Parallel processing and image analysis in the eyes of mantis shrimps

The compound eyes of mantis shrimps, a group of tropical marine crustaceans, incorporate principles of serial and parallel processing of visual information that may be applicable to artificial imaging systems. Their eyes include numerous specializations for analysis of the spectral and polarizational properties of light, and include more photoreceptor classes for analysis of ultraviolet light, color, and polarization than occur in any other known visual system. This is possible because receptors in different regions of the eye are anatomically diverse and incorporate unusual structural features, such as spectral filters, not seen in other compound eyes. Unlike eyes of most other animals, eyes of mantis shrimps must move to acquire some types of visual information and to integrate color and polarization with spatial vision. Information leaving the retina appears to be processed into numerous parallel data streams leading into the central nervous system, greatly reducing the analytical requirements at higher levels. Many of these unusual features of mantis shrimp vision may inspire new sensor designs for machine vision.     

Use and role of invertebrate models in endocrine disruptor research and testing

Historically, invertebrates have been excellent models for studying endocrine systems and for testing toxic chemicals. Some invertebrate endocrine systems are well suited for testing chemicals and environmental media because of the ease of using certain species, their sensitivity to toxic chemicals, and the broad choice of models from which to choose. Such assays will be useful in identifying endocrine disruptors to protect invertebrate populations and as screening systems for vertebrates. Hormone systems are found in all animal phyla, although the most simple animals may have only rudimentary endocrine systems. Invertebrate endocrine systems use a variety of types of hormones, including steroids, peptides, simple amides, and terpenes. The most well-studied hormone systems are the molting and juvenile hormones in insects, the molting hormones in crustaceans, and several of the neurohormones in molluscs and arthropods. These groups offer several options for assays that may be useful for predicting endocrine disruption in invertebrates. A few invertebrate phyla offer predictive capabilities for understanding vertebrate endocrine-disrupting chemicals. The echinoderms, and to a lesser extent molluscs, have closer evolutionary relationships with the vertebrates than the arthropods and these phyla. The recently identified estrogen receptor structure within the genome of the marine gastropod, Aplysia, indicates that the estrogens, and probably the basic steroid receptor, are quite old evolutionarily. This review of the recent literature confirms the effects of some endocrine-disrupting chemicals on invertebrates--tributyltin on snails, pesticides on insects and crustaceans, and industrial compounds on marine animals.     

Invertebrate models for biomedical research, testing, and education

Invertebrate animals have been used as medicinals for 4,000 years and have served as models for research and teaching since the late 1800s. Interest in invertebrate models has increased over the past several decades as the research community has responded to public concerns about the use of vertebrate animals in research. As a result, invertebrates are being evaluated and recognized as models for many diseases and conditions. Their use has led to discoveries in almost every area of biology and medicine--from embryonic development to aging processes. Species range from terrestrial invertebrates such as nematodes and insects to freshwater and marine life including planarians, crustaceans, molluscs, and many others. The most often used models are the fruit fly Drosophila melanogaster and the minuscule nematode Caenorhabditis elegans. Topics in this article are categorized by biologic system, process, or disease with discussion of associated invertebrate models. Sections on bioactive products discovered from invertebrates follow the models section, and the article concludes with uses of invertebrates in teaching. The models reviewed can serve as references for scientists, researchers, veterinarians, institutional animal care and use committees (IACUCs), and others interested in alternatives to vertebrate animals.     

