Evidence for woodlice-specialization in Dysdera spiders: behavioural versus developmental approaches

Abstract.  The dietary specialization in a woodlouse-eating spider Dysdera hungarica Kulczyñski (Araneae: Dysderidae) is studied using two types of laboratory experiments. In the first experiment, the rate of development of spiderlings reared on one of three diets: pure woodlice [composed of two species Oniscus asellus Linnaeus and Armadillidium vulgare (Latreille)]; pure flies ( Drosophila melanogaster Meigen); and a mixed woodlouse-fly diet, is studied. Spiders develop significantly faster on the woodlice-containing diets (i.e. pure woodlice and mixed diet) than on the fly diet. In the second experiment, the prey-choice for two woodlice species ( O.   asellus and A.   vulgare ) and a fly ( D.   melanogaster ) is investigated. Dysdera hungarica spiders capture significantly more often flies than woodlice. These contrasting results reveal the different value of developmental and behavioural experiments. The dietary studies are assumed to provide better evidence of specialization than behavioural experiments, which might be misleading due to unnatural conditions. It is concluded that D.   hungarica is a metabolically adapted woodlice specialist. The present study thus provides the first evidence of nutritional specialization on woodlice.     

Encoding of sound duration by neurons in the auditory cortex of the little brown bat, Myotis lucifugus

Responses of 117 single- or multi-units in the auditory cortex (AC) of bats (Myotis lucifugus) to tone bursts of different stimulus durations (1– 400 ms) were studied over a wide range of stimulus intensities to determine how stimulus duration is represented in the AC. 36% of AC neurons responded more strongly to short stimulus durations showing short-pass duration response functions, 31% responded equally to all pulse durations (i.e., all-pass), 18% responded preferentially to stimuli having longer durations (i.e., long-pass), and 15% responded to a narrow range of stimulus durations (i.e., band-pass). Neurons showing long-pass and short-pass duration response functions were narrowly distributed within two horizontal slabs of the cortex, over the rostrocaudal extent of the AC. The effects of stimulus level on duration selectivity were evaluated for 17 AC neurons. For 65% of these units, an increase in stimulus intensity resulted in a progressive decrease in the best duration. In light of the unusual intensity-dependent duration responses of AC neurons, we hypothesized that the response selectivities of AC neurons is different from that in the brainstem. This hypothesis was validated by results of study of the duration response characteristics of single neurons in the inferior colliculus. Accepted: 8 November 1996     

Suitability of woodlice prey for generalist and specialist spider predators: a comparative study

1. Predators select a prey according to its energetic and nutritional composition. Generalist predators avoid, whereas specialists often specialise on well‐defended prey. The aim of this study was to find the suitability of woodlice prey for generalist and specialist predators by comparing their handling efficiency. 2. Laboratory experiments were performed in which specialist and generalist predators were reared on monotypic diets comprising one or other of two woodlice species that differ in their defensive strategies: rollers (Armadillidium) and clingers (Porcellio). A control group was reared on a mixture of arthropods (excluding woodlice). Three spider predators were used that differ in their adaptations to deal with woodlice prey: a woodlice specialist, Dysdera crocata; an oligophagous generalist, Pholcus phalangioides, that also captures woodlice; and a euryphagous generalist, Tegenaria domestica, that does not feed on woodlice. The frequency of capture was recorded and various fitness parameters were measured, namely survival, growth rate, and ontogenetic development. 3. It was found that the specialist, D. crocata, performed best on the Porcellio diet, and similarly well on Armadillidium and mixed diets. The two generalists, P. phalangioides and T. domestica, had poor performance on both woodlice diets but performed well on the mixed diet. 4. The results show that woodlice are unsuitable prey for both oligophagous and euryphagous generalist predators.     

