Preface: feeding ecology of elasmobranchs

Elasmobranchs are apical predators in most marine communities where they occur, often playing a substantial role in the food web dynamics of those communities. However, despite their high trophic status they are often poorly studied compared to most commercially important teleosts. Furthermore, despite efforts towards ecosystem-based management, elasmobranchs are still often lumped into generic categories referred to as ??shark?? or ??skate?? unclassified, with limited effort to identify individual species. The role of elasmobranchs in ecosystems has never been more important to our understanding of marine ecology due to high levels of exploitation of many species. Similar to other high trophic level predators, many elasmobranchs have life-history characteristics that make them vulnerable to over-exploitation. Elasmobranch populations are now heavily targeted in many fisheries throughout the world. Increasing exploitation of this group is especially alarming because their feeding ecology is poorly studied and by extension their influence in shaping ecosystems. Given recent increased attention on elasmobranchs in the scientific literature, management and conservation circles, and the general news media, researchers over the past decade have begun to more closely examine the ecological role of this important taxon of fishes. Due to this increasing awareness, and the development of new and innovative methods and analytical techniques, it prompted us to organize an international symposium on the ??Feeding Ecology of Elasmobranchs??. The symposium was held on 10 July 2010, in conjunction with the 27^th annual meeting of the American Elasmobranch Society meetings in Providence, Rhode Island.     

Functional ecology of feeding in elasmobranchs

Feeding behavior in the species of captive chondrichthyans is studied to clarify the functional mechanisms responsible for feeding ecology. Kinematics and pressure in the buccal, hyoid and pharyngeal regions were quantified in Squalus acanthias, Chiloscyllium plagiosum and Leucoraja erinacea using sonomicrometry and pressure transducers. Means and coefficients of variation were analyzed by species and by behavior to test for stereotypy and flexibility in the feeding mechanism. Several instances of mechanical stereotypy as well as flexibility were found in the feeding kinematics and pressure of the three chondrichthyan species. In general, Squalus acanthias shows more stereotyped feeding behavior than C. plagiosum and L. erinacea. Different aspects of feeding behavior stand out among the three species. Chiloscyllium plagiosum generates lowest pressures, S. acanthias achieves the greatest area changes, and L. erinacea has longer durations for manipulating prey. Capture events are functionally and behaviorally stereotyped while processing events are functionally and behaviorally flexible with the ability to use suction or compression to process the same food item. Squalus acanthias is a functional specialist and C. plagiosum is functionally a generalist, with both species exhibiting behavioral flexibility. Leucoraja erinacea is a functional and behavioral generalist. Using functional morphology to explain mechanical stereotypy and flexibility in the feeding behavior of three suction feeding chondrichthyan species has allowed a better understanding of specialist and generalist trophic behaviors.     

Evolution and ecology of feeding in elasmobranchs

Paleozoic chondrichthyans had a large gape, numerous spike-liketeeth, limited cranial kinesis, and a non-suspensory hyoid,suggesting a feeding mechanism dominated by bite and ram. Modernsharks are characterized by a mobile upper jaw braced by a suspensoryhyoid arch that is highly kinetic. In batoids, the upper jawis dissociated from the cranium permitting extensive protrusionof the jaws. Similar to actinopterygians, the evolution of highlymobile mandibular and hyoid elements has been correlated withextensive radiation of feeding modes in elasmobranchs, particularlythat of suction. Modern elasmobranchs possess a remarkable varietyof feeding modes for a group containing so few species. Biting,suction or filter-feeding may be used in conjunction with ramto capture prey, with most species able to use a combinationof behaviors during a strike. Suction-feeding has repeatedlyarisen within all recent major elasmobranch clades and is associatedwith a suite of morphological and behavioral specializations.Prey capture in a diverse assemblage of purported suction-feedingelasmobranchs is investigated in this study. Drop in water pressuremeasured in the mouth and at the location of the prey showsthat suction inflow drops off rapidly with distance from thepredator's mouth. Elasmobranchs specializing in suction-feedingmay be limited to bottom associated prey and because of theirsmall gape may have a diet restricted to relatively small prey.Behavior can affect performance and overcome constraints imposedby the fluid medium. Suction performance can be enhanced byproximity to a substrate or by decreasing distance from predatorto prey using various morphological and/or behavioral characteristics.Benthic suction-feeders benefit by the increased strike radiusdue to deflection of water flow when feeding close to a substrate,and perhaps require less accuracy when capturing prey. Suctionand ram-suction-feeding elasmobranchs can also use suction inflowto draw prey to them from a short distance, while ram-feedingsharks must accelerate and overtake the prey. The relationshipbetween feeding strategy and ecology may depend in part on ecological,mechanistic or evolutionary specialization. Mechanistic suction-feedingspecialist elasmobranchs are primarily benthic, while most epibenthicand pelagic elasmobranchs are generalists and use ram, suction,and biting to catch a diversity of prey in various habitats.Some shark species are considered to be ecological specialistsin choosing certain kinds of prey over others. Batoids are evolutionaryspecialists in having a flattened morphology and most are generalistfeeders. Filter-feeding elasmobranchs are ecological, mechanistic,and evolutionary specialists.     