Tuning of photoreceptor function in three mantis shrimp species that inhabit a range of depths. I. Visual pigments

Visual pigments in many animal species, including stomatopod crustaceans, are adapted to the photic environments inhabited by that species. However, some species occupy a diversity of environments as adults (such as a range of depths in the ocean), and a single set of visual pigments would not be equally adaptive for all habitats in which individuals live. We characterized the visual pigment complements of three species of stomatopod crustaceans, Haptosquilla trispinosa, Gonodactylellus affinis, and Gonodactylopsis spongicola, which are unusual for this group in that each lives at depths from the subtidal to several tens of meters. Using microspectrophotometry, we determined the visual pigments in all classes of main rhabdoms in individuals of each species from shallow or deep habitats. Each species expressed the typical diversity of visual pigments commonly found in stomatopods, but there was little or no evidence of differential expression of visual pigments in animals of any species collected from different depths. Vision in these species, therefore, is not tuned to spectral characteristics of the photic environment by varying the assemblages of visual pigments appearing in their retinas.     

On the influence of fishes on the evolution of benthic crustaceans

Though the interweavement of trophic chains in marine ecosystems of the past cannot be reconstructed, as the abundance, nutrition, reproductive rates and other aspects of the biology of fossil species are not known, it is possible to correlate trends in the evolution of single groups of animals, in the presented case the Crustacea, with the influence of other evolving organisms. The evolution of carnivorous, buoyant, well swimming modern fishes probably induced profound changes in marine ecosystems and influenced the evolution of crustaceans. Indications for these interactions are a. the absence or rare occurrence of defenseless archaic crustaceans in habitats, which are populated by teleostean fishes and the survival of some of these forms in refuges like caves and subterranean waters, and b. the reduction of the pleon in the course of the mesozoic evolution of the Decapoda, which occurs parallel to the radiation of the Teleostei, namely in the period between the early Jurassic and the Tertiary. The shortening of the pleon is interreted as a consequence of the stepwise change from a hyperbenthic to a more benthic Iife-stJe and the abandonment of the caridoid escape reaction. Other adaptations are also construed as results from the selective pressure produced by predators. Extant crustaceans which are able to coexist with fishes, among the macrozoobenthos especially the Decapoda and Peracarida, have a variety of protective adaptations, which help to reduce predation.     

Case studies on decapod crustaceans from the Philippines reveal deep,steep underwater slopes as prime habitats for ‘rare’ species

Relatively few studies have been done to define or assess rarity in the marine environment. Published studies have focused on shallow-water and intertidal habitats, and the available information appears to reflect the same pattern observed in terrestrial environments, i.e., that there are many rare species and few common species in any one given area. However, our studies of the abundance of new and/or supposedly rare taxa of decapod crustaceans from the deep, steep slopes of the island of Balicasag, in the central Philippines, have raised questions on how rarity should be defined in marine invertebrates. Examples of such supposedly rare species of crabs and lobsters (Crustacea: Decapoda) are presented here. That these animals come from deep, steep slopes, a relatively under-studied habitat, highlights the major gaps in current knowledge of marine biodiversity that are in part due to the inadequacy of both traditional and high technology sampling methodologies and the limited habitat types that the former can target. Low-technology, artisanal tangle nets have proved to be an optimal capture technique for deep-water decapod crustaceans on deep, steep slopes; many new taxa have been discovered and, in other cases, perceptions of rarity and endemicity have been corrected.     

Adaptive signaling behavior in stomatopods under varying light conditions

Stomatopod crustaceans (mantis shrimp) are aggressive benthic marine predators with extraordinary color vision. When communicating with conspecifics, many stomatopods display conspicuously colored body areas, often in combination with other types of signals such as motion and chemical cues. Some species occupy wide depth ranges (>30 m), where changing light conditions can influence color perception. To test the potential effects of differing ambient lights on signaling behavior, stomatopods (Gonodactylus smithii) interacted with conspecifics in aquaria, under full-spectrum, high intensity light or light restricted in either spectrum or intensity. During intrasexual and intersexual trials in full-spectrum, high intensity light, animals performed more aggressive acts using colored body parts (meral spread, lunge, strike). Stomatopods used significantly more antennular flicking, and performed aggressive acts at reduced distances under restricted light conditions. To compare the use of antennules in visual and chemical communication, additional experiments showed more antennular flicking in response to chemical stimuli from food or conspecifics compared to seawater controls. This response ceased immediately after ablation of antennular chemoreceptors but returned to pre-treatment levels after 5 days of recovery. These findings suggest that stomatopods can vary their use of signals during conspecific interactions under different photic conditions. These inducible, plastic behavioral responses can potentially improve signal transfer in varying light environments.     

