Exposure to environmentally relevant levels of settleable atmospheric particulate matter (SePM) limits the aerobic performance of inactive healthy adult fish. We evaluated the gill morpho-functional impacts (gill structure and ionic balance) caused by SePM exposure (96h) in Nile tilapia when fish is impelled to maximal aerobic swimming effort. In control fish, swimming reduced epithelial filament thickness (EFT, 25%), increased the distance between lamellae (DL, 21%), and reduced the plasma Na+ (5%). In resting fish, SePM exposure increased EFT (11%), reduced DL (30%), lamellar height (LH, 18%), and plasma Na+ (4%), whereas increased Cl- and K+ (8% and 20%). In SePM-exposure fish, swimming reduced EFT (40%), increased DL (30%) and LH (30%), while reduced Na+ and Cl- (9% and 18%), and increased K+ when compared to inactive SePM-exposed fish. These morpho-functional limitations may compromise whole-body maintenance and swimming recovery, affecting the fish ability to perform at higher activity levels.
Atmospheric particulate matter (APM) produced by the steel industry comprises a complex mixture of particles that includes a wide variety of metals and metallic nanoparticles. These particles settle out onto areas sur-rounding the industries. There is evidence that this 'settleable' APM (SePM) may cause air-to-water cross-contamination with significant effects on aquatic biota. Recent investigations have reported sublethal impacts on the gill structure and blood oxygen-carrying capacity of fishes, which raises the hypothesis that there will be consequences for gas exchange capacity and ability to support aerobic activities. Therefore, we investigated the effects of an environmentally relevant level of SePM contamination on swimming performance and associated aerobic metabolic rates in Nile tilapia, Oreochromis niloticus. Short-term exposure (96 h) to SePM reduced critical swimming speed, energetic efficiency of aerobic swimming, standard metabolic rate, maximum metabolic rate, and aerobic scope. The compromised swimming performance could have adverse ecological effects by limiting foraging ability, predator evasion, territorial protection, and migration. The impairments to aerobic capacity could also affect overall fish performance by influencing long-term energy balance and allocation to growth and reproduction. Thus, despite being sublethal, SePM contamination is considerably debilitating, and if its limiting effects are not compensated for in the longer term, this may reduce the survival and fitness of fish populations.
Atmospheric particulate matter (APM) emitted by iron ore processing industries has a complex composition, including diverse metallic particles and nanoparticles. Settleable APM (SePM) causes air to water cross-contamination and has recently been demonstrated to have harmful sublethal impacts on fish, eliciting stress responses, affecting the immune system, and reducing blood oxygen-carrying capacity. These findings imply potential consequences for fish aerobic performance and energy allocation, particularly in their ability to tolerate respiratory challenges such as aquatic hypoxia. To assess that potential limitation, we analyzed metabolic, cardiorespiratory, and morphological alterations after exposing tilapia, Oreochromis niloticus, to an environmentally relevant concentration of SePM (96 h) and progressive hypoxia. The contamination initiated detectable gill damage, reducing respiratory efficiency, increasing ventilatory effort, and compromising fish capacity to deal with hypoxia. Even in normoxia, the resting respiratory frequency was elevated and limited respiratory adjustments during hypoxia. SePM increased O(2)crit from 26 to 34% of O-2 (1.84 to 2.76 mg O-2.L (-1)). Such ventilatory inefficacy implies higher ventilatory cost with relevant alterations in energy allocation. Progression in gill damage might be problematic and cause: infection, blood loss, ion imbalance, and limited cardiorespiratory performance. The contamination did not cause immediate lethality but may threaten fish populations due to limitations in physiological performance. This was the first investigation to evaluate the physiological responses of fish to hypoxia after SePM contamination. We suggest that the present level of environmental SePM deserves attention. The present results demonstrate the need for comprehensive studies on SePM effects in aquatic fauna.
The inaugural Kjell Johansen Lecture in the Zoophysiology Department of Aarhus University (Aarhus, Denmark) afforded the opportunity for a focused workshop comprising comparative cardiovascular physiologists to ponder some of the key unanswered questions in the field. Discussions were centred around three themes. The first considered function of the vertebrate heart in its various forms in extant vertebrates, with particular focus on the role of intracardiac shunts, the trabecular (spongy') nature of the ventricle in many vertebrates, coronary blood supply and the building plan of the heart as revealed by molecular approaches. The second theme involved the key unanswered questions in the control of the cardiovascular system, emphasizing autonomic control, hypoxic vasoconstriction and developmental plasticity in cardiovascular control. The final theme involved poorly understood aspects of the interaction of the cardiovascular system with the lymphatic, renal and digestive systems. Having posed key questions around these three themes, it is increasingly clear that an abundance of new analytical tools and approaches will allow us to learn much about vertebrate cardiovascular systems in the coming years.