The pit organs of elasmobranchs: a review

Elasmobranchs have hundreds of tiny sensory organs, called pit organs, scattered over the skin surface. The pit organs were noted in many early studies of the lateral line, but their exact nature has long remained a mystery. Although pit organs were known to be innervated by the lateral line nerves, and light micrographs suggested that they were free neuromasts, speculation that they may be external taste buds or chemoreceptors has persisted until recently. Electron micrographs have now revealed that the pit organs are indeed free neuromasts. Their functional and behavioural role(s), however, are yet to be investigated.     

The physiological control of feeding in corals: a review

Little is known about the factors that control food capture by the coral polyp and the way that internal physiological events may coordinate, or even modify, this behaviour. A total understanding of the nutritional dynamics of corals demands detailed study of their feeding behaviour, and recent advances in behavioural and electrophysiological techniques are starting to provide some insights into how polyps control this behaviour. This review describes our present understanding of the physiological control of feeding behaviour in corals and suggests new studies on food recognition, food capture, and ingestion.     

Freshwater elasmobranchs: a review of their physiology and biochemistry

Only 5% of elasmobranch species live in freshwater (FW) compared to more than 40% of known teleost species. The factors affecting the poor penetration of elasmobranchs into FW environments are currently unknown, however, an important consideration may be the high urea requirement of many proteins in marine elasmobranchs. Urea is an important osmolyte in marine elasmobranchs and must be reduced in dilute environments. There are three identifiable stages in the successful colonization of FW. The euryhaline marine species freely entering and leaving FW represent the initial stage of FW colonization. In this group, there is an apparent inability to eliminate all urea due to protein integrity issues and this results in energy and nitrogen losses that may constrain growth and reproduction. The second stage is represented by those species that live entirely in FW but must also retain some urea. This group also suffers from the same constraints as the first group. These two groups have kidneys and sensory organs that more closely resemble strictly marine forms. The third and final stage is represented by the Potamotrygonid stingrays where the need for urea in FW has been eliminated. Consequently nitrogen and energy losses are reduced and those sections of the kidney needed for urea conservation have been eliminated. The driving force for such modifications is a reduction in urea levels and the concomitant saving of energy needed for urea synthesis. Other physiological adaptations associated with survival in FW include giving birth to live young, the capacity of sperm to be activated in freshwater and modifications of the electrosensory system to function in a low conductivity environment. The need for many anatomical, metabolic and physiological modifications for FW existence may constrain the rapidity and hence the frequency of FW colonization, compared to the situation in the more advanced osmoregulating teleosts. Once optimally adapted to FW, recolonization of sea water by elasmobranchs is problematic due to the loss of urea synthetic capacity and renal structures for urea retention.     

