工蜂的形态显微图（上）

本文讲述了工蜂的形态，通过显微镜从各个角度进行拍摄讲解，每张图片为一个角度，图片下面有相应的解说。

WORKER HEAD ( FRONTAL VIEW )

WORKER EYES

TOP. Three ocelli (two lateral and one medial) on the dorsal portion of the head between the two compound eyes. Branched body hairs are abundant around these three photoreceptor organs. Each of the biconvex lenses refracts light to about 800 well-packed photoreceptor cells. As light monitors, these organs may regulate the start and finish of foraging activity as well as visually measure horizon versus ground to assist in maintaining level flight. (x 167)

BOTTOM. Compound eye with its thousands of facets. Under each facet are nine photoreceptor cells, all of which are color receptors and some of which are capable of analyzing polarization patterns in the skylight. The conspicuous, long, interfacetal hairs are mechanoreceptors that inform the worker of flight speed and wind direction. (x 270)

WORKER FACE

TOP RIGHT Photomontage of frontal view of the head showing the elbowed character of the antennae. The three occlli are seen at the vertex; to the right of that cluster is the hairy compound eye. The anterior tentorial pit is near the bottom (arrow) ( x 31)

MIDDLE LEFT Higher magnification of the antenna] bases. The antennal socket and sclerite enclose the bulbous base of the elongated scape (first segment) of the antenna. The basal surface is covered with rows of short (probably mechanoreceptor) hairs. Extending forward from the antennal base,; is the broad triangular clypeus. The anterior tentorial pit is visible (arrow) (x 50)

TOP LEFT Close-up of the scape-pedicellus junction (the antennal elbow). The short hairs at the base of the pedicellus may be external proprioceptors that monitor the varying angle of antennal flexion as different groups of these hairs contact and are bent against the edge of the scape at given degrees of flexion. Johnston's organ (an auditory organ) is located within the pedicel. ( x 186)

BOTTOM LEFT Anterior tentorial pit. These cavities lie in the epistomal sulcus and are difficult to discern at low magnification (arrows in upper right and middle left micrographs). The pits are external manifestations of the anterior roots of the cannular tentorium, the endoskeleton of the head. (x 409)

BOTTOM LEFT Close-up of the corrugated cuticular floor and periphery of the anterior tentorial pit. ( x 1,650)

WORKER ANTENNA

BOTTOM LEFT. Survey of the medial surface of the ultimate six (flagellar) segments. Even at this low magnification, very bright foci (denoting clustered pore sensilla) are apparent on each segment. ( x 65)

BOTTOM MIDDLE. Higher magnification of the sixth antennal segment. the three bright foci are areas of' far greater secondary electron capture bv the microscope, and each bright area delimits a pore sensillum. In addition, numerous trichoid sensilla are visible on the right and many placoid sensilla are on the left. (x 702)

BOTTOM RIGHT Antenna, posterior surface. Robust trichoid sensilla are sparsely distributed among slender short trichoid sensilla. (x 65)

TOP LEFT Terminal antennal segment, dorsal view. All sensilla types known for the antenna are found here. Four pore organs are easily visible (the bright, circular spots) in the midst of many plate and trichoid seisilla (at least three types of trichoid sensilla). Electron beam damage may have caused the curled or bent trichoid sensilla. ( x 390)

TOP RIGHT. Terminal antennal segment, ventral view. On this surface are 'at least three morphologically different forms of trichoid sensilla. (x 312)

WORKER MOUTHPARTS

TOP Tangential view of the ventral aspect of the head showing the right and left compound eyes. Most prominent are the crossed mandibles, which are laterally articulated and lie immediately beneath the oval platelike labrum. The mandibles appear to embrace the proximal portion of the glossa and surrounding galeae (of maxillae), which are folded back beneath the head. (x '03)

BOTTOM. Close-up of the sensilla and their distribution on the labrum, mandibles, and maxillae. The lateral articulation of the left mandible with the ventral margin of the gena is visible. These spatulate jaws are drawn in (toward the insect) and out. ( x 85)

WORKER HEAD LATERAL VIEW

TOP. Left profile of the head. The compound eye is in the tipper right corner. The relatively glabrous bladelike maxilla extends diagonally from lower left to center. Behind the makilla are the labial palps and glossa. The arrow indicates an area further magnified in the bottom micrograph. (x 55)