Development within the cleidoic egg of birds and reptiles presents the embryo with the problem of accumulation of wastes from nitrogen metabolism. Ammonia derived from protein catabolism is converted into the less toxic product urea or relatively insoluble uric acid. The pattern of nitrogen excretion of the green iguana, Iguana iguana, was determined during embryonic development using samples from allantoic fluid and from the whole homogenized egg, and in hatchlings and adults using samples of blood plasma. Urea was the major excretory product over the course of embryonic development. It was found in higher concentrations in the allantoic sac, suggesting that there is a mechanism present on the allantoic membrane enabling the concentration of urea. The newly hatched iguana still produced urea while adults produced uric acid. The time course of this shift in the type of nitrogen waste was not determined but the change is likely to be related to the water relations associated with the terrestrial habit of the adult. The green iguana produces parchment-shelled eggs that double in mass during incubation due to water absorption; the eggs also accumulate 0.02 mM of urea, representing 82% of the total measured nitrogenous residues that accumulate inside the allantois. The increase in egg mass and urea concentration became significant after 55 days of incubation then were unchanged until hatching.
Fish characteristically breathe continuously with a respiratory rhythm originating in the brain. This neural generator has a diffuse and, as yet, poorly defined location within the medulla oblongata. Its output to gill musculature is modulated by inputs from the midbrain and stabilized by feedback from gill mechanoreceptors. Fish also may breathe episodically and even arrest rhythmic breathing when ram-ventilating their gills during forward swimming. During periods of high oxygen demand, feeding muscles can be recruited into the respiratory cycle. These same muscles are used to gulp air by air-breathing fishes. The respiratory rhythm generator for air breathing has yet to be clearly defined.
Fish and amphibians utilise a suction/force pump to ventilate gills or lungs, with the respiratory muscles innervated by cranial nerves, while reptiles have a thoracic, aspiratory pump innervated by spinal nerves. However, fish can recruit a hypobranchial pump for active jaw occlusion during hypoxia, using feeding muscles innervated by anterior spinal nerves. This same pump is used to ventilate the air-breathing organ in air-breathing fishes. Some reptiles retain a buccal force pump for use during hypoxia or exercise. All vertebrates have respiratory rhythm generators (RRG) located in the brainstem. In cyclostomes and possibly jawed fishes, this may comprise elements of the trigeminal nucleus, though in the latter group RRG neurons have been located in the reticular formation. In air-breathing fishes and amphibians, there may be separate RRG for gill and lung ventilation. There is some evidence for multiple RRG in reptiles. Both amphibians and reptiles show episodic breathing patterns that may be centrally generated, though they do respond to changes in oxygen supply. Fish and larval amphibians have chemoreceptors sensitive to oxygen partial pressure located on the gills. Hypoxia induces increased ventilation and a reflex bradycardia and may trigger aquatic surface respiration or air-breathing, though these latter activities also respond to behavioural cues. Adult amphibians and reptiles have peripheral chemoreceptors located on the carotid arteries and central chemoreceptors sensitive to blood carbon dioxide levels. Lung perfusion may be regulated by cardiac shunting and lung ventilation stimulates lung stretch receptors.
Control of the heart rate and cardiorespiratory interactions (CRI) is predominantly parasympathetic in all jawed vertebrates, with the sympathetic nervous system having some influence in tetrapods. Respiratory sinus arrhythmia (RSA) has been described as a solely mammalian phenomenon but respiration-related beat-to-beat control of the heart has been described in fish and reptiles. Though they are both important, the relative roles of feed-forward central control and peripheral reflexes in generating CRI vary between groups of fishes and probably between other vertebrates. CRI may relate to two locations for the vagal preganglionic neurons (VPN) and in particular cardiac VPN in the brainstem. This has been described in representatives from all vertebrate groups, though the proportion in each location is variable. Air-breathing fishes, amphibians and reptiles breathe discontinuously and the onset of a bout of breathing is characteristically accompanied by an immediate increase in heart rate plus, in the latter two groups, a left-right shunting of blood through the pulmonary circuit. Both the increase in heart rate and opening of a sphincter on the pulmonary artery are due to withdrawal of vagal tone. An increase in heart rate following a meal in snakes is related to withdrawal of vagal tone plus a non-adrenergic-non-cholinergic effect that may be due to humoral factors released by the gut. Histamine is one candidate for this role.