Sources and timing of calcium intake during reproduction in flycatchers

Calcium availability may limit the reproductive output of birds and snail shells are considered to be the main source of calcium in many passerine species. This study of collared (Ficedula albicollis) and pied (F. hypoleuca) flycatchers evaluates calcium intake of a natural diet in Central Europe, and sex differences in the utilization of experimentally supplemented sources of calcium during the entire breeding period in aviary birds. The study provides the first evidence that successful reproduction of these species depends on the availability of woodlice (Isopoda) and millipedes (Diplopoda). Each of these two components provided about 3 times more calcium than the snail shells contained in a natural nestling diet. The breeding performance of aviary birds was poor when only snail shells and the fragments of eggshells were provided in food, i.e., irregular laying, smaller clutches, eggshell defects (25 of 53 eggs), and eggs dried-up during incubation. In contrast, no defective eggshell or dried-up eggs were found and the overall breeding performance increased 2–3 times when woodlice were added to the food. Females increased their intake of woodlice during both the pre-laying and laying periods, and both sexes did so during the nestling period. Both sexes took more woodlice in the evening than in the morning, independent of the nesting stage. Intake by females was low until 4 days before laying the first egg, then increased to the highest level, dropping immediately after laying the last egg. Intake of woodlice by both sexes increased steadily from hatching until the nestling age of about 10–12 days decreasing thereafter, which corresponds with the period of rapid skeletal growth. In contrast, the intake of mealworms increased until the nestling age of 13–14 days leveling off thereafter which corresponds with the growth curve of nestling body mass.Due to an error in the citation line, this revised PDF (published in December 2003) deviates from the printed version, and is the correct and authoritative version of the paper.     

Facilitation and interference components in the Simon effect.

In four experiments we investigated whether the Simon effect consists of both facilitation and interference. In Experiment 1 subjects had to press one of two lateralized keys in response to one of two stimuli (a rectangle or a square). The stimuli were presented at fixation or to the right or left of it. The stimulus-response mappings could be corresponding (i.e., right key--rectangle or square on the right side), non-corresponding (i.e., right key--rectangle or square on the left side), or neutral (i.e., stimulus in the center). Results showed both facilitation and interference effects, but interference was three times greater than facilitation. To test whether the neutral position was favored by visual acuity, Experiments 2, 3 and 4 used slight different displays where visual acuity was the same at every stimulus position. Results indicated that the Simon effect is comprised of facilitation and interference components of the same magnitude. These findings were discussed with reference to dual-route models of the Simon effect.     

Behavioural Contagion Explains Group Cohesion in a Social Crustacean

In gregarious species, social interactions maintain group cohesion and the associated adaptive values of group living. The understanding of mechanisms leading to group cohesion is essential for understanding the collective dynamics of groups and the spatio-temporal distribution of organisms in environment. In this view, social aggregation in terrestrial isopods represents an interesting model due to its recurrence both in the field and in the laboratory. In this study, and under a perturbation context, we experimentally tested the stability of groups of woodlice according to group size and time spent in group. Our results indicate that the response to the disturbance of groups decreases with increases in these two variables. Models neglecting social effects cannot reproduce experimental data, attesting that cohesion of aggregation in terrestrial isopods is partly governed by a social effect. In particular, models involving calmed and excited individuals and a social transition between these two behavioural states more accurately reproduced our experimental data. Therefore, we concluded that group cohesion (and collective response to stimulus) in terrestrial isopods is governed by a transitory resting state under the influence of density of conspecifics and time spent in group. Lastly, we discuss the nature of direct or indirect interactions possibly implicated.     

Forward conditioning with wheel running causes place aversion in rats

Backward pairings of a distinctive chamber as a conditioned stimulus and wheel running as an unconditioned stimulus (i.e., running-then-chamber) can produce a conditioned place preference in rats. The present study explored whether a forward conditioning procedure with these stimuli (i.e., chamber-then-running) would yield place preference or aversion. Confinement of a rat in one of two distinctive chambers was followed by a 20- or 60-min running opportunity, but confinement in the other was not. After four repetitions of this treatment (i.e., differential conditioning), a choice preference test was given in which the rat had free access to both chambers. This choice test showed that the rats given 60-min running opportunities spent less time in the running-paired chamber than in the unpaired chamber. Namely, a 60-min running opportunity after confinement in a distinctive chamber caused conditioned aversion to that chamber after four paired trials. This result was discussed with regard to the opponent-process theory of motivation.     

