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Animals deliver and withstand physical impacts in diverse behavioral contexts, from competing rams clashing their antlers together to archerfish impacting prey with jets of water. Though the ability of animals to withstand impact has generally been studied by focusing on morphology, behaviors may also influence impact resistance. Mantis shrimp exchange high-force strikes on each other's coiled, armored telsons (tailplates) during contests over territory. Prior work has shown that telson morphology has high impact resistance. I hypothesized that the behavior of coiling the telson also contributes to impact energy dissipation. By measuring impact dynamics from high-speed videos of strikes exchanged during contests between freely moving animals, I found that approximately 20% more impact energy was dissipated by the telson as compared with findings from a prior study that focused solely on morphology. This increase is likely due to behavior: because the telson is lifted off the substrate, the entire body flexes after contact, dissipating more energy than exoskeletal morphology does on its own. While variation in the degree of telson coil did not affect energy dissipation, proportionally more energy was dissipated from higher velocity strikes and from strikes from more massive appendages. Overall, these findings show that analysis of both behavior and morphology is crucial to understanding impact resistance, and suggest future research on the evolution of structure and function under the selective pressure of biological impacts., Competing Interests: Competing interests The author declares no competing or financial interests., (© 2024. Published by The Company of Biologists Ltd.)

Larval stomatopods have generally been described as having a typical larval crustacean compound eye, which lacks the visual pigment diversity and morphological specializations of the well-studied stomatopod adult eye. However, recent work has suggested that larval stomatopod eyes are more complex than previously described. In this study, we provide physiological and behavioral evidence of at least three distinct photoreceptor classes in three species of larval stomatopods: Gonodactylellus n. sp., Gonodactylaceus falcatus and Pullosquilla n. sp. First, electroretinogram recordings were used to measure the spectral sensitivity of each species. Evidence for at least three spectral classes were identified in each: an ultraviolet, peaking at 340-376 nm; a short-wavelength blue, peaking at 455-464 nm; and a long-wavelength orange, peaking at 576-602 nm. Next, the behavioral response to light was investigated. We found that each species demonstrated positive phototactic responses to monochromatic stimuli across the UV-visible spectrum. In wavelength preference trials, distinct preferences among species were identified when different colored light stimuli were presented simultaneously. All species displayed a strong response to the UV stimulus, as well as responses to blue and orange stimuli, although at different response strengths, but no response to green. The results of this study demonstrate that larval stomatopods not only have multiple physiologically active spectral classes but they also display clear and distinct responses to wavelengths across the spectrum. We propose that the spectral classes demonstrated in each are related to visually guided ecological tasks of the larvae, which may differ between species., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2023. Published by The Company of Biologists Ltd.)

Stomatopod crustaceans, or mantis shrimps, are known for their extensive range of spectral sensitivity but relatively poor spectral discrimination. Instead of the colour-opponent mechanism of other colour vision systems, the 12 narrow-band colour channels they possess may underlie a different method of colour processing. We investigated one hypothesis in which the photoreceptors are proposed to act as individual wave-band detectors, interpreting colour as a parallel pattern of photoreceptor activation, rather than a ratiometric comparison of individual signals. This different form of colour detection has been used to explain previous behavioural tests in which low-saturation blue was not discriminated from grey, potentially because of similar activation patterns. Results here, however, indicate that the stomatopod Haptosquilla trispinosa was able to easily distinguish several colours, including blue of both high and low saturation, from greys. The animals did show a decrease in performance over time in an artificially lit environment, indicating plasticity in colour discrimination ability. This rapid plasticity, most likely the result of a change in opsin (visual pigment) expression, has now been noted in several animal lineages (both invertebrate and vertebrate) and is a factor we suggest needs attention and potential re-examination in any colour-based behavioural tests. As for stomatopods, it remains unclear why they achieve poor colour discrimination using the most comprehensive set of spectral sensitivities in the animal kingdom and also what form of colour processing they may utilise., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2022. Published by The Company of Biologists Ltd.)

Stomatopod crustaceans have among the most complex eyes in the animal kingdom, with up to 12 different color detection channels. The capabilities of these unique eyes include photoreception of ultraviolet (UV) wavelengths (<400 nm). UV vision has been well characterized in adult stomatopods but has not been previously demonstrated in the comparatively simpler larval eye. Larval stomatopod eyes are developmentally distinct from their adult counterpart and have been described as lacking the visual pigment diversity and morphological specializations found in adult eyes. However, recent studies have provided evidence that larval stomatopod eyes are more complex than previously thought and warrant closer investigation. Using electroretinogram recordings in live animals we found physiological evidence of blue- and UV-sensitive photoreceptors in larvae of the Caribbean stomatopod species Neogonodactylus oerstedii. Transcriptomes of individual larvae were used to identify the expression of three distinct UV opsin mRNA transcripts, which may indicate the presence of multiple UV spectral channels. This is the first paper to document UV vision in any larval stomatopod, expanding our understanding of the importance of UV sensitivity in plankton. Larval stomatopod eyes are more complex and more similar to adult eyes than expected, showing previously uncharacterized molecular diversity and physiological functions., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2022. Published by The Company of Biologists Ltd.)

Mantis shrimp strikes are one of the fastest animal movements, despite their occurrence in a water medium with viscous drag. Since the strike is produced by a latch-mediated spring-actuated system and not directly driven by muscle action, we predicted that strikes performed in air would be faster than underwater as a result of reduction in the medium's drag. Using high-speed video analysis of stereotyped strikes elicited from Squilla mantis , we found the exact opposite: strikes are much slower and less powerful in air than in water. S. mantis strikes in air have a similar mass and performance to latch-mediated spring-actuated jumps in locusts, suggesting a potential threshold for the energetics of a 1-2 g limb rotating in air. Drag forces induced by the media may be a key feature in the evolution of mantis shrimp strikes and provide a potential target for probing the braking system of these extremely fast movements., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2020. Published by The Company of Biologists Ltd.)

Measurements of energy use, and its scaling with size, are critical to understanding how organisms accomplish myriad tasks. For example, energy budgets are central to game theory models of assessment during contests and underlie patterns of feeding behavior. Clear tests connecting energy to behavioral theory require measurements of the energy use of single individuals for particular behaviors. Many species of mantis shrimp (Stomatopoda: Crustacea) use elastic energy storage to power high-speed strikes that they deliver to opponents during territorial contests and to hard-shelled prey while feeding. We compared the scaling of strike kinematics and energetics between feeding and contests in the mantis shrimp Neogonodactylus bredini We filmed strikes with high-speed video, measured strike velocity and used a mathematical model to calculate strike energy. During contests, strike velocity did not scale with body size but strike energy scaled positively with size. Conversely, while feeding, strike velocity decreased with increasing size and strike energy did not vary according to body size. Individuals most likely achieved this strike variation through differential compression of their exoskeletal spring prior to the strike. Post hoc analyses found that N. bredini used greater velocity and energy when striking larger opponents, yet variation in prey size was not accompanied by varying strike velocity or energetics. Our estimates of energetics inform prior tests of contest and feeding behavior in this species. More broadly, our findings elucidate the role behavioral context plays in measurements of animal performance., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2019. Published by The Company of Biologists Ltd.)