Ocypode saratan breathing humid air has a postpulmonary PO2 of 15.7 kPa and PCO2 of 1.0 kPa with a gas exchange ratio over the lung of 0.42. When in air, however, O. saratan perfuses the lung in parallel with the gills to generate a mixed postbranchial PaO2 of 10.5 kPa and PaCO2 of 1.7 kPa in humid conditions with access to water and a PaO2 of 2.7 kPa and PaCO2 of 1.1 kPa in dry conditions with no access to water. When submerged O. saratan does not perfuse the lungs, to give a postbranchial PO2 of 2.7 kPa and a PCO2 in both post and prebranchial haemolymph of approximately 1 kPa. Submerged crabs accumulate lactate ions after 5–6 h which appears to be buffered by either HCO‐ 3 uptake or H+ excretion during passage of the haemolymph through the branchial circulation. The gas exchange ratio across the gills is approximately 1. These data suggest that the main site for oxygen and CO2 exchange in O. saratan is the lungs and that respiratory gas exchange is impaired during desiccation and during submersion, whe...
Live Norway lobsters (Nephrops norvegicus L.) were trawled at depths of 30 to 55 m off the coast of Jutland (Denmark) in late winter (March) and in summer (August) in 2006. Water temperatures at the bottom and surface of the sea were 7 degrees C and 2 degrees C during the winter, and 12 degrees C and 21 degrees C in the summer, respectively. The recovery of specific physiological and metabolic variables from the intense stresses associated with capture (trawling and air-exposure during sorting) was followed in seawater at 5 degrees C in winter or 18 degrees C in summer. Recovery was compared in lobsters held individually in two different live-storage positions, either resting vertically on the tail or sitting horizontally. In winter, many animals were alive when brought on board and approximately 86% were still alive at the end of experimentation (96 h). In summer very few animals were alive when brought on board and, of these, approximately 95% were dead at 24 h. When compared with values measured in laboratory controls, the stresses of capture elicited very high haemolymph lactate contents in both seasons, although levels recovered within 24 h. Trawling also caused very high haemolymph glucose concentrations, which differed with season. In winter, haemolymph glucose was elevated for 24 h to levels significantly higher than in summer. In summer, glucose had returned to control levels by 4 h. At 4 h after trawling, haemolymph O-2 status was not markedly influenced in either season, but there were significant disturbances of acid-base status. In winter, a potential metabolic lactic acidosis was compensated by a marked respiratory alkalosis, with significantly increased haemolymph pH and decreased CO2 total content and partial pressure. These effects disappeared gradually over 96 h. Summer lobsters showed combined metabolic and respiratory acidosis at 4 h, although this had recovered to control values in the small number of survivors sampled at 24 h. The capture stresses elicited very high haemolymph crustacean hyperglycaemic hormone (CHH) titres, significantly higher in summer than in winter. In winter, CHH titre had declined significantly at 24 h, whereas it exhibited a further significant increase at 24 h in summer. Live-storage position had no significant effect on survival or recovery from capture stresses in either season. The results demonstrate that Nephrops were much more stressed by trawling at high summer temperatures and had difficulty recovering from this, with pronounced negative effects on their survival, irrespective of their live-storage position. (C) 2009 Elsevier B.V. All rights reserved.