In vitro assays for repellents and deterrents for ticks: differing effects of products when tested with attractant or arrestment stimuli

Most in vivo and in vitro tests with repellents or deterrents against ticks have not considered which sensory channel is being targeted. We have recorded the responses of two hard tick species (Acari: Ixodidae) in vitro to determine if such products can disrupt the perception of an attractant in a repellent assay or the perception of an arrestment stimulus in a deterrent assay. Ethyl butylacetylaminopropionate (EBAAP), N,N-diethyl-methyl-benzamide (deet), permethrin and indalone were chosen to test their capacity to inhibit the attraction of Amblyomma variegatum Fabricius to its aggregation-attachment pheromone. Vapours of each test product plus those from a synthetic blend of the pheromone were delivered to the walking tick in an air stream on a locomotion compensator. Neither EBAAP, deet, permethrin nor indalone could inhibit attraction of A. variegatum even when each of the test products was delivered at 106 times the pheromone. Indalone did decrease the attraction of A. variegatum to the pheromone and induced repulsion of A. variegatum when presented on its own in the air stream. The effect of permethrin, a sodium channel blocker, was also tested in a deterrent assay measuring the arrestment of Ixodes ricinus (L.) adults on its own faeces and faecal constituents. Permethrin deterred arrestment at doses of 670 fg/cm2 to 67 ng/cm2, i.e. at levels five times lower than the dose of chemostimuli present in the arrestment stimulus. This sensitivity to permethrin suggests that it acts via the contact chemoreception channel.     

Temporal changes in effectiveness of an inspiratory inhibitory electrical pontine stimulus

We determined the temporal changes in effectiveness of inspiratory-shortening expiratory-prolonging stimulus trains delivered in the region of the nucleus parabrachialis medialis and compared the responses to those observed during trains delivered to the vagus in the same animals (pentobarbital, sodium-anesthetized paralyzed cats). The inspiratory inhibitory effect of the pontine stimulus was assessed from the effect the stimulus has on threshold for terminating inspiration. Stimulus effect increased gradually, reached a peak at 0.2-0.4 s, and declined thereafter. The time of occurrence of peak effect was different from that observed in the course of vagal stimulus trains. With long stimulus trains (19-40 s), the initial effect on inspiratory duration (TI) (i.e., shortening) rapidly subsided and, in six of eight animals, was replaced by TI prolongation. The initial effect on expiratory duration (TE) (i.e., prolongation) also gradually declined with time but TE remained above control throughout. The time constant of adaptation was very similar with vagal and pontine stimulus trains (12.2 and 11.0 s, respectively), but the gain of the adapting response was much more pronounced with pontine stimuli, resulting in a paradoxical effect while stimulation continued. We conclude that the response to pontine stimuli, as with vagal stimuli, displays both integrative and adaptive characteristics. The similarity of the time constants for vagal and pontine adaptation responses suggests that these two inputs share common processing pathways.     

Dynamics of heart rhythm in chimpanzees during the perception of the emotionally colored human voice

The performed study allowed to observe in two chimpanzees both common and individual changes of the heart rate under the action of human voice with emotionally positive or negative intonation, as well as under the action of preferred (sweet) or rejected (quinine solution) natural stimulus. The common property consists in a significant slowing down of the heart rate in response to negative vocal signal and rejected natural stimulus and in only slight change of the heart rate in response to positive signals. Individual properties comprise differences in reactions of the cardiovascular system of both animals to negative action: a greater efficiency of the vocal signal for one chimpanzee and of the direct (natural) stimulus for the second one.     

Electrophysiologic study of the reactions of the brain to regular and skipped acoustic stimuli in man

Human electrical cerebral reactions to acoustic stimuli and to their omission at fixed moments of a restricted rhythmic sequence repeatedly presented in conditions of distraction and attraction of attention to stimuli, were recorded on the vertex. By special methods of bioelectrical signals processing on a computer, the potentials are singled out in response to stimulus omission (SO) with each set ordinal number, which, as well as the potentials, evoked by the actual stimulus (AS), possessed a main negative-positive complex N1P2. In conditions of attention distraction, the negative component of this complex had a latency of 144 ms and the positive one--about 207 ms. In the situation of the attraction of attention to stimuli a decrease of N1 latency up to 125 ms, took place, as well as complication of SO configuration due to the appearance of a later positive oscillation with a latency of about 560 ms, and parametres stabilization of SO as a whole. The analogous dynamics of N1 latency was characteristic of AS in both situations. The mechanisms of SO formation are discussed in connection with trace processes in the central nervous system.     

