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Stomatopods have an elaborate visual system served by a retina that is unique to this class of pancrustaceans. Its upper and lower eye hemispheres encode luminance and linear polarization while an equatorial band of photoreceptors termed the midband detects color, circularly polarized light and linear polarization in the ultraviolet. In common with many malacostracan crustaceans, stomatopods have stalked eyes, but they can move these independently within three degrees of rotational freedom. Both eyes separately use saccadic and scanning movements but they can also move in a coordinated fashion to track selected targets or maintain a forward eyestalk posture during swimming. Visual information is initially processed in the first two optic neuropils, the lamina and the medulla, where the eye's midband is represented by enlarged regions within each neuropil that contain populations of neurons, the axons of which are segregated from the neuropil regions subtending the hemispheres. Neuronal channels representing the midband extend from the medulla to the lobula where populations of putative inhibitory glutamic acid decarboxylase-positive neurons and tyrosine hydroxylase-positive neurons intrinsic to the lobula have specific associations with the midband. Here we investigate the organization of the midband representation in the medulla and the lobula in the context of their overall architecture. We discuss the implications of observed arrangements, in which midband inputs to the lobula send out collaterals that extend across the retinotopic mosaic pertaining to the hemispheres. This organization suggests an integrative design that diverges from the eumalacostracan ground pattern and, for the stomatopod, enables color and polarization information to be integrated with luminance information that presumably encodes shape and motion.

Crustaceans and insects share many similarities of brain organization suggesting that their common ancestor possessed some components of those shared features. Stomatopods (mantis shrimps) are basal eumalacostracan crustaceans famous for their elaborate visual system, the most complex of which possesses 12 types of color photoreceptors and the ability to detect both linearly and circularly polarized light. Here, using a palette of histological methods we describe neurons and their neuropils most immediately associated with the stomatopod retina. We first provide a general overview of the major neuropil structures in the eyestalks lateral protocerebrum, with respect to the optical pathways originating from the six rows of specialized ommatidia in the stomatopod's eye, termed the midband. We then focus on the structure and neuronal types of the lamina, the first optic neuropil in the stomatopod visual system. Using Golgi impregnations to resolve single neurons we identify cells in different parts of the lamina corresponding to the three different regions of the stomatopod eye (midband and the upper and lower eye halves). While the optic cartridges relating to the spectral and polarization sensitive midband ommatidia show some specializations not found in the lamina serving the upper and lower eye halves, the general morphology of the midband lamina reflects cell types elsewhere in the lamina and cell types described for other species of Eumalacostraca.

Mantis shrimps (Stomatopoda) possess in common with other crustaceans, and with Hexapoda, specific neuroanatomical attributes of the protocerebrum, the most anterior part of the arthropod brain. These attributes include assemblages of interconnected centers called the central body complex and in the lateral protocerebra, situated in the eyestalks, paired mushroom bodies. The phenotypic homologues of these centers across Panarthropoda support the view that ancestral integrative circuits crucial to action selection and memory have persisted since the early Cambrian or late Ediacaran. However, the discovery of another prominent integrative neuropil in the stomatopod lateral protocerebrum raises the question whether it is unique to Stomatopoda or at least most developed in this lineage, which may have originated in the upper Ordovician or early Devonian. Here, we describe the neuroanatomical structure of this center, called the reniform body. Using confocal microscopy and classical silver staining, we demonstrate that the reniform body receives inputs from multiple sources, including the optic lobe's lobula. Although the mushroom body also receives projections from the lobula, it is entirely distinct from the reniform body, albeit connected to it by discrete tracts. We discuss the implications of their coexistence in Stomatopoda, the occurrence of the reniform body in another eumalacostracan lineage and what this may mean for our understanding of brain functionality in Pancrustacea.

Stomatopods (mantis shrimps) possess one of the most complex eyes in the world with photoreceptors detecting up to 12 different colors. It is not yet understood why stomatopods have almost four times the number of spectral photoreceptors compared with most other animals. It has, however, been suggested that these seemingly redundant photoreceptors could encode color through a new mechanism. Here we compare the spectral sensitivities across five species of stomatopods within the superfamily Gonodactyloidea using intracellular electrophysiological recordings. The results show that the spectral sensitivities across species of stomatopods are remarkably similar apart from some variation in the long-wavelength receptors. We relate these results to spectral sensitivity estimates previously obtained using microspectrophotometry and discuss the variation in the spectral sensitivity maxima (λmax) of the long-wavelength receptors in regard to the previous findings that stomatopods are able to tune their spectral sensitivities according to their respective light environment. We further discuss the similarities of the spectral sensitivities across species of stomatopods in regard to how color information might be processed by their visual systems.

