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This chapter looks at Lady Gregory’s Grania (published 1912) and Marina Carr’s The Mai (1994), in which the central women live in exile, yet attempt to negotiate expression of their embodied subjectivity. The capacity to generate new forms is pursued through attention to the process of corporeal change: how the metamorphic body might refute and escape its unhomeliness. These protean bodies can draw attention to, and undermine, the relationship between form and the limitations it imposes, and furthermore evoke a female morphology which demands expression through an alternative cultural imaginary. While the chapter draws on Luce Irigaray’s advocation of a creative female corporeality which evokes a female imaginary, it also looks to Judith Butler’s work on those bodies which fail to signify or matter, and are delegitimated and abjected at the boundaries of the dominant social order. The chapter proposes that Grania’s and The Mai’s metamorphoses question interpretations of both plays’ endings as an act of despair and submission to abjection, instead offering performative confrontations with cultural representations of viable bodies.
Dietary delivery of bacterially expressed double-stranded RNA (dsRNA) has a great potential for management of Leptinotarsa decemlineata. An important first step is to discover possible RNA-interference (RNAi)-target genes effective against larvae, especially the old larvae. In the present paper, five putative Broad-Complex (BrC) cDNAs (Z1-Z4, and Z6) were identified in L. decemlineata. The expression of the five LdBrC isoforms was suppressed by juvenile hormone signaling, whereas the transcription was upregulated by 20-hydroxyecdysone signaling at the fourth (final) instar larval stage. Feeding of bacterially expressed dsBrC (derived from a common fragment of the five LdBrC variants) in the third- and fourth-instar larvae successfully knocked down the target mRNAs. For the fourth-instar LdBrC RNAi hypomorphs, they had a higher larval mortality compared with the controls. Moreover, most dsBrC-fed beetles did not pupate normally. After removal of the apolysed larval cuticle, a miniature adult was found. The adult head, compound eyes, prothorax, mesothorax, metathorax were found on the dorsal view. Distinct adult cuticle pigmentation was seen on the prothorax. The mouthparts, forelegs, midlegs, and hindlegs could be observed on the ventral view of the miniature adults. For the third-instar LdBrC RNAi specimens, around 20% moribund beetles remained as prepupae and finally died. Therefore, LdBrC is among the most attractive candidate genes for RNAi to control the fourth-instar larvae in L. decemlineata.
This article reviews several books published since the turn of the millennium that explore the role and representation of animals in different areas of ancient Greek and Roman culture. Despite differences in focus and outlook, these books herald the arrival within classical studies of the questions, concepts and methods of human/animal studies as an emerging field of enquiry. This article takes their publication as an opportunity to take stock and to outline the relationship between these disciplines. I explore how current research on ancient animals resonates both in existing debates in classical scholarship and within the context of the larger interdisciplinary debate. I also suggest how this debate can point to productive avenues for further enquiry in classical studies. More specifically, I argue that the interdisciplinary debate sets an important agenda, which should be embraced more fully by classical studies. Classical scholarship on the role, function and perception of animals in different areas of ancient Greek and Roman life can provide important insights into one aspect of the heritage – Western conceptions of humanity and the place of the animal within it – which has not yet received the attention it deserves. I conclude that classical scholarship can make a significant contribution to the interdisciplinary debate, helping it deliver on its stated goal of examining and challenging Western concepts of self, as well as the ideologies of ‘the other’ underpinning them.
Shumardiid trilobites had a small, unique morphology, and formed a key constituent in trilobite faunas during the Cambrian–Ordovician. Because of their unusual morphology, they have been the subject of research s, into various aspects such as their life habit, functional morphology, evolutionary origin and ontogeny. Originally, a flat, adult-like protaspid morphology was suggested for shumardiids, but subsequently a bulbous protaspid morphology interpreted to be associated with metamorphosis was also suggested for this unique trilobite group. This article documents the ontogeny of the two co-occurring shumardiid trilobites, Akoldinioidia latus Park and Kihm and Koldinioidia choii Park and Kihm, from the middle Furongian Hwajeol Formation, Taebaeksan Basin, Korea. Interestingly, protaspides of the two shumardiids have a bulbous morphology. Given the stratigraphic occurrences of the two shumardiids, it can be inferred that commutavi protaspis appeared quite early in the shumardiid evolution. The co-occurrence of the two closely related trilobites is reminiscent of sexual dimorphism, although further evidence is required to prove it. The appearance of metamorphosis-undergoing protaspides in the Furongian shumardiids may have been due to the onset of increasing ecological pressure in the early phase of the Great Ordovician Biodiversification Event.
