Host discrimination modulates brood guarding behaviour and the adaptive superparasitism in the parasitoid wasp Trissolcus semistriatus (Hymenoptera: Scelionidae)

Because hosts utilized by parasitoids are vulnerable to further oviposition by conspecifics, host guarding benefits female wasps. The present study aims to test whether female adults regulate brood guarding behaviour by host discrimination in a solitary parasitoid Trissolcus semistriatus by presenting an intact or parasitized host egg mass to a female adult. Virgin females without oviposition experience have host discrimination ability, which enables them to adjust the number of eggs laid in the hosts. Mating experience increases superparasitism by female adults, whereas mated females achieve a higher discrimination ability as a result of oviposition experience and show a lower superparasitism rate. As expected, females exhibit brood guard after parasitizing an intact host egg mass, whereas those females visiting a previously parasitized host egg mass, do not. Because the survival of eggs in superparasitized hosts is relatively low, regulating brood guarding behaviour by host discrimination is adaptive for female wasps.     

Courtship role reversal and deceptive signals in the long-tailed dance fly, Rhamphomyia longicauda

We examined the function of secondary sexual characters in the role-reversed, lekking behaviour of female long-tailed dance flies, Rhamphomyia longicauda Loew (Empididae), to test the hypothesis that the degree of abdominal distention is an honest female signal about the state of egg development. Female Rhamphomyia cannot hunt for prey and they receive all of their protein from males by exchanging copulations for nuptial prey gifts. Females compete for male gifts within leks that are organized for a brief period each evening before dark. Before hovering within leks, females swallow air, inflating expandable pouches on the pleural margins of the abdomen. The result is a large saucer-like abdomen which is further exaggerated by wrapping scaled pro-, meso- and metathoracic legs along its pleural margins. Male preference for an enlarged abdomen was confirmed by suspending plastic models of varying size from monofilament lines and recording which models attracted the most males. There was a positive relationship between egg development and abdominal distention in a related species, R. sociabilis (Williston), which lacks inflatable abdominal pouches. Multiple regression showed that in R. longicauda, abdominal inflation completely masks the state of egg development. We conclude that female R. longicauda deceive mate-seeking males with the unreliable message that eggs are nearing maturation in order to obtain a protein meal in exchange for copulation. Males that fail to identify a female bearing mature eggs risk near-certain cuckoldry and an increased probability that the female will die before oviposition. Copyright 2000 The Association for the Study of Animal Behaviour.     

Egg performance on an egg-carrying bug. Experiments in the field

Selection of oviposition sites has direct influence on female fitness. Differences in offspring survival among sites should favour females to select oviposition sites with the highest survival. Golden egg bug females ( Phyllomorpha laciniata; Coreidae, Heteroptera) use conspecifics as oviposition substrates. Most eggs are laid on the back of the bug but they can be found on all body parts. Females never carry their own eggs, and males commonly carry eggs received without copulation with the donor. We examined differences in egg survival relative to paternity to the egg, host sex and body size and attachment of an egg on a host's body in the field. We also studied which bugs received eggs. Egg loss was surprisingly common: 30–80% of recaptured individuals lost eggs during a six-day period. Unexpectedly, host's paternity to the eggs did not affect egg survival. Also, egg loss did not differ among hosts in respect to other parameters studied. Males received more eggs than females, but survival of the eggs was not better on a male's back than on a female's back. Since egg loss is common in the golden egg bug, we suggest that female oviposition strategy to lay eggs on conspecifics is not perfect. This is due to active behaviour of potential hosts (for example, rejected oviposition attempts) or lack of suitable oviposition substrates (conspecifics) in the field. Since eggs do not survive unless carried, females may be acting opportunistically, doing the best job possible by laying eggs on available conspecifics.     

