Page:EB1911 - Volume 13.djvu/441

EMBRYOLOGY] Great difference of opinion exists as to the hypopharynx, which has even been thought to represent a distinct segment, or the pair of appendages of a distinct segment. Heymons considers that it represents the sternites of the three trophal segments, and that the gula is merely a secondary development. Folsom looks on the hypopharynx as a secondary development. Riley holds that the hypopharynx belongs to the mandibular and maxillary segments, while the cervical sclerites or gula represent the sternum of the labial segment. The ganglia of the nervous system offer some important evidence as to the morphology of the head, and are alluded to below.

Thoracic Segments.—These are always three in number. The three pairs of legs appear very early as rudiments. Though the thoracic segments bear the wings, no trace of these appendages exists till the close of the embryonic life, nor even, in many cases, till much later. The thoracic segments, as seen in an early stage of the ventral plate, display in a well-marked manner the essential elements of the insect segment. These elements are a central piece or sternite, and a lateral field on each side bearing the leg-rudiment. The external part of the lateral field subsequently grows up, and by coalescence with its fellow forms the tergite or dorsal part of the segment.

Abdominal Segments and Appendages.—We have already seen that in numerous lower insects the abdomen is formed from twelve divisions placed in linear fashion. Eleven of these may perhaps be considered as true segments, but the twelfth or terminal one is different, and is called by Heymons a telson; in it is placed the anal orifice, and the mass subsequently becomes the upper and lower laminae anales. In Hemiptera this telson is absent, and the anal orifice is placed quite at the termination of the eleventh segment. Moreover, in this order the abdomen shows at first a division into only nine segments and a terminal mass, which last subsequently becomes divided into two. The appendages of the abdomen are called cerci, stylets and gonapophyses. They differ much according to the kind of insect, and in the adult according to sex. Difference of opinion as to the nature of the abdominal appendages prevails. The cerci, when present, appear in the mature insect to be attached to the tenth segment, but according to Heymons they are really appendages of the eleventh segment, their connexion with the tenth being secondary and the result of considerable changes that take place in the terminal segments. It has been disputed whether any true cerci exist in the higher insects, but they are probably represented in the Diptera and in the scorpion-flies (Mecaptera). In those insects in which a median terminal appendage exists between the two cerci this is considered to be a prolongation of the eleventh tergite. The stylets, when present, are placed on the ninth segment, and in some Thysanura exist also on the eighth segment; their development takes place later in life than that of the cerci. The gonapophyses are the projections near the extremity of the body that surround the sexual orifices, and vary extremely according to the kind of insect. They have chiefly been studied in the female, and form the sting and ovipositor, organs peculiar to this sex. They are developed on the ventral surface of the body and are six in number, one pair arising from the eighth ventral plate and two pairs from the ninth. This has been found to be the case in insects so widely different as Orthoptera and Aculeate Hymenoptera. The genital armature of the male is formed to a considerable extent by modifications of the segments themselves. The development of the armature has been little studied, and the question whether there may be present gonapophyses homologous with those of the female is open.

In the adult state no insect possesses more than six legs, and they are always attached to the thorax; in many Thysanura there are, however, processes on the abdomen that, as to their position, are similar to legs. In the embryos of many insects there are projections from the segments of the abdomen similar, to a considerable extent, to the rudimentary thoracic legs. The question whether these projections can be considered an indication of former polypody in insects has been raised. They do not long persist in the embryo, but disappear, and the area each one occupied becomes part of the sternite. In some embryos there is but a single pair of these rudiments (or vestiges) situate on the first abdominal segment, and in some cases they become invaginations of a glandular nature. Whether cerci, stylets and gonapophyses are developed from these rudiments has been much debated. It appears that it is possible to accept cerci and stylets as modifications of the temporary pseudopods, but it is more difficult to believe that this is the case with the gonapophyses, for they apparently commence their development considerably later than cerci and stylets and only after the apparently complete disappearance of the embryonic pseudopods. The fact that there are two pairs of gonapophyses on the ninth abdominal segment would be fatal to the view that they are in any way homologous with legs, were it not that there is some evidence that the division into two pairs is secondary and incomplete. But another and apparently insuperable objection may be raised—that the appendages of the ninth segment are the stylets, and that the gonapophyses cannot therefore be appendicular. The pseudopods that exist on the abdomen of numerous caterpillars may possibly arise from the embryonic pseudopods, but this also is far from being established.

Nervous System.—The nervous system is ectodermal in origin, and is developed and segmented to a large extent in connexion with the outer part of the body, so that it affords important evidence as to the segmentation thereof. The continuous layer of cells from which the nervous system is developed undergoes a segmentation analogous with that we have described as occurring in the ventral plate; there is thus formed a pair of contiguous ganglia for each segment of the body, but there is no ganglion for the telson. The ganglia become greatly changed in position during the later life, and it is usually said that there are only ten pairs of abdominal ganglia even in the embryo. In Orthoptera, Heymons has demonstrated the existence of eleven pairs, the terminal pair becoming, however, soon united with the tenth. The nervous system of the embryonic head exhibits three ganglionic masses, anterior to the thoracic ganglionic masses; these three masses subsequently amalgamate and form the sub-oesophageal ganglion, which supplies the trophal segments. In front of the three masses that will form the sub-oesophageal ganglion the mass of cells that is to form the nervous system is very large, and projects on each side; this anterior or “brain” mass consists of three lobes (the prot-, deut-, and tritencephalon of Viallanes and others), each of which might be thought to represent a segmental ganglion. But the protocerebrum contains the ganglia of the ocular segment in addition to those of the procephalic lobes. These three divisions subsequently form the supra-oesophageal ganglion or brain proper. There are other ganglia in addition to those of the ventral chain, and Janet supposes that the ganglia of the sympathetic system indicate the existence of three anterior head-segments; the remains of the segments themselves are, in accordance with this view, to be sought in the