The role of the parasympathetic nervous system, operating via the vagus nerve, in determining heart rate (f(H)) and cardiorespiratory interactions was investigated in the neotropical fish Piaractus mesopotamicus. Motor nuclei of branches of cranial nerves VII, IX and X, supplying respiratory muscles and the heart, have an overlapping distribution in the brainstem, while the Vth motor nucleus is more rostrally located. Respiration-related efferent activity in the cardiac vagus appeared to entrain the heart to ventilation. Peripheral stimulation of the cardiac vagus with short bursts of electrical stimuli entrained the heart at a ratio of 1:1 over a range of frequencies, both below and sometimes above the intrinsic heart rate. Alternatively, at higher bursting frequencies the induced f(H) was slower than the applied stimulus, being recruited by a whole number fraction (1:2 to 1:6) of the stimulus frequency. These effects indicate that respiration-related changes in f(H) in pacu are under direct, beat-to-beat vagal control. Central burst stimulation of respiratory branches of cranial nerves VII, IX and X also entrained the heart, which implies that cardiorespiratory interactions can be generated reflexly. Central stimulation of the Vth cranial nerve was without effect on heart rate, possibly because its central projections do not overlap with cardiac vagal preganglionic neurons in the brainstem. However, bursts of activity recorded from the cardiac vagus were concurrent with bursts in this nerve, suggesting that cardiorespiratory interactions can arise within the CNS, possibly by irradiation from a central respiratory pattern generator, when respiratory drive is high.
The functional anatomy of the respiratory and cardiovascular systems and the neuro-anatomy and neuro-physiology of the systems that are implicated in the co-ordination of cardiac output with ventilation are reviewed in fish, including air-breathing fish, amphibians and reptiles. Recent data is reviewed in the light of previous studies on mammals. This account focuses on the roles of the autonomic nervous system in both feed-forward and feedback control of the respiration-related, beat-to-beat changes in heart rate that accompany continuous, rhythmical breathing as well as the marked changes in heart rate associated with bouts of discontinuous breathing. The control of cardiac shunting in species with incompletely separated systemic and pulmonary blood flow is also described.
SUMMARY Autonomic control of the cardiovascular system in reptiles includes sympathetic components but heart rate (fH), pulmonary blood flow (Q̇pul) and cardiac shunt patterns are primarily controlled by the parasympathetic nervous system. The vagus innervates both the heart and a sphincter on the pulmonary artery. The present study reveals that whereas both the left and right vagi influence fH, it is only the left vagus that influences pulmonary vascular resistance. This is associated with the fact that rattlesnakes, in common with some other species of snakes, have a single functional lung, as the other lung regresses during development. Stimulation of the left cervical vagus in anaesthetised snakes slowed the heart and markedly reduced blood flow in the pulmonary artery whereas stimulation of the right cervical vagus slowed the heart and caused a small increase in stroke volume (VS) in both the systemic and pulmonary circulations. Central stimulation of either vagus caused small (5–10%) reductions in systemic blood pressure but did not affect blood flows or fH. A bilateral differentiation between the vagi was confirmed by progressive vagotomy in recovered snakes. Transection of the left vagus caused a slight increase in fH (10%) but a 70% increase in Q̇pul, largely due to an increase in pulmonary stroke volume (VS,pul). Subsequent complete vagotomy caused a 60% increase in fH accompanied by a slight rise in Q̇pul, with no further change in VS,pul. By contrast, transection of the right vagus elicited a slight tachycardia but no change in VS,pul. Subsequent complete vagotomy was accompanied by marked increases in fH, Q̇pul and VS,pul. These data show that although the heart receives bilateral vagal innervation, the sphincter on the pulmonary artery is innervated solely by the left vagus. This paves the way for an investigation of the role of the cardiac shunt in regulating metabolic rate, as chronic left vagotomy will cause a pronounced left–right shunt in recovered animals, whilst leaving intact control of the heart, via the right vagus.
The role of the vagus nerve in determining heart rate (f H) and cardiorespiratory interactions was investigated in a neotropical fish, Piaractus mesopotamicus. During progressive hypoxia f H initially increased, establishing a 1:1 ratio with ventilation rate (f R). Subsequently there was a hypoxic bradycardia. Injection of atropine abolished a normoxic inhibitory tonus on the heart and the f H adjustments during progressive hypoxia, confirming that they are imposed by efferent parasympathetic inputs via the vagus nerve. Efferent activity recorded from the cardiac vagus in lightly anesthetized normoxic fish included occasional bursts of activity related to spontaneous changes in ventilation amplitude, which increased the cardiac interval. Restricting the flow of aerated water irrigating the gills resulted in increased respiratory effort and bursts of respiration-related activity in the cardiac vagus that seemed to cause f H to couple with f R. Cell bodies of cardiac vagal pre-ganglionic neurons were located in two distinct groups within the dorsal vagal motor column having an overlapping distribution with respiratory motor-neurons. A small proportion of cardiac vagal pre-ganglionic neurons (2%) was in scattered positions in the ventrolateral medulla. This division of cardiac vagal pre-ganglionic neurons into distinct motor groups may relate to their functional roles in determining cardiorespiratory interactions.