Many predators fracture strong mollusk shells, requiring specialized weaponry and behaviors. The current shell fracture paradigm is based on jaw- and claw-based predators that slowly apply forces (high impulse, low peak force). However, predators also strike shells with transient intense impacts (low impulse, high peak force). Toward the goal of incorporating impact fracture strategies into the prevailing paradigm, we measured how mantis shrimp ( Neogonodactylus bredini ) impact snail shells, tested whether they strike shells in different locations depending on prey shape ( Nerita spp., Cenchritis muricatus , Cerithium spp.) and deployed a physical model (Ninjabot) to test the effectiveness of strike locations. We found that, contrary to their formidable reputation, mantis shrimp struck shells tens to hundreds of times while targeting distinct shell locations. They consistently struck the aperture of globular shells and changed from the aperture to the apex of high-spired shells. Ninjabot tests revealed that mantis shrimp avoid strike locations that cause little damage and that reaching the threshold for eating soft tissue is increasingly difficult as fracture progresses. Their ballistic strategy requires feed-forward control, relying on extensive pre-strike set-up, unlike jaw- and claw-based strategies that can use real-time neural feedback when crushing. However, alongside this pre-processing cost to impact fracture comes the ability to circumvent gape limits and thus process larger prey. In sum, mantis shrimp target specific shell regions, alter their strategy depending on shell shape, and present a model system for studying the physics and materials of impact fracture in the context of the rich evolutionary history of predator-prey interactions., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2018. Published by The Company of Biologists Ltd.)

Light is a primary environmental factor used by aquatic invertebrates for depth selection behavior. Many branchiopod crustaceans live in ephemeral aquatic habitats. All branchiopod crustaceans studied to date express four or more visual opsins in their compound eyes. We asked whether two branchiopods, Triops longicaudatus and Streptocephalus mackini , use multiple spectral channels to regulate their position in the water column. At the lowest intensities that elicited photonegative behavior, both species had broad spectral bandwidths, suggesting they use multiple spectral photoreceptor classes. Male S. mackini were more likely to maintain a vertical position 8.0-12.0 cm below the surface than females, independently of whether females were present. Male photopositive behavior at low intensity was restricted to a narrow bandwidth centered at 532 nm, suggesting a single photoreceptor class is used to maintain position above females. We compared ephemeral pools from two regions in Arizona and found that diffuse light attenuation coefficients were two orders of magnitude greater than the most heavily attenuating coastal waters. At less than 1 m of depth, pools were often dimmer than terrestrial habitats under starlight. Soil particle size distribution in each region affected spectral light environments, and behavioral responses of field-caught shrimp were adapted to the spectral properties of their region. The results suggest that branchiopods predominantly use luminance vision summed from multiple spectral photoreceptor classes for depth selection in dim, spectrally variable environments. The neuroanatomical basis for summation is described in a companion paper., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2018. Published by The Company of Biologists Ltd.)

Animals that have true color vision possess several spectral classes of photoreceptors. Pancrustaceans (Hexapoda+Crustacea) that integrate spectral information about their reconstructed visual world do so from photoreceptor terminals supplying their second optic neuropils, with subsequent participation of the third (lobula) and deeper centers (optic foci). Here, we describe experiments and correlative neural arrangements underlying convergent visual pathways in two species of branchiopod crustaceans that have to cope with a broad range of spectral ambience and illuminance in ephemeral pools, yet possess just two optic neuropils, the lamina and the optic tectum. Electroretinographic recordings and multimodel inference based on modeled spectral absorptance were used to identify the most likely number of spectral photoreceptor classes in their compound eyes. Recordings from the retina provide support for four color channels. Neuroanatomical observations resolve arrangements in their laminas that suggest signal summation at low light intensities, incorporating chromatic channels. Neuroanatomical observations demonstrate that spatial summation in the lamina of the two species are mediated by quite different mechanisms, both of which allow signals from several ommatidia to be pooled at single lamina monopolar cells. We propose that such summation provides sufficient signal for vision at intensities equivalent to those experienced by insects in terrestrial habitats under dim starlight. Our findings suggest that despite the absence of optic lobe neuropils necessary for spectral discrimination utilized by true color vision, four spectral photoreceptor classes have been maintained in Branchiopoda for vision at very low light intensities at variable ambient wavelengths that typify conditions in ephemeral freshwater habitats., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2018. Published by The Company of Biologists Ltd.)

A combination of behavioural and electrophysiological experiments have previously shown that two species of stomatopod, Odontodactylus scyllarus and Gonodactylaceus falcatus , can differentiate between left- and right-handed circularly polarized light (CPL), and between CPL and linearly polarized light (LPL). It remains unknown if these visual abilities are common across all stomatopod species, and if so, how circular polarization sensitivity may vary between and within species. A subsection of the midband, a specialized region of stomatopod eyes, contains distally placed photoreceptor cells, termed R8 (retinular cell number 8). These cells are specifically built with unidirectional microvilli and appear to be angled precisely to convert CPL into LPL. They are mostly quarter-wave retarders for human visible light (400-700 nm), as well as being ultraviolet-sensitive linear polarization detectors. The effectiveness of the R8 cells in this role is determined by their geometric and optical properties. In particular, the length and birefringence of the R8 cells are crucial for retardation efficiency. Here, our comparative studies show that most species investigated have the theoretical ability to convert CPL into LPL, such that the handedness of an incoming circular reflection or signal could be discriminated. One species, Haptosquilla trispinosa , shows less than quarter-wave retardance. Whilst some species are known to produce circularly polarized reflections (some Odontodactylus species and G. falcatus , for example), others do not, so a variety of functions for this ability are worth considering., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2017. Published by The Company of Biologists Ltd.)

The eyes of two glacial-relict populations of opossum shrimp Mysis relicta inhabiting the different photic environments of a deep, dark-brown freshwater lake and a variably lit bay of the Baltic Sea differ in their susceptibility to functional depression from strong light exposures. The lake population is much more vulnerable than the sea population. We hypothesized that the difference reflects physiological adaptation mechanisms operating on long time scales rather than genetically fixed differences between the populations. To test this, we studied how acclimation to ultra-slowly increased illumination (on time scales of several weeks to months) affected the resilience of the eyes to bright-light exposures. Light responses of whole eyes were measured by electroretinography, the visual-pigment content of single rhabdoms by microspectrophotometry and the structural integrity of photoreceptor cells by electron microscopy (EM). Slow acclimation mitigated and even abolished the depression of photoresponsiveness caused by strong light exposures, making a dramatic difference especially in the lake animals. Still, acclimation in the sea animals was faster and the EM studies suggested intrinsic differences in the dynamics of microvillar membrane cycling. In conclusion, we report a novel form of physiological adaptation to general light levels, effective on the time scale of seasonal changes. It explains part but not all of the differences in light tolerance between the lake and sea populations., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2017. Published by The Company of Biologists Ltd.)

Stomatopods have an extraordinary visual system, incorporating independent movement of their eyes in all three degrees of rotational freedom. In this work, we demonstrate that in the peacock mantis shrimp, Odontodactylus scyllarus , the level of ocular independence is task dependent. During gaze stabilization in the context of optokinesis, there is weak but significant correlation between the left and right eyes in the yaw degree of rotational freedom, but not in pitch and torsion. When one eye is completely occluded, the uncovered eye does not drive the covered eye during gaze stabilization. However, occluding one eye does significantly affect the uncovered eye, lowering its gaze stabilization performance. There is a lateral asymmetry, with the magnitude of the effect depending on the eye (left or right) combined with the direction of motion of the visual field. In contrast, during a startle saccade, the uncovered eye does drive a covered eye. Such disparate levels of independence between the two eyes suggest that responses to individual visual tasks are likely to follow different neural pathways., (© 2017. Published by The Company of Biologists Ltd.)

To circumvent the limits of muscle, ultrafast movements achieve high power through the use of springs and latches. The time scale of these movements is too short for control through typical neuromuscular mechanisms, thus ultrafast movements are either invariant or controlled prior to movement. We tested whether mantis shrimp (Stomatopoda: Neogonodactylus bredini) vary their ultrafast smashing strikes and, if so, how this control is achieved prior to movement. We collected high-speed images of strike mechanics and electromyograms of the extensor and flexor muscles that control spring compression and latch release. During spring compression, lateral extensor and flexor units were co-activated. The strike initiated several milliseconds after the flexor units ceased, suggesting that flexor activity prevents spring release and determines the timing of strike initiation. We used linear mixed models and Akaike's information criterion to serially evaluate multiple hypotheses for control mechanisms. We found that variation in spring compression and strike angular velocity were statistically explained by spike activity of the extensor muscle. The results show that mantis shrimp can generate kinematically variable strikes and that their kinematics can be changed through adjustments to motor activity prior to the movement, thus supporting an upstream, central-nervous-system-based control of ultrafast movement. Based on these and other findings, we present a shishiodoshi model that illustrates alternative models of control in biological ballistic systems. The discovery of feed-forward control in mantis shrimp sets the stage for the assessment of targets, strategic variation in kinematics and the role of learning in ultrafast animals., (© 2016. Published by The Company of Biologists Ltd.)