BOTTOM. Higher magnification of the abductor apodeme and hinge of the mandibles, which is on the dorsal margin of the mandible (arrow in top micrograph). On the right side is the gena. The posterior margin of the gena is juxtaposed with the mandible. This apodeme is seen externally as a small cuticular cavity at the lateral margin of the mandible. The cranial abductor muscle inserts into the internal cuticular inflection formed by this apodeme. Socketed proprioceptor hairs are visible on the base of the mandible. (x 335)

WORKER MOUTHPARTS

TOP LEFT. Photomontage of the glossa. The glossa is extended when feeding here it is extending between the pendulous galeae, the terminal segment of the underlying labial palps are nearly covered by the galeae, and only the tips are exposed. The arrow indicates the area further magnified in the top right micrograph. ( x 34)

TOP RIGHT. External opening of the mandibular gland (arrow) exposed when the mandibles are wide open. (x 100)

MIDDLE RIGHT Higher magnification of the cutting edge of the right mandible. Trichoid and basiconic sensilla are present on the exterior surface of the mandibles; these putative chemoreceptors may be gustatory in nature and assist in tasting materials that are being manipulated and chewed by the mandibles. The longer socketed hairs may be mechanoreceptors that, on contact with the other mandible, provide proprioceptive information about the degree of mandibular retraction. (x 110)

WORKER GLOSSA

MIDDLE. Survey of the glossa projecting outward from the partial clasping of the two maxillary galea. (x 53)

TOP Bilobed glossa covered with hairs that arise from a series of cuticular rings. The intersegmental membrane intervenes between, and is overlaid by, the hairs. (x 680)

BOTTOM LEFT. Flabellum (tongue tip). The terminus, attached to the glossa by a single ventral hinge, is spatulate and tipped with branching hairs. (x 600)

BOTTOM RIGHT. Close-up of the flabellum showing its spoonlike geometry and the hair fringe around the "spoon . " The precise function of this struc- ture is unknown although it may serve as a filter for the nectar, which apparently must flow into the "spoon" before it can be imbibed by the bee. (x 650)

WORKER MOUTHPARTS, POSTERIOR VIEW

LEFT Photomontage of the posterior view of the mouthparts. The occipital foramen (top of the field is still occupied by the severed tissues that pass from head to thorax. Flanking the foramen are two posterior tentorial pits. Projecting directly clown from the foramen, in the hollow (fossa of the proboscis) behind the head, are the labium and paired maxillae. The triangular piece immediately below the foramen is the postmentum. The maxillae gape, revealing the long, hairy glossa, which is half clasped by the labial palps. (Compare this micrograph with Plate 1.10, the queen mouthparts.) Lastly, the "shoulders" behind and on either side of the labium (prementum) are the mandibles. The top arrow indicates an area that is further magnified in the top right micrograph, the bottom arrow an area in the bottom right micrograph. The middle arrow points to the apical papilla of the maxillary palp, shown at higher magnification in the center right micrograph. (x 34)

TOP RIGHT Higher magnification of the prementum-mandible sector showing the branched body hairs along with three kinds of trichoid sensilla (lop arrow In left micrograph (x 230)

MIDDLE RIGHT. Apical papilla of the maxillary palp. One slender sensilla is seen near its tip (middle, arrow in the left micrograph). (x 450)

BOTTOM RIGHT Array of spine-tipped cuticular scales (bottom arrow in left micrograph on the interior surface of the maxilla (galeae). On the lateral margin of the maxilla, relatively large socketed hairs (sensilla chaetica) arise at regular intervals. These sensors are known to be mechanoreceptors and chemoreceptors (the latter for sugar and salts). ( x 1,200)

WORKER MOUTHPARTS, ANTERIOR VIEW

TOP Survey of worker mouthparts (the compound eye and vertex of the head are cropped away at the top). Maxillae and labium appear as a twotiered awning (maxillae over labium) over the glossa. Two major segments of the maxillae are prominent. On the right is the stipes with its fringe of setae. The leftmost segments, connecting with the stipes, are the galeae with their bare-ringed (basal) and then setae-fringed margins. A pair of labial palps flanks the decumbent, elongate glossa. When not in use the glossa is retracted, merged with the labium, enclosed by the maxillae, and then folded back at the hinges (arrows) Compare this micrograph with Plate 3.8, the retracted drone mouthparts. (x60)