Visual physiology of the Antarctic amphipod Abyssorchomene plebs

Although the visual systems of animals living in the cold, dark water of the deep sea have been investigated for some time, little is known about vision in animals inhabiting polar oceans, where temperatures are even colder and irradiance fluctuates dramatically with ice cover and season. Physiology of the compound eye of the amphipod Abyssorchomene plebs (Gammaridea: Lysianassoidea), a common Antarctic benthic scavenger, was studied electrophysiologically by electroretinography. A. plebs has a monochromatic visual system with a spectral sensitivity maximum at 487 nm, and higher sensitivity at ultraviolet wavelengths than predicted by a visual pigment template. While irradiance sensitivity determined from V/log I curves is comparable to that of mesopelagic crustaceans, temporal resolution calculated from response waveform dynamics and as determined by critical flicker fusion frequency suggest that the A. plebs eye is slower than that of crustaceans from the deep sea. A. plebs photoreceptors are physiologically adapted for a slow lifestyle in a low-light environment, where maximizing photon capture occurs at the expense of detecting fast events in the visual scene.     

Do saline taxa evolve faster? Comparing relative rates of molecular evolution between freshwater and marine eukaryotes

The major branches of life diversified in the marine realm, and numerous taxa have since transitioned between marine and freshwaters. Previous studies have demonstrated higher rates of molecular evolution in crustaceans inhabiting continental saline habitats as compared with freshwaters, but it is unclear whether this trend is pervasive or whether it applies to the marine environment. We employ the phylogenetic comparative method to investigate relative molecular evolutionary rates between 148 pairs of marine or continental saline versus freshwater lineages representing disparate eukaryote groups, including bony fish, elasmobranchs, cetaceans, crustaceans, mollusks, annelids, algae, and other eukaryotes, using available protein‐coding and noncoding genes. Overall, we observed no consistent pattern in nucleotide substitution rates linked to habitat across all genes and taxa. However, we observed some trends of higher evolutionary rates within protein‐coding genes in freshwater taxa—the comparisons mainly involving bony fish—compared with their marine relatives. The results suggest no systematic differences in substitution rate between marine and freshwater organisms.     

Distribution of planktonic crustaceans in Lake Balkhash in relation to environmental factors

The quantitative development of planktonic crustaceans in Lake Balkhash has been studied in relation to water transparency, pH, mineralization, ion concentrations, ion ratios, content of easily oxidizable organic matter, nutrients, and heavy metals. It is shown that the decrease in the abundance of animals in a brackish part of the lake area is caused by a high concentration of K⁺ at a relatively low amount of Ca²⁺ and Na⁺ ions. The dependence of quantitative development of planktonic crustaceans on the content of nutrients, mineralization, concentrations of the main ions (except for alkali metals) and their ratio is not manifested for all species. Pollution of the lake by heavy metals has caused either a sharp decrease or an increase in the range of variations of the abundance of crustaceans.     

Underwater linear polarization: physical limitations to biological functions

Polarization sensitivity is documented in a range of marine animals. The variety of tasks for which animals can use this sensitivity, and the range over which they do so, are confined by the visual systems of these animals and by the propagation of the polarization information in the aquatic environment. We examine the environmental physical constraints in an attempt to reveal the depth, range and other limitations to the use of polarization sensitivity by marine animals. In clear oceanic waters, navigation that is based on the polarization pattern of the sky appears to be limited to shallow waters, while solar-based navigation is possible down to 200-400 m. When combined with intensity difference, polarization sensitivity allows an increase in target detection range by 70-80% with an upper limit of 15 m for large-eyed animals. This distance will be significantly smaller for small animals, such as plankton, and in turbid waters. Polarization-contrast detection, which is relevant to object detection and communication, is strongly affected by water conditions and in clear waters its range limit may reach 15 m as well. We show that polarization sensitivity may also serve for target distance estimation, when examining point source bioluminescent objects in the photic mesopelagic depth range.     

