Morphogenesis of opal teeth in calanoid copepods

Opal teeth of calanoid copepods develop early in the premolt phase of the molt cycle. They form in the apolysis space beneath the old tooth row on the mandibular gnathobase. We examined stages of tooth formation inNeocalanus spp. andCalanus pacificus. Apolysis occurs early in the distal gnathobase, then several epidermal cell types participate sequentially in formation of new teeth. Fibrous molds in the shapes of the new teeth are extruded onto the epidermal surface, then additional materials, probably proteinaceous, are secreted into them. Tooth molds next are linked to a gland in the proximal part of the gnathobase by ducts of an unusual type, lamellar-walled ducts. Silicification follows, apparently using highly osmiophilic material supplied by the proximal gland. Opal is laid down at the outer periphery of the mold then thickens toward the attachment of the mold to newly formed chitin at its base. During apolysis the epidermal cells move proximally without breaking ducts that connect small dermal glands with pores in the sides of the opal teeth. The molds for the new teeth form surrounding these ducts, and new pores result. The glands are like the ciliary exocrine glands seen in other Crustacea, with the duct deriving from fusion of the axonemes of a cilium. Presumably the glands secrete a substance into food newly broken by the teeth. This could be a toxin or a digestive enzyme.     

Structure and function of the eversible glands of the aquatic firefly <Emphasis Type="Italic">Luciola leii</Emphasis> (Coleoptera: Lampyridae)

Summary. When attacked or otherwise disturbed, larvae of the aquatic firefly Luciola leii display fork-shaped glands laterally from the meso- and metathorax as well as each abdominal segment. Glandular eversion is accompanied by a strong pine oil-like odour, thanatosis and glowing from paired larval light organs. Observations by SEM and TEM revealed that there are numerous, almost spherical protuberances, measuring 9 μm in diameter on the surface of each gland. Each protuberance is connected to a well developed secretory cell via a 0.1 μm thick and 0.2 μm long stalk and bears three to six 7 μm long spines on its apex. The convoluted glandular walls measure 0.2 μm. The cytoplasm of the secretory cells is characterized by the presence of numerous mitochondria and an extensive system of cisternae and tubular endoplasmic reticulum. Preliminary GC-MS analysis of the glandular secretion revealed two volatile terpenoids: terpinolene and γ-terpinene. Choice and no-choice bioassays involving fish and ants as well as other predators demonstrate that the secretions serve as an effective deterrent against a range of ecologically relevant enemies. The larval postural adjustments, light emission, everted glands, and glandular release of distasteful or repellent compounds, appear to function as a multi-modal, aposematic larval defence. The tandem evolution of glandular secretions and conspicuousness in larval fireflies could partially explain their successful radiation into both aquatic and terrestrial habitats.     

A scanning electron microscope study of the mandibular morphology of boreal copepods

Morphological details of the mandibular blades of 11 species of copepods were examined with scanning electron microscopy. The micrographs improved our understanding of the complex toothed edge of the mandibular blade, which in turn aids in identification of the copepod prey of chaetognaths by gut-content analysis. Beklemishev's discovery of siliceous tooth crowns in calanoid copepods was confirmed for most of the species examined. Long, sharp projections on the crowns of herbivorous species, and the deep grooves into which they fit on the teeth of the opposite mandible, suggest a cracking rather than a grinding function for these teeth.     

Identification of trail pheromone compounds from the labial glands of the stingless bee Geotrigona mombuca

Foragers of several species of stingless bees deposit pheromone spots in the vegetation to guide recruited nestmates to a rich food source. Recent studies have shown that Trigona and Scaptotrigona workers secrete these pheromones from their labial glands. An earlier report stated that species within the genus Geotrigona use citral from their mandibular glands for scent marking. Since convincing experimental proof for this conjecture is lacking, we studied the glandular origin of the trail pheromone of Geotrigona mombuca. In field bioassays, newly recruited bees were diverted by artificial scent trails that branched off from the natural scent trail deposited by their nestmates only when they were baited with extracts from the foragers’ labial glands. Compounds extracted from the mandibular glands, however, did not release trail following behavior. This demonstrates that the trail pheromone of G. mombuca is produced in the labial glands, as in Trigona and Scaptotrigona. Furthermore, in chemical analyses citral was identified exclusively in the foragers’ mandibular glands, which disproves its supposed role as a trail pheromone. The labial glands contained a series of terpene- and wax type esters, with farnesyl butanoate as major constituent. We, therefore, postulate that the trail pheromone of G. mombuca is composed of a blend of esters.     