Mushroom bodies are the iconic learning and memory centers of insects. No previously described crustacean possesses a mushroom body as defined by strict morphological criteria although crustacean centers called hemiellipsoid bodies, which serve functions in sensory integration, have been viewed as evolutionarily convergent with mushroom bodies. Here, using key identifiers to characterize neural arrangements, we demonstrate insect-like mushroom bodies in stomatopod crustaceans (mantis shrimps). More than any other crustacean taxon, mantis shrimps display sophisticated behaviors relating to predation, spatial memory, and visual recognition comparable to those of insects. However, neuroanatomy-based cladistics suggesting close phylogenetic proximity of insects and stomatopod crustaceans conflicts with genomic evidence showing hexapods closely related to simple crustaceans called remipedes. We discuss whether corresponding anatomical phenotypes described here reflect the cerebral morphology of a common ancestor of Pancrustacea or an extraordinary example of convergent evolution., eLife digest With more than four million species, arthropods are the largest and most diverse group of animals on the planet and include, for example, crustaceans, insects and spiders. They are defined by their segmented bodies, hard outer skeletons and jointed limbs. All arthropods share a common ancestor that lived more than 550 million years ago. Exactly how this ancestral arthropod gave rise to the myriad species that exist today is unclear but we know that at some point the arthropod family tree split into branches, one of which went on to become the crustaceans. The crustacean branch then split again, giving rise to a line of descendants that would become the insects. But although insects evolved from crustaceans, the brains of insects possess structures that those of crustaceans do not. Known as mushroom bodies, these structures help to form and store memories. Their absence in crustaceans has therefore been an enduring mystery. Wolff et al. now add a piece to the puzzle by showing that one group of modern-day crustaceans, the mantis shrimps, does in fact possess mushroom bodies. By visualizing cells and pathways within the brains of mantis shrimps, and also a number of closely related species, Wolff et al. show that only these shrimps possess true mushroom bodies. However, some of the mantis shrimp’s close relatives possess a few attributes of these structures. This suggests that mushroom bodies are evolutionarily ancient structures that arose in a common ancestor of insects and crustaceans, before being lost or radically modified in most of the crustaceans. So why did this happen? Mantis shrimps are top predators with excellent vision that hunt over considerable distances, requiring them to evaluate and memorize complex features of their environment. These cognitive demands, which might not be shared by other crustaceans, may have led to the mantis shrimps retaining their mushroom bodies. Further research into the brains and behavior of the mantis shrimp may provide insights into how mushroom bodies construct memories of a complex sensory world.

One approach to investigating functional attributes of the central complex is to relate its various elaborations to pancrustacean phylogeny, to taxon-specific behavioral repertoires and ecological settings. Here we review morphological similarities between the central complex of stomatopod crustaceans and the central complex of dicondylic insects. We discuss whether their central complexes possess comparable functional properties, despite the phyletic distance separating these taxa, with mantis shrimp (Stomatopoda) belonging to the basal branch of Eumalacostraca. Stomatopods possess the most elaborate visual receptor system in nature and display a fascinating behavioral repertoire, including refined appendicular dexterity such as independently moving eyestalks. They are also unparalleled in their ability to maneuver during both swimming and substrate locomotion. Like other pancrustaceans, stomatopods possess a set of midline neuropils, called the central complex, which in dicondylic insects have been shown to mediate the selection of motor actions for a range of behaviors. As in dicondylic insects, the stomatopod central complex comprises a modular protocerebral bridge (PB) supplying decussating axons to a scalloped fan-shaped body (FB) and its accompanying ellipsoid body (EB), which is linked to a set of paired noduli and other recognized satellite regions. We consider the functional implications of these attributes in the context of stomatopod behaviors, particularly of their eyestalks that can move independently or conjointly depending on the visual scene.