Age- and density-related oviposition behavior of the European corn borer,Ostrinia nubilalis (Lepidoptera: Pyralidae)

Age-related oviposition patterns ofOstrinia nubilalis were studied at three population densities in the laboratory by releasing newly eclosed adults in wire-screened cages and analyzing their oviposition throughout the adult stage with digital analysis. Oviposition sequences of individual females depositing egg masses were documented on the third and seventh nights after eclosion with a video camcorder. During a sequence, a female produced an egg in an average time of 15 or 26 s on the third and seventh nights, respectively, and completed depositing an egg mass the size of 20–39 eggs in an average time of 316 and 525 s, respectively. Females were not easily disturbed during egg mass deposition and pulsated their abdomen before deposition of each egg. Females produced few egg masses the first night after eclosion. Oviposition increased on nights 2 and 3 but declined steadily thereafter as females matured. Females older than 6 nights produced fewer egg masses; the proportion of egg masses with fewer than 20 eggs increased gradually. By the end of the adults' lifetime, nearly 100% of the egg masses had fewer than 20 eggs. The data are fundamental to our research to define the role of phytochemicals in modifying oviposition behavior of the European corn borer.     

Egg and time limitation mediate an egg protection strategy

The number of mature eggs remaining in the ovaries and the time left for oviposition determine the reproductive decisions of the hyperdiverse guild of insects that require discrete and potentially limiting resources for oviposition (such as seeds, fruits or other insects). A female may run out of eggs before all available oviposition sites are used (egg limitation), or die before using all of her eggs (time limitation). Females are predicted to change clutch size depending on whether eggs or time is the limiting resource. We extend this framework and ask whether the same constraints influence a strategy in which females modify eggs into protective shields. In response to egg parasitism cues, female seed beetles (Mimosestes amicus) lay eggs in vertical groups of 2–4, modifying the top 1–3 eggs into shields in order to protect the bottom egg from attack by parasitoids. We made contrasting predictions of how egg and time limitation would influence egg size and the incidence and level of egg protection. By varying access to seed pods, we manipulated the number of remaining eggs a female had at the time she received a parasitism cue. Although egg size was not affected, our results confirm that egg‐limited females protected fewer eggs and time‐limited females protected more eggs. Female body size explained the number of eggs in a stack rather than host deprivation or the timing of parasitoid exposure. Our results clearly show that host availability relative to female age influences the incidence of egg protection in M. amicus. Furthermore, our study represents a novel use of life history theory to explain patterns in an unusual but compelling defensive behaviour.     

Different oviposition behaviour in Chrysomelid beetles: Characterisation of the interface between oviposition secretion and the plant surface

The behavioural sequence of the oviposition process can be quite distinct in herbivorous leaf beetles: eggs are either connected to the plant surface or laid into mesophyll tissue. Females of two beetle species were recorded with a digital camera in order to analyse their oviposition behaviour. The morphology of the plant-egg-interface was investigated in three species by histological studies and scanning electron microscopy. The lily leaf beetle, Lilioceris lilii, oviposits directly on the plant cuticle without damaging the plant tissue. Several eggs are released together with secretion and are laid in a row. The secretion sorbes in the plant cuticle. In the close relative, Lilioceris merdigera, the secretion can enter the leaf tissue via the stomata. The mustard leaf beetle, Phaedon cochleariae, gnaws a characteristic hole into the epidermis and part of the mesophyll. Into this cavity, it releases in an alternating sequence secretion, a single egg and again secretion. The egg, which is usually laid on plants growing in wetlands, is surrounded by an extrachorion that might possess a respiratory function. The potential eco-physiological relevance of the specific nature of the placement of eggs and secretion on or in the plant tissue is discussed.     

Deposition and longevity of oviposition-deterring pheromone in the cabbage seed weevil

Abstract The cabbage seed weevil ( Ceutorhynchus assimilis Payk.) lays eggs singly into pods of oilseed rape ( Brassica napus L.) through punctures bored with the mouthparts, preferring pods not recently used for oviposition. A simple new choice test has been used to test individual components of egg-laying behaviour for their effect on oviposition site selection. It is confirmed that an oviposition-deterring pheromone (ODP) is deposited during abdomen brushing of the pod which follows egg-laying. Neither pin punctures, weevil feeding punctures, oviposition punctures nor eggs had any deterrent effect. Pods walked on by female weevils were not avoided by those laying eggs. Observations suggest that the ODP is sensed by contact chemoreceptors on the antennae. The deterrent effect lasted only 1–2 h. The implications of these findings on the adaptive significance of the pheromone and its possible use in pest control are discussed.     

