Simultaneous chemical and electrical stimulation of labellar taste hairs of the blowfly Calliphora vicina

When recording from the tip of insect taste hairs, responses to chemical stimulation may be influenced by electrical currents, such as the preamplifier's input bias current. The effect of electrical currents on firing frequency of the salt receptor cell to KCl and NaCl stimulation was determined in labellar ‘aboral’ and ‘adoral’ taste hairs of the blowfly Calliphora vicina. Negative currents always decreased spike frequency, whereas positive currents either increased it, or did not change it significantly. Spike frequency changed less than 1% per 5 × 10^?11 A.A consistent picture of the electrophysiology of blowfly taste hairs is given. It includes a distal pore, present in the dendrite-free lumen of the hair. It abandons the concept of a generator current that transmits excitation from the distal, chemoreceptive part of the taste cell dendrite to the action potential generator in or near the taste cell body. The experimental results are interpreted on the basis of this picture. It is concluded that the ‘electrophoretic effect’ of the electrical current is very small. Thus, the measured effect should mainly be due to a ‘direct effect’ of electrical current on electrically excitable structures in the salt receptor cell, particularly in its dendrite.     

Fine structure of the tip of the labellar taste hair of the blow flies,Phormia regina (Mg.) and Calliphora vicina R.-D. (Diptera,Calliphoridae)

Summary The labellar taste hairs of the blow flies, Phormia regina and Calliphora vicina, have an opening mechanism at the tip which consists of two stump cuticular prongs and a funnel-like cuticular pouch. Opening and folding of these structures are regulated by the pressure within the dendrite-free lumen of the hair. The extrusion of viscous substance at the tip of the taste hair is possible through spongy cuticle and one pore in each prong; it seems likewise to depend on the pressure within the dendrite-free lumen and results in regional collapsing of this lumen. Described and discussed are: The cuticle and pores of the structures at the hair tip, pore filaments which extend from the dendrites, and the number and arrangement of the dendrites.This work was supported by a grant from the 7USDA, Entomology Research Division, Beltsville, Md., and the grant GB-13500 from the National Science Foundation.We thank Dr. J. F. Worley, USDA, Plant Science Research Division, Beltsville, Md., for his collaboration in fluorescence microscopy.     

Electrical impedance of the labellar taste hair of the blowfly,Calliphora erythrocephala MG.

Summary The impedance characteristics (resistance and phase angle) were measured of the labellar taste hair of Calliphora erythrocephala by recording the values between the base and tip of intact and tip-amputated hairs of young and old flies.Measurements made at 30 Hz to 20 kHz indicated frequency independent values with changes in phase angle up to about 1 kHz and complete bypassing of the insulating cuticle at 15 to 20 kHz.Measurements made at 0.9 kHz with intact and amputated hairs showed a low change in phase angle and thus a negligible reactive component. Subsequently these values were treated as the resistive component of the impedance. Amputation of the tip always caused a drop in impedance of about 30 MOhm.Calculations based on known morphologic data allowed the conclusion that the measurements of intact hairs at slight dipping of the tip have been made through both channels and not the dendrite-containing channel alone; consequently the pore distad of the dendritic endings must have an electrolytic connection to both channels of the taste hair. Slightly deeper dipping of the hair tip (5 or less) resulted in 4 to 7 MOhm reduced impedance which indicates a second pore or opening in the greater tip region.This work was partially supported by National Science Foundation Grant GB-13500.     

The proboscis extention response and tarsal and labellar chemosensory hairs in the blowfly

Summary Four types of tarsal chemosensory hairs proposed by Grabowski and Dethier (1954) in the blowfly,Phormia regina, were reclassified into the following six types:A-,B ₁-,B ₂-,B ₃-,C- andD-type, based on the electrophysiological results. As for the labellar hairs, the results obtained by Wilczek (1967) agreed well with those in this work. The concentration-response curves of sugar receptors inB ₂-, andD-type of tarsal hairs and all types of labellar hairs for sugar stimulation were compared in relation to the regression of the acceptance threshold in the proboscis extention response of the blowflies starved for twenty four hours. The initial step of this response could be considered to be triggered mainly by about twelve impulses of every sugar receptor unD-type hairs during initial 0.1 sec after stimulus.     