BOTTOM RIGHT Posterior (central) sector of the head. The principal feature is the duckbill-shaped postmentum of the labium. This sclerite points rearward toward the cervix (not visible). Above the postmentum and joining it is the prementum with its few short cuticular hairs. The cuticular "cradle" into which the postmentum fits is the lorum. (x 175)

BOTTOM LEFT Higher magnification of the near-contact zone between lorum (on the left) and postmentum (on the right). The interesting, pebblegrained inner surface of the postmentum is visible. (x 270)

WORKER MAXILLARY PALP

BOTTOM. Survey of the anteriolateral portion of the head (upper arrow in top micrograph, Plate 2.10). The thumblike cuticular appendage in the center is the maxillary palp, which is to the right of the labium. Behind this palp (extreme lower right) is the interior surface of the mandible with its numerous hairs projecting inward. ( x 432)

TOP Close-up of a hair plate containing about two dozen mechanoreceptor hairs located at the base of the maxillary palp. The hairs at the curved margin (upper right) are bent whenever the palp is moved medially so these mechanoreceptors can signal its movement and position relative to the other mouthparts. (x 2,520)

WORKER MAXILLA AND LABIUM

TOP LEFT. Two labial palps (each partly covered by the galea of corresponding side maxilla) on each side of the very hairy glossa. (x 119)

TOP RIGHT. Four-segmented labial palp. (x 119)

BOTTOM RIGHT. Frontal view of the labial palp showing the ultimate segment with numerous sensilla. (x 372)

BOTTOM LEFT. Close-up of an annulated galea with its robust elongate trichoid sensilla. On the right is the scaly surface of the labial palp. (x 480)

WORKER ORAL CAVITY

BOTTOM Base of the extended proboscis, posterior view. Angling off and extending to the lower left corner is the stipes. At center is the interior surface of the left mandible. To the right of center is the right mandible, which is further magnified in the center right micrograph. (x 75)

MIDDLE RIGHT Interior surface of the partially closed mandibles. The arrows indicate areas further magnified in the top and bottom left micrographs. (x 3,200)

BOTTOM LEFT. Cuticular scales in the oral cavity (lower arrow in center right micrograph). (x3,200)

TOP Close-up of the interior surface of the oral cavity (upper arrow in center right micrograph). ( x 3,200)

WORKER POSTGENAL AREA

BOTTOM Ventral view of the occipital region of the head capsule. A portion of the cervix (neck) (center top) is flanked by the posterior tentorial pits. The "Gothic arch" is formed by the two postgenal inflections that surround the fossa of the proboscis. A membranous region with parallel folds Joins the postmentum (the "duckbill" at the bottom of the field). (x 108)

TOP Cuticular spines on the intersegmental membrane in the area immediately below the tip of the postmentum. (x 4,320)

WORKER CERVIX, VENTRAL VIEW

TOP Posterioventral view of head where the membranous cervix (neck) (covered with ranks of short spines) Joins the head. On either side of the cervix are the posterior tentorial pits. The occipital foramen (where the cervix joins the head) is flanked by two hair plates, each consisting of 30 short, trichoid sensilla (arrow) which are bent when the head turns laterally toward the cervix. These sensilla are gravity receptors. The pre- mentum extends upward, pointing to the cervix. (x 80)

TOP LEFT Close-up of the 30 socketed hairs (known mechanoreceptors that make up the left lateral occipital hair plate. These are adjacent to the cervix (arrow in top right micrograph). (x 720)

BOTTOM Posterior of the head. The hair plate is located in the ventromedial region of the head (arrow in top right micrograph). These trichoid sensilla are probably gravity receptors (and external proprioceptors) that inform the central nervous system about the attitude of the head and/or mouthparts relative to the rest of the body. Signaling may be accomplished when the hairs contact and are bent by the postmenturn or the cervical sclerite. (x 720)