Larval life-styles and oviposition behaviour of parasites and grazers

Summary We apply a theory for the computation of reproductive success for insects whose offspring can move from the host on which they hatch to compute the expected reproductive success associated with two kinds of oviposition behaviour. In the first type, accept or reject (AR), the ovipositing female either accepts a host for oviposition and lays her entire egg complement or rejects the host for oviposition. In the second type, clutch optimization (CO), the ovipositing female adjusts clutch according to host properties. The strongest prediction of our theory is that parasites which are both pro-ovigenic (emerge with essentially a full complement of eggs) and use the AR behaviour will be very rare.     

Egg adhesion of the codling moth Cydia pomonella L. (Lepidoptera, Tortricidae) to various substrates: I. Leaf surfaces of different apple cultivars

Codling moths, Cydia pomonella L. (Lepidoptera, Tortricidae), of the first generation deposit eggs on apple leaves in the vicinity of small fruits. The choice of the suitable oviposition sites and proper fixation of eggs are expected to be crucial factors for the survival of the offspring. In this study, we investigated egg adhesion of the codling moth to leaf surfaces of different cultivars of the domestic apple, Malus domestica Borkh., by measuring the pull-off force required to detach the eggs from leaves. Since surface features may influence insect egg adhesion, morphological and physicochemical properties (wettability, free surface energy) of these leaf surfaces were analyzed. Furthermore, eggs and their adhesives covering leaf surfaces were visualized. Eggs on the smooth upper leaf surfaces of all tested cultivars required significantly similar pull-off forces to be detached, at a total average of 6.0?mN. Up to 2?C3 times stronger pull-off forces had to be applied to detach eggs from trichome-covered lower leaves, and these forces differed significantly between cultivars. The role of leaf surface properties is discussed in the context of egg adhesion, oviposition site choice, female attachment, as well as neonate locomotion speed and survival. The obtained results shed light on the susceptibility of various apple cultivars and leaf surfaces to the infestation of apple trees by first-generation codling moths.     

Brochosome influence on parasitisation efficiency of Homalodisca coagulata (Say) (Hemiptera: Cicadellidae) egg masses by Gonatocerus ashmeadi Girault (Hymenoptera: Mymaridae)

Abstract. 1. Many cicadellid females in the tribe Proconiini (Hemiptera: Cicadellidae) cover their egg masses with specialised, usually rod‐shaped, brochosomes as the eggs are being laid. The brochosomes are produced in Golgi complexes in the Malpighian tubules of Cicadellidae. In contrast to the gravid females, adult males, pre‐reproductive adult females, and nymphal males and females produce specialised, usually spherically shaped brochosomes. Brochosomes are also used to cover the external surfaces of nymphs and newly moulted adult males and females. 2. The function of the brochosome covering the egg masses is unknown but various hypotheses have been suggested, including protecting the eggs against pathogens, predators, and parasitoids. Based on preliminary observations of Gonatocerus ashmeadi Girault (Hymenoptera: Mymaridae) parasitising the eggs of the cicadellid, Homalodisca coagulata (Say), it is speculated here that brochosomes covering an egg mass hinder parasitisation of eggs by G. ashmeadi. This hypothesis was tested by observing G. ashmeadi females foraging on leaves with H. coagulata egg masses heavily covered with rod‐shaped brochosomes vs. those lacking brochosomes. 3. Cox's proportional hazards model was used to evaluate the probability, per unit time, that a female G. ashmeadi displayed the sequence of behaviours that ended in successful oviposition as influenced by five variables: (a) presence or absence of brochosomes on an egg mass, (b) the leaf surface, upper or lower, being searched by the parasitoid (the egg masses are laid in the parenchyma on the lower leaf surface), (c) the parasitoid's previous ovipositional experience, (d) egg mass size, and (e) the parasitoid's age. 4. Brochosomes significantly decreased oviposition efficacy of G. ashmeadi females. Scanning electron microscopy showed that females exposed to brochosome‐covered egg masses had brochosomes adhering to their tarsi, legs, antennae, and eyes, all of which prompted extensive bouts of grooming.     