Electrical resistance of labellar taste hairs in the blowfly, Phormia regina M., as a function of age and sex

The electrical resistance of the labellar taste hairs of Phormia regina M. is enhanced by aging in both males and females; this increase is greater in males. Resistance variations, related to changes in response of stimulated chemoreceptors, may explain, at least in part, the reduction in reflex food intake, O2 consumption, weight increase and, on the whole, survival capabilities of males vis-à-vis female insects.     

The morphogenesis of cell hairs on Drosophila wings

We describe in this paper details of morphogenesis of wing hairs in Drosophila pupae. The ultimate objective is to relate specific protein components used in hair construction to specific components produced in the rapidly changing patterns of gene expression that are characteristic for the period of hair differentiation in wing cells (H. K. Mitchell and N. S. Petersen, 1981, Dev. Biol. 85, 233-242). Hair extrusion to essentially full size occurs quite suddenly at about 34 hr (postpupariation) and this is followed by deposition of a double-layer of cuticulin during the next 4 to 5 hr. Extreme changes in shape of cells and hairs, probably related to actin synthesis, then occur for the next 5 to 6 hr. Deposition of fibers within the hairs and on hair pedestals follows. Formation of cuticle on the cell surface begins and continues until some time in the 60-hr range. It appears that cuticle is formed only on the cell surface and not in hairs or on the top of hair pedestals. The protein synthesis patterns associated with these events are described.     

Structure of galeal sensory complex in adults of the red turnip beetle,Entomoscelis americana brown (Coleoptera,Chrysomelidae)

Summary The internal and external structure of the galeae of the adult red turnip beetle, Entomoscelis americana, was studied using SEM and TEM. The galea broadens from base to truncated tip and its sides are of thick, sculpted cuticle invested with pores and coarse spines. The tip is of thinner, flexible cuticle covered with 8–12 uniporous, blunt-tipped apical pegs and a single, aporous, sharply-pointed apical hair.The coarse spines are singly innervated probable mechanosensilla owing to the tubular body at the distal end of the dendrite. These sensilla likely act as tactile hairs monitoring galeal-effected movements of food particles into the functional mouth. The pores are associated with glands within the galea. The function of the presumed secretion is not known but may be to keep objects and dried saliva from sticking to the mouthparts.The apical pegs are innervated by five neurons, each producing a single dendrite. Four dendrites enter the single peg lumen and communicate with the terminal pore. The fifth differentiates into a tubular body that inserts into the peg base. These are typical insect contact chemosensilla that, because of their location, would taste incoming food.The apical hair has no pores but is innervated by two neurons, each extending a dendrite into the hair lumen in chemosensillar fashion. The sensory mode of this sensillum is unknown but is probably not mechanoor chemoreception. Many of its features, reminiscent of taste hairs, lead us to hypothesize that it represents a one-time chemosensillum recently modified to a new form and sensory mode.Because larval and adult E. americana share similar food plant requirements, we hypothesize that similarities will be seen in their mouthpart sensilla. Comparisons of the adults and larvae show the common features between their respective galeal taste hairs are only those of insect contact chemosensilla in general. However, the adult apical hair and the larval medial sensillum show striking specific structural similarities. We propose that these are true structural and functional homologues.     

Sugar cell responses to lactose and sucrose in labellar and tarsal taste hairs of Musca domestica

Receptor cell responses in the largest labellar (LL) and tarsal (D) taste hairs of the housefly Musca domestica were investigated electrophysiologically using the tip-recording technique. In LL hairs, test series with lactose in concentrations of 12.5–400 mmol · l⁻¹ yielded a threshold concentration around 12 mmol · l⁻¹ and a calculated concentration eliciting half-maximal response of around 40 mmol · l⁻¹, the maximal response varying between 18 and 30 impulses/300 ms. D hairs are more sensitive towards lactose, indicated by a slightly lower threshold and a by 60% higher response to 400 mmol · l⁻¹ lactose. The high variation in the relative stimulating effectiveness of lactose and sucrose and experiments with sugar mixtures imply that these sugars bind to different receptor sites without noticeable cross affinity. A comparison of the concentration response characteristics for sucrose and lactose in LL and D hairs suggests that sucrose can combine with more than one site type, expressed in different proportions in both hair types. Results obtained with p-nitrophenyl-β-galactoside as stimulus indicate that a β-galactoside link is not sufficient for a substance to interact specifically with the lactose binding site. The exceptional lactose sensitivity of the sugar cell in M. domestica is discussed in the context of food acquirement and digestion. Accepted: 14 November 1997     

