Context Of Anatomical Study

Terms of Orientation and Conventions

Terms to describe orientation are not intuitive for insects. Most orientation terms are derived from the study of the human body—a body that stands upright—and their application to insects causes confusion. Some standard terms used with insects include anterior (in front), posterior (behind), dorsal (above), ventral (below), medial (middle), and lateral (side). Anatomical description usually follows in the same order, hence, we begin our discussion with the head, move on to the thorax and then the abdomen, and finish with the genitalia. Description of the relative placement of anatomical features can be cumbersome, but they are critical elements in the study of anatomical structure because relative position is one of the three basic tenets of homology, including size and shape, and embryology.

Measures of Success

The design of the insect body can be described as successful for many reasons: there are millions of species, they range in size over four orders of magnitude, their extensiveness of terrestrial and aquatic habitat exploitation (the diversity of resources), and once a successful form has been developed, there appears to be relatively little change over evolutionary time (Fig. 1). The basic insect design allows for adaptation to a variety of environmental requirements. The success of the design is rooted in the nature of the main material used for its construction.

The Building Material

When we look at an insect, it is the integument that we see. Structurally, the integument is a multiple-layered, composite organ that defines body shape, size, and color. The ultrastructure of the integument is composed of living cells and the secretory products of those cells. Each layer is of a different thickness and chemical composition, and each displays physical properties different from those of the surrounding

FIGURE 1 Fossil insects are easily recognizable today, indicating an early establishment of a successful design. Left to right: Heplagenes (Late Jurassic 150 mya, Liaoning, China); cricket (Eocene, 50 mya, Green River formation, Utah); fulgorid (Eocene, 50 mya, Green River formation, Utah).

layers. Perhaps more importantly, the integument also is the organ with the greatest diversity of structure and function.

There are two common misconceptions about the integument. First, some believe chitin is responsible for integument hardness. Actually, there is proportionally more chitin found in the soft and flexible membranous parts of the integument than in the hard, sclerotized plates. Integument hardness is attributed to an increased number of cross-linkages between protein chains contained in the integument layers. Second, some believe that the integument is rigid and that growth is incremental and limited to expansion during molting; yet some endopterygote insects are able to grow continuously between molts.

The integument determines the shape of the insect body and its appendages. One of the most captivating features of insects is their seemingly infinite variation in body shapes— everything from a simple bag (Hymenoptera grub) to a mimic of orchid flowers (Mantidae). Similarly, appendage shape is exceedingly plastic. Terms such as "pectinate," "flabbate," and "filiform" are among more than 30 terms taxonomists have proposed to describe antennal shapes. Leg shapes are similarly highly variable and express functional modifications. Among these shapes are "cursorial," "gressorial," "raptorial," "fosso-rial," and "scansorial." Again, these modifications of shape reflect the function of structure. Finally, wing shapes are highly variable among insects and are determined by body size and shape as well as by aerodynamic considerations.

Tagmata

Most people recognize the three tagmata—head, thorax, and abdomen—as characteristic of insects. The way they appear is rooted in a division of responsibilities. The head is for orientation, ingestion, and cognitive process; the thorax for locomotion; and the abdomen for digestion and reproduction. But even casual observations reveal further divisions of these body regions.

Segmentation of Tagmata

Two types of segmentation are evident among arthropods, primary and secondary. Primary segmentation is characteristic of soft-bodied organisms such as larval holometabolans. The body wall in these organisms is punctuated by grooves or rings that surround the anterior and posterior margin of each somite. These rings represent intersegmental lines of the body wall and define the limits of each somite. Internally, the grooves coincide with the lines of attachment of the primary longitudinal muscles. From a functional standpoint, this intrasegmental, longitudinal musculature permits flexibility and enables the body to move from side to side.

More complex plans of body organization exhibit structural modifications. Secondary segmentation is characteristic of hard-bodied arthropods, including adult and nymphal insects. Secondary body segmentation is an evolutionarily

FIGURE 2 Secondary segmentation. Top: diagram of sagittal section of dorsal sclerites of thorax. Bottom: ventral view of abdominal sternites showing overlap due to secondary segmentation (Coleoptera: Scarabaeidae).

derived anatomical feature. The musculature we see in secondary segmentation is intersegmental, or between segments (Fig. 2). The acquisition of secondary segmentation represents a major evolutionary step in the development of the Arthropoda. The soft-bodied arthropod has primary segmentation and muscles that are intrasegmental, or within each segment. Movement of the body and its parts is relatively simple because the body wall is flexible. However, when the body wall becomes hardened, flexibility is restricted to the articulation between hardened parts or the extension provided by intersegmental membranes. The arthropod is, in a metaphorical sense, clad in a suit of armor; most movement is possible only if soft and flexible membranes are positioned between inflexible (hardened) body parts. Exceptions may be seen in the indirect flight mechanism of pterygote insects.

In all probability secondary segmentation evolved many times, and it probably continues to evolve in response to specific problems confronting insects today. Secondary segmentation is most evident and most readily appreciated in the insect abdomen. It is less apparent in the thorax and almost totally obscured in the head.

Sclerites

The hardening of the body wall contributes significantly to the external features observed in insects. Sclerites are hardened areas of the insect body wall that are consequences of the process of sclerotization. Sclerites, also called "plates," are variable in size and shape. Sclerites do not define anatomical areas and do not reflect a common plan of segmentation. Sclerites develop as de novo hardening of membranous areas of the body wall, as de novo separations from larger sclerotized areas of the body, and in other ways.

The hardened insect body displays many superficial and internal features that are a consequence of hardening. Understanding the distinction between these conditions and the terms applied to them is critical in understanding insect anatomy and its application in taxonomic identification. These features are of three types. First, sutures (Latin, sutura = seam), in the traditional sense of vertebrate anatomists, provide seams that are produced by the union of adjacent sclerotized parts of the body wall. On the insect body, sutures appear as etchings on the surface of the body and form lines of contact between sclerites. Second, sulci (Latin, sulcus = furrow) represent any externally visible line formed by the inflection of cuticle. Biomechanically, a sulcus forms a strengthening ridge. In contrast, lines of weakness are cuticular features that are used at molting. Lines of weakness are frequently named as if they were sutures, but they should not be viewed as such. For instance, the ecdysial cleavage line is a line of weakness that is sometimes considered to be synonymous with the epicranial suture. The two features are similar in position and appearance, but structurally they may have been derived from different conditions. Finally, apodemes (Greek, apo = away; demas = body) are hardened cuticular inflections of the body wall that are usually marked externally by a groove or pit. Structures called apophyses (Greek, apo = away; phyein = to bring forth) are armlike apodemes. Apodemes have been defined as a hollow invagination or inflection of the cuticle and an apophysis as a solid invagination. Functionally, apodemes strengthen the body wall and serve as a surface for muscle attachment.

FIGURE 3 (A) Anterior view of the head of a grasshopper (Orthoptera: Acrididae). (B) Larval pterygote head showing epicranial and frontal sutures (Lepidoptera: Noctuidae). (C) Posterior aspect of the head (Orthoptera: Stenopelmatidae).

Sclerites receive different names depending upon the region of the body they are located. Tergites (Latin, tergum = back) are sclerites that form a subdivision of the dorsal part of the body wall (tergum). Latrotergites are sclerites that form as a subdivision of the lateral portion of the tergum. Sternites (Latin, sternum = breast bone) are sclerites that form as a subdivision of the ventral part of the body wall (sternum), or any of the sclerotic components of the definitive sternum. Pleurites (Greek, pleura = side) are sclerites in the pleural region of the body wall that are derived from limb bases.

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