Extrication,the primary event in eclosion,and its neural control in Sarcophaga bullata

The flesh fly, Sarcophaga bullata, employs the same “extrication-behaviour” programme for adult eclosion and, as the newly eclosed fly, for digging its way out of the medium in which pupariation occurred. Studies reveal that the thoracic ganglion probably contains the neural circuits required for local extrication movements, with the ganglia in the head being essential for coordinated programme execution.The onset and cessation of the programme appear to be controlled by sensory feedback. Information generated by normal leg activity, e.g. walking, resting without restraint on a surface, or manipulating a polystyrene ball, can stop an ongoing programme, whereas restriction of leg movement releases the behaviour.     

Eclosion behavior in tsetse (Diptera: Glossinidae): Extrication from the puparium and expansion of the adult

The tsetse adult extricates itself from the puparium and surrounding substrate by a series of muscular contractions that generate a Stereotypic pattern of changes in hemolymph pressure. The digging action of the fly can be distinguished from a second pattern of hemocoelic pulsations that is used to remove obstacles from its path. When the fly is restrained extrication behavior will persist for over 10 h. If the adult's legs are freed while the remainder of the body remains encased in the puparium, the fly fails to engage in extrication behavior, a result which suggests that freedom of the legs switches off extrication behavior and permits the onset of expansion of the body to its final adult size and shape. Expansion behavior includes walking, grooming, pumping air into the gut, and contracting the abdominal muscles to generate rhythmic pulses of hemocoelic pressure. A barographic record of internal pressure changes reflects the dynamics of this morphogenetic process. Results from tsetse are compared with previous observations recorded in flesh flies.     

Eclosion and hatching in cockroach first instar larvae: A stereotyped pattern of behaviour

Cockroach pharate first instar larvae (Periplaneta americana) spontaneously initiate eclosion and hatch from the oötheca after 30 days' incubation at 29°C. The caudal-to-rostral peristaltic movements involved in both eclosion and hatching are initiated even when the first instar larvae have been removed from the oötheca and incubated separately in organ culture dishes. Therefore, environmental stimuli and conditions in situ are not necessary for the onset of eclosion. However, environmental factors associated with the cuticle control the termination and/or duration of the eclosion behaviour sequence. Larvae which had cuticles glued to their bodies had longer than normal eclosion episodes while larvae which experienced premature cuticle removal immediately ceased the movements. Cuticle removal immediately ‘switched’ behaviour from that of the larva to that of the adult with the characteristic walking gait. Eclosion in larvae removed from the oötheca could be initiated by tactile stimulation. A rôle for this response in synchronizing the eclosion-hatching movements of the many larvae within an oötheca is suggested.     

The expansion of Schistocerca gregaria at the imaginal ecdysis: The mechanical properties of the cuticle and the internal pressure

(1) At the imaginal ecdysis of Schistocerca, the cuticle is viscoelastic rather than elastic. (2) The cuticle becomes softer just before emergence. It is suggested that this is due to an ‘eclosion hormone’. The softening permits the extrication of the appendages and an increase in the size of the locust. (3) The stiffness of the cuticle increases transiently at the end of emergence. (4) In the later part of wing expansion, the cuticle becomes elastic and its stiffness again increases. (5) Maximum pressures are recorded about 10 min into emergence, and pressures in the gut are greater than those in the tracheal system. (6) From these results it is concluded that the expansion of the wings initially depends on the high haemolymph pressure and low stiffness. Only in the last 15 min of wing expansion do the processes involved in autonomous expansion become important.     

A ‘simple clock’ approach of circadian rhythms: An easy way to predict the clock's singularity

A graphical method is presented which allows the prediction of phase resetting curves of circadian rhythms for both type 1 and type 0 resetting, starting from one experimentally determined phase resetting curve. Calculations were based on literature data for the pupal eclosion rhythm of Drosophila pseudoobscura. The method is based on the assumption that for all practical purposes the rhythm may be approached as a “simple clock”, i.e. an oscillator with only one state variable, namely its phase or circadian time, CT. Besides predicting both “types” of phase resetting the method is capable to locate the “position” of the phase singularity and thus the transition from type 1 to type 0 resetting. This graphical method is an elaboration of the “transformation method”, developed in 1972 by A. Johnsson and H. G. Karlsson, which was effective in predicting phase resetting by “strong” stimuli, but failed in the case of “weak” stimuli. Predictions made using the extended transformation method are in good agreement with experimental results obtained with Drosophila. Also for the flesh fly, Sarcophaga argyrostoma, a prediction is made of the position of the phase singularity of the eclosion rhythm and compared with experimental results.     