A new look at embryonic development of the visual system in decapod crustaceans: neuropil formation, neurogenesis, and apoptotic cell death

In recent years, comparing the structure and development of the central nervous system in crustaceans has provided new insights into the phylogenetic relationships of arthropods. Furthermore, the structural evolution of the compound eyes and optic ganglia of adult arthropods has been discussed, but it was not possible to compare the ontogeny of arthropod visual systems, owing to the lack of data on species other than insects. In the present report, we studied the development of the crustacean visual system by examining neurogenesis, neuropil formation, and apoptotic cell death in embryos of the American lobster, Homarus americanus, the spider crab, Hyas araneus, and the caridean shrimp, Palaemonetes argentinus, and compare these processes with those found in insects. Our results on the patterns of stem cell proliferation provide evidence that in decapod crustaceans and hemimetabolous insects, there exist considerable similarities in the mechanisms by which accretion of the compound eyes and growth of the optic lobes is achieved, suggesting an evolutionary conservation of these mechanisms.     

Ecological inferences about marine mammals from passive acoustic data

Monitoring on the basis of sound recordings, or passive acoustic monitoring, can complement or serve as an alternative to real-time visual or aural monitoring of marine mammals and other animals by human observers. Passive acoustic data can support the estimation of common, individual-level ecological metrics, such as presence, detection-weighted occupancy, abundance and density, population viability and structure, and behaviour. Passive acoustic data also can support estimation of some community-level metrics, such as species richness and composition. The feasibility of estimation and certainty of estimates is highly context dependent, and understanding the factors that affect the reliability of measurements is useful for those considering whether to use passive acoustic data. Here, we review basic concepts and methods of passive acoustic sampling in marine systems that often are applicable to marine mammal research and conservation. Our ultimate aim is to facilitate collaboration among ecologists, bioacousticians, and data analysts. Ecological applications of passive acoustics require one to make decisions about sampling design, which in turn requires consideration of sound propagation, sampling of signals, and data storage. One also must make decisions about signal detection and classification and evaluation of the performance of algorithms for these tasks. Investment in the research and development of systems that automate detection and classification, including machine learning, are increasing. Passive acoustic monitoring is more reliable for detection of species presence than for estimation of other species-level metrics. Use of passive acoustic monitoring to distinguish among individual animals remains difficult. However, information about detection probability, vocalisation or cue rate, and relations between vocalisations and the number and behaviour of animals increases the feasibility of estimating abundance or density. Most sensor deployments are fixed in space or are sporadic, making temporal turnover in species composition more tractable to estimate than spatial turnover. Collaborations between acousticians and ecologists are most likely to be successful and rewarding when all partners critically examine and share a fundamental understanding of the target variables, sampling process, and analytical methods.     

Some studies on the biosynthesis of ubiquinone, isoprenoid alcohols, squalene and sterols by marine invertebrates

The ability of fourteen marine invertebrates to utilize [(14)C]mevalonate for the biosynthesis of isoprenoid compounds was investigated. Several of the animals, in particular crustaceans, bivalve molluscs, a coelenterate and a sponge, were unable to synthesize squalene and sterols, whereas gastropod molluscs, echinoderms, an annelid and a sponge could. Regardless of sterol-synthesizing ability the animals (with the exception of a sponge) always made dolichol and ubiquinone, and thus a specific block in squalene and sterol synthesis was indicated in some animals. Radioactivity accumulated in relatively large amounts in farnesol and geranylgeraniol in those animals incapable of making sterols.     