Importance of plant physical cues in host acceptance for oviposition by Busseola fusca

Plant and surrogate stems exhibiting specific combinations of physical cues were used to determine which plant‐related stimuli influence the oviposition of Busseola fusca (Fuller) (Lepidoptera: Noctuidae). The number of eggs and egg batches laid per female increased with an increase in diameter of both natural and artificial stems. Direct observations of the oviposition behaviour (walking, antennating, and sweeping with the ovipositor) indicated that the female moths preferred oviposition supports with a large diameter and non‐pubescent or smooth surfaces over pubescent or rough ones. Pubescence and rough surfaces significantly affected the behavioural steps leading to oviposition by interfering with the ovipositor sweep process necessary to find a suitable oviposition site. Furthermore, more eggs and egg batches were laid on soft than rigid supports. The rigidity of the support affected the proper insertion of the ovipositor for egg deposition. Our results underline the importance of physical stimuli in B. fusca's choice of an oviposition site, which may facilitate the identification of potential host plants or preferred oviposition sites on a plant for this species.     

Reduced realised fecundity in the pine looper Bupalus piniarius caused by host plant defoliation

1. Whether the potential fecundity of herbivorous insects is realised or not may depend on female behaviour, which in turn may be influenced by host plant acceptability. Female Bupalus piniarius were observed to discriminate against needles growing out the year following defoliation (current‐year needles) of its host plant Pinus sylvestris. 2. It was hypothesised that the discriminatory behaviour was due to current‐year needles being less secure as a substrate. Field and laboratory experiments were designed to test this hypothesis and to estimate the discrepancy between potential and realised fecundity when females were offered defoliated branches. 3. In a laboratory oviposition experiment, B. piniarius females were exposed to branches bearing either current‐year needles only or both mature and current‐year needles. Daily oviposition rate, egg batch size, longevity, and mature eggs and fat retained at death were recorded for each female. In field experiments, the rate at which eggs dropped from expanding needles and the capacity of neonate larvae hatching from the dropped eggs to colonise a tree were assessed. 4. Significantly fewer eggs were laid when females were exposed to defoliated branches. 5. Twenty‐six and 16% of the eggs laid on current‐year needles dropped from the needles in 1998 and 1999 respectively, whereas no eggs dropped from mature needles in 1998 and only one egg (< 1%) dropped in 1999. 6. A very small proportion of larvae hatching on the forest floor (simulated egg drop) was able to recolonise host trees. 7. These results emphasise the importance of oviposition behaviour on realised fecundity when analysing insect population dynamics. In the case of B. piniarius, egg placement, although a minor detail during the normal course of events, became of key importance when defoliation deprived females of their preferred egg attachment site.     

Host selection by Homalodisca vitripennis: the interplay between feeding, egg maturation, egg load, and oviposition

For some phytophagous insects, egg maturation may be dependent on adult feeding. Accordingly, rates of egg maturation may be dependent on the quality and quantity of available food sources. In turn, oviposition behavior could be affected by diet quality via changes in egg load (number of mature eggs carried by a female). Experiments were conducted to determine whether adult feeding may affect oviposition behavior of the glassy-winged sharpshooter, Homalodisca vitripennis. No-choice tests demonstrated that eggs accumulated in glassy-winged sharpshooter abdomens as time since last oviposition increased largely as a function of feeding plant species. In choice tests, glassy-winged sharpshooter females were observed most frequently on the plant species that imparted the greatest egg maturation rate in no-choice tests. Direct tests of the effects of egg load on glassy-winged sharpshooter oviposition behavior found that females were more likely to deposit eggs as egg load increased. Similarly, acceptance of a low-ranked oviposition plant species by female glassy-winged sharpshooters increased with egg load and time since last oviposition. The results indicate that adult feeding affected glassy-winged sharpshooter egg maturation, plant species varied in quality for providing nutrients for egg maturation, and egg load affected oviposition behavior. Thus, the quantity and quality of available feeding plant species may affect glassy-winged sharpshooter egg maturation rates, which in turn may affect the plant species female glassy-winged sharpshooters select for oviposition.     

Experience‐ and age‐mediated oviposition behaviour in the yellow fever mosquito Stegomyia aegypti (=Aedes aegypti)