Ultrastructural observations on prey capture and digestion in the scyphomedusa Aurelia aurita

Aurelia aurita medusae are able to catch their prey with their entire body surface. Catch efficiency in medusae caught in Kiel harbour in May 1985 was found to be highest at the tentacles and lowest at the subumbrella. Surface structures of the medusa as well as the cnidom are described by SEM observations. Microbasic heterotrichous euryteles and atrichous isorhizas were found. Discharged nematocysts on the prey's skin indicate different functions of the two types. The villi in the gastral cavity show a characteristic morphological differentiation that consists of a ciliated distal and a basal area covered by vesicles. Four types of glandular cells were identified by TEM observations. Mucous cell types preferably occur in densely ciliated areas. The presence of serous cells is restricted to the basal region of the gastral villi and gastral cavity where the extracellular predigestion takes place. The time of food passage in young medusae of A. aurita decreases from 19 h at 4°C to 4 h at 22°C.     

Cuvierian tubules of the holothuroid Holothuria forskali (Echinodermata): a morphofunctional study

Quiescent Cuvierian tubules of Holothuria forskali (Delle Chiaje, 1823) have a pseudostratified mesothelium made of ciliated cells that cover elongated cells containing lipoproteie granules. Together these cells form conspicuous transversal folds that penetrate the underlaying connective tissue. Circular and longitudinal muscle cells occur under the tips of the mesothelial folds. These cells are presumably myoepithelial cells; they divide the connective tissue into a thin outer layer (where collagen fibers are directed perpendicularly to the tubule long axis) and a thick inner layer (where collagen fibers are arranged in helices parallel to the tubular long axis). Nerve processes are associated with the longitudinal muscles; they also occur between the spirally arranged collagen fibers. A spiral nerve runs along the tubules; it innervates the circular muscles. The tubule lumen is narrow with a highly convoluted limiting epithelium. Epithelial cells are not ciliated; they have basal processes that contain enlarged spherules including mucosubstances and proteins. While having the same diameter as quiescent tubules, elongated tubules have a much thiner wall surrounding an enlarged cylindrical lumen. The mesothelium has no adluminal cells and consists of unfolded granular cells. In contact with a solid surface, granules become a sticky secretion. Longitudinal muscle fibers disintegrate and circular fibers form conspicuous rings along the tubules. Collagen helices still occur, but are considerably elongated and flattened. Inner epithelial cells dissociate from each other and typically lack basal processes and spherules, the latter being secreted into the lumen of elongating tubules. Discharge of Cuvierian tubules results from tubule expulsion and elongation. Tubule expulsion is triggered by the opening of the anus of a contracted holothuroid and results from the rupture of the cloaca with subsequent expulsion of coelomic fluid. Tubule elongation results both from the mechanical stimulation of the tubule sphincter by hydrostatic pressure and the entry of water into the tubule. Mechanical stimulation of the sphincter results in a rapid contraction of the circular muscles that causes the disintegration of adluminal cells, the unfolding of granular cells and the stretching of inner epithelial cells. Entry of water immediately follows sphincter stimulation and complete tubule elongation. Water pressure causes the irreversible elongation of collagen helices, which results both in a considerable thinning of the tubule wall and the formation of enlarged water-filled lumen. It is suggested that slippage of collagen fibers in elongating tubules is nervously mediated.     