Recent developments in the study of pain in animals have demonstrated the potential for pain perception in a variety of wholly aquatic species such as molluscs, crustaceans and fish. This allows us to gain insight into how the ecological pressures and differential life history of living in a watery medium can yield novel data that inform the comparative physiology and evolution of pain. Nociception is the simple detection of potentially painful stimuli usually accompanied by a reflex withdrawal response, and nociceptors have been found in aquatic invertebrates such as the sea slug Aplysia. It would seem adaptive to have a warning system that allows animals to avoid life-threatening injury, yet debate does still continue over the capacity for non-mammalian species to experience the discomfort or suffering that is a key component of pain rather than a nociceptive reflex. Contemporary studies over the last 10 years have demonstrated that bony fish possess nociceptors that are similar to those in mammals; that they demonstrate pain-related changes in physiology and behaviour that are reduced by painkillers; that they exhibit higher brain activity when painfully stimulated; and that pain is more important than showing fear or anti-predator behaviour in bony fish. The neurophysiological basis of nociception or pain in fish is demonstrably similar to that in mammals. Pain perception in invertebrates is more controversial as they lack the vertebrate brain, yet recent research evidence confirms that there are behavioural changes in response to potentially painful events. This review will assess the field of pain perception in aquatic species, focusing on fish and selected invertebrate groups to interpret how research findings can inform our understanding of the physiology and evolution of pain. Further, if we accept these animals may be capable of experiencing the negative experience of pain, then the wider implications of human use of these animals should be considered., (© 2015. Published by The Company of Biologists Ltd.)

The polarization of light provides information that is used by many animals for a number of different visually guided behaviours. Several marine species, such as stomatopod crustaceans and cephalopod molluscs, communicate using visual signals that contain polarized information, content that is often part of a more complex multi-dimensional visual signal. In this work, we investigate the evolution of polarized signals in species of Haptosquilla, a widespread genus of stomatopod, as well as related protosquillids. We present evidence for a pre-existing bias towards horizontally polarized signal content and demonstrate that the properties of the polarization vision system in these animals increase the signal-to-noise ratio of the signal. Combining these results with the increase in efficacy that polarization provides over intensity and hue in a shallow marine environment, we propose a joint framework for the evolution of the polarized form of these complex signals based on both efficacy-driven (proximate) and content-driven (ultimate) selection pressures., (© 2014. Published by The Company of Biologists Ltd.)

The polarisation of light is used by many species of cephalopods and crustaceans to discriminate objects or to communicate. Most visual systems with this ability, such as that of the fiddler crab, include receptors with photopigments that are oriented horizontally and vertically relative to the outside world. Photoreceptors in such an orthogonal array are maximally sensitive to polarised light with the same fixed e-vector orientation. Using opponent neural connections, this two-channel system may produce a single value of polarisation contrast and, consequently, it may suffer from null points of discrimination. Stomatopod crustaceans use a different system for polarisation vision, comprising at least four types of polarisation-sensitive photoreceptor arranged at 0, 45, 90 and 135 deg relative to each other, in conjunction with extensive rotational eye movements. This anatomical arrangement should not suffer from equivalent null points of discrimination. To test whether these two systems were vulnerable to null points, we presented the fiddler crab Uca heteropleura and the stomatopod Haptosquilla trispinosa with polarised looming stimuli on a modified LCD monitor. The fiddler crab was less sensitive to differences in the degree of polarised light when the e-vector was at -45 deg than when the e-vector was horizontal. In comparison, stomatopods showed no difference in sensitivity between the two stimulus types. The results suggest that fiddler crabs suffer from a null point of sensitivity, while stomatopods do not., (© 2014. Published by The Company of Biologists Ltd.)

Traditionally, physiologists have estimated the ability of organisms to withstand lower partial pressures of oxygen by estimating the partial pressure at which oxygen consumption begins to decrease (known as the critical PO2 or Pc). For almost 30 years, the principal way in which Pc has been estimated has been via piecewise 'broken stick' regression (BSR). BSR was a useful approach when more sophisticated analyses were less available, but BSR makes a number of unsupported assumptions about the underlying form of the relationship between the rate of oxygen consumption and oxygen availability. The BSR approach also distils a range of values into a single point with no estimate of error. In accordance with more general calls to fit functions to continuous data, we propose the use of nonlinear regression (NLR) to fit various curvilinear functions to oxygen consumption data in order to estimate Pc. Importantly, our approach is back-compatible so that estimates using traditional methods in earlier studies can be compared with data estimates from our technique. When we compared the performance of our approach relative to the traditional BSR approach for real world and simulated data, we found that under realistic circumstances, NLR was more accurate and provided more powerful hypothesis tests. We recommend that future studies make use of NLR to estimate Pc, and also suggest that this approach might be more appropriate for a range of physiological studies that use BSR currently.

The climate variability hypothesis assumes that the thermal tolerance breadth of a species is primarily determined by temperature variations experienced in its environment. If so, aquatic invertebrates living in thermally buffered environments would be expected to exhibit narrow thermal tolerance breadths (stenothermy). We tested this prediction by studying the thermal physiology of three isopods (Asellidae, Proasellus) colonizing groundwater habitats characterized by an annual temperature amplitude of less than 1°C. The species responses to temperature variation were assessed in the laboratory using five physiological variables: survival, locomotor activity, aerobic respiration, immune defense and concentrations of total free amino acids and sugars. The three species exhibited contrasted thermal physiologies, although all variables were not equally informative. In accordance with the climate variability hypothesis, two species were extremely sensitive even to moderate changes in temperature (2°C) below and above their habitat temperature. In contrast, the third species exhibited a surprisingly high thermal tolerance breadth (11°C). Differences in response to temperature variation among Proasellus species indicated that their thermal physiology was not solely shaped by the current temperature seasonality in their natural habitats. More particularly, recent gene flow among populations living in thermally constant yet contrasted habitats might explain the occurrence of eurytherm species in thermally buffered environments.

Intertidal zone organisms naturally experience daily fluctuations in pH, presently reaching values beyond what is predicted for open ocean surface waters from ocean acidification (OA) by the year 2100, and thus present an opportunity to study the pH sensitivity of organisms that are presumably adapted to an acidified environment. The intertidal zone porcelain crab, Petrolisthes cinctipes, was used to study physiological responses to low pH in embryonic, larval and newly recruited juvenile life-history stages. In these crabs, embryonic development occurs in the pH-variable intertidal zone (pH 6.9-9.5), larvae mature in the more stable pelagic environment (pH 7.9-8.2), and juvenile crabs settle back into the pH-variable intertidal zone. We examined survival, cardiac performance, energetics and morphology in embryonic, larval and juvenile crabs exposed to two pH conditions (pH 7.9 and 7.6). Embryos and larvae were split by brood between the pH treatments for 9 days to examine brood-specific responses to low pH. Hatching success did not differ between pH conditions, but ranged from 30% to 95% among broods. Larval survival was not affected by acidification, but juvenile survival was reduced by ~30% after longer (40 days) exposure to low pH. Embryonic and larval heart rates were 37% and 20% lower at low pH, and there was a brood-specific response in embryos. Embryos did not increase in volume under acidified conditions, compared with a 15% increase in ambient conditions. We conclude that sustained exposure to low pH could be detrimental to P. cinctipes embryos and larvae despite the fact that embryos are regularly exposed to naturally fluctuating hypercapnic water in the intertidal zone. Importantly, our results indicate that early life-history stage responses to OA may be brood specific through as yet undetermined mechanisms.