Labellar lobe spreading in the blowfly: Regulation by taste and satiety

Summary Lobe spreading behavior was studied by recording electromyograms from the muscles which spread the labellar lobes, the retractors of the furca (RF) inPhormia regina. RF responses and lobe spreading could be elicited by stimulating labellar, but not tarsal, taste hairs with sucrose (Fig. 3). RF activity was important to spread the lobes at the beginning of a meal, but was not necessary for continued feeding (Fig. 4).Temporal summation between sugar receptor spikes was necessary to elicit RF responses. Central response decrement occurs independently for different labellar hairs and may participate in the termination of motor responses.RF responses were more probable and more intense when either the sucrose concentration of the stimulus or the number of hairs stimulated was increased (Fig. 7). Stimulation with NaCl had no effect on the response to simultaneous sucrose stimulation of other hairs (Table 1).Feeding caused decreases in the probability and intensity of motor responses, but did not alter chemosensory responses (Figs. 8 and 9). Section of either the recurrent or median abdominal nerves prevented this postingestional inhibition of lobe spreading (Fig. 9).These results are discussed with regard to the possible role that regulation of lobe spreading may play in the control of food intake.This work was supported by United States Public Health Service Training Grant 5T01 GM 00457-13S2 and by a grant from the National Science Foundation to Dr. Vincent G. Dethier. I wish to thank Dr. Dethier for his support and encouragement.     

The pollen-collecting hairs of Campanula (Campanulaceae). I. Morphological variation and the retractive mechanism

The pollen-collecting hairs (PCHs) of Campanula have been a subject of intense debate for the past two centuries. Although several morphological studies have been made on these hairs, detailed comparative studies among species are still lacking, their function and adaptive significance being an unsolved question. The present study comprises two microscopy techniques: scanning electron microscopy and confocal scanning laser microscopy. The aim of the present study is to elucidate: 1) the variation in morphology of the PCHs, 2) the variation in presence/absence of the PCHs by the time of spreading of the stigmatic lobes, 3) the variation in the retractive mechanism of the PCHs, and 4) the correlation between pollination and the retraction of the PCHs. In several species PCHs of various lengths are found. Despite the variations in length of the hairs, the same retractive mechanism is found in all species studied. In most species the hairs retract into basal cavities within the style late in anthesis. The cells into which the hairs retract differ in length among species. Pollen grains are often found within the cavities together with the retracted hairs, a mechanism considered to prevent self-pollination. Pollen germination within the cavities was not observed. In a few species, the PCHs are still present at stigma receptivity. Differences in the shape and size of the cells surrounding the PCHs are documented. The diameter of the pits and the pollen grains vary among species. Other types of hairs on the style are recognized in some species, being of various lengths. These other types do not retract at stigma development and should not be regarded as pollen-collectors. They possibly facilitate for visiting insects to reach the nectar glands, present at the top of the ovary.     

The shape of wind-receptor hairs of cricket and cockroach

We examined the exact shapes of the thread-like wind-receptor hairs in the cricket and cockroach. The diameters of hairs at various distances from the hair tip as measured by scanning electron microscopy revealed unexpected hair shapes. We had expected, a priori, that the shape of the hair would be a slender linearly tapered cone, but the measurements revealed hairs in the form of extremely elongated paraboloids. The diameter of the wind-receptor hairs varies with the square root of the distance from the hair tip, i.e., the diameter rapidly increases with the distance from the tip and is asymptotic to the base diameter. Both the cricket, Gryllus bimaculatus, and the cockroach, Periplaneta americana, showed the same hair shape. In both insects, the formation of the wind-receptor hair during metamorphosis seems to be controlled by a common cytological program. The shape of the hair constrains the mobility of the wind-receptor hair, because both the drag force caused by moving air and the moment of inertia of motion dynamics are functions of shaft diameter. The shape of the hair is a biological trait which affects the sensory information transmitted to the central nervous system. Accepted: 24 February 1998     