Multiple-front iron-mineralisation in chiton teeth (<Emphasis Type="Italic">Acanthopleura</Emphasis><Emphasis Type="Italic">echinata</Emphasis>: Mollusca: Polyplacophora)

The mineralisation of iron in the major lateral teeth of the chiton Acanthopleura echinata has been examined using light and electron microscopy, together with energy dispersive and Raman spectroscopy. The deposition of magnetite in the posterior region of the tooth cusp has been shown to occur simultaneously on two fronts, from both the anterior and posterior sides. In contrast, in the lepidocrocite region, the mineral is formed by aggregation over the whole region. In addition, evidence is presented for the existence of a thin veneer of ferrihydrite over the surface of the magnetite region. The veneer preserves this region from oxidation prior to use in the mature tooth. Data also suggest that the junction zone plays a vital role in the overall biomineralisation process, contributing large amounts of iron at critical stages of development.     

Mating behavior of Labidocera aestiva (Copepoda: Calanoida)

The calanoid copepod Labidocera aestiva Wheeler exhibits a ritualized mating behavior. Sensory hairs and spines and integumental pores on the male and female are believed to play a regulatory role in the mating encounter. Scanning electron microscopy (SEM) revealed approximately 20 unusual pore structures (pit-pores) on the right ventrolateral surface of the female's abdominal segment, the area where the spermatophore coupling device is attached. Prior to spermatophore transfer, the male vigorously strokes this area with the modified endopod of his left fifth leg to prepare the pit-pore region for attachment of a new spermatophore. Transmission electron microscopy (TEM) indicated that the cells underlying each pit-pore may have a secretory function. We hypothesize that the pit-pores secrete a substance which dissolves the cement binding the coupling device to the abdomen and aids the female to remove a discharged spermatophore from her urosome.Contribution No. 116, Harbor Branch Foundation, Inc.     

Feeding in dominant Antarctic copepods—does the morphology of the mandibular gnathobases relate to diet?

This study should clarify the importance of morphology and stability of the mandibular gnathobases for the diet of Antarctic copepod species. The gnathobase morphology of the dominant copepod species Calanoides acutus, Calanus propinquus, Ctenocalanus citer, Rhincalanus gigas, Metridia gerlachei, Stephos longipes, Microcalanus pygmaeus and Paraeuchaeta antarctica from the Southern Ocean was investigated in detail by means of a scanning electron microscope. The mandibular gnathobases of C. acutus, C. propinquus and C. citer have relatively short and compact teeth. These species feed mainly on diatoms and are able to crack the silicious diatom frustules with their mandibular gnathobases by directed pressure. In contrast the teeth of the mandibular gnathobases of P. antarctica are very long and pointed. The nutrition of this species consists predominantly of other smaller copepod species. The motile prey can be held by skewering, using the gnathobases, and then eventually minced. The mandibular gnathobases of P. antarctica have notably more small bristles than those of the other investigated copepod species. These bristles are probably associated with receptors and could serve to locate the prey. The morphology of the gnathobases of R. gigas and M. gerlachei is between that of P. antarctica on the one side and that of C. acutus, C. propinquus and C. citer on the other. Based on the morphology of its gnathobases the copepod species S. longipes, which has to date been found to feed primarily on phytoplankton, mainly ice algae, must also be considered a zooplankton feeder. The investigation showed that M. pygmaeus has gnathobases with surprisingly long and pointed teeth, indicating that this species very probably feeds both on phyto- and on zooplankton organisms. While the mandibular gnathobases of the males of C. propinquus, R. gigas, M. gerlachei and S. longipes have the same morphology as the females of the respective species, in the other four investigated copepod species the males have reduced (C. acutus, C. citer and M. pygmaeus) or completely missing mandibular gnathobases (P. antarctica). The teeth of the gnathobases of all studied species with the exception of M. gerlachei consist of a different material than the remaining parts of the gnathobases. This material seems to be silicate, which probably enhances the stability of the gnathobase teeth.Communicated by O. Kinne, Oldendorf/Luhe     

Functional mouthpart morphology of the squat lobster Munida sarsi, with comparison to other anomurans