Scalable rules for coherent group motion in a gregarious vertebrate

Individuals of gregarious species that initiate collective movement require mechanisms of cohesion in order to maintain advantages of group living. One fundamental question in the study of collective movement is what individual rules are employed when making movement decisions. Previous studies have revealed that group movements often depend on social interactions among individual members and specifically that collective decisions to move often follow a quorum-like response. However, these studies either did not quantify the response function at the individual scale (but rather tested hypotheses based on group-level behaviours), or they used a single group size and did not demonstrate which social stimuli influence the individual decision-making process. One challenge in the study of collective movement has been to discriminate between a common response to an external stimulus and the synchronization of behaviours resulting from social interactions. Here we discriminate between these two mechanisms by triggering the departure of one trained Merino sheep (Ovis aries) from groups containing one, three, five and seven naïve individuals. Each individual was thus exposed to various combinations of already-departed and non-departed individuals, depending on its rank of departure. To investigate which individual mechanisms are involved in maintaining group cohesion under conditions of leadership, we quantified the temporal dynamic of response at the individual scale. We found that individuals'' decisions to move do not follow a quorum response but rather follow a rule based on a double mimetic effect: attraction to already-departed individuals and attraction to non-departed individuals. This rule is shown to be in agreement with an adaptive strategy that is inherently scalable as a function of group size.     

Individual preferences and social interactions determine the aggregation of woodlice

Background

The aggregation of woodlice in dark and moist places is considered an adaptation to land life and most studies are focused on its functionality or on the behavioural mechanisms related to the individual''s response to abiotic factors. Until now, no clear experimental demonstration was available about aggregation resulting from inter-attraction between conspecifics.

Methodology/Main Findings

We present the dynamics of aggregation, not previously described in detail in literature, as being independent of the experimental conditions: homogeneous and heterogeneous environments with identical or different shelters. Indeed whatever these conditions, the aggregation is very quick. In less than 10 minutes more than 50% of woodlice were aggregated in several small groups in the homogeneous environment or under shelters in the heterogeneous environment. After this fast aggregation, woodlice progressively moved into a single aggregate or under one shelter.

Conclusions/Significance

Here we show for the first time that aggregation in woodlice implies a strong social component and results from a trade-off between individual preferences and inter-attraction between individuals. Moreover, our results reveal that the response to the heterogeneities affects only the location of the aggregates and not the level of aggregation, and demonstrate the strong inter-attraction between conspecifics which can outweigh individual preferences. This inter-attraction can lead to situations that could seem sub-optimal.     

Quantification of sensory information transmission using timeseries decorrelation techniques

To estimate the information transmitted across a neuronal sensory system one has to deal with serial dependence among consecutive samples of the stimulus and the response signal. Common methods usually require a huge amount of data, or are restricted to Gaussian stimuli. Here, we describe stimulus and response as stochastic processes, i.e. as sequences of random variables, in the same coordinate system. Stimulus-response pairs of these random variables must not be considered independently because otherwise the transinformation is overestimated. To account for the linear fraction of the serial dependence, we present two decorrelation techniques based on coordinate transformation. They provide a representation of the processes with uncorrelated random variables and yield a more precise estimate of the transinformation.     

Renewal and reinstatement of the conditioned but not the unconditioned response following habituation of the unconditioned stimulus

Research on the inhibition of learned fear currently relies almost exclusively on one specific procedure, namely extinction of the conditioned stimulus (CS). Importantly, however, learned fear responses can be reduced by a number of other procedures, including habituation of the unconditioned stimulus (US). We recently demonstrated that reductions in learned fear following US habituation, like CS extinction, were subject to both renewal and reinstatement (Storsve et al., 2010). The present study further investigates the associative and non-associative processes shared between habituation and extinction. Given that habituation is typically context-independent (Mackintosh, 1987), in the present study we directly compared renewal and reinstatement of both a conditioned response (CR; freezing) and an unconditioned response (UR; startle) following habituation. It was found that the reduction in conditioned freezing resulting from habituation was context specific (i.e., a change in context led to a renewal of the conditioned fear response; Experiment 1) and was attenuated when a pre-test shock was given (i.e., reinstatement of conditioned fear was observed; Experiment 2). In contrast, habituation of an unconditioned response elicited by the US (i.e., a startle response) was unaffected by either a change in test context or administration of a pre-test shock. This dissociation in the effects of habituation on learned and unlearned responses is discussed in relation to theories of fear extinction.     