A new look at an old visual system: structure and development of the compound eyes and optic ganglia of the brine shrimp Artemia salina Linnaeus, 1758 (Branchiopoda,anostraca)

Compared to research carried out on decapod crustaceans, the development of the visual system in representatives of the entomostracan crustaceans is poorly understood. However, the structural evolution of the arthropod visual system is an important topic in the new debate on arthropod relationships, and entomostracan crustaceans play a key role in this discussion. Hence, data on structure and ontogeny of the entomostracan visual system are likely to contribute new aspects to our understanding of arthropod phylogeny. Therefore, we explored the proliferation of neuronal stem cells (in vivo incorporation of bromodeoxyuridine) and the developmental expression of synaptic proteins (immunohistochemistry against synapsins) in the developing optic neuropils of the brine shrimp Artemia salina Linnaeus, 1758 (Crustacea, Entomostraca, Branchiopoda, Anostraca) from hatching to adulthood. The morphology of the adult visual system was examined in serial sections of plastic embedded specimens. Our results indicate that the cellular material that gives rise to the visual system (compound eyes and two optic ganglia) is contributed by the mitotic activity of neuronal stem cells that are arranged in three band-shaped proliferation zones. Synapsin-like immunoreactivity in the lamina ganglionaris and the medulla externa initiated only after the anlagen of the compound eyes had already formed, suggesting that the emergence of the two optic neuropils lags behind the proliferative action of these stem cells. Neurogenesis in A. salina is compared to similar processes in malacostracan crustaceans and possible phylogenetic implications are discussed.     

Endocrine mechanisms in marine invertebrates

The objective of this minireview is to summarize the recent advances made in the area of marine invertebrate endocrinology, with special emphasis on the literature published in 1972 and 1973. Most of the pertinent publications in these two years have dealt with crustaceans. Nevertheless, significant contributions have also been made with representatives of other groups of marine invertebrates.     

First record of ostracods (Crustacea) in Baltic amber

Ostracods were found in two pieces of Baltic amber for the first time. The animals belong to the freshwater genus Cyclocypris. Since Baltic amber was formed during Eocene times in southern Scandinavia, the age of the specimens is estimated at 42–54 million years. As aquatic organisms, ostracods are seldom trapped in amber; it is considered that one specimen, already dead and dried, was either blown onto sticky resin on a plant stem, or washed there by a high flood level, while the other was probably splashed, alive, in a drop of water against the flowing resin, again probably during a flood. Associated faunal remains, including other crustaceans, suggest a coastal palaeoenvironment with some marine as well as freshwater influence.     

Disease will limit future food supply from the global crustacean fishery and aquaculture sectors

Seafood is a highly traded food commodity. Farmed and captured crustaceans contribute a significant proportion with annual production exceeding 10 M metric tonnes with first sale value of $40bn. The sector is dominated by farmed tropical marine shrimp, the fastest growing sector of the global aquaculture industry. It is significant in supporting rural livelihoods and alleviating poverty in producing nations within Asia and Latin America while forming an increasing contribution to aquatic food supply in more developed countries. Nations with marine borders often also support important marine fisheries for crustaceans that are regionally traded as live animals and commodity products. A general separation of net producing and net consuming nations for crustacean seafood has created a truly globalised food industry. Projections for increasing global demand for seafood in the face of level or declining fisheries requires continued expansion and intensification of aquaculture while ensuring best utilisation of captured stocks. Furthermore, continued pressure from consuming nations to ensure safe products for human consumption are being augmented by additional legislative requirements for animals (and their products) to be of low disease status. As a consequence, increasing emphasis is being placed on enforcement of regulations and better governance of the sector; currently this is a challenge in light of a fragmented industry and less stringent regulations associated with animal disease within producer nations. Current estimates predict that up to 40% of tropical shrimp production (>$3bn) is lost annually, mainly due to viral pathogens for which standard preventative measures (e.g. such as vaccination) are not feasible. In light of this problem, new approaches are urgently required to enhance yield by improving broodstock and larval sourcing, promoting best management practices by farmer outreach and supporting cutting-edge research that aims to harness the natural abilities of invertebrates to mitigate assault from pathogens (e.g. the use of RNA interference therapeutics). In terms of fisheries losses associated with disease, key issues are centred on mortality and quality degradation in the post-capture phase, largely due to poor grading and handling by fishers and the industry chain. Occurrence of disease in wild crustaceans is also widely reported, with some indications that climatic changes may be increasing susceptibility to important pathogens (e.g. the parasite Hematodinium). However, despite improvements in field and laboratory diagnostics, defining population-level effects of disease in these fisheries remains elusive. Coordination of disease specialists with fisheries scientists will be required to understand current and future impacts of existing and emergent diseases on wild stocks. Overall, the increasing demand for crustacean seafood in light of these issues signals a clear warning for the future sustainability of this global industry. The linking together of global experts in the culture, capture and trading of crustaceans with pathologists, epidemiologists, ecologists, therapeutics specialists and policy makers in the field of food security will allow these issues to be better identified and addressed.     