Using new collecting techniques with the Johnson-Sea-Link submersible, eight species of deep-sea benthic crustaceans were collected with intact visual systems. Their spectral sensitivities and temporal resolutions were determined shipboard using electroretinography. Useable spectral sensitivity data were obtained from seven species, and in the dark-adapted eyes, the spectral sensitivity peaks were in the blue region of the visible spectrum, ranging from 470 to 497 nm. Under blue chromatic adaptation, a secondary sensitivity peak in the UV portion of the spectrum appeared for two species of anomuran crabs: Eumunida picta (λ(max)363 nm) and Gastroptychus spinifer (λ(max)383 nm). Wavelength-specific differences in response waveforms under blue chromatic adaptation in these two species suggest that two populations of photoreceptor cells are present. Temporal resolution was determined in all eight species using the maximum critical flicker frequency (CFF(max)). The CFF(max) for the isopod Booralana tricarinata of 4 Hz proved to be the lowest ever measured using this technique, and suggests that this species is not able to track even slow-moving prey. Both the putative dual visual pigment system in the crabs and the extremely slow eye of the isopod may be adaptations for seeing bioluminescence in the benthic environment.

Luminescent signals can be used by animals for a number of purposes, including courtship and defense, sometimes by the same individual. However, the relative costs of producing these different behaviors are largely unknown. In the marine ostracod Photeros annecohenae, males utilize extracellular luminescence for complex courtship displays, and both males and females luminesce as a predation defense. We compared the relative luminescent output of courtship with that of defensive displays and also with respect to their total luminescent stores. Courtship displays are relatively inexpensive compared with defensive displays, with an average defensive display releasing 50 times more luminescence than the average courtship display. Furthermore, in order to completely exhaust its stores, a male would have to produce 450 typical courtship displays or approximately 10 average defensive displays. Both courtship pulses and defensive displays show first-order decay kinetics, yet courtship pulses decay three times faster than defensive displays, suggesting that there is differential release of the luciferin, luciferase and mucus in order to control the reaction kinetics.

Stomatopod crustaceans have complex and diverse visual systems. Among their many unique features are a specialized ommatidial region (the midband) that enables the eye to have multiple overlapping visual fields, as well as sets of spectral filters that are intercalated at two levels between tiers of photoreceptors involved in polychromatic color vision. Although the physiology and visual function of stomatopod eyes have been studied for many years, how these unique visual features originated and diversified is still an open question. In order to investigate how stomatopods have attained the current complexity in visual function, we have combined physiological and morphological information (e.g. number of midband rows, number of filters in the retina, and the spectral properties of filters) with new phylogenetic analyses of relationships among species based on nucleotide sequence data from two nuclear (18S and 28S rDNA) and two mitochondrial [16S and cytochrome oxidase I (COI)] genes. Based on our recovered phylogenetic relationships among species, we propose two new superfamilies within the Stomatopoda: Hemisquilloidea and Pseudosquillodea. Maximum likelihood ancestral state reconstructions indicate that ancestral stomatopod eyes contained six midband rows and four intrarhabdomal filters, illustrating that the visual physiological complexity originated early in stomatopod evolutionary history. While the two distal filters contain conservative sets of filter pigments, the proximal filters show more spectral diversity in filter types, particularly in midband row 2, and are involved in tuning the color vision system to the photic environment. In particular, a set of related gonodactyloid families (Gonodactylidae, Protosquillidae, Takuidae) inhabiting shallow, brightly lit coral reef waters contain the largest diversity of filter pigments, which are spectrally placed relative to the underlying photoreceptors to take advantage of the broad spectrum of light available in the environment.

A possible role of the insect stomatogastric nervous system (STNS) in ecdysis was first implied in early studies reporting on internal air pressure build-up in the digestive tract and air swallowing during ecdysis. The frontal ganglion, a major component of the insect STNS, was suggested to play an important part in this behaviour. Recent neurophysiological studies have confirmed the critical role of the STNS in the successful completion of both larval and adult moults in insects. In aquatic arthropods, though much less studied, the STNS plays an equally important and probably very similar role in water swallowing. Water uptake is instrumental in splitting the crustacean cuticle and allowing successful ecdysis. Current data are presented in a comparative view that contributes to our understanding of the role of the STNS in arthropod behaviour. It also sheds light on the question of homology of the STNS among the different arthropod groups. New insights into the neurohormonal control of ecdysis, related to the STNS in both insects and crustaceans, are also presented and comparatively discussed.

In the western Caribbean Sea, about an hour after the sun sets, a complex and ritualized light show of precise, vertically placed luminescent pulses erupts over shallow grassbeds. These are among the most complex displays known in marine systems. Displays consist of repeated trains of secreted bioluminescent pulses in a specific pattern ejected into the water column as courtship signals by male Vargula annecohenae, which are small (<2 mm) myodocopid ostracod crustaceans. Although these animals display in near darkness, we have used image intensification and infrared videography and three-dimensional analysis in the lab to demonstrate that each luminescent display train, which can be up to 60 cm long, consists of two distinct luminescent and swimming phases. The first, or 'stationary,' phase consists of three (usually) bright, longer pulses placed close together, with the male swimming in a looping pattern. We hypothesize that this pattern acts as an attention-grabbing signal for receptive females. The stationary phase is followed by the 'helical phase,' which consists of about a dozen evenly placed dimmer, shorter pulses secreted by an individual male rapidly spiraling upward in a helical pattern. We hypothesize that this phase, which has very uniform interpulse intervals and distances, helps an approaching female target and intercept the rapidly moving male. Here we provide details of these two phases, and produce a three-dimensional model of a multiply-displaying male.

Gonodactyloid stomatopod crustaceans possess polarization vision, which enables them to discriminate light of different e-vector angle. Their unusual apposition compound eyes are divided by an equatorial band of six rows of enlarged, structurally modified ommatidia, the mid-band (MB). The rhabdoms of the two most ventral MB rows 5 and 6 are structurally designed for polarization vision. Here we show, with electrophysiological recordings, that the photoreceptors R1-R7 within these two MB rows in Gonodactylus chiragra are highly sensitive to linear polarized light of two orthogonal directions (PS=6.1). They possess a narrow spectral sensitivity peaking at 565 nm. Unexpectedly, photoreceptors within the distal rhabdomal tier of MB row 2 also possess highly sensitive linear polarization receptors, which are in their spectral and polarization characteristics similar to the receptors of MB rows 5 and 6. Photoreceptors R1-R7 within the remainder of the MB exhibit low polarization sensitivity (PS=2.3). Outside the MB, in the two hemispheres, R1-R7 possess medium linear polarization sensitivity (PS=3.8) and a broad spectral sensitivity peaking at around 500 nm, typical for most crustaceans. Throughout the retina the most distally situated UV-sensitive R8 cells are not sensitive to linear polarized light.

Crystallization temperature (T(c)), resistance to inoculative freezing (IF), ice contents, bound water, protein and glycogen body contents were measured in the aquatic subterranean crustacean Niphargus rhenorhodanensis and in the morphologically close surface-dwelling aquatic crustacean Gammarus fossarum, both acclimated at 12 degrees C, 3 degrees C and -2 degrees C. Cold acclimation induced an increase in the T(c) values in both species but no survival was observed after thawing. However, after inoculation at high sub-zero temperatures, cold-acclimated N. rhenorhodanensis survived whereas all others, including the 3 degrees C and -2 degrees C acclimated G. fossarum died. In its aquatic environment, N. rhenorhodanensis is likely to encounter inoculative freezing before reaching the T(c) and IF tolerance appears as a highly adaptive trait in this species. Bound water and glycogen were found to increase in the 3 degrees C and -2 degrees C acclimated N. rhenorhodanensis, whereas no variation was observed in G. fossarum. Considering the hydrophilic properties of glycogen, such a rise may be correlated with the increased bound water measured in cold-acclimated N. rhenorhodanensis, and may be linked to the survival of this species when it was inoculated. The ecological significance of the survival of the aquatic subterranean crustacean to inoculative freezing is paradoxical, as temperature is currently highly buffered in its habitat. However, we assume that past geographical distribution and resulting life history traits of N. rhenorhodanensis are key parameters in the current cold-hardiness of the species.