Hyoid and pharyngeal arch function during ventilation and feeding in elasmobranchs: Conservation and modification in function

The hypothesis that the mandibular and hyoid arches evolved from anterior pharyngeal arches to increase ventilation performance and subsequently became adapted for feeding is widely accepted. As jaws evolved, the morphology of the hyoid arch changed notably from that of a pharyngeal arch. Furthermore, hyoid arch morphology varies considerably among elasmobranch taxa and has been shown to be related to feeding style. The goal of this study is to determine whether the function (direction of movement or change in cavity cross‐section) of the hyoid arch is altered from that of the pharyngeal arch, and whether function is altered between ventilation, the basal behavior, and feeding, the derived behavior. Similar effects and associations of the pharyngeal arches by orientation to feeding or ventilation are also investigated. The kinematics of the hyoid and second pharyngeal arch during ventilation and feeding are quantified using sonomicrometry and hyomandibular angle measured in five shark and one skate species representing widely divergent hyomandibular morphologies. Hyoid and pharyngeal cavity width follows the same pattern of movement during ventilation; therefore the hyoid arch retains the ancestral function of the pharyngeal arches. The orientation of the hyomandibular cartilage appears to influence the pattern of arch movement during ventilation: anterior directed elements decrease in cavity width; laterally directed elements increase in cavity width; while posterior directed elements increase in cavity width or do not change; while cavity depth increases in all species. Hyoid and pharyngeal cavity width movement differs among the species during feeding and also appears to be related to hyoid arch orientation as well as feeding style. There appears to be a division between those species with hyomandibular angles less than 110° from those that are greater between feeding mode and hyoid cavity width movement. Primarily suction feeding species decrease hyoid cavity width whereas primarily bite feeding species increase hyoid cavity width during feeding while all species increase hyoid cavity depth.     

Sperm storage in male elasmobranchs: A description and survey

Two basic types of spermatozoan aggregates, spermatophores and spermatozeugmata, found in 14 different species of sharks, one species of skate, and one species of chimaera (holocephalan), were investigated using light and scanning electron microscopy. Spermatophores, aggregates (usually 1,000–6,000 μm in diameter and larger) of randomly clumped sperm embedded in and surrounded by an eosinophilic matrix, were found in Alopias superciliosus, Odontaspis taurus, Carcharodon carcharias, Isurus oxyrinchus, and Lamna nasus. Three types of spermatozeugmata, sperm structures without a surrounding capsule or matrix, are described. The first, clumps of 60–200 sperm unbound in a supporting matrix, are found in Squalus acanthias and Hydrolagus colliei. In the second type, single-layered spheres are formed of sperm clumps with the sperm heads bound in a common supporting matrix. These are found in Carcharhinus limbatus and Carcharhinus plumbeus. The third type of spermatozeugmata are large multilayered, compound structures formed by the accretion of several single-layered aggregates. These multilayered structures characteristically are found in Carcharhinus falciformis, C. limbatus, Carcharhinus obscurus, C. plumbeus, Carcharhinus porosus, Prionace gluaca, Rhizoprionodon terraenovae, and Sphyrna lewini. Sperm aggregates of all types are stored between the septa and in the lumen of the terminal ampulla of the epididymis. In their various forms they are the final product of the mature male elasmobranch reproductive tract. In a male with mature claspers, the presence of sperm aggregates is a more reliable indicator of maturity and sexual activity than is clasper condition alone. © 1994 Wiley-Liss, Inc.     

A model for neural control of gradation of muscle force

A mathematical muscle model is presented that relates neural control signals linearly to muscle force without violating important known physiological constraints, such as the size-principle (Henneman and Mendell 1981) and non-linear twitch summation (Burke et al. 1976). This linearity implies that the neural control signals (defined as a weighted sum of activities in a nerve bundle) can be interpreted as the internal representation of total muscle force. The model allows for different relative contributions from the two force-grading mechanisms, i.e. the recruitment of motor units and the modulation of their firing frequency. It can therefore be applied to a variety of (distal and proximal) muscles. Furthermore, it permits simple mechanisms for controlling muscle force, e.g. in superposed motor tasks. The model confirms our intuitive notion that a weighted sum of activities in a nerve bundle can directly represent an external controlled variable, which in this case is exerted muscle force.     