Complex life cycles are common in free-living and parasitic organisms alike. The adaptive decoupling hypothesis postulates that separate life cycle stages have a degree of developmental and genetic autonomy, allowing them to be independently optimized for dissimilar, competing tasks. That is, complex life cycles evolved to facilitate functional specialization. Here, I review the connections between the different stages in parasite life cycles. I first examine evolutionary connections between life stages, such as the genetic coupling of parasite performance in consecutive hosts, the interspecific correlations between traits expressed in different hosts, and the developmental and functional obstacles to stage loss. Then, I evaluate how environmental factors link life stages through carryover effects, where stressful larval conditions impact parasites even after transmission to a new host. There is evidence for both autonomy and integration across stages, so the relevant question becomes how integrated are parasite life cycles and through what mechanisms? By highlighting how genetics, development, selection and the environment can lead to interdependencies among successive life stages, I wish to promote a holistic approach to studying complex life cycle parasites and emphasize that what happens in one stage is potentially highly relevant for later stages.
In this paper, metamorphosis of laboratory-reared cyprids of the pedunculate cirripede, Capitulum mitella was successfully induced. A timeline and a detailed description of morphological events during metamorphosis were derived from light microscopy and scanning electron microscopy analyses. The metamorphic process of cyprids involves five main events. (1) Triggering of metamorphosis: after 0–6 h of reagent exposure, the epidermis ventral to the compound eyes gradually invaginates. (2) Invagination and separation of epidermis: after 6–36 h, the invagination bypasses the compound eyes, the cypris epidermis separates slightly from the carapace and the nauplius eye gradually migrates ventrally. (3) Formation of juvenile rudiment: after 36–72 h, the metamorphosing specimen shape slightly resembles that of a juvenile. (4) Shedding of thoracopodal exuvium: after 72–96 h, the cyprid finally sheds the thoracopodal exuvium and metamorphoses into a juvenile barnacle. (5) Early development of juvenile barnacle: the juvenile shed the cirrum cuticle for the first, second, third and fourth time at 1.5–2, 1.5–2, 4–5 and 5–6 days, respectively. The newly formed juvenile completely assumes the shape and armature of all shell-plates found in adult barnacle. Rostro-carnal length of the juvenile is 240–260 μm. Tergum, scutum and upper latus are paired, and carina and rostrum are unpaired. Uniform circular depressions are dispersed on the surface of the tergum, scutum and carina. The rostrum and upper latus, which have the shape of small triangle, are not clear. The entire surface of the peduncle is covered with convoluted cuticular ridges. The metamorphosis of C. mitella is similar to that of Lepas.
The application of the sterile insect technique to fruit flies involves the mass-production of the pest insects using an artificial diet, irradiation during a narrow time window at the late pupal or early imaginal stage to inhibit reproduction without affecting reproductive capacity, and then release into the target area where the sterile insects compete reproductively with their wild counterparts. The timing of irradiation is important to enable the release of males that are sterile but of good quality and exhibit an acceptable sexual performance. In this study, we examined the pupal development of 12 tephritid (Diptera: Tephritidae) species: Anastrepha fraterculus (Wiedemann), A. ludens (Loew), A. obliqua (Macquart), A. serpentina (Wiedemann), Bactrocera cucurbitae (Coquillett), B. dorsalis (Hendel), B. invadens (Drew, Tsuruta & White), B. oleae (Rossi), B. philippinensis (Drew & Hancock), B. tryoni (Froggatt), B. zonata (Saunders) and Ceratitis capitata (Wiedemann). The insects were reared at various temperatures, in the laboratory (15–28 °C) and under fluctuating natural conditions (20–35 °C). The gradual colour changes of the insect eyes during metamorphosis were observed and photographed, measuring the specific eye colour parameters of each species and matching them with the colour scale of the Munsell Soil Color Charts. The duration of pupal development and the time to emergence in Anastrepha species were longer than those in C. capitata and Bactrocera species at all the holding temperatures. The data obtained can be used by mass-rearing facilities to manage pupal holding conditions and as indicators for optimizing the timing of irradiation.