To determine whether neuronal function in Antarctic crustaceans is adapted to the low and narrow range of temperatures at which these animals live, we have compared conduction velocities in the peripheral nervous systems of two temperate species, the decapod Carcinus maenas and the isopod Ligia oceanica, and two Antarctic species, the isopod Glyptonotus antarcticus and the amphipod Paraceradocus gibber. Neuronal conduction velocity differs among the species in the order C. maenas > G. antarcticus > P. gibber > L. oceanica. When measured at the normal environmental temperatures characteristic of each species, conduction velocity of the Antarctic peracarid P. gibber is greater than that of its similar sized temperate relative L. oceanica, demonstrating complete thermal compensation. The temperate decapod C. maenas has a higher thermal dependence of neuronal conduction velocity than either of the Antarctic species, G. antarcticus and P. gibber, but the temperate L. oceanica does not. These data, when collated with published values, indicate that peracarid crustaceans (L. oceanica, G. antarcticus and P. gibber) have lower neuronal conduction velocities and a lower thermal dependence of neuronal conduction velocity than do other arthropods, irrespective of habitat. There is a linear dependence of conduction velocity on temperature down to -1.8 degrees C in all three species. Our data extend by more than 10 degrees the lower range of temperatures at which conduction velocities have been tested systematically in previous studies. The upper thermal block of neuronal conduction is similar in C. maenas, G. antarcticus, P. gibber and L. oceanica at 24.5, 19.5, 21.5 and 19.5 degrees C, respectively. This suggests that failure to conduct action potentials is not what determines the mortality of Antarctic invertebrates at approximately 10 degrees C. The excitability of axons in the leg nerve of G. antarcticus is not affected by temperatures ranging from -1.8 to +18 degrees C. The responses of sensory neurones activated by movements of spines on the leg, however, are strongly modulated by temperature, with maximal responses at 5-10 degrees C; well above the normal environmental temperature range for the species. The responses fail at 20-22 degrees C. The number of large diameter axons (which produce the fast action potentials recorded in this study) is the same in L. oceanica and G. antarcticus, but the median axon diameter is greater in L. oceanica than G. antarcticus. In G. antarcticus, however, there are glial wrappings around some large (>5 microm diameter) axons that may increase their conduction velocity. Such wrappings are not found in L. oceanica.

Hemoglobin (Hb) is a highly conserved protein that provides a vital link between environmental oxygen and its use and/or storage within an organism. While ubiquitous among vertebrates, Hb occurs frequently in invertebrate phyla as well. Many arthropod species use the copper-binding pigment hemocyanin, but unique in this phylum are the branchiopod crustaceans, which express Hb. Branchiopod Hb concentration and structure are exquisitely sensitive to environmental oxygen availability. Hemoglobin concentration and oxygen-binding affinity increase with decreasing oxygen tension in Daphnia, Artemia and Triops. The change in binding affinity is attributed to differential Hb subunit expression in Daphnia and Artemia but remains unclear for Triops. This is the first study to demonstrate developmental plasticity of Hb subunit expression in a notostracan, Triops longicaudatus, reared under conditions of varying oxygen availability. In response to variable oxygen environments, T. longicaudatus differentially express four primary Hb subunits ranging between 30 and 34 kDa, with normoxic-reared animals expressing primarily the heavier subunits, and hypoxic-reared animals expressing increased proportions of the lower molecular mass subunits. Moreover, differential Hb subunit expression is induced upon transfer of normoxic-reared adults to a hypoxic environment, such that the distribution of Hb subunits in the transferred adults becomes similar to that of hypoxic-reared animals. Two-dimensional gel electrophoresis and follow-up analyses revealed several isoforms of Hb subunits that may represent differential gene expression and/or post-translational modification. Unlike Daphnia and Artemia, the Hb hypoxic response in Triops is not reversible in that there was no significant decrease in Hb concentration or change in Hb subunit expression pattern when hypoxic-reared adults were transferred to a normoxic environment.

We studied basic principles of respiratory physiology in Cylindroleberididae, Cylindroleberis mariae Baird 1850, which are millimetre-sized crustaceans (myodocop ostracod) having a fossil record dating back to about 425 millions years ago. Facing experimental changes of O2 partial pressures in the range 2-40 kPa (normoxia is 21 kPa), C. mariae lack any regulatory mechanism to adapt their ventilatory and circulatory activity. Thus, the oxygenation status of their internal milieu must follow, as a dependent variable, the ambient oxygenation. Freely behaving C. mariae exhibit a marked diurnal activity rhythm. They are actively swimming in the water column during night, where they inspire in normoxic-normocapnic water. They are resting in self-made nests during daytime, where they are rebreathing in a confined and hypoxic environment. By analogy to extensive previous literature data, we suggest that these changes of respiratory gas content, and the associated tissue gas status, participate to the shaping of their metabolic activity and behaviour. To conclude, as Cylindroleberididae are early crustaceans exhibiting a remarkable stasis since the Palaeozoic, present data illustrates how principles of tissue oxygenation strategy can cover an impressive time scale.

An adaptation strategy whereby O(2) partial pressure, P(O(2)), in the tissues is maintained within a low, narrow range of 1-3 kPa, largely independent of the inspired P(O(2)), has been reported in water- and air-breathing poikilotherms and in homeotherms. Based on the postulate that this basic cellular mechanism has been established since the early stages of evolution, it has been hypothesized that it could be the consequence of an early adaptation strategy to maintain cellular oxygenation within the same low and primitive range. To test this hypothesis we studied the basic mechanisms of oxygen regulation in podocopid ostracods, minute crustaceans that have existed on earth for at least 500 million years. Podocopids lack any regulatory mechanism for adapting their ventilation to cope with changes in water oxygenation, and instead adjust their tissue oxygenation status by migrating through the O(2) gradient to sediment layers where the P(O(2)) of the water is 3-5 kPa. Experimental manipulation of the O(2) profile induced their vertical migration to follow this precise water P(O(2)) and demonstrates the existence of a regulation strategy. This strategy must be associated with the lower P(O(2)) values within the animal's carapace valves, showing that podocopids can actively regulate their tissue P(O(2)) at constant but even lower values than the water. In conclusion, the low tissue P(O(2)) strategy could have existed in early crustaceans and, by extension, in early animals.

The emphasis in this review will be on Na(+) absorption across the skin and gills of vertebrates and the gills of crustaceans. However, some recent studies of Cl(-) uptake, especially in crustaceans, will also be described.

The heart of Bathynomus doederleini gives rise to an anterior median artery (AMA), one pair of anterior lateral arteries (ALAs) and five pairs of lateral arteries (LAs). Cardioarterial valves are located at the junctions between the heart and arteries, each composed of a pair of muscular flaps. All valves of the AMA and the ALAs receive valve excitatory (constrictor) nerves (VEs). The valves of the ALAs receive dual innervation from both constrictor and inhibitor (dilator) nerves, while the valves of the AMA receive innervation from a constrictor nerve alone. The effects of candidate neurohormones on cardioarterial valves were examined by measuring the pressure in each artery at which haemolymph flows out of the heart through the valve. Serotonin, octopamine, norepinephrine, glutamate (Glu) and proctolin constricted the cardioarterial valves and thus decreased the arterial pressure in all the arteries. Dopamine also decreased the arterial pressure of arteries except for the ALAs, in which pressure was increased. Among the neurohormones exerting excitatory effects on the valves, only Glu depolarized the membrane potential of valve muscle cells. The glutamatergic agonists kainate and quisqualate also depolarized the valve muscle cells of the AMA. Excitatory junctional potentials produced in the valves of the AMA in response to the stimulation of a VE were blocked by the glutamatergic antagonists Joro spider toxin and MK-801. Glu is the likeliest candidate for a neurotransmitter for the VEs.