 The mouthparts of Munida sarsi Huus, 1935, including the labrum and paragnaths, are described using LM and SEM, with special attention to the setal composition and its function. We also used high-resolution macrovideo equipment to record the feeding processes and monitor setae in action. This combination of morphological and preliminary behavioral data enabled us to arrive at a more comprehensive understanding of the functional morphology. M. sarsi was found to be an opportunist and capable of handling and eating most kinds of animal tissue. The species can also be a selective deposit feeder, sorting the sediment by quality and not by size. This sorting is performed by the setae on the medial rim of the bases of maxilla 1, maxilla 2 and maxilliped 1. Many of the other mouthpart functions ranging from food detection to ingestion are mapped with comparison to what has been found for other anomurans. Grooming was shown to be a significant function of many mouthparts, especially the endopod of maxilliped 3. In general the mouthparts closest to the mouth have the least complex movements and functional scheme, with the mandibles being able to perform only one movement, i.e. cutting or crunching food objects. Some functions are attributed to the paragnath, the endopod of maxilla 2 and the epipod of maxilliped 1. Received: 3 October 1999 / Accepted: 22 March 2000     

Influence of activity and salinity on the weight-dependent oxygen consumption of the rainbow trout Salmo gairdneri

Two types of secretory cell have been found in the cement glands of the cypris larva of Balanus balanoides (L.). Histochemical tests show the presence of proteins, phenols and the enzyme polyphenol oxidase in both the glands and secreted cement. It is proposed that cyprid cement is composed of tanned protein, all the components of which are derived from the cement glands.     

Predators, parasitoids, and pathogens: a cross-cutting examination of intraguild predation theory

Although the canonical concept of intraguild predation evokes images of predators and prey, several subdisciplines within ecology have developed theory not specifically framed in terms of predation and competition and often using system-specific terminology, yet functionally quite similar. Here, we formulate models combining exploitation and competition in predator-prey, host-parasitoid, and host-pathogen communities to compare dynamics, food web structure, and coexistence criteria for these disparate communities. Although dynamic stability in the coexistence region varies strongly among systems, in all cases coexistence of two consumers on a single resource occurs only if the intraguild prey species is more efficient than the intraguild predator at suppressing the abundance of the basal resource, and if the intraguild predator accrues a sufficient gain from attacking the intraguild prey. In addition, equilibrial abundances of all species in all three formulations respond similarly to increases in productivity of the basal resource. Our understanding of predator-prey and parasitoid-host communities has benefited from explicit examination of intraguild predation (IGP) theory, and we suggest that future research examining pathogen communities, in particular, will benefit substantially from explicit recognition of predictions from IGP theory.     

Protective skin structures in shell-less marine gastropods

Skin and stomach epithelia of the four suborders of nudibranch gastropods (Mollusca, Opisthobranchia) are characterized by large intracellular ovoid disks, here called spindles. These spindles are an evolutionary novelty in the nudibranchs; in shell-less sacoglossan species they are missing. We here examined whether the distribution and occurrence of the spindles is consistent with the assumption of a protective role against discharging nematocysts of co-habiting and prey Cnidaria. Spindles were abundant in epidermal cells of regions exposed to nematocysts, such as the cerata, the lips, the edges of lamellate rhinophores, the surfaces of non-retractile gill-like organs, as well as in the stomach epithelium of eolid and dendronotacean species which feed on Cnidaria. While cells packed with spindles almost exclusively formed the epidermis of eolid and arminacean species, they were less numerous in the skin of dendronotacean and dorid species, where glandular cells predominated. The preponderance of either glandular or spindle cells suggests a dual complementary defense strategy, on the one side the production of mucus coats and aversive glandular secretions, on the other structural defensive devices that are cushion-like entities filled with inert grains.     

Nuchal glands: a novel defensive system in snakes

Of the various chemical defensive adaptations of vertebrates, nuchal glands are among the most unusual. First described in a Japanese natricine snake, Rhabdophis tigrinus, in 1935, these organs are embedded under the skin of the neck region as a series of paired glands that have neither lumina nor ducts. The major chemical components of the glandular fluid are bufadienolides, which are cardiotonic steroids also found in the skin secretion of toads. Here we review early studies of nuchal glands and briefly introduce our recent findings on the sequestration of bufadienolides from consumed toads and the maternal provisioning of those sequestered compounds. We summarize behavioral studies associated with the antipredator function of the nuchal glands, which have been conducted during our more than decade-long collaboration. Results of preliminary analyses on the possible costs of toad-eating and on the ultrastructure of the nuchal glands are also presented. Finally, we discuss the evolutionary origin of the nuchal glands and suggest future directions designed to understand the biological importance of these novel vertebrate organs, which have evolved in a limited number of snake species.     