The tradeoff between reproductive and food resources in the crayfish Orconectes virilis

The conflict between mating and foraging is one of the many tradeoffs associated with reproduction. The relative attraction to sex pheromone and food stimuli by male virile crayfish (Orconectes virilis) were tested using two experiments. In the first experiment, the responses by males to pheromone and food stimuli alone were assessed. In the second experiment, males were exposed to both pheromone and food stimuli in a choice maze. To determine the effects of feeding motivation, the males in the choice experiment were fed either one day or seven days prior to observation. Male crayfish exhibited similar responses to the pheromone and food stimuli individually. Males exhibited a preference for the pheromone stimulus over the food stimulus in the choice trials, and that preference that was not influenced by starvation.     

Ultrasound sensitive neurons in the cricket brain

1. The aim of this study was to identify neurons in the brain of the cricket, Teleogryllus oceanicus, that are tuned to high frequencies and to determine if these neurons are involved in the pathway controlling negative phonotaxis. In this paper we describe, both morphologically and physiologically, 20 neurons in the cricket brain which are preferentially tuned to high frequencies. 2. These neurons can be divided into two morphological classes: descending brain interneurons (DBINs) which have a posteriorly projecting axon in the circumesophageal connective and local brain neurons (LBNs) whose processes reside entirely within the brain. All the DBINs and LBNs have processes which project into one common area of the brain, the ventral brain region at the border of the protocerebrum and deutocerebrum. Some of the terminal arborizations of Int-1, an ascending ultrasound sensitive interneuron which initiates negative phonotaxis, also extend into this region. 3. Physiologically, ultrasonic sound pulses produce 3 types of responses in the DBINs and LBNs. (1) Seven DBINs and 6 LBNs are excited by ultrasound. (2) Ongoing activity in one DBIN and 5 LBNs is inhibited by ultrasound, and (3) one cell, (LBN-ei), is either excited or inhibited by ultrasound depending on the direction of the stimulus. 4. Many of the response properties of both the DBINs and LBNs to auditory stimuli are similar to those of Int-1. Specifically, the strength of the response, either excitation or inhibition, to 20 kHz sound pulses increases with increasing stimulus intensity, while the response latency generally decreases. Moreover, the thresholds to high frequencies are much lower than to low frequencies. These observations suggest that the DBINs and LBNs receive a majority of their auditory input from Int-1. However, the response latencies and directional sensitivity of only LBN-ei suggest that it is directly connected to Int-1. 5. The response of only one identified brain neuron, DBIN8, which is inhibited by 20 kHz sound pulses, is facilitated during flight compared to its response at rest. This suggests that suppression of activity in DBIN8 may be associated with ultrasound-induced negative phonotactic steering responses in flying crickets. The other DBINs and LBNs identified in this paper may also play a role in negative phonotaxis, and possibly in other cricket auditory behaviors influenced by ultrasonic frequencies.     

Social spacing in the mormyrid fish Gnathonemus petersii (Pisces): A multisensory approach