Many marine animals track odor plumes to their source. Although studies of plume-tracking behavior have been performed in unidirectional flow, benthic animals such as crustaceans live in coastal habitats characterized by waves. We compared signal encounters by odor-plume-tracking stomatopods (mantis shrimp) in wave-affected and unidirectional flow in a flume. Stomatopods are small enough that we can study their natural behavior in a flume. They sample odors by flicking their antennules. A thin sheet of laser light illuminating an odor plume labeled with dye [planar laser induced fluorescence (PLIF) technique] permitted us to measure the instantaneous odor concentration encountered by the animal's chemosensory organs (antennules) while it tracked the plume. We simultaneously measured behavior and the high-resolution odor signal at the spatial and temporal scale of the animal. We found that the navigating animal encountered odor filaments more often in wave-affected flow than in unidirectional flow. Odor filaments along the animals' antennules were significantly wider and of higher concentration in waves than in unidirectional flow.

Some stomatopod crustacean species that inhabit a range of habitat depths have color vision systems that adapt to changes in ambient light conditions. To date, this change in retinal function has been demonstrated in species within the superfamily Gonodactyloidea in response to varying the spectral range of light. Intrarhabdomal filters in certain ommatidia within the specialized midband of the eye change spectrally, modifying the sensitivity of underlying photoreceptors to match the spectrum of available light. In the present study, we utilized Pullosquilla litoralis, a member of the superfamily Lysiosquilloidea that also has a wide depth range. Individuals were placed within one of three light treatments: (1) full-spectrum, high-intensity 'white' light, (2) narrow-spectrum 'blue' light and (3) full-spectrum, reduced-intensity 'gray' light. After 3 months, the intrarhabdomal filters in Row 3 ommatidia of the midband in blue- and gray-light-treated animals were short-wavelength shifted by 10-20 nm compared with homologous filters in animals in white-light treatments. These spectral changes increase the relative sensitivity of associated photoreceptors in animals that inhabit environments where light spectral range or intensity is reduced. The adaptable color vision system of stomatopods may allow animals to make the best use of the ambient light occurring at their habitat regardless of depth. The major controlling element of the plasticity in lysiosquilloid stomatopod color vision appears to be light intensity rather than spectral distribution.

The origin of arthropod sound-producing morphology typically involves modification of two translating body surfaces, such as the legs and thorax. In an unusual structural rearrangement, I show that one lineage of palinurid lobsters lost an antennal joint articulation, which transformed this joint from moving with one degree of freedom into a sliding joint with multiple degrees of freedom. With this sliding joint, 'stick-and-slip' sounds are produced by rubbing the base of each antenna against the antennular plate. To understand the musculo-skeletal changes that occurred during the origin and evolutionary variation of this sound-producing mechanism, I examined joint morphology and antennal muscle anatomy across sound-producing and non-sound-producing palinurids. Plectrum movement and antennal muscle activity were measured in a sound-producing species, Panulirus argus. The promotor muscle pulls the plectrum over the file during sound-producing and non-sound-producing movements; a higher intensity of muscle activity is associated with sound production. The promotor muscle is larger and attaches more medially in sound-producing palinurids than in non-sound producers. In Panulirus argus, each shingle on the file has an additional ridge; in Palinurus elephas, the shingle surfaces are smooth. These differences in shingle surface features suggest variation in the stick-and-slip properties of the system. Translational motion permitted by the sliding joint is necessary for sound production; hence, the construction of a sliding joint is a key modification in the origin of this sound-producing mechanism.

The relationship between segmental mobility and the organisation of thoracic stretch receptors was examined in the deep-sea isopod Bathynomus doederleini, which shows a developed adaptive behaviour during digging. The movements of segments during digging were analysed from video recordings, which showed that a large excursion occurred in the anterior thoracic segments. Dye-fills of axons revealed four types of thoracic stretch receptor (TSR): an N-cell type (TSR-1), a differentiated N-cell type (TSR-2), a muscle receptor organ (MRO)-type with a long, single receptor muscle (TSR-3) and an MRO-type with a short, single receptor muscle (TSR-4 to TSR-7). Physiologically, TSR-1 and TSR-2 are tonic-type stretch receptors. TSR-3 to TSR-7 show two kinds of stretch-activated responses, a tonic response and a phasico-tonic response in which responses are maintained as long as the stretch stimulus is delivered. Both TSR-2, with a long muscle strand, and TSR-3, with a single, long receptor muscle, have a wide dynamic range in their stretch-activated response. In addition, TSR-2 is controlled by an intersegmental inhibitory reflex from TSR-3. These results suggest that, although TSR-1 has no receptor muscle and TSR-2 has a less-differentiated receptor-like muscle, they are fully functional position detectors of segmental movements, as are the MRO-type receptors TSR-3 to TSR-7.

The stomatogastric nervous system (STNS) controls the movements of the foregut and the oesophagus of decapod crustaceans and is a good example for demonstrating that peptides are ubiquitously distributed chemical mediators in the nervous system. The stomatogastric ganglion (STG), one of the four ganglia of the STNS, contains the most intensively investigated neuronal circuits. The other ganglia, including the two commissural ganglia (CoGs) and the oesophageal ganglion (OG), are thought to be modulatory control centres. Peptides reach the STNS either as neurohormones or are released as transmitters. Peptide neurohormones can be released either from neurohaemal organs or from local neurohaemal release zones located on the surface of nerves and connectives. There were thought to be no peptidergic neurones with cell bodies in the STG itself. However, some have recently been described in adults of four species, in addition to a transient expression of peptides during development in two species. None of these peptidergic neurones has been investigated physiologically, in contrast to peptidergic neurones that project to the STG and have cell bodies in either the CoGs or the OG. It has been shown that neurones containing the same peptide elicit different motor patterns, that the peptide transmitter and the classical transmitter are not necessarily co-released and that the effect of a peptidergic neurone depends on its firing frequency and on which other modulatory neurones are co-active. The activity of modulatory projection neurones can be elicited by sensory neurones, and their activity can depend on the firing frequency of the sensory neurone. In addition to being found within the neuropile of ganglia, peptides are present in neuropile patches located within the nerves of the STNS, suggesting that these nerves can integrate as well as transfer information. Furthermore, sensory neurones and muscles exhibit peptide-like immunoreactivity and are modulated by peptides. Bath-applied peptides elicit peptide-specific motor patterns within the STG by targeting subsets of neurones. This divergence is contrasted by a convergence at the level of currents: five different peptides modulate a single current. Peptides not only induce motor patterns but can also switch the alliance of neurones from one network to another or are able to fuse different networks. In general, peptides are the most abundant group of modulators within the STNS; they are ubiquitously present, indicating that they play multiple roles in the plasticity of neural networks.

Our understanding of calcium homeostasis during the crustacean moulting cycle derives from research on intermoult animals that has been extrapolated to other stages. In terms of transepithelial Ca(2+) flux, the more interesting stages are those surrounding ecdysis since crustaceans experience a sizeable negative calcium balance in immediate premoult and a significant positive calcium balance in immediate postmoult. These stages are elusive in the sense that larger species such as lobsters are rarely captured at this time, and smaller species such as blue crabs and crayfish are seldom synchronized in their moulting cycle. The reductionist approaches employed in cellular physiology, such as vesicle techniques, employ pooling of fresh tissues from many organisms. Examination of the elusive moulting stages requires more sensitive approaches that can utilize tissue from an individual crustacean to characterize Ca(2+) pumps (Sarco/Endoplasmic Reticulum Ca(2+)-ATPase, SERCA; Plasma Membrane Ca(2+)-ATPase, PMCA) and the Na(+)/Ca(2+) eXchanger (NCX). An emerging subcellular approach described in this paper is to use flow cytometry as a technique to monitor Ca(2+) uptake into Fluo-3-loaded membrane vesicles. This paper illustrates the utility of this technique for measuring ATP-dependent Ca(2+) uptake into hepatopancreatic basolateral membrane vesicles. Obstacles to progress in molecular studies have not been limited by synchronization of moulting since tissue can be snap-frozen and collected from many animals over time. Here, the problem has been the lack of specific antibodies that hybridize with the Ca(2+) transporters of interest so that they can be localized within epithelia. In this paper, we introduce polyclonal antibodies raised in rabbits against crayfish SERCA, PMCA and NCX. Immunocytochemistry of SERCA in muscle, PMCA in antennal gland and NCX in heart confirms the specificity of the antibodies.