Major primary stages of biomineralization in radular teeth of the limpet Lottia gigantea

The radula (feeding organ) of the limpet Lottia gigantea is a membraneous ribbon to which are attached up to 150 transverse rows of teeth. The organic matrix of the teeth is impregnated with inorganic salts (Fe, Si, Ca). In its posterior end, the radula originates from a radular gland which is coiled dorsally, where long odontoblast cells secrete the tooth and the radular membrane. Teeth mature gradually. The organic matrix of this complex was stained homogeneously in the first five rows (hematoxylin-eosin). From Row 6 onward, the tooth became demarcated from the radular membrane by forming segments within the matrix of the radular membrane. The cusp's organic matrix milieu was histologically divided into two main domains, internal and external, each one consisting of fibrillar milieu. Teeth are embedded within the tall columnar superior epithelium, which superimposes the cusps. Iron-bearing granules started to appear in these cells beginning at Rows 7 to 10, increasing in number to Row 20 and continuing to at least Row 65. Iron proceeded to infiltrate first into the internal part of the cusp at Row 15. The outer part of the cusp was mineralized beginning at Rows 22 to 27. At about Row 30, the cusp was heavily mineralized by iron. Calcium was deposited into the radular membrane and tooth bases from the most posterior part. It is concluded that radula biomineralization is subject to highly complex but precisely controlled cytological-biochemical processes and that different parts of each young tooth are subjected simultaneously to different biomineralization pathways. Specimens used in this study were collected in 1984 from the intertidal zone at La Jolla, California.     

Mineralization of major lateral teeth in the radula of a deep-sea hydrothermal vent limpet (Gastropoda:Neolepetopsidae)

Radular teeth of a neolepetopsid patellogastropod limpet, Paralepetopsis ferrugivora, from a deep-sea hydrothermal vent site have similar elemental composition (O, Si, P, S, Cl, K and Fe) to radular teeth of the shallow-waters patellogastropod limpets. However, in contrast to shallow waters limpets, the fully mature teeth of P. ferrugivora do not show any crystalline phases. Amorphous silica was found in the cusp of the teeth and amorphous iron oxide in the junction zone and base of the teeth. Ferritin-rich vesicles were observed in cells adjacent to the junction zone of the early mature teeth, suggesting that these vesicles can mediate the delivery of iron to the tooth matrix. The similarity in the elemental composition between the hydrothermal limpet P. ferrugivora and shallow-waters patellogastropod limpets shows that the extreme hydrothermal environment did not alter the elemental composition of the radular teeth in the deep-sea species.Communicated by P. W. Sammarco, Chauvin     

Symmetry, male dominance and female mate preferences in the Iberian rock lizard, Lacerta monticola

Female preference for dominant males is widespread and it is generally assumed that success in male-male competition reflects high quality. However, male dominance is not always attractive to females. Alternatively, relatively symmetric individuals may experience fitness advantages, but it remains to be determined whether males with more symmetrical secondary sexual traits experience advantages in both intra- and intersexual selection. We analysed the factors that determine dominance status in males of the lizard Lacerta monticola, and their relationship to female mate preference, estimated by the attractiveness of males' scents to females. Sexually dimorphic traits of this lizard (head size and femoral pores) appear to be advanced by different selection pressures. Males with relatively higher heads, which give them advantage in intrasexual contests, were more dominant. However, head size was unimportant to females, which preferred to be in areas marked by relatively heavier males, but also by males more symmetric in their counts of left and right femoral pores. Chemicals arising from the femoral pores and other glands might honestly indicate quality (i.e. related to the symmetry levels) of a male to females and may result from intersexual selection. Females may use this information because the only benefit of mate choice to female lizards may be genetic quality. Chemical signals may be more reliable and have a greater importance in sexual selection processes of lizards than has previously been considered.     