One of the most complex eyes in the animal kingdom can be found in species of stomatopod crustaceans (mantis shrimp), some of which have 12 different photoreceptor types, each sampling a narrow set of wavelengths ranging from deep ultraviolet to far red (300 to 720 nanometers) ( 1 – 3 ). Functionally, this chromatic complexity has presented a mystery ( 3 – 5 ). Why use 12 color channels when three or four are sufficient for fine color discrimination? Behavioral wavelength discrimination tests (Δλ functions) in stomatopods revealed a surprisingly poor performance, ruling out color vision that makes use of the conventional color-opponent coding system ( 6 – 8 ). Instead, our experiments suggest that stomatopods use a previously unknown color vision system based on temporal signaling combined with scanning eye movements, enabling a type of color recognition rather than discrimination.

As for many deep-sea animals, the red colouration of the two amphipods Eurythenes gryllus and Anonyx sp. has an important function providing camouflage, as the attenuation of the red wavelengths in seawater is higher than other colours within the visible range. Variation in colouration between different stages of colour intensity (related to size) is evident in both species. The red colour is caused by carotenoids, and the carotenoid composition was identified and quantified using spectral optical density signatures, high-performance liquid chromatography (HPLC) and liquid chromatography–mass spectrometry (LC–MS). The carotenoid astaxanthin was identified as the major carotenoid in both amphipods, both in pure and in esterified forms. In addition, minor amounts of lutein-like, canthaxanthin-like and several unidentified carotenoids were found in E. gryllus, while diatoxanthin, β,β-carotene and canthaxanthin-like carotenoids were detected in Anonyx sp. Generally, both species displayed an increase in the amount of carotenoids as a function of colour intensity and size. Shifts in λmax in the OD (Optical density; dimensionless, acronym absorbance) spectra were evident in both species between the different colour stages in both the in vivo and the in vitro material, probably caused by changes in pigment composition. Similar shifts in λmax were observed between the in vivo and in vitro pigment raw extracts in general, most likely caused by pigment-binding proteins. The differences in pigment composition and wavelength shifts suggest large intra- and inter-specific differences between the two species. Probable reasons for changes in pigment composition could be related to diet, season, moulting patterns, metabolic pathways and reproduction.

Stomatopods (commonly known as mantis shrimps) have one of the most complex visual systems in the animal kingdom with up to 20 different photoreceptor types and the ability to see both linear and circular polarized light. The eye of a stomatopod is divided into three different parts; a dorsal and a ventral hemisphere divided up by 6 distinct rows of specialised ommatidia (visual units) termed the midband. The midband contains a complex array of up to 12 spectral receptors with maximum sensitivities ranging from the ultraviolet (UV) to the far-red (300-700 nm), in addition to achromatic receptors of which several are sensitive to different forms of polarized light. The abundance of spectral receptors has puzzled researchers, as it seemed excessive even for an animal living in a spectrally rich environment such as the coral reef. Furthermore, theoretical analyses suggest that there is really no need to have more than 4-7 receptors to have satisfactory colour vision in the 300-400 nm spectral range. Our increasing knowledge of polarization sensitivity is rapidly expanding the theme of photoreceptor proliferation with up to six channels of polarization information from a combination of midband and hemisphere receptors. The aim of this thesis was to investigate how stomatopods process and analyse chromatic and polarization information using a complementary approach of behavioural, electrophysiological and neuroanatomical techniques. A comparison of spectral sensitivities across species in the superfamily Gonodactyloidea was carried out in Chapter 2. This study showed that their spectral sensitivities remain very similar between species, with narrow, steeply shaped sensitivity curves spread evenly throughout the spectrum, suggesting there is a benefit of having uniform sampling of all wavelengths. Behavioural experiments were performed to test stomatopod spectral discrimination in Chapter 3, and this was found to be surprisingly poor, both compared to other animals and to theoretical modelling. To investigate if the poor discrimination was due to the spectral shape of the stimuli, more natural-shaped stimuli (with step-shaped spectra instead of the conventional peak-shaped spectra) were tested in similar experiments (Chapter 4). However, these experiments yielded similar results to the ones obtained in Chapter 3. The electrophysiological and behavioural results suggest that the stomatopod visual system may use a simpler, serial processing system unlike the qconventionalq opponency system known from other animals and may be the reason for why the spectral and polarization space must be covered by several channels. While such as system may appear exceptional, there are similarities between this system and they way primates process colour, only at different steps in the processing pathway (Chapter 5). Using an interval-decoding scheme coupled with the narrow, binned spectral sensitivities of stomatopods could allow quick and precise colour judgements but at the cost of very fine discrimination. As very little was known about the neural architecture of the stomatopods visual system, and to investigate possible answers to questions posed in Chapter 2-5, a range of methods was employed to examine the various neuronal structures. These included: immunohistochemistry experiments and 3D-reconstructions to visualise the gross morphology of the optic lobes, Bodian staining and fluorescent tracer injections to investigate the neuronal pathways, and Golgi impregnations to identify single neuronal types (Chapter 6). First, observations from previous studies were confirmed, in that the midband information pathway remains visible throughout the three first optic lobes as a distinct swelling (Kleinlogel et al., 2003). Chapter 6.2 describes the cell types found in the first optic neuropil, the lamina, where despite size and shape differences between some cells, no clear specializations were observed in the midband region compared to the hemispheral regions of the lamina. The stomatopod medulla was described in Chapter 6.3 and was shown to be clearly stratified having at least 12 main layers, with the midband pathway distinctly visible as a swelling on the distal side of the medulla. The chromatic information appears to be integrated with the achromatic information in the lobula (the third optic lobe, Chapter 6.4) with lateral collaterals forming in the midband pathway and projecting into the hemispheral regions of the lobula. Further projections from the lobula terminate in optic glomeruli in the lateral protocerebrum, which may be involved in refining and sharpening the signals from the lobula columnar neurons. Finally, the stomatopod central complex in the cephalic brain was described in Chapter 6.5, revealing that stomatopods have a central complex more similar to that of an insect than to other crustaceans, possibly shedding new light of to the evolutionary relationship between crustaceans and insects. Overall, this study has given new insights into how stomatopods process chromatic and polarization information. It has revealed a system that at the photoreceptor level appears very complex, but that not necessarily requires high levels of complex processing. These findings may provide inspiration to the development of new optic and camera technologies, and propose a sparse signal-processing scheme, which could provide useful in artificial imaging technologies that require fast and low-power processing speeds. e