Work in the last decade has shown that crustacean open circulatory systems are highly efficient and controlled in a complex manner. Control occurs at several levels. Myocardial contraction is initiated in the cardiac ganglion but constantly modulated by the central nervous system, both directly via the cardioregulatory nerves and indirectly via the neurohormonal system. Heart rate and stroke volume can be controlled independently and measurements of both are needed to assess cardiac output accurately. Haemolymph outflow from many arthropod hearts is via a complex multiarterial distribution system, and the regional distribution of cardiac output is tightly controlled via cardioarterial valves at the base of each artery. These valves contain innervated muscle, and differential contraction serves to regulate the efflux of oxygenated haemolymph into a particular system. The major influence on both the evolution and control of arthropod open blood vascular systems is efficiency of oxygen uptake and delivery. This influence is illustrated by reference to a variety of crustacean and other arthropod types.

The structural organization of the retinula cells in the eye of Ligia exotica changes diurnally. At night, the microvilli elongate, losing the regular and parallel alignment characteristic of the day condition. Crystalline cones and distal rhabdom tips are not pushed into each other during the day, but at night the rhabdoms protrude into the crystalline cones by up to 5 microm. Screening pigment granules in the retinula cells disperse during the night, but migrate radially towards the vicinity of the rhabdom during the day. No such displacements of the pigment granules of either distal or proximal screening pigment cells were observed. The sensitivity of the eye, monitored by electroretinogram (ERG) recordings, changes diurnally: values at midnight are, on average, 10 times those occurring during the day. However, intracellular recordings from single retinula cells (50 during the day and 50 at night) indicate that the difference between night and day sensitivities is only 2.5-fold. Two-dimensional angular sensitivity curves, indicative of a single unit's spatial sensitivity, had considerably less regular outlines at night than during the day. If based on the 50 % sensitivity level, day and night eyes possessed receptive fields of almost identical width (approximately 2 degrees), but if sensitivities below the 50 % limit were included, then receptive fields at night were significantly more extensive. We suggest that the morphological adaptations and diurnal changes in chromophore content seen in the apposition eye of L. exotica allow this animal to improve its photon capture at night while preserving at least some of the spatial resolving power characteristic of the light-adapted state. This would explain why this animal is capable of performing complex escape behaviours in the presence of predators both in bright and in very dim light.

Many crustaceans detect odors from distant sources using chemosensory sensilla (aesthetascs) on their antennules. The greater the flow of water through arrays of aesthetascs, the faster the access of odorant to receptors inside the aesthetascs. Stomatopods facilitate odorant access by flicking their antennules, thus increasing the relative velocity of the water reaching their aesthetascs. We used dynamically scaled physical models to investigate how aesthetasc size and spacing and antennule flicking velocity affect flow penetration into the simple aesthetasc arrays of the stomatopod Gonodactylaceus mutatus. Particle image velocimetry of flow fields near models of juvenile and adult antennules revealed that velocity gradients around the aesthetascs are steeper during the outward part of the flick than during the return stroke and that the velocity gradients are steeper at the aesthetasc tips than at their bases. More fluid per unit time flows between aesthetasc rows during the outward stroke than during the return stroke, ensuring that odor sampling is pulsatile. During flicking, velocity gradients are steeper near adult aesthetascs than near juvenile aesthetascs, and adults process more fluid per unit time than juveniles. The resulting differences in odorant access can be related to size- and age-dependent changes in stomatopod ecology.

The salmon louse, Lepeophtheirus salmonis, is an ectoparasitic copepod of salmonid fishes whose life cycle involves two broadly defined, free-living larval stages, the nauplius and the copepodid. After settling on a host, the copepodid goes through various transformations to become a mobile adult. We recorded swimming responses of free-swimming salmon lice at the naupliar, copepodid and adult stages to the onset (ON) and offset (OFF) of lights of varying spectral irradiance and polarization. Nauplii showed a prominent swim-up OFF response across the spectrum 352-652 nm, but no ON response. Copepodids exhibited a swim-up ON response and a passive (sinking) OFF response across the same spectral range. Adults showed active swim-up responses to both ON and OFF stimuli, although the OFF response was proportionately stronger. The spectral range of the adult ON and OFF responses was the same as that of the copepodids and slightly greater than that of the nauplii, which did not exhibit responses at 652 nm. The absolute sensitivity of copepodids under white light (approx. 10(-13) photons m(2) s(1)) was higher than that of nauplii (approx. 10(-17) photons(-1) m(2 )s, OFF response) and that of adult female lice (approx. 10(-14) photons(-1)m(2)s). This suggests that the naupliar visual system is best suited for detection of shadows (e.g. the host) under a bright light field (daylight hours), while copepodids and adults may be more specialized for host detection at crepuscular periods and during the night, when light levels are low. None of the developmental stages responded to the rotation of the plane of polarized light or exhibited any difference in directed response when polarized light was used in place of diffuse light.

We present several lines of evidence for the occurrence of graded synaptic transmission in addition to impulse-mediated transmission at the neuromuscular junction between cardiac ganglion (CG) neurones and the myocardium in the isopod crustacean Ligia exotica. In the heart of adult Ligia exotica, the CG acts as a primary pacemaker for the heartbeat by generating periodic bursts of impulses and entrains the myogenicity of the myocardium via impulse-mediated excitatory junctional potentials. When impulse generation was blocked by tetrodotoxin (TTX; 50 nmol l(-)(1)), the CG neurones and the myocardium periodically exhibited synchronized slow depolarizing potentials. The association between the slow depolarizing potentials in the neurone and the myocardium was eliminated by application of Joro spider toxin (JSTX), a specific glutamate antagonist. When the CG neurone was made quiescent by a higher dose of TTX (1.0 micromol l(-)(1)), sinusoidal current injected into the CG neurone induced similar sinusoidal membrane potential responses in the myocardium. The sinusoidal muscle responses were eliminated by application of either JSTX or low-Ca(2+) saline. Under voltage-clamp conditions, the myocardium exhibited periodic inward current responses to sinusoidal current stimuli applied to the CG neurone. The reversal potential for the current response of the myocardium was similar to that of the impulse-mediated excitatory junctional current (EJC). Extracellular macropatch recordings of EJCs made at the neuromuscular junctional site revealed the spontaneous appearance of miniature EJCs asynchronous with the CG spikes in addition to large spike-evoked EJCs. The miniature EJCs were present in saline containing TTX, and their frequency was strongly affected by the slow membrane potential change in the CG neurone. These results suggest that the CG neurones drive the myocardium by graded neuromuscular transmission in addition to impulse-mediated transmission in the heart of Ligia exotica.

