, Hyphantria, and Acronycta oblinita) rested on the white parts, while on the darker, reddish portions sat Catocalæ and other dark or reddish moths. Gross observed that house-flies would frequent a bluish green ring on the ceiling of his chamber; but if it were covered by white paper, the flies would leave the spot, though they would return as soon as the paper ring was removed (Kolbe). We have observed that house-flies prefer green paper to the yellowish wall of a kitchen, but were not attracted to sheets of a Prussian blue paper, attached to the same wall and ceiling.
It is generally supposed that the shape and high colors of flowers attract insects; but Plateau has made a number of ingenious experiments which tend to disprove this view. He used in his investigations the dahlia, with its central head of flowerets, which contrast so strongly with the corolla. He finds (1) that insects frequent flowers which have not undergone any mutilation, but whose form and colors are hidden by green leaves. (2) Neither the shape nor lively colors of the central head (capitulum) seem to attract them. (3) The gayly colored peripheral flowerets of simple dahlias and, consequently, of the heads of other composite flowers, do not play the rôle of signals, such as has been attributed to them. (4) The insects are evidently guided by another sense than that of sight, and this sense is probably that of smell.