The main cell types of the adult bee fat body are trophocytes and oenocytes; however, in pupae of some newly emerged bees, trophocytes are modified into cells called urocytes, which possibly function as a substitute for Malpighian tubules during metamorphosis when larval tubules are not functional and/or storage of urate salts is required. This study evaluated the morphology of urocytes in the stingless bee Melipona quadrifasciata and the possibility of maintaining these cells in primary culture. The urocytes M. quadrifasciata are white spherical cells with an irregular surface as observed by stereomicroscopy. They may be found individually or in groups associated with tracheae. Urocytes have a single, small, and spherical nucleus and cytoplasm rich in neutral polysaccharides, lipid droplets, protein, and granules containing calcium and urate salts. Our findings suggest that urocytes play a role in storage of neutral polysaccharides and calcium in M. quadrifasciata pupae and that these cells can be cultured for 72 h.
A large often greenish-appearing type of leptocephalus larva has been observed and photographed or filmed by divers at 4 different locations within the Indonesian Archipelago from 2008 to 2011, and this paper documents these sightings and evaluates what can be learned about this type of unusual eel larvae. These leptocephali (~300–400 mm long) appear to be exceptionally large metamorphosing larvae of some type of moray eel of the family Muraenidae. Because of the morphological similarities between these larvae and ribbon eels, Rhinomuraena quaesita, they are hypothesized to be their larvae, which have never been distinguished among leptocephali of the Muraenidae. These leptocephali were observed at Sangeang Island, in Lembeh Strait adjacent to north-eastern Sulawesi Island, in two different years at Ambon Island, and on two consecutive days at Bali, between December and June of 4 different years. All observations of the large leptocephali were at depths shallower than about 23 m in areas with mixed coral rubble or coarse sand and silt substrates during daylight hours. The larvae used various styles of anguilliform swimming and swam near the bottom or in the lower water column while being observed. One larva tried unsuccessfully several times to enter small crevices. They were all at the metamorphosing stage and may have been entering coastal habitats to find a hiding place where they could transform into juvenile eels. Further studies are needed to clearly identify these large leptocephali and to learn about their life history.
Developmental plasticity is the property of a given genotype to produce different phenotypes in response to the environmental conditions experienced during development. Chordates have two basic modes of development, direct and indirect. Direct development (mode of humans) was derived evolutionarily from indirect development (mode of many amphibians), the major difference being the presence of a larval stage with indirect development; larvae undergo metamorphosis to the juvenile adult. In amphibians, environmental conditions experienced during the larval stage can lead to extreme plasticity in behaviour, morphology and the timing of metamorphosis and can cause variation in adult phenotypic expression (carry-over effects, or developmental programming). Hormones of the neuroendocrine stress axis play pivotal roles in mediating environmental effects on animal development. Stress hormones, produced in response to a deteriorating larval habitat, accelerate amphibian metamorphosis; in mammals, stress hormones hasten the onset of parturition and play an important role in pre-term birth caused by intra-uterine stress. While stress hormones can promote survival in a deteriorating larval or intra-uterine habitat, costs may be incurred, such as reduced growth and size at metamorphosis or birth. Furthermore, exposure to elevated stress hormones during the tadpole or foetal stage can cause permanent neurological changes, leading to altered physiology and behaviour later in life. The actions of stress hormones in animal development are evolutionarily conserved, and therefore amphibians can serve as important model organisms for research on the mechanisms of developmental plasticity.
During their complex life history, anguilliform eels go through a major metamorphosis when developing from a fresh water yellow eel into a deep-sea silver eel. In addition to major changes in body morphology, the visual system also adapts from a fresh water teleost duplex retina with rods and cones, to a specialized deep-sea retina containing only rods. The history of the rods is well documented with an initial switch from a porphyropsin to a rhodopsin (P5232 to P5011) and then a total change in gene expression with the down regulation of a “freshwater” opsin and its concomitant replacement by the expression of a typical “deep-sea” opsin (P5011 to P4821). Yellow eels possess only two spectral classes of single cones, one sensitive in the green presumably expressing an RH2 opsin gene and the second sensitive in the blue expressing an SWS2 opsin gene. In immature glass eels, entering into rivers from the sea, the cones contain mixtures of rhodopsins and porphyropsins, whereas the fully freshwater yellow eels have cone pigments that are almost pure porphyropsins with peak sensitivities at about 540–545 nm and 435–440 nm, respectively. However, during the early stages of metamorphosis, the pigments switch to rhodopsins with the maximum sensitivity of the “green”-sensitive cone shifting to about 525 nm, somewhat paralleling, but preceding the change in rods. During metamorphosis, the cones are almost completely lost.