Postnatal development of subterranean habits in tuco-tucos <Emphasis Type="Italic">Ctenomys talarum</Emphasis> (Rodentia,Caviomorpha, Ctenomyidae)

Postnatal development of subterranean habits was investigated in Los Talas’ tuco-tuco Ctenomys talarum, a subterranean caviomorph rodent endemic from South America. Since in this species, some key morpho-functional traits related to scratch-digging behaviour—a form of underground progression—are already present during early ontogeny and develop progressively, we predicted that this behaviour expresses early during postanatal development and its performance enhances gradually from pups to adults. The process of acquisition of different behaviours associated to the construction of a burrow system was recorded in 11 individuals, each one coming from different litters, inside a terrarium filled with natural soil. We found that scratch-digging and burrowing behaviours expressed early during postnatal development, particularly, during lactancy. The digging of a “true burrow” clearly preceded the dispersal age, with a high inter-individual variability, from 18 (lactancy) to 47 (post-weaning) postnatal days. Pups could lose the soil using their foreclaws and remove the accumulated substrate using their hindfeet as adults do. During lactancy individuals could construct a simple burrow to shelter, and first burrow construction occurred in the absence of either a burrowing demonstrator or an early subterranean environment (a natal burrow). However, certain features of the complex burrow system that characterize this species, such as lateral branches and nest chamber, just appeared after weaning. The time elapsed until animals started to dig and the time dedicated to underground activities varied with age, decreasing and increasing, respectively. In sum, our results show that—in C. talarum—immature digging behaviour gets expressed early during ontogeny, and develops progressively. The role of the early ability to build its own burrow and its possible function influencing the development of musculoskeletal traits and on efficiency for such conduct is discussed.     

Neuromuscular and hormonal control of post-eclosion processes in flies

Flies (exemplified by Sarcophaga bullata) expand after eclosion from the puparium by processes of “pulsing” (slow rhythmical abdominal contractions) and “pumping” of air (fast rhythmical contractions of the cibarial pump). Pulsing and pumping are inhibited if a newly eclosed fly is kept in an enclosed space (sand, a glass tube, an empty puparium). This inhibition no longer applies if such flies are injected with either hemolymph from flies 10–15 min old or cAMP, or are confined at the age of 10 min. This suggests a hormonal control of pulsing and pumping. Pumping alone, without pulsing, occurs in flies treated with certain paralyzing agents like ether, tetrodotoxin, bee venom, or “FlyNap,” or have the connectives in the neck cut or interrupted by cauterization. Application of FlyNap or neck cauterization leads to excessive pumping which results in bloating. Expansion by bloating is confined to the soft membranes, leaving sclerites and wings largely unexpanded. The function of pulsing is probably that of “plasticizing” and stretching the cuticle to make it respond to increased steady pressure by air-pumping. Flies ligated at the proboscis show almost regular pulsing and pumping, but without intake of air, and consequently no expansion. Cuticle (sclerites) and wings, however, become plasticized. Some plasticization occurs even in the absence, or reduction, of pulsing (in a neck-cauterized fly), brought about by a hormonal process. Eclosion from the puparium is also initiated by a hormonal action. Thus, the following processes during fly emergence are controlled by hormones: eclosion proper, pulsing, pumping, plasticization, and tanning. These hormones are separate entities, with the possible exception of the pulsing/pumping hormone(s).     