Pathogenic fungi of marine animals: A taxonomic perspective

Fungi cause diseases in a variety of marine animal hosts. After a thorough review of published literature, we identified 225 fungal species causing infections of 193 animal species, for a total of 357 combinations of pathogenic fungi and marine animal hosts. Among the 193 animal host species, Chordata (100 species, 51.8 %) and Arthropoda (68 species, 35.2 %) were discovered to be the most frequently reported hosts of fungal pathogens. Microsporidia (111 species, 49.3 %) constitutes over half of the described pathogenic fungal species of marine animals, followed by Ascomycota (85 species, 37.8 %), Mucoromycota (22 species, 9.8 %), Basidiomycota (6 species, 2.7 %) and Chytridiomycota (1 species, 0.4 %). Microsporidia primarily parasitize marine arthropods and Teleostei fish, while Basidiomycota are primarily known to cause respiratory diseases of marine mammals. Ascomycota has a diverse host range, from mammals, fish, crustaceans, soft corals and sea turtle. Few Mucoromycota and Chytridiomycota were reported to infect marine animals. Fungal diseases documented in this review likely represent a fraction of fungal diseases in the ocean, where it was estimated to be inhabited by 2.15 million animal species. Intensification of aquaculture practices, global warming and marine pollution may increase fungal disease outbreak of marine animals. All the topics mentioned above will be discussed in greater details in this review.     

Ensuring crustacean product quality in the post-harvest phase

Recent studies of the fisheries for the Norway lobster, Nephrops norvegicus (L.), have illustrated the negative effects of pathogens and of the physiological stresses of capture processes on the exploitation of live animals and their products, and have identified mitigating measures. Firstly, having established that trawl capture of N. norvegicus is highly stressful, but that these animals have powerful physiological mechanisms of recovery, procedures for on-board recovery of animals destined for vivier transport to distant European markets have been implemented commercially, with significant improvements in survival rates. Such procedures also mitigate against the initiation of a stress-induced muscle necrosis. Secondly, measurements of post-mortem autolytic and spoilage processes have identified the existence of a post-capture 'handling window' of several hours which allows the whole or tailed products to be preserved, by icing or freezing, without detriment to quality. Commercial consortia of Scottish fishermen are exploiting this opportunity to extend product shelf-life by freezing at sea within this handling window. Thirdly, the well-documented infections of Scottish N. norvegicus populations by the dinoflagellate Hematodinium sp. not only provide examples of pathogen-induced mortality, but also have effects on post-harvest products including vivier transport losses and changes in post-mortem degradation leading to earlier organoleptic rejection. Under commercial conditions these effects can be mitigated by post-capture visual screening, but only during the periods of peak patent infection when parasitised animals are visually identifiable. Wider implementation of such mitigating procedures during the harvesting of wild-caught crustaceans will contribute to a more sustainable exploitation of these valuable marine resources.