WORKER CERVIX AND VENTRAL POSTOCCIPUT

MIDDLE LEFT. Posterior view of the head showing the cervix (neck). Oil the left is the left tentorial pit. In the right half of the field the cervical membrane connects the occipital foramen of the head to the prescutum of the thorax. The upper arrow points to the area further magnified in the bottom right micrograph; the lower arrow points to the area shown in the top micrograph. (x 68)

BOTTOM LEFT Close-up of the left tentorial pit. The large "hole" is the external manifestation of an inwardly directed posterior tentorial arm, an important constituent of the endoskeleton, or tentorium. The small cavity above leads to the cannular tentorial bridge; the smaller one below is a tube that extends ventrally inward and terminates at the occipital foramen. (x 220)

BOTTOM RIGHT Array of body hairs on the ventral portion of the cervix. These may engage the cervical hair plate on the posterior portion of the head (upper arrow in middle left micrograph) to monitor head movement (X 980)

TOP The ventral cervical hair plate on the prescutum (lower arrow in middle left micrograph). These three dozen or so socketed hairs constitute a major external proprioceptor organ that monitors head flexion. Specifically, these mechanoreceptors record the degree of ventral (downward) projection of the head as the lower part of the head touches these hairs on the "throat." (x875)

WORKER CERVIX, VENTRAL VIEW, CLOSE-UP

TOP RIGHT Ventral cervix with an unidentified mass that may be a congealed secretion. (See also Plate 2.15, top right micrograph.) ( x 910)

BOTTOM Ventral view of the cervix with the congealed mass removed The area beneath the mass is quite unlike the surface of the rest of the cervical membrane. Numerous unoriented, unsocketed, short hairs arise from nonsclerotized cuticle. (x3,060)

TOP LEFT. Ventral view showing several ranks of these triangular cuticular scales, from each of which issue about a dozen hairs of varying lengths. (x 1,320)

MIDDLE LEFT Close-up of several triangular scales showing that the longest hairs emanate fron-i the triangular apex. A slight film (probably honey) is visible at the base of some of the scales. ( x 2,750)

WORKER CERVIX, DORSAL VIEW

BOTTOM Dorsal cervix (neck). The flexible, nonsclerotized membranous nature of the cervix (neck) is evident. (The neck has been stretched on this specimen.) A pair of lateral hair plates (one per side) is visible. Certain mechanoreceptor hairs in each plate are stimulated when the\- engage the back of the head (arrow) ( x 94)

TOP RIGHT Lateral hair plate (the head is to the top, the thorax at the bottom). (x 225)

TOP LEFT Head, posterior view. The vaulted area in the upper region of this field is the back of the head, the basal portion the thorax. Sensory hairs situated on the curved plate engage the back of the head, (x240)

MIDDLE LEFT. Surface relief of the cervical membrane, dorsal view. The rows of small rounded tubercles have small, discrete projections arising from them. (x 1,000)

WORKER WINGS

RIGHT. Photomontage of worker wings, a survey of most of the metathoracic (hind) wing covering a sizable fraction of the mesothoracic (fore-) wing. Most of the wing sclerites (plates at the wing base) are shrouded with hairs. ( x 2 7)

TOP LEFT. Wing bases with wings folded back over the abdomen. The hairy, roughly rectangular pleurite (at 10 o'clock) is the supraepimeron; immediately below it is the intraepimeron. (x 55)

MIDDLE LEFT. Pit marking the apodeme on the pleural sulcus between the intraepimeron and episternum. (x 240)

BOTTOM LEFT. Dorsolateral view of wing sclerites with wings extended (the head is to the left). The posterior notal wing process is prominent (at 8 o clock) with the radius (wing vein) approaching the side of the process. With their wings "disengaged" and folded back over the body, bees are able to vibrate their wing muscles to produce heat and also sounds, such as those created while dancing. ( x 50)

WORKER WING SURFACE, DORSAL VIEW

TOP Leading edge and upper surface of the forewing. The costal ribs are covered with short and rnedium-length hairs. Hairs arising from ribs may be innervated because the hollow rib can acconuriodatc nerves and heiriolymph. (x 170)

BOTTOM Leading edge and upper surface of the hind wing showing the bank of hooks that connects with the curled posterior margin of the forewing. The hind-wing hooks of the worker are somewhat smaller and more angulated than those of the drone. Rows of microtrichia stud the surface of the hind wing. (x 318)