Estimating risk probabilities for sickness from behavioural patterns to identify health challenges in dairy cows with multivariate cumulative sum control charts

Dairy cattle housing is characterised by increasing herd sizes and the need for assisting technical tools to monitor the cows’ health. This study investigated the combination of logistic regression models with multivariate cumulative sum (MCUSUM) control charts in healthmonitoring of dairy cattle. Sensor information of 618 cows with 791 lactations (138 438 cow days), nine behavioural variables were included as parts of the behavioural patterns: physical activity (“neck activity”, “leg activity”, “walking duration”), resting (“lying duration”, “standing duration”, “transitions from lying to standing”) and feeding (“feeding duration”, “rumination duration”, “inactivity duration”) behaviour. For each of these behavioural patterns, a logistic regression model with the health status (sick vs not sick) as a dependent variable was designed after a variable selection (herd level) based on the herd dataset with 618 cows (618 lactations; 115 547 cow days), which included the variables of each behaviour pattern and the stage of lactation nested in the number of lactations as explanatory variables. The explanatory variables were added stepwise to the model, with the final model being selected with respect to the lowest values of Akaike’s and Bayes’ information criteria. Each model was then applied to a dataset with 173 cows (22 891 cow days) at cow level, resulting in individual daily risk probabilities for getting sick. Thus, risk probabilities of each behavioural pattern were estimated and included in the MCUSUM control charts to identify cows at risk of disease. The performance of the MCUSUM control charts was cross-validated to identify the best fitting reference value k and the threshold value h. Alerts given within 5 days prior to diagnosis were counted as detected sicknesses. The performance resulted in a block sensitivity of 70.9–81.4%, specificity of 87.9–94.2% and a false-positive rate of 5.8–12.1%. The performance was confirmed while testing the entire algorithm resulting in a mean area under the receiver operating characteristics curve of 0.89. Calculating precision and the F₁-score resulted in a precision of 49.0–60.9% (training: 48.8–63.5%) and an F₁-score of 61.1–65.7% in testing (training: 61.0–67.0%). The precision-recall curve (PRC) was derived from precision and recall with an area under the PRC of 0.70 in training and testing. In summary, the present study was able to develop an algorithm showing good classification potential for the online monitoring of sickness behaviour.     

Re-evaluation of the role of corpora cardiaca in calling and oviposition behaviour of giant silk moths

Re-investigation of the role of the corpora cardiaca in the reproductive behaviour of the giant silkmoths, Hyalophora cecropia and Antheraea polyphemus, showed that this pair of glands plays no essential role, either in “calling” behaviour by virgin females or in increased oviposition due to mating. Removal of corpora cardiaca-corpora allata complexes, either from diapausing pupae or from freshly eclosed adult females, had no effect on the calling behaviour or on its timing in either species. Moreover after mating, these operated females laid eggs in the typical mated oviposition pattern. Furthermore, females in which there was only a nervous connection between the brain and the abdomen but no haemolymph circulation called normally and oviposited after mating.Although the corpora cardiaca were not essential for calling behaviour, hormogenates of corpora cardiaca-corpora allata complexes and blood from calling or ovipositing females induced a typical “calling” response in 30–60% of the isolated virgin H. cecropia abdomens tested. This activity was not species-specific as it was also found in Manduca sexta, but the restriction of major activity to corpora cardiaca extracts and haemolymph suggested that a neurosecretory factor may modulate the normal neural control of calling behaviour.     

Eclosion hormone activity in developing embryos of the silkworm, Bombyx mori

The ultimate timing of hatching in the silkworm, Bombyx mori, is controlled by a circadian oscillator. The presence of eclosion hormone in developing embryos of the silkworm is demonstrated. Eclosion hormone activity first becomes detectable in embryos which have developed almost to the stage of the differentiation of the neuroendocrine system. Hormonal activity increases sharply to a maximum level 1 day before hatching and falls by about a half in the newly hatched larvae. Eclosion hormone was partially purified from the pharate first-instar larvae and approx, a 2100-fold purification was achieved. The molecular weight of the embryo eclosion hormone is estimated to be 7000 ～ 9000 Daltons by gel-filtration on Sephadex G-50 (superfine). The role of eclosion hormone on hatching behaviour of the silkworm, Bombyx mori, is discussed.     