The mechanism of active, osmoregulatory ion uptake was investigated in the pleopods of the marine isopod Idotea baltica (Pallas). Using isolated split half-podites of isopods acclimated to brackish water (20 salinity) mounted in a micro-Ussing chamber and symmetrically superfused with identical haemolymph-like salines, a mean short-circuit current I(sc) of -445 microA cm(-)(2) was measured in endopodites 3-5, corresponding to an inwardly directed transcellular movement of negative charge. Application of ouabain (5 mmol l(-)(1)) to the basolateral superfusate resulted in the almost total abolition of the I(sc) (reduced from -531 to -47 microA cm(-)(2)), suggesting that the Na(+)/K(+)-ATPase is the driving force for active, electrogenic uptake of NaCl. In contrast, mean I(sc) values close to zero were found in preparations of all exopodites and in endopodites 1 and 2. The specific activities of Na(+)/K(+)-ATPase corresponded with these results. Specific activities were highest in posterior endopodites 3-5 and depended on ambient salinity. In all other rami, the activities were much lower and independent of ambient salinity. Activities in posterior endopodites 3-5 were lowest in isopods acclimated to 30 salinity (2-4 micromol P(i )mg(-)(1 )protein h(-)(1)), increased in individuals kept in 20 salinity (8.4 micromol P(i )mg(-)(1 )protein h(-)(1)) and were highest in isopods acclimated to 15 salinity (18.2 micromol P(i )mg(-)(1 )protein h(-)(1)). When specimens were transferred from 30 to 40 salinity, Na(+)/K(+)-ATPase activity increased in the posterior endopodites. The electrophysiological and Na(+)/K(+)-ATPase activity measurements show that active electrogenic ion transport in this species occurs almost exclusively in posterior endopodites 3-5. The endopodite of the fifth pleopod of I. baltica exhibited a microscopic structure remarkably similar to that described for the lamellae of the phyllobranchiae of brachyurans. It is composed of two opposed epithelial monolayers of ionocytes, each covered by cuticle. Bundles of pillar cells are located within the ionocyte layers, which are separated by a fenestrated lamellar septum of connective tissue. The results obtained in this study indicate that endopodites 3-5 play the main role in osmoregulatory ion uptake of the isopod I. baltica. Moreover, the Na(+)/K(+)-ATPase is the only driving force behind active electrogenic ion uptake across the epithelial cells.

Reptant decapod crustaceans are almost absent from the Southern Ocean south of the Antarctic Convergence. We tested the hypothesis that this may be due to the reduced ability of this group to regulate Mg(2+) levels in the haemolymph ([Mg(2+)](HL)). Mg(2+) acts as an anaesthetic in marine invertebrates and its level is higher in Reptantia (crabs such as Cancer spp., Chionoecetes spp., Maja spp., 30-50 mmol l(-)(1)) than in Natantia (prawns such as Pandalus spp., Palaemon spp., Crangon spp., 5-12 mmol l(-)(1)). We varied [Mg(2+)](HL) in three species of reptant decapod crustaceans, Carcinus maenas, Hyas araneus and Eurypodius latreillei, and investigated heart rate, the rate of oxygen consumption and levels of spontaneous and forced activity at different temperatures. The rate of oxygen consumption and heart rate increased significantly with reduction in [Mg(2+)](HL) over the entire temperature range investigated in E. latreillei. In H. araneus, an increase in metabolic and heart rates compared with control values was found only at temperatures below 2 degrees C. Forced and spontaneous activity levels increased significantly in the group of [Mg(2+)](HL)-reduced animals below 0 degrees C, at which control animals were mostly inactive. At a reduced [Mg(2+)](HL) of 5-12 mmol l(-)(1), which is the [Mg(2+)](HL) of caridean shrimps in the Southern Ocean, Q(10) and activation energy were reduced for all these variables and extended the temperature range over which physiological functions were maintained. We suggest that the high [Mg(2+)](HL) in Reptantia causes relaxation of the animals and reduces their scope for activity, especially at temperatures below 0 degrees C. The hypothesis that the synergistic effects of high [Mg(2+)](HL) and low temperature probably prevented the Reptantia from recolonizing the permanently cold water of polar areas is discussed.

The mantis shrimp Squilla mantis shows a graded series of avoidance/escape responses to visual and mechanical (vibration and touch) rostral stimuli. A low-threshold response is mediated by the simultaneous protraction of the thoracic walking legs and abdominal swimmerets and telson, producing a backwards 'lurch' or jump that can displace the animal by up to one-third of its body length, but leaves it facing in the same direction. A stronger response starts with similar limb protraction, but is followed by partial abdominal flexion. The maximal response also consists of limb protraction followed by abdominal flexion, but in this case the abdominal flexion is sufficiently vigorous to pull the animal into a tight vertical loop, which leaves it inverted and facing away from the stimulus. The animal then swims forward (away from the stimulus) and rights itself by executing a half-roll. A bilaterally paired, large-diameter, rapidly conducting axon in the dorsal region of the ventral nerve excites swimmeret protractor motoneurons in several ganglia and is likely to be the driver neuron for the limb-protraction response. The same neuron also excites unidentified abdominal trunk motoneurons, but less reliably. The escape response is a key feature of the malacostracan caridoid facies, and we provide the first detailed description of this response in a group that diverged early in malacostracan evolution. We show that the components of the escape response contrast strongly with those of the full caridoid reaction, and we provide physiological and behavioural evidence for the biological plausibility of a limb-before-tail thesis for the evolution of the escape response.

The primary determinants of muscle force throughout a shortening-lengthening cycle, and therefore of the net work done during the cycle, are (1) the shortening or lengthening velocity of the muscle and the force-velocity relationship for the muscle, (2) muscle length and the length-tension relationship for the muscle, and (3) the pattern of stimulation and the time course of muscle activation following stimulation. In addition to these primary factors, there are what are termed secondary determinants of force and work output, which arise from interactions between the primary determinants. The secondary determinants are length-dependent changes in the kinetics of muscle activation, and shortening deactivation, the extent of which depends on the work that has been done during the preceding shortening. The primary and secondary determinants of muscle force and work are illustrated with examples drawn from studies of crustacean muscles.

Developmental changes in heartbeat pacemaker mechanisms were examined electrophysiologically in the isopod crustacean Ligia exotica. The heartbeat of embryos and early juveniles was myogenic. The heart muscle cells were coupled electrically, and no localized pacemaker activity was found in the heart. In newly hatched juveniles, the cardiac ganglion exhibited no spontaneous activity, although stimulation of the cardiac ganglion produced excitatory junctional potentials (EJPs) in the heart muscle. The myogenic activity of the heart was reset and entrained by the EJPs evoked by ganglionic stimulation. During juvenile development, spontaneous EJPs appeared irregularly in the heart muscle. Later in development, the cardiac ganglion started rhythmic bursting, and each muscle response followed a ganglionic burst discharge and overlapped the EJPs evoked by ganglionic activity. At this point, the activity of the cardiac ganglion was suppressed by application of tetrodotoxin (TTX); however, even in old adults, both muscle activity and the heartbeat continued following TTX application. Heartbeat frequency was lower in TTX-containing saline than in normal saline. These results show that, during juvenile development, the heart pacemaker is transferred from the heart muscle to the cardiac ganglion, which becomes the primary pacemaker and entrains the heart muscle activity to a higher frequency via EJPs.

1. In the neurogenic heart of the isopod crustacean Porcellio dilatatus, repetitive electrical stimulation of the cardiac nerves elicted either cardio-acceleratory or cardio-inhibitory effects depending on the stimulation parameters. 2. Acceleratory effects were accompanied by a decrease of membrane potential and by changes in the contour of the spontaneous electrical responses: increase in the speed of the rising phase and enhancement of the plateau phase. 3. Inhibitory stimulation acted on rhytjmicity and/or contour of spontaneous responses. At stimulation pulse frequencies beyond 25/s a hyperpolarization appeared after the cessation of the inhibitory train. 4. Inhibitory stimulation elicted IJPs in the myocardium. Their reversal potential was found to be close to the value of the resting membrane potential. During inhibitory stimulation, the membrane resistance of the heart muscle was frequently decreased. 5. The effects of changing the external chloride content, and of adding GABA and picrotoxin support the hypothesis that the inhibitory impulses increased the myocardium permeability to CL-. 6. On the basis of these findings it is assumed that cardio-inhibitory fibres act on both cardiac ganglion and myocardium. 7. Comparisons are established between the wood-louse's heart and the skeletal or heart muscle of some arthropods. The functional significance of peripheral inhibition is further discussed in relation to the nature of the spontaneous electrical responses and to contraction.