Mantis shrimp and primates both possess good color vision, but the neural implementation in the two species is very different, a reflection of the largely unrelated evolutionary lineages of these creatures. Mantis shrimp have scanning compound eyes with 12 classes of photoreceptors, and have evolved a system to decode color information at the front-end of the sensory stream. Primates have image-focusing eyes with three classes of cones, and decode color further along the visual-processing hierarchy. Despite these differences, we report a fascinating parallel between the computational strategies at the color-decoding stage in the brains of stomatopods and primates. Both species appear to use narrowly tuned cells that support interval decoding color identification.

Reactive organic chemicals comprise a large number of compounds with a variety of reactive moieties. While most assays for reactive toxicity focus on DNA damage, reactivity towards proteins can also lead to irreparable damage, but reactivity towards proteins is typically not included in any test battery for water quality assessment. Glutathione (GSH) is a small tripeptide whose cysteine moiety can serve as a model for nucleophilic sites on proteins. GSH is also an important indicator of detoxification processes and the redox status of cells and due to its protective role, depletion of GSH ultimately leads to adverse effects. A bioassay based on genetically modified Escherichia coli strains was used to quantify the specific reactivity towards the protein-like biological nucelophile GSH. The significance of GSH for detoxification was assessed by comparing the growth inhibition induced by reference chemicals or water samples in a GSH-deficient strain to its fully functional parent strain. The GSH deficient strain showed the same sensitivity as the GSH proficient strain to non-reactive and DNA damaging chemicals, but was more sensitive to chemicals that attack cysteine in proteins. The difference in effect concentrations for 50% inhibition of growth assessed as biomass increase (EC(50)) between the two strains indicates the relevance of GSH conjugation as a detoxification step as well as direct reactivity with cysteine-containing proteins. Seven reference compounds serving as positive and negative controls were investigated. The E. coli strain that lacks GSH was four times more sensitive towards the positive control Sea-Nine, while negative controls benzo[a]pyrene, 2-aminoanthracene, phenol, t-butylhydroquinone, methyl methane sulfonate and 4-nitroquinoline oxide showed equal effect concentrations in both strains. Water samples collected across an indirect potable reuse scheme representing the complete water cycle from sewage to drinking water in South East Queensland, Australia were used to evaluate the applicability of the E. coli assay for reactive toxicity in water samples. While the EC(50) values of the GSH+ strain showed similar trends as in other biological endpoints over the various treatment chains, the specific response indicative of protein damage was only observed in samples that had undergone chlorination as a disinfection process. High natural organic matter or other matrix components disturbed the bioassay so much that we recommend it for future routine testing only in tertiary treated water or drinking water.