Defensive behaviour and its inheritance in the anabantoid fish, Macropodus opercularis and Macropodus opercularis concolor

In this preliminary study defense behaviour patterns (fear responses) are described in two closely related, behaviourally different inbred labyrinth fish subspecies and in their F₁ generation. The subspecies M. opercularis (characterized briefly by “active escape”) and M. opercularis concolor (characterized by “passive escape”) showed specific differences in the manifestation of certain defense behaviour patterns. In the F₁ hybrid generation dominance and overdominance of M. opercularis was found in most defense behaviour patterns. Analysing the frequencies and sequences of movement patterns it could be shown that defensive behaviour is not a random or entirely “plastic” process but that there is sequential linkage between the patterns and they form characteristic clusters. Our results suggest that manifestations of different patterns are under genetic control and presumably, genetic determination of certain patterns is not very complex. Attempts were made to determine whole brain noradrenaline, serotonine and dopamine levels of the two subspecies and a significant difference was found in the noradrenaline content.     

Azadirachtin,ecdysteroids and ecdysis in Locusta migratoria

1. The injection of azadirachtin into eight-day-old 5th instar nymphs of Locusta migratoria elicits different morphological effects dependent upon the timing of the injection relative to the major haemolymph ecdysteroid peak.2. Ecdysteroid levels can be drastically reduced, delayed and extended or not significantly affected by azadirachtin treatment dependent upon the timing of injection.3. Air swallowing to inflate the gut during the moult is significantly impaired in azadirachtin-treated insects resulting in both a reduced rate of inflation and a reduction in total air volume.4. The significance of azadirachtin treatment to the attainment of a “critical body mass”, the release of eclosion hormone and successful ecdysis is discussed.     

Concepts of macro- and microevolution as related to the problem of origin and global expansion of the plague pathogen Yersinia pestis

The relationship between macro- and microevolutionary processes is considered with reference to the ecological scenario of the origin of the plague pathogen and its subsequent natural and anthropogenic global expansion. The macroevolutionary transformation of the ancestral pseudotuberculosis microbe clone into the initial plague microbe Yersinia pestis tarbagani occurred in Central Asia at the end of the Late Pleistocene by a “vertical” Darwinian way in an inadaptive heterothermal continual intermediate environment—the Mongolian marmot Marmota sibirica—flea Oropsylla silantiewi system—via a sequence of unstable and currently extinct intermediate forms. Its natural geographic expansion on the “oil spot” principle in the postglacial time led to the microevolutionary formation of 20–30 hostal subspecies circulating in populations of the background species of burrowing rodents and pikas in arid areas of Eurasia. The intercontinental spread of the “marmot” and “rat” pathogen subspecies in the past few centuries has been exclusively anthropogenic, with the involvement of synanthropic (ship) rats.     

Antlions (Neuroptera,Myrmeleontidae) from ornithological traps on the Curonian Spit: A three-species community containing a new species

A mixed antlion community is recorded at the “Fringilla” Research Station, “Rybachii” Biological Station, Curonian Spit, Baltic Sea, Kaliningrad Province, Russia; the adults were captured using ornithological traps and the larvae were found on sand dunes around. The ratio of the larval numbers in the mixed colonies of Myrmeleon tschernovi sp. n., Myrmeleon formicarius L., and Euroleon nostras (Geoffr.) is 100: 3: 2. The new species is described, the other two are recorded in Kaliningrad Province for the first time. Morphologically, Myrmeleon tschernovi sp. n. is similar to Myrmeleon bore (Tjed.), being its neighbor in the Baltic Region and occupying its econiche. The most characteristic distinctions between these species are found in the male genitalia and in the larval head chaetotaxy and color pattern.     