Researchers are becoming interested in testing whether investment in growth and/or development trades off against investment in parasite defence. We tested this idea by examining relations between development of Wood Frogs (Rana sylvatica) and susceptibility to lung nematodes (Rhabdias ranae). Male and female frogs reared in outdoor mesocosms were the same length and mass at metamorphosis. However, males metamorphosed sooner than females. Lung nematodes were no more likely to penetrate male versus female metamorphs following controlled exposures, but males had higher intensities of adult female worms and the largest worms per host were, on average, of larger size in male metamorphs. Males that took longer to metamorphose carried higher numbers of worms in their lungs than males that metamorphosed early. In comparison, females that developed faster harboured more worms in their lungs than females that took longer to reach metamorphosis. Our results suggest that variation in susceptibility to lung nematodes is influenced by host sex and possibly also by sex-specific relations with developmental rate. Further, male hosts might prove to be a more important source of infective stages of worms than female hosts.
The development of the skeleton of the small temnospondyl Acanthostomatops vorax is described, based on 32 specimens from the Lower Rotliegend of Niederhäslich in Saxony. Both individually and ontogenetically, the skull table, length of gape, and dentition of the parasphenoid are broadly variable. The stapes has a short and straight shaft and a well-defined two-headed proximal region. Small specimens have three or four pairs of ossified ceratobranchials in the gill region, while the adult hyobranchium is composed of four pairs of hypobranchials and probably some additional elements. The postcranium is characterised by a stout and well-ossified humerus that is already large in small larvae, a minute rectangular interclavicle, a robust but short femur, a relatively short trunk with 21–22 presacrals and a moderate tail with 28 vertebrae. In adults, the trunk skeleton became proportionally shorter and the femur smaller with respect to skull length. Metamorphosis is best indicated by the disappearance of the larval gill skeleton, which was probably resorbed, while the structure of the limbs and axial skeleton as well as the absence of lateral line sulci suggest a terrestrial existence for adults.
Recent models argue that thermal environments are the major cause of ectotherm life-history clines. However, elevational clines in body size in the mountain dusky salamander Desmognathus ocoee (family Plethodontidae) shift from positive at hatching, to negative at metamorphosis to positive again as adults, and so are not consistent with this explanation. The clinal shift from hatching to metamorphosis was investigated by examining the clinal and seasonal feeding patterns of larval salamanders at high and low elevation sites in rockface and woodland habitats. Repeated cohort sampling was also used to examine clinal and seasonal patterns in body size and to estimate average growth rates. Larval growth in both rockface and woodland habitats was tightly correlated with feeding activity. Although temperature was found to vary between high and low elevation sites, the greatest growth occurred in a cold woodland habitat with a high elevation, and the lowest growth occurred in an adjacent rockface habitat. Because this difference in growth cannot be attributed to thermal differences, we conclude that local food resource levels are the predominant source of local differences in growth. These findings, clinal patterns of variation in other predatory salamanders, and experimental analyses in which both food and temperature are orthogonally manipulated, indicate that general models that single out temperature as the principle cause of ectotherm life-history clines should be viewed with caution.
Effects of ultraviolet-B (UV-B) radiation on amphibian embryonic and larval development have been investigated in a number of studies, but the effects on later life-stages (metamorphosis) have received less attention. Hence, the effects of UV-B radiation treatments (control (no UV-B), normal and 26% enhanced levels of UV-B) on the development of the common frog Rana temporaria embryos from fertilization until metamorphosis were investigated. Survival until metamorphosis was significantly higher among individuals exposed to normal, as compared to individuals sheltered from, UV-B or those exposed to enhanced levels of UV-B radiation. There were no effects of normal or enhanced levels of UV-B radiation on the frequency of developmental anomalies. However, UV-B radiation delayed the timing of metamorphosis in a dose-dependent fashion, and the individuals from the normal and enhanced UV-B treatments metamorphosed at smaller size than those raised in the absence of UV-B radiation. These results suggest that UV-B radiation experienced through embryonic and larval stages can have negative effects on the growth and development of R. temporaria. Delayed metamorphosis at reduced size is also likely to lower the fitness during later life-stages.