The sensory basis of group cohesion in the weak-electric fish Gnathonemus petersii was investigated in a circular tank with groups of four fish each, interacting through a wide-meshed plastic screen with intact or operated conspecifics, or with other stimulus objects. We confined these stimuli to one or two peripheral holding compartments. The response measures were obtained from the free swimming fish and included (1) the time the fish spent together as a group, (2) the time they spent in front of the holding compartments, (3) the circular distribution of the fish's positions, and (4) the mean nearest neighbour distances. Under empty compartment conditions, four different groups were tested, consisting of either (1) intact, electrically active fish, or (2) electrically ‘silent’ fish (with their electric organ surgically rendered inoperative), or (3) blind, or (4) ‘silent’ and blind animals. The loss of either sensory modality, vision or feedback from electric organ discharge, led to changes of comparable size, decreasing the time spent as a group and increasing the mean nearest neighbour distance. In fish lacking both modalities, group cohesion was further impaired. With stimuli present in one or both holding compartments, the strength of social attraction depended on the nature of the stimulus: the more intact stimulus conspecifics were present, the more densely did the fish group in front of the stimulus compartment. ‘Wired-in’ electric organ discharges (simulating waveform and intensity) and electrically ‘silent’ fish were equally attractive, but only half as attractive as intact fish. Blind free swimming fish aggregated with intact and also with ‘silent’ conspecifics. Under dim light conditions, group cohesion was predominantly, though not exclusively, affected by electrosensory feedback from the electric organ discharge and visual input. Mechanical and olfactory cues may also be involved.     

Auditory stimulus intensity gradients and response to methylphenidate in ADD children

Using an auditory stimulus intensity paradigm, we obtained both event related potentials (ERPs) and press and release reaction times (RT) from a large sample of children with Attention Deficit Disorder (ADD). The ERP gradients to three tone intensities were used to classify the children as augmenters (steep gradients), moderates, or reducers (shallow or negative gradients). The RT data were used to classify the children as strong or sensitive, following neo-Pavlovian guidelines. The children were then cross-classified on these two dimensions and compared on cognitive, behavioral, and performance measures. The groups were also compared in response to two dosage levels of methylphenidate. Based on prior studies, we hypothesized that: 1. ERP augmenters would respond as well to the low as high dose but that reducers would respond better to high than low dose; and II. sensitive types (RT measure) would do better on the higher dose and strong types on the low dose. The first hypothesis was confirmed on a performance task but not on behavioral ratings. At the lower dose, augmenters improved most and reducers least on a 10-minute coding task presumed to require sustained attention. There was no support for the second hypothesis either in ratings or performance. The ERP augmentation measure was significantly related to teacher rated attentiveness; i.e., reducers and moderates were rated more adversely. The RT sensitivity measure tended to be related to achievement; i.e., strong types had lower reading and spelling scores. The ERP and RT sensitivity measures were not significantly correlated.     

Behavioral characterization of attractin,a water-borne peptide pheromone in the genus aplysia

Pheromones play a significant role in coordinating reproductive activity in many animals, including opisthobranch molluscs of the genus APLYSIA: Although solitary during most of the year, these simultaneous hermaphrodites gather into breeding aggregations during the reproductive season. The aggregations contain both mating and egg-laying animals and are associated with masses of egg cordons. The egg cordons are a source of pheromones that attract other Aplysia to the area, reduce their latency to mating, and induce egg laying. One of these water-borne egg cordon pheromones ("attractin") has been characterized and shown to be attractive in T-maze assays. Attractin is the first water-borne peptide pheromone characterized in invertebrates. In the current studies, behavioral assays were used to better characterize the attraction, and to examine whether attractin can induce mating. Although the two activities could be related (i.e., attraction occurring because animals were looking for a partner), this was not tested. T-maze assays showed that attractin works as part of a bouquet of odors: the peptide is attractive only when Aplysia brasiliana is part of the stimulus. The animal does not need to be a conspecific, perhaps explaining why multiple species may be associated with one aggregation. Native and recombinant attractin are equally attractive, verifying that N-glycosylation at residue 8 is not required for attraction. Mating studies showed that both native and recombinant attractin reduce the latency to mating. The effects are larger when hermaphroditic mating is considered: in addition to reducing latency, attractin doubles the number of pairs mating as hermaphrodites. The effect may result from attractin stimulating both animals to mate as males and would be consistent with behaviors previously seen in the T-maze. Attractin may thus be contributing to the formation of copulatory chains and rings seen in aggregations in the field. These results may be interpreted in two ways: (1). attractin has multiple activities that contribute to the establishment and maintenance of the aggregation; or (2). the induced desire to mate may make attractin attractive when it is presented in conjunction with an animal. In either case, the results open the door for cellular and molecular studies of mechanism of action.