A note on the specific cuticle structure of wool hairs in otters (Lutrinae)

The cuticle structure of the wool hairs (secondary hairs) of six otter species was examined by scanning electron microscopy to clarify the specific function of this hair type in the Lutrinae. The species studied were chosen according to the different genera, climatic regions, and degrees of association to water of the Lutrinae. Independent of their preferred habitats, the cuticle of every wool hair examined exhibited in all animals a rather similar shape and arrangement of the scales. This specific adaptive feature allows a flexible interlocking of adjacent wool hairs, which also helps to form thin wool hair bundles that surround small oval shaped spaces. Thus, the trapping of an effective insulating air layer is facilitated and heat loss from the body is reduced.     

Atypical pseudopollen-forming hairs in Maxillaria (Orchidaceae)

Pseudopollen is found on the labella of several species of Maxillaria and is formed by the fragmentation of uniseriate, multicellular, moniliform trichomes. The resultant component cells are rich in protein. Since flowers of pseudopollen-forming species generally lack nectar, it is probable that pseudopollen gradually replaced nectar as the pollinator reward. However, direct evidence for this hypothesis is lacking. The present paper examines the labellar micromorphology of five Brazilian species of Maxillaria assigned to the M. discolor alliance. The flowers of two of these species, namely M. violaceopunctata and M. villosa , produce both food hairs and a lipoidal, labellar secretion which is rich in aromatic amino acids. Moreover, in the case of M. violaceopunctata , the secretion was found to contain reducing sugars. This may represent an intermediate stage in the transition from nectar-producing to pseudopollen-forming flowers. SEM studies indicate how pseudopollen-forming, moniliform trichomes, such as those found in the Maxillaria grandiflora complex, may have evolved from simple, uniseriate, multicellular hairs.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 143 , 151−158.     

The innervation of some proboscis structures involved in feeding behavior of the blowfly (Calliphora vicina)

The microtopology of the motoneurons involved in protraction and retraction of the proboscis of the blowfly (Calliphora vicina) has been studied. In addition, taste input from the labellar hairs was investigated. As a result of this study it appears that protraction movements are controlled by two while retraction movements are guided by three motoneurons on each side. The neurons in each group apear to be in ipsicontralateral communication with each other. The musculi protractores fulcri (MPF) probably contain a proprioceptive cell group which projects to the MPF motoneurons. It is proposed that the proboscis motor system can be modulated by proprioception as well as by chemosensory labellar input. Neurosecretory cells may be involved in adjusting muscle power.     

The development of taste and tactile hairs in the pharate fly Protophormia terraenovae (Diptera,Calliphoridae) and in the embryonal cricket Acheta domestica (Orthopteroidea,Ensifera)

Summary 1. The development of taste hairs and tactile hairs of the fly Protophormia terraenovae is described using light microscope, scanning, and transmission electron microscope methods.2. The development of taste hairs proceeds in the same way on tarsi, labella, and wings. First the dendritic outer segments of ciliary origin become visible above the hypodermal cell surface [2 days after pupariation (AP) at 19° C]; then the dendritic sheath starts growing out and finally the trichogen process follows. In a typical intermediate stage (stage C) the distal sections of the dendrites float freely in the fluid surrounding the pharate adult. The more proximal sections are enclosed by the dendritic sheath around which the trichogen process is wrapped (4 days AP). The protruding dendrites disappear when the cuticle starts being deposited on the fully grown trichogen process, and the sheath vanishes later (9–10 days AP or 1 day before eclosion). The development is discussed with respect to the known structural organization of the adult hair.3. In the tactile hairs the single dendrite which grows outwards is completely covered by the dendritic sheath and lies beside the trichogen process [stage C(m)].4. The taste and tactile hair development proceeds in the same way on legs isolated from the pupa after disc eversion in an artificial medium containing ecdysterone.5. To check that both these patterns of development are widespread the development of taste and tactile hairs of the first instar cercus of the cricket Acheta domestica was studied with the light microscope: Both hair types pass through identical early stages.     