A spring model and equivalent neural network for arm posture control

A model is introduced for the motor program which controls final position. The first part of the model relates the biomechanical properties of the muscles to the EMG activities of the extensor and flexor muscles and thereby generates quatitative predictions for the relationships between the EMGs, final position, external forces, muscle stiffness, and muscle tension. To the extent that comparable data exist, the model is shown to give correct quantitative predictions. When only qualitative comparisons can be made, the model is consistent with the data in the literature. The model is complete and can be tested quantitatively in detail in the future. An equivalent circuit for the neural network that innervates the muscles is given. It is shown to have the advantages of making the programming of final position simple to either compute or lookup in a table. In addition, new situations, such as adapting to a force, or an unusual viewing angle, lead to very simple changes in the basic program in terms of the equivalent circuit.     

Infant feeding practices and breastfeeding duration in Japan: A review

The Japanese health system places great emphasis on healthy development. However, the prevalence of Exclusive Breastfeeding at one month postpartum between 1980 and 2005 has remained unchanged, fluctuating between 42% and 49%. At the same time, the Any Breastfeeding prevalence has gradually increased from about 80% to 95%. In 2010, the latest national breastfeeding report showed that ‘exclusive’ and ‘any’ breastfeeding rates have improved. However, as the World Health Organization (WHO) definition of breastfeeding practices was not used in this study or in other national surveys, it is difficult to interpret these latest results. While the Japanese government has launched several promotion projects, there have been few studies and reviews of risk factors that influence breastfeeding duration. The objectives of this review were to summarise the factors that have influenced the duration of breastfeeding in Japan to provide information relevant to breastfeeding promotion programs. A search of electronic databases in Japanese and English was undertaken up to 2011. The inclusion criteria for this review were studies that focused on infant feeding practices and targeted Japanese mothers, fathers, or health professionals, but excluded mothers’ friends and peer groups. In total, 12 articles were selected for the final analysis. Smoking status, low birth weight of infants and maternal perceptions of insufficient breast milk supply were negative influences on breastfeeding duration, while support from husbands/partners is associated with continued breastfeeding. Some factors that have been found to be associated with breastfeeding in other countries, including maternal age, family income, maternal educational levels, and living with grandparents of infants have not been confirmed in Japan. While the national breastfeeding rates were higher than other countries of similar health status, inconsistent knowledge of breastfeeding benefits and inappropriate hospital practices remain in Japan may be associated with increased the use of infant formula and reduced breastfeeding duration. Most of the studies reviewed were cross-sectional in design, with only a limited number of cohort studies. Also many published studies used small sample sizes. Cohort studies of infant feeding practices with larger sample sizes are required to monitor trends in rates and risk factors for breastfeeding outcomes.     

Gonadotropin-releasing hormone (GnRH) pathways and reproductive control in elasmobranchs

Synopsis Immunoreactive (ir) gonadotropin-releasing hormone (GnRH) is localized in many neurons of the terminal nerve (TN) and midbrain tegmentum, while few ir-cells are observed in the preoptic area and ventral hypothalamus. The paucity of preoptic ir-cells may relate to an unusual feature of the elasmobranch pituitary, i.e. a lack of portal control of gonadotropin-producing cells. TN and midbrain GnRH-ir neurons may be major sources of GnRH used to modulate or otherwise control both pituitary and brain cells via delivery through the systemic circulation. These ir-nuclei also appear to directly innervate CNS regions (the preoptic area, habenula and clasper control area of the spinal cord) involved in sexual functions. Important regulatory mechanisms, represented by interactions between GnRH pathways and sex-steroid concentrating neurons, are likely to occur in the preoptic area, habenula and midbrain tegmentum.     