In repeated behaviours such as those of feeding and reproduction, past experiences can inform future behaviour. By altering their behaviour in response to environmental stimuli, insects in highly variable landscapes can tailor their behaviour to their particular environment. In particular, female mosquitoes may benefit from plasticity in their choice of egg‐laying site as these sites are often temporally variable and clustered. The opportunity to adapt egg‐laying behaviour to past experience also exists for mosquito populations as females typically lay eggs multiple times throughout their lives. Whether experience and age affect egg‐laying (or oviposition) behaviour in the mosquito Stegomyia aegypti (=Aedes aegypti) (Diptera: Culicidae) was assessed using a wind tunnel. Initially, gravid mosquitoes were provided with a cup containing either repellent or well water. After ovipositing in these cups, the mosquitoes were blood‐fed and introduced into a wind tunnel. In this wind tunnel, an oviposition cup containing repellent was placed in the immediate vicinity of the gravid mosquitoes. A cup containing well water was placed at the opposite end of the tunnel so that if the females flew across the chamber, they encountered the well water cup, in which they readily laid eggs. Mosquitoes previously exposed to repellent cups became significantly more likely to later lay eggs in repellent cups, suggesting that previous experience with suboptimal oviposition sites informs mosquitoes of the characteristics of nearby oviposition sites. These results provide further evidence that mosquitoes modify behaviour in response to environmental information and are demonstrated in a vector species in which behavioural plasticity may be ecologically and epidemiologically meaningful.     

Context-Dependent Plastic Response during Egg-Laying in a Widespread Newt Species

Previous research on predator-induced phenotypic plasticity mostly focused on responses in morphology, developmental time and/or behaviour during early life stages, but the potential significance of anticipatory parental responses has been investigated less often. In this study I examined behavioural and maternal responses of gravid female smooth newts, Lissotriton vulgaris, in the presence of chemical cues originating from invertebrate predators, Acilius sulcatus water beetles and Aeshna cyanea dragonfly larvae. More specifically, I tested the extent of oviposition preference, plasticity in egg-wrapping behaviour and plasticity in egg size when females had the possibility to lay eggs at oviposition sites with and without predator cues during overnight trials. I found that individuals did not avoid laying eggs in the environment with predator cues; however, individuals that deposited eggs into both environments adjusted the size of the laid eggs to the perceived environment. Females deposited larger eggs earlier in the season but egg size decreased with time in the absence of predator cues, whereas individuals laid eggs of average size throughout the investigated reproductive period when such cues were present. Also, egg size was found to be positively related to hatching success. Individuals did not adjust their wrapping behaviour to the presence of predator cues, but females differed in the extent of egg-wrapping between ponds. Females’ body mass and tail depth were also different between ponds, whereas their body size was positively associated with egg size. According to these results, female smooth newts have the potential to exhibit activational plasticity and invest differently into eggs depending on temporal and environmental factors. Such an anticipatory response may contribute to the success of this caudate species under a wide range of predator regimes at its natural breeding habitats.     

Distribution of second generation European corn borer,Ostrinia nubilalis, egg masses in field corn and relationship to subsequent tunneling damage

The relationship between second generation European corn borer (Ostrinia nubilalis Hübner) egg mass numbers and subsequent field corn damage, as measured by stalk cavity numbers, was studied in 79 fields in northeastern North Carolina over three years. A mean of 0.028 egg masses per plant (645 egg masses/23400 plants) was found over the course of the study. Significant differences in oviposition rate were detected between fields and years. Ca. 85% of egg masses were deposited in a five leaf zone surrounding the primary ear; of these, 89% were found on the lower four leaves in this zone. Egg masses appeared to be distributed randomly within fields but at low rates of incidence, and oviposition was relatively uniform between sampling areas within individual fields. Under moderate to high oviposition pressure (mean number of egg masses per plant over the duration of the oviposition period >ca. 0.02), eggs laid during the early phases of the oviposition period account for more subsequent stalk damage than eggs laid during the later phases of the oviposition period. Variations in second generation egg mass numbers accounted for ca. 70% of variation in stalk cavity numbers.     

Oviposition‐induced plant cues: do they arrest Trichogramma wasps during host location?