Chemosensory control of pollen ingestion in the hoverfly Eristalis tenax by labellar taste hairs

The labellar gustatory system of the dronefly Eristalis tenax L. (Syrphidae; Diptera) that enables the fly to discriminate between pollen and nectar is investigated, and the triggering of pollen ingestion is examined. In behavioural preference tests, exhaustively extracted pollen of the sunflower Helianthus annuus is consumed in smaller amounts than untreated pollen, indicating that water-soluble substances are important for acceptance. Dry pollen is preferred over moist pollen in which the grains stick together, suggesting that mechanical properties of the pollen also play a role in its sensory assessment. Electrophysiological studies of the labellar taste hairs reveal that aqueous extracts of pollen (2% w/v) stimulate the salt receptor cell, but not the sugar receptor cell. The response of the water receptor cell remains the same as to pure water (or standard electrolyte, 10 mmol · l⁻¹ KCl). Of the 20 amino acids tested, the salt cell is sensitive only to proline in a submillimolar range. Behavioural experiments support the electrophysiological findings. When KCl is applied at concentrations eliciting salt-cell spike frequencies equal to those produced by pollen extract (which is often accepted), the water receptor cell is inhibited and a pronounced rejection behaviour occurs. This rejection of concentrated salt solution in Eristalis is therefore mainly mediated by the inhibition of the water cell. Accepted: 27 November 1999     

Cell surface expansion in polarly growing root hairs of Medicago truncatula

Fluorescent microspheres were used as material markers to investigate the relative rates of cell surface expansion at the growing tips of Medicago truncatula root hairs. From the analysis of tip shape and microsphere movements, we propose three characteristic zones of expansion in growing root hairs. The center of the apical dome is an area of 1- to 2- microm diameter with relatively constant curvature and high growth rate. Distal to the apex is a more rapidly expanding region 1 to 2 microm in width exhibiting constant surges of off-axis growth. This middle region forms an annulus of maximum growth rate and is visible as an area of accentuated curvature in the tip profile. The remainder of the apical dome is characterized by strong radial expansion anisotropy where the meridional rate of expansion falls below the radial expansion rate. Data also suggest possible meridional contraction at the juncture between the apical dome and the cell body. The cell cylinder distal to the tip expands slightly over time, but only around the circumference. These data for surface expansion in the legume root hair provide new insight into the mechanism of tip growth and the morphogenesis of the root hair.     

Proboscis extension in the blowfly: directional responses to stimulation of identified chemosensitive hairs

Summary We studied the direction of proboscis extension elicited by stimulating each of an identified array of gustatory sensilla, the largest hairs, on the labellum of the flyPhormia regina. Individual hairs, or pairs of hairs, were stimulated with sucrose or water and the angle of the ensuing extension of the proboscis was recorded. The direction of the response was graded and depended on the identities of the hairs stimulated. Hairs situated on the anterior region of the labellum elicited anterior extensions, mid-level hairs elicited lateral extensions and posterior hairs resulted in posterior extensions. Previous studies of the labellar hairs have been concerned with their encoding of the chemical nature of the stimulus. Our findings show that each hair also relays information about the location of the stimulus. The positional information provided by these sensilla may help the fly to orient itself with respect to a food source.Abbreviation CES central excitatory state     

Behavioral responses of Phormia regina (Meigen) to labellar stimulation with amino acids

Behavioral responses to labellar stimulation with 19 L-amino acids were predicted on the basis of electrophysiological responses of largest labellar hairs. With the exceptions alanine, aspartic and glutamic acids, and valine tests of these predictions confirmed that Phormia can discriminate amino acids, and that these acids may be grouped according to their effects. Electrophysiological investigation of the four exceptions was repeated and results were consistent with the behavioral data. In particular, these acids elicited previously unreported responses from the salt receptor. The discrepancies between this and earlier studies may be explainable, in part, on methodological grounds. There was evidence for response differences among hairs of different sizes and among the largest labellar hairs themselves. The significance of amino acid discrimination for the problem of protein recognition can only be speculated upon until more complete electrophysiological and nutritional information is available.