A model for control of breathing in mammals: coupling neural dynamics to peripheral gas exchange and transport

A new model for aspects of the control of respiration in mammals has been developed. The model integrates a reduced representation of the brainstem respiratory neural controller together with peripheral gas exchange and transport mechanisms. The neural controller consists of two components. One component represents the inspiratory oscillator in the pre-Bötzinger complex (pre-BötC) incorporating biophysical mechanisms for rhythm generation. The other component represents the ventral respiratory group (VRG), which is driven by the pre-BötC for generation of inspiratory (pre)motor output. The neural model was coupled to simplified models of the lungs incorporating oxygen and carbon dioxide transport. The simplified representation of the brainstem neural circuitry has regulation of both frequency and amplitude of respiration and is done in response to partial pressures of oxygen and carbon dioxide in the blood using proportional (P) and proportional plus integral (PI) controllers. We have studied the coupled system under open and closed loop control. We show that two breathing regimes can exist in the model. In one regime an increase in the inspiratory frequency is accompanied by an increase in amplitude. In the second regime an increase in frequency is accompanied by a decrease in amplitude. The dynamic response of the model to changes in the concentration of inspired O₂ or inspired CO₂ was compared qualitatively with experimental data reported in the physiological literature. We show that the dynamic response with a PI-controller fits the experimental data better but suggests that when high levels of CO₂ are inspired the respiratory system cannot reach steady state. Our model also predicts that there could be two possible mechanisms for apnea appearance when 100% O₂ is inspired following a period of 5% inspired O₂. This paper represents a novel attempt to link neural control and gas transport mechanisms, highlights important issues in amplitude and frequency control and sets the stage for more complete neurophysiological control models.     

Diet and feeding habits of two endemic demersal bycatch elasmobranchs: Trygonorrhina fasciata & Dentiraja australis

This study describes diet and feeding habits of two common bycatch elasmobranchs in East Australia: The Eastern Fiddler Ray Trygonorrhina fasciata (Muller & Henle, 1841) and the Sydney Skate Dentiraja australis (Macleay, 1884). Using stomach content analysis, we describe diet composition, food preferences and trophic levels of these batoids. Crustaceans were major components in the diet of both species which included teleosts and cephalopods. There was no indication of sex-related distinction in the diet of either species. The estimated trophic level (TL) of the T. fasciata was TL = 3.36 and the D. australis was TL = 3.43.     

SIRT1 and AMPK in regulating mammalian senescence: a critical review and a working model

Ageing in mammals remains an unsolved mystery. Anti-ageing is a recurring topic in the history of scientific research. Lifespan extension evoked by Sir2 protein in lower organisms has attracted a large amount of interests in the last decade. This review summarizes recent evidence supporting the role of a Sir2 mammalian homologue, SIRT1 (Silent information regulator T1), in regulating ageing and cellular senescence. The various signaling networks responsible for the anti-ageing and anti-senescence activity of SIRT1 have been discussed. In particular, a counter-balancing model involving the cross-talks between SIRT1 and AMP-activated protein kinase (AMPK), another stress and energy sensor, is suggested for controlling the senescence program in mammalian cells.     

A review of the bacterial flora of teleosts and elasmobranchs, including methods for its analysis

Techniques for the quantitative and qualitative analysis of the commensal bacterial flora of fish are reviewed. The literature concerning the commensal bacterial flora of fish indicates that teleost skin often features 10²–0⁷ organisms per cm² of skin, representing the following taxa, in descending numerical order, Achromobacter, Pseudomonas, Flavo-bacterium and Cytophaga species also coryneform organisms. Actively feeding teleosts appear to have an alimentary canal bacterial flora similar to that of the skin and gills, often there are from 10³–10⁸ organisms per 1.0 g, wet weight, of tissue. According to the limited data available for elasmobranchs Gram–positive bacteria are the predominant component of the bacterial flora on the skin of sharks, though the bacterial flora of North Sea skate ( Raja spp.) is reported as being similar to that of teleosts. The commensal flora of fish, particularly of teleosts, is related to their environment and is influenced by environmental changes that affect the quality of the water. Regular monitoring of the water of commercial fish ponds and aquaria for bacterial quality, temperature, dissolved oxygen concentration and pH, combined with a base–line knowledge of the water's quality, may provide an advanced indication of the presence of bacteria potentially able to produce diseased conditions, and factors that will enhance their growth.