In the present study, norepinephrine (NE), epinephrine (E), dopamine (DA) and 5-hydroxy-tryptamine (5HT) have been quantified by high performance liquid chromatography (HPLC) in Pecten maximus during larval and post-larval development. With average values ranging from 2 to 5 pg μg–1 of protein, NE remained low through the whole larval life, while epinephrine (E) was undetected. DA and 5HT were the most abundant monoamines with significant variations between larval, pre-metamorphic and post-larval stages. During the first 20 days, corresponding to D larval and umboned larval stages, levels of DA and 5HT increased from 15 to 30 and 10 to 15 pg μg–1 of protein, respectively. Then during the last week of larval life, at the approach of metamorphosis, DA rose sharply from 30 to 50 pg μg–1 of protein and 5HT from 15 to 50 pg μg–1 of protein. Lastly during the first week of post-larval life (day 27 to day 34) DA and 5HT contents declined to levels similar to those detected in the first days of larval life reaching progressively 1 pg μg–1 of protein during the second week of post-larval life (day 34 to day 55). These rapid and transient variations in monoamine contents (5HT and DA) around metamorphosis, present a great interest. However, this relation between neurochemical changes and metamorphosis must be confirmed with future studies in order to verify if such monoamines might be used as indicators of larval competence in P. maximus, a commercially important species.
Can variation in egg size explain the local variation in tadpole growth and development of the common frog Rana temporaria or are other mechanisms, e.g. microevolution, involved? To study this, tadpoles were raised in outdoor tanks. Each tank housed tadpoles from one clutch. Eggs were collected during 2 years from six different ponds and their sizes were measured. Large eggs gave rise to tadpoles that grew and developed faster than those hatched from small eggs. Tadpoles from large eggs thus metamorphosed earlier but, because they grew for a shorter time, size at metamorphosis was not affected by initial egg size. Tadpoles from different ponds differed in strategy; after correcting for egg size effects, tadpoles from some ponds tended to metamorphose earlier and at a larger size than those from others. These ponds occasionally dry out during or before the tadpoles' period of metamorphosis. I argue that the detected pond differences are genetically based and represent a case of microevolution.
The habitat occupied by larval winter flounder (Pseudopleuronectes americanus) differs considerably in light regime from that of the adult. To understand how the visual system has adapted to such changes, photoreceptor spectral absorbance was measured microspectrophotometrically in premetamorphic and postmetamorphic specimens of winter flounder. Before metamorphosis, larval flounder retinas contain only one kind of photoreceptor which is morphologically cone-like with peak absorbance at 519 nm. After metamorphosis, the adult retina has three types of photoreceptors: single cones, double cones, and rods. The visual pigment in single cones has a peak absorbance at λmax = 457 nm, the double cones at λmax = 531 and 547 nm, and the rod photoreceptors at λmax = 506 nm. Double cones were morphologically identical, but the two members contained either different (531/547 nm) or identical pigments (531/531 nm). The latter type were found only in the dorsal retina. The measured spectral half-bandwidths (HBW) were typical of visual pigments with chromophores derived from vitamin A1 with the possible exception of the long-wavelength absorbing pigment in double cones which appeared slightly broader. Because the premetamorphic pigment absorbance has a different λmax than those of the postmetamorphic pigments, different opsin genes must be expressed before and after metamorphosis.
The lowest weight at which a final instar larva of the leafroller, Cnephasia jactalana (Walker) (Lepidoptera: Tortricidae) can initiate pupation, the larval critical weight, was unaffected by diet quality. Use of a non-nutritive artificial diet showed that the larval-pupal metamorphosis was possibly triggered by stretch receptors, and was not related to nutrition per se. However, a final-instar larva was not able to moult until it had ingested and assimilated the quantity of food required to attain the larval critical weight. The period between attaining the larval critical weight and the larval maximum weight is the latent feeding period which was found to be longer on higher-quality diets. The decrease from larval maximum weight to pupal weight depended on diet quality: optimum-quality diets gave the smallest decreases. The decrease from pupal to adult weight was not affected by diet. Reproductive performance was strongly affected by the quality of the diet during the latent feeding period.
The juvenile hormone analogue (JHA) 6′, 7′,-epxy-3, 7′-dimethyl-undec-2′-enyl 6-ethyl-3-pyridyl ether at 10 ppm (v/v) applied to whole wheat grains was not toxic to adult insects, but effectively reduced the F1 generation of the lesser grain borer, Rhizopertha dominica (F.). However, topical applications of 4 ppm (v/v) of the JHA prevented normal metamorphosis of the larvae and pupae; all the test insects died due to morphogenetic abnormalities induced by treatment with the JHA.