A review of the chemical ecology of the Cerambycidae (Coleoptera)

Summary. This review summarizes the literature related to the chemical ecology of the Cerambycidae and provides a brief overview of cerambycid biology, ecology, economic significance, and management. Beetles in the family Cerambycidae have assumed increasing prominence as pests of forest and shade trees, shrubs, and raw wood products, and as vectors of tree diseases. Exotic species associated with solid wood packing materials have been notable tree killers in North American urban and peri-urban forests. In forested ecosystems native species respond to disturbances such as fires and windstorms, and initiate the biodeterioration of woody tissue. Eggs are laid by females, on or through the bark surface of stem and branch tissue of moribund, recently killed or decomposing woody plants; larval cerambycids (roundheaded woodborers) typically feed in the phloem and later in the xylem. Females will also, in some cases, select living hosts, e.g. adult conifer and angiosperm trees, for oviposition. Research on the chemical ecology of over 70 species has revealed many examples of attractive kairomones (such as floral volatiles, smoke volatiles, trunk and leaf volatiles, and bark beetle pheromones), repellents and deterrents, oviposition stimulants, short- and long-range sex pheromones, and defensive substances. Emerging generalities are that attractants tend to be monoterpenoids and phenolic esters; oviposition stimulants are monoterpenoids and flavanoids; short-range sex pheromones are female-produced, methyl-branched cuticular hydrocarbons; and long-range sex pheromones are male-produced -hydroxy ketones and (,)-diols ranging in length from 6 to 10 carbons. The latter compounds appear to originate from glands in the male thorax; putative defensive substances originate from metasternal secretory pores or mandibular glands. In one unusual case, a flightless, subterranean female that attacks sugar cane produces a sex pheromone that is derived from the amino acid isoleucine. With significantly more than 35,000 species of Cerambycidae worldwide, these generalities will be subject to change as more species are examined. Addendum The authors would like to point out that the electronic version is more accurate than the printed version.     

Feinstruktur des Trichterepithels von Rhinomuraena ambonensis (Teleostei,Anguilliformes)

The ultrastructure of the funnel-like tubes of the olfactory organ in Rhinomuraena ambonensis was investigated by electron microscopy. Frontal sections of the tubes demonstrate a symmetrical arrangement of an inner epithelium, connective tissue, and an outer epithelium. The surface of each peripheral cell of the epithelia bears a system of small ridges which does not correspond with that of the adjacent cells. In case of a rectilinear course of the epithelial cell borders, numerous desmosomes are located along the cell membranes. The fine structure of the desmosomes was analysed. Number and size of the intercellular spaces increases with increasing depth of the epithelium cell layers. The fine structure of epithelial, basal and secretory cells is described. The outer region of each epithelial cell contains densely packed tonofilaments. The fine structure of sensory “buds” which are sparsely distributed within the outer epithelium of the funnels is described. The basal border of the epithelia consists of a rather thick basal membrane. Blood vessels, nerve bundles, chromatophores and storage cells with cristalline inclusions are embedded in the connective tissue. The fine structure of the chromatophores and storage cells is described. The importance of desmosomes, tonofilaments and intercellular spaces, as well as the basal membrane and the amount of collagen is discussed in relation to the reinforcement of the funnels.     

Effect of mercuric chloride on terrestrial slug Semperula maculata and histopathology of reproductive organs: a comprehensive study

Since the last few decades, terrestrial diversity is found to be under stress due to the contamination by various chemicals and gases of aquatic and agricultural media. The bioaccumulation of these compounds at certain concentrations in different tissues produces histopathological changes leading to behavioral changes in target animals. This study focused on terrestrial molluscan slug, Semperula maculata, against lethal concentrations of mercury chloride (HgCl₂). Bioaccumulation of Hg was highest in ovotestis, while lowest in the molluscan penis. Histopathological changes were observed in the cellular arrangement of reproductive organs. Alterations in the penis and the dart gland included dilated unicellular, multicellular glands, and muscular dystrophy, dilated peripheral glandular cells, respectively; dilation being maximal in the prostate gland. Ovotestis showed depleted number of gametocytes. Vacoulized pre-vitellogenic oocytes with early vitellogenic oocytes were noted under light microscopy. Mercury produced toxicity on secretory functions and lowered the rate of reproduction in animals. Evidence indicates that Hg produced gamete degeneration, impacted reproductive capacity and survival of this species in the ecosystem.