Endocrine manipulations,juvenile hormone and ontogenesis of male sexual behaviour in locusts

By injection of azadirachtin (AZA) to 5th instar hoppers of Locusta migratoria migratorioides, “over-aged” nymphs were produced, some of which survived for long periods without moulting to adults. Such over-aged male nymphs exhibited mating behaviour belatedly, though at a lower intensity than normal adults. Juvenile hormone (JH) markedly intensifies male sexual behaviour in crowded adult (6th instar) Locusta; thus, the lower intensity of sexual behaviour of the over-aged nymphs might have been caused by a low endogenous JH titre, or by a weaker response to the hormone. To clarify this point, chemically allatectomized over-aged 5th instar nymphs were produced by combined treatment with precocene and AZA. These exhibited a very low intensity of mating behaviour. Injections of exogenous JH III intensified the mating behaviour of the chemically allatectomized over-aged 5th instar male nymphs in a dose-dependent way, but the effect was weaker than in chemically allatectomized adult males. AZA-induced over-aged 4th instar male Locusta hoppers also exhibited mating behaviour. By injection of AZA, 5th instar over-aged hoppers of Schistocerca gregaria were also produced. Males showed mating behaviour belatedly, despite the fact that JH completely controls male sexual behaviour in this species. The intensity of this behaviour was again lower than in adults. Interrelations between integumental morphogenesis, “ethogenesis” of male sexual behaviour, and endocrine factors in locusts are discussed.     

The taxonomy,chronostratigraphy and paleobiogeography of glyptosaurine lizards (Glyptosaurinae,Anguidae)

Glyptosaurine lizards (Glyptosaurinae, Anguidae) are an extinct group of heavily armored lizards known from North America, Europe and Asia. Glyptosaurine lizards, taxa that possess fully developed tuberculated dermal armor, appear to have been established in North America by late early Puercan time (To3). “Proxestops,” a taxon distinguished by a combination of vermiculate and tuberculated osteoderm sculpturing, is considered to be a non-glyptosaurine, a sister taxon of the Glyptosaurinae. Known from only fragmentary remains, its wide chronostratigraphic distribution suggests that “Proxestops” is a form genus that, in all probability, represents more than one taxon, that ranges from the middle Paleocene to the early Eocene of North America. Moreover, the taxa Odaxosaurus piger, Parodaxosaurus sanjuanensis and “Proxestops” are best considered “proto-glyptosaurines”. “Melanosaurins” and glyptosaurins were well-established by the early Eocene, especially in North America, and are here documented by their type species and chronostratigraphic levels. Both tribes are present in Europe (MP7), too, but the record is not as estensive as that of North America. The North American taxon Gaultia silvaticus (Wa0) is transitional between a “melanosaurin” and glyptosaurin. Because it lacks the well-defined hexagonal osteoderms that characterize the Glyptosaurini, it is removed from that group and considered to be a “melanosaurin”. The “melanosaurin” taxon “Xestops” savagei (Wa4–Wa6) cannot be referred to Xestops (Br2) based on non-corresponding elements and because superficial similarity does not justify assignment to this taxon. Arpadosaurus sepulchralis (Wa6?), whose holotype is a fragmentary right frontal, is considered a subjective junior synonym of A. gazinorum, based on minor differences in the epidermal scale pattern that probably represent individual variation. “Glyptosaurus” agmodon (Wa6?), based on a partial right maxilla, cannot be referred to Glyptosaurus (sensu stricto), and the material upon which this taxon is based bears strong resemblance to material identified as cf. “?Paraglyptosaurus” yatkolai (Wa5–Wa6). “Glyptosaurus” rhodinos (Wa5) is based on an incomplete parietal, and its reference to Glyptosaurus is considered problematic. Eoglyptosaurus donohoei (Wa7) is probably valid and is re-established here. Glyptosaurus (sensu stricto) is known solely from the middle Eocene (Br2) by G. sylvestris. Dimetoposaurus wyomingensis (Br3) is removed from Xestops vagans because its synonymy was based on superficial similarities. Helodermoides tuberculatus, the largest and last glyptosaurin (Ch3), is restricted to the Chadronian of North America. Only the “melanosaurin” Peltosaurus granulosus (Or2–Or3), which includes the species P. abbotti, seems to have crossed the Eocene-Oligocene boundary, and appears to be largely restricted to the Orellan, but extended into the Arikareean. European glyptosaurines are also represented by both glyptosaurins and “melanosaurins” early in the Eocene (MP7). Placosauriops-like “melanosaurins” are known from Dormaal (MP7), and the glyptosaurin taxon?Placosaurus ragei occurs at the same level. “Placosauriops abderhaldeni” has been identified from the Grube Messel (MP11), but this assignment remains dubious because the species has not been adequately diagnosed, and the holotype species is from the Geiseltal (MP13), which is some 4.5 million years younger. Placosauriops weigelti (MP13) is the only valid species of this genus. Paraxestops stehlini (MP14) is not referable to the North American taxon Xestops, and its relationship to Placosauriops has not been studied. The late Eocene glyptosaurins Placosaurus estesi (MP17) and P. rugosus (MP18) are the last glyptosaurines known from Europe and appear to have gone extinct at the Eocene-Oligocene boundary, casulties perhaps of the “Grande Coupure”. Asian glyptosaurines are known solely from one species, Stenoplacosaurus mongoliensis, from the middle Eocene (Sharamurunian) of China. Glyptosaurines most likely originated in North America, diversified by late Paleocene time, and rapidly spread across the North Atlantic into Europe by the early Eocene. Both “melanosaurins” and glyptosaurins took a foothold in Europe by the early Neustrian, but the glyptosaurins, aside from one occurrence (Dormaal, MP7), were conspicuously absent for most of Neustrian through early Robiacian time. In North America, glyptosaurins diversified during the early and middle Eocene, while in Europe small “melanosaurins” were a prominent part of the paleoherpetofauna, and glyptosaurins are unknown for most of the Neustrian through the Geiseltalian, in both the fossilferous Lagerstätten of Messel and Geiseltal. Stenoplacosaurus is the only known glyptosaurin glyptosaurine from Asia, and its abrupt appearance during the late Eocene suggests the possiblity of a Beringian dispersal from North America into Asia.     