Plants can defend themselves against herbivorous insects before the larvae hatch from eggs and start feeding. One of these preventive defence strategies is to produce plant volatiles, in response to egg deposition, which attract egg parasitoids that subsequently kill the herbivore eggs. Here, we studied whether egg deposition by Pieris brassicae L. (Lepidoptera: Pieridae) induces Brussels sprouts plants to produce cues that attract or arrest Trichogramma brassicae Bezdeko (Hymenoptera: Trichogrammatidae). Olfactometer bioassays revealed that odours from plants with eggs did not attract or arrest parasitoids. However, contact bioassays showed that T. brassicae females were arrested on egg‐free leaf squares excised from leaves with 72 h‐old egg masses, which are highly suitable for parasitisation. We tested the hypothesis that this arresting activity is due to scales and chemicals deposited by the butterflies during oviposition and which are thus present on the leaf surface in the vicinity of the eggs. Indeed, leaf squares excised from egg‐free leaves, but contaminated with butterfly deposits, arrested the wasps when the squares were tested 1 day after contamination. However, squares from egg‐free leaves with 72 h‐old butterfly deposits had no arresting activity. Thus, we exclude that the arresting activity of the leaf area near 72 h‐old egg masses was elicited by cues from scales and other butterfly deposits. We suggest that egg deposition of P. brassicae induces a change in the leaf surface chemicals in leaves with egg masses. A systemic induction extending to an egg‐free leaf neighbouring an egg‐carrying leaf could not be detected. Our data suggest that a local, oviposition‐induced change of leaf surface chemicals arrests T. brassicae in the vicinity of host eggs.     

柳蓝叶甲的生物学特性室内观察

在实验室条件下(27±0.5℃,12 D:12 L)研究了柳蓝叶甲Plagiodera versicolora的生物学特性。结果表明,幼虫每隔2~3 d蜕皮1次,共蜕皮2次。成虫产卵前期4 d,世代周期17 d。产卵具有一定的周期性,约10 d为1个周期。叶龄对幼虫的生长发育和成虫的产卵行为有极显著的影响。当幼虫以成熟叶为食料时,其生长发育速率明显减慢,各龄期延长,蛹重较轻;成虫以成熟叶或老叶为食料时产卵几乎完全被抑制,而以幼叶为食料时几乎每天均能产卵。此外,柳蓝叶甲产出卵表面化合物对产卵行为有极为显著的影响。卵表面的有机提取物对成虫产卵具有显著的引诱作用;相反,卵表面的水提取物对产卵有一定的驱避作用。     

Biochemical Composition of the Eggs of the Freshwater Snail Lymnaea stagnalis and Oviposition-induced Restoration of Albumen Gland Secretion

Summary

Galactogen and protein form the main constituents of the eggs of Lymnaea stagnalis. The amount of galactogen per egg is fairly constant, irrespective of the size of the egg mass or the age of the snail.

The restoration of the albumen gland, which produces the perivitelline fluid for the eggs, was studied in long-day (16 hr light-8 hr dark) snails after spontaneous oviposition. The wet wt of the gland and its galactogen and protein contents are markedly increased within 8 hr and reach a maximum at 32 hr after oviposition. These maxima correspond to the levels determined in snails that did not lay eggs for at least 1 to 2 days. The amounts of galactogen and of protein in the albumen gland are linearly related to the wet wt of this gland.

The restoration period of the albumen gland almost covers the mean egglaying interval. This implies synchronized cycles of albumen storage and egg formation.

The estimated amount of galactogen, released by the albumen gland during egg mass formation, is in accordance with that deposited in the eggs. In contrast, the wet wt of the eggs is 4.6 times higher than that of the released secretory material. Since after oviposition water uptake by the eggs in the egg mass is negligible, the perivitelline fluid, which is released by the albumen gland and surrounds the egg cell, must be diluted in the reproductive tract of the snail prior to oviposition.     

The role of internal pressure and muscle activation during locust oviposition

The oviposition of female locusts is a complex behaviour that includes a dramatic extension of the abdomen. The role of internal pressure during oviposition was investigated by monitoring the intra-tracheal pressure and the activity of selected longitudinal muscles, while movements of the abdomen were visualised with a video imaging system. Locust oviposition consists of a sequence of four distinct phases: (i) probing the substrate and digging without elongation of the abdomen, (ii) longitudinal extension of the abdomen up to four times its normal length, (iii) laying packages of eggs while (iv) gradually withdrawing the abdomen. During extension, neurograms and myograms of selected longitudinal muscles revealed a decreased level of activity. When the abdomen retracted to its normal length, muscle activity re-appeared. In phases two and three, rising internal pressure prevented the abdomen from slipping back when the valves released their lateral grip from the substrate. Locking the genital segments in the hole by relative bending kept the abdomen in place when producing foam or laying eggs. Intra-abdominal pressure, therefore, is not the main cause of abdominal extension, but rather maintains extension when no mechanical locking in the hole prevents the abdomen from elastic retraction.