Page:EB1911 - Volume 13.djvu/440

Rh the greater part of the digestive tract, arise from ectodermal invaginations (stomodaeum and proctodaeum respectively) at the positions of the future mouth and anus. The origin of the mid-gut (mesenteron), that has no chitinous lining in the developed insect, is the disputed point. According to the classical researches of A. Kowalevsky (1871 and 1887) on the embryology of the water-beetle Hydrophilus and of the muscid flies, an anterior and a posterior endoderm-rudiment both derived from the “endoblast” become apparent at an early stage, in close association with the stomodaeum and the proctodaeum respectively. These two endoderm-rudiments ultimately grow together and give rise to the epithelium of the mid-gut. These results were confirmed by the observations of K. Heider and W. M. Wheeler (1889) on the embryos of two beetles—Hydrophilus and Doryphora respectively. V. Graber, however (1889), stated that in the Muscidae, while the anterior endoderm-rudiment arises as Kowalevsky had observed, the posterior part of the “mid-gut” has its origin as a direct outgrowth from the proctodaeum. The recent researches of R. Heymons (1895) on the Orthoptera, and of A. Lécaillon (1898) on various leaf beetles, tend to show that the whole of the “mid-gut” arises from the proliferation of cells at the extremity of the stomodaeum and of the proctodaeum. On this view the entire food-canal in most Hexapoda must be regarded as of ectodermal origin, the “endoblast” represents mesoderm only, and the median furrow whence it arises can be no longer compared with the blastopore. According to Heymons, the yolk-cells must be regarded as the true endoderm in the hexapod embryo, for he states (1897) that in the bristle-tail Lepisma and in dragon-flies they give rise to the mid-gut. These views are not, however, supported by other recent observers. J. Carrière’s researches (1897) on the embryology of the mason bee (Chalicodoma) agree entirely with the interpretations of Kowalevsky and Heider, and so on the whole do those of F. Schwangart, who has studied (1904) the embryonic development of Lepidoptera. He finds that the endoderm arises from an anterior and a posterior rudiment derived from the “endoblast,” that many of the cells of these rudiments wander into the yolk, and that the mesenteric epithelium becomes reinforced by cells that migrate from the yolk. K. Escherich (1901), after a new research on the embryology of the muscid Diptera, claims that the fore and hind endodermal rudiments arise from the blastoderm by invagination, and are from their origin distinct from the mesoderm. On the whole it seems likely that the endoderm is represented in part by the yolk, and in part by those anterior and posterior rudiments which usually form the mesenteron, but that in some Hexapoda the whole digestive tract may be ectodermal. It must be admitted that some or the later work on insect embryology has justified the growing scepticism in the universal applicability of the “germ-layer theory.” Heider has suggested, however, that the apparent origin of the mid-gut from the stomodaeum and proctodaeum may be explained by the presence of a “latent endoderm-group” in those invaginations.

Embryonic Membranes.—A remarkable feature in the embryonic development of most Hexapoda is the formation of a protective membrane analogous to the amnion of higher Vertebrates and known by the same term. Usually there arises around the edge of the germ band a double fold in the undifferentiated blastoderm, which grows over the surface of the embryo, so that its inner and outer layers become continuous, forming respectively the amnion and the serosa (fig. 16, A, S). The embryo of a moth, a dragon-fly or a bug is invaginated into the yolk at the head end, the portion of the blastoderm necessarily pushed in with it forming the amnion. The embryo thus becomes transferred to the dorsal face of the egg, but at a later stage it undergoes reversion to its original ventral position. In some parasitic Hymenoptera there is only a single embryonic membrane formed by delamination from the blastoderm, while in a few insects, including the wingless spring-tails, the embryonic membranes are vestigial or entirely wanting. In the bristle-tails Lepisma and Machilis, an interesting transitional condition of the embryonic membranes has lately been shown by Heymons. The embryo is invaginated into the yolk, but the surface edges of the blastoderm do not close over, so that a groove or pore puts the insunken space that represents the amniotic cavity into communication with the outside. Heymons believes that the “dorsal organ” in the embryos of the lower Arthropoda corresponds with the region invaginated to form the serosa of the hexapod embryo. Wheeler, however, compares with the “dorsal organ” the peculiar extra embryonic membrane or indusium which he has observed between serosa and amnion in the embryo of the grasshopper Xiphidium.

Metameric Segmentation.—The segments are perceptible at a very early stage of the development as a number of transverse bands arranged in a linear sequence. The first segmentation of the ventral plate is not, however, very definite, and the segmentation does not make its appearance simultaneously throughout the whole length of the plate; the anterior parts are segmented before the posterior. In Orthoptera and Thysanura, as well as some others of the lower insects, twenty-one of these divisions—not, however, all similar—may be readily distinguished, six of which subsequently enter into the formation of the head, three going to the thorax and twelve to the abdomen. In Hemiptera only eleven and in Collembola only six abdominal segments have been detected. The first and last of these twenty-one divisions are so different from the others that they can scarcely be considered true segments.

Head Segments.—In the adult insect the head is insignificant in size compared with the thorax or abdomen, but in the embryo it forms a much larger portion of the body than it does in the adult. Its composition has been the subject of prolonged difference of opinion. Formerly it was said that the head consisted of four divisions, viz. three segments and the procephalic or prae-oral lobes. It is now ascertained that the procephalic lobes consist of three divisions, so that the head must certainly be formed from at least six segments. The first of these, according to the nomenclature of Heymons (see fig. 17), is the mouth or oral piece; the second, the antennal segment; the third, the intercalary or prae-mandibular segment; while the fourth, fifth, and sixth are respectively the segments of the mandibles and of the first and second maxillae. These six divisions of the head are diverse in kind, and subsequently undergo so much change that the part each of them takes in the formation of the head-capsule is not finally determined. The labrum and clypeus are developed as a single prolongation of the oral piece, not as a pair of appendages. The antennal segment apparently entirely disappears, with the exception of a pair of appendages it bears; these become the antennae; it is possible that the original segment, or some part of it, may even become a portion of the actual antennae. The intercalary segment has no appendages, nor rudiments thereof, except, according to H. Uzel (1897), in the thysanuran Campodea, and probably entirely disappears, though J. H. Comstock and C. Kochi believe that the labrum belongs to it. The appendages of the posterior three or trophal segments become the parts of the mouth. The appendages of the two maxillary segments arise as treble instead of single projections, thus differing from other appendages. From these facts it appears that the anterior three divisions of the head differ strongly from the posterior three, which greatly resemble thoracic segments; hence it has been thought possible that the anterior divisions may represent a primitive head, to which three segments and their leg-like appendages were subsequently added to form the head as it now exists. This is, however, very doubtful, and an entirely different inference is possible. Besides the five limb-bearing somites just enumerated, two others must now be recognized in the head. One of these is the ocular segment, in front of the antennal, and behind the primitive pre-oral segment. The other is the segment of the maxillulae (see above, under Jaws), behind the mandibular somite; the presence of this in the embryo of the collembolan Anurida has been lately shown (1900) by J. W. Folsom (fig. 18, v. 5), who terms the maxillulae “superlinguae” on account of their close association with the hypopharynx or lingua. In reference to the structure of the head-capsule in the imago, it appears that the clypeus and labrum represent, as already said, an unpaired median outgrowth of the oral piece. According to W. A. Riley (1904) the epicranium or “vertex,” the compound eyes and the front divisions of the genae are formed by the cephalic lobes of the embryo (belonging to the ocular segment), while the mandibular and maxillary segments form the hinder parts of the genae and the hypopharynx.