Social interactions in a group of free-ranging sows

Earlier studies on social behaviour in the domestic pig have only been performed on groups housed artificially. The present study was undertaken to investigate the social behaviour of free-ranging sows in a well-established flock, so that it could be compared with the behaviour of sows housed indoors that had been studied earlier. The free-ranging flock, which contained 13 animals, 5 of which were sows, lived in an enclosure, approximately 1.1 ha in area. The interaction sequences were registered during 1 h after feeding and during foraging in the enclosure. With sequence analysis, the effect of different behaviours on the behaviour of the receiver animal was measured. This allowed the behaviour patterns to be classed after function. “Aiming”, a behaviour pattern not previously seen, was interpreted as a threat behaviour, being “statistically strongly directive” towards submissive and flight behaviour. The functions of the other 7 behaviours included in the study were found to be the same as in the indoor groups studied earlier. The dominance order was not found to be better established than in the most extensively housed indoor group, but the level of aggression was very low, indicating that a stable dominance order does not automatically lead to a low level of aggression. It was found that the dominance order was mainly maintained through the behaviour of the subordinate animals, supporting the use of an “avoidance order” instead of an “aggression order” as a measurement of social dominance. On the whole, no dramatic differences were found between the behaviour of the free-ranging sows and that of the most extensively housed of the indoor groups studied earlier, although great differences were found between free-ranging sows and sows kept in confinement indoors.     

The role of eclosion hormone in the larval ecdyses of Manduca sexta

Each larval moult in Manduca sexta consists of an identical series of developmental and behavioural events leading up to ecdysis. Injections of eclosion hormone into staged larvae in any instar resulted in the premature elicitation of the larval pre-ecdysis behaviour, comprising a rhythmic sequence of muscle contractions, followed by the larval ecdysis behaviour.A marked depletion of eclosion hormone stores form the ventral chain of ganglia coincided with each larval ecdysis and in the moult to the fifth instar, eclosion hormone activity appeared in the blood at the onset of the pre-ecdysis behaviour.Responsiveness to eclosion hormone for pre-ecdysis and ecdysis behaviour developed about 12 and 6 hr before normal ecdysis, respectively. Elicitation of ecdysis behaviour by exogenous hormone inhibited both subsequent behavioural responses to eclosion hormone and endogenous hormonal release.In conclusion, the behavioural programme involved in each larval ecdysis appears to be controlled by the eclosion hormone.