Fish control the relative flow rates of water and blood over the gills in order to optimise respiratory gas exchange. As both flows are markedly pulsatile, close beat-to-beat relationships can be predicted. Cardiorespiratory interactions in fish are controlled primarily by activity in the parasympathetic nervous system that has its origin in cardiac vagal preganglionic neurons. Recordings of efferent activity in the cardiac vagus include units firing in respiration-related bursts. Bursts of electrical stimuli delivered peripherally to the cardiac vagus or centrally to respiratory branches of cranial nerves can recruit the heart over a range of frequencies. So, phasic, efferent activity in cardiac vagi, that in the intact fish are respiration-related, can cause heart rate to be modulated by the respiratory rhythm. In elasmobranch fishes this phasic activity seems to arise primarily from central feed-forward interactions with respiratory motor neurones that have overlapping distributions with cardiac neurons in the brainstem. In teleost fish, they arise from increased levels of efferent vagal activity arising from reflex stimulation of chemoreceptors and mechanoreceptors in the orobranchial cavity. However, these differences are largely a matter of emphasis as both groups show elements of feed-forward and feed-back control of cardiorespiratory interactions.
Protein synthesis is a major determinant of growth and yet little is known about the environmental factors that influence protein synthesis rates in farmed freshwater prawns. To this end, post-larvae and juveniles of Macrobrachium rosenbergii were exposed to various salinities (0, 14, 30 per thousand) to determine whole-animal rates of fractional protein synthesis (k(s)) and oxygen uptake. In the post-larvae that migrate upstream from brackish to freshwater areas, whole-animal k(s) was unaffected by salinity, but rates of oxygen uptake were significantly lower at 14 per thousand. In the freshwater juveniles, a different response was observed, as mean k(s) was significantly higher at 14 per thousand compared with 0 per thousand, but rates of oxygen uptake remained unchanged. Such differences are thought to be related to the energetic costs of osmoregulation and to the ability to maintain osmotic gradients in freshwater. In an additional experiment, acclimation temperature (20, 26, 30 degrees C) had a direct effect on k(s) in juveniles held at 0 per thousand. In all cases, changes in k(s) resulted from alterations in RNA activity at constant RNA capacity. In juveniles at least, whole-animal rates of protein synthesis were highest at 14 per thousand and 30 degrees C which corresponds to the optimal salinity and temperature recommended for the growth and culture of M. rosenbergii.
The present paper reports the synthesis and characterization photoluminescence and thermoluminescence studies of Gd3+ doped Y2O3 phosphors. The effect of variable concentration of europium on photoluminescence (PL) and thermoluminescence (TL) behavior are also studied. The samples were prepared by solid state synthesis technique which is suitable for large scale production of phosphors. The starting materials used for sample preparation are ZrO2 and Gd2O3 and CaF2 used as a flux. The prepared sample was characterized by X-ray diffraction technique (XRD). The surface morphology of prepared phosphor was determined by field emission gun scanning electron microscopy (FEGSEM) technique. The diffraction pattern was measured by transmission electron microscopy (TEM) with selected area diffraction pattern. All prepared phosphor with variable concentration of Gd3+ (0.2–2 mol%) was studied by photoluminescence analysis it is found that the excitation spectra of prepared phosphor shows broad excitation centered at 249 and 254 nm with few shoulder weak peaks at 275, 308 and 315 nm. The excitation spectra with variable concentration of Gd3+ show strong peaks at 613 nm for 254 nm excitation. For 275 nm excitation strong peaks found at 468, 567, 578 and 608 nm. For recording TL glow curve every time 2 mg phosphor was irradiated by UV 254 nm source and fixed the heating rate at 6.7 °C s−1. Sample shows well resolved peak at 97 °C for 2 mol% of Gd3+. Trapping parameters are calculated for every recorded glow curve.
Complex physiological traits, such as routine aerobic metabolic rate or exercise performance, are indicators of the functional integrity of fish that can reveal sub-lethal toxicological effects of aquatic pollutants. These traits have proved valuable in laboratory investigations of the sub-lethal effects of heavy metals, ammonia and various xenobiotics. It is not known, however, whether they can also function as biomarkers of the complex potential range of effects upon overall functional integrity caused by exposure to mixtures of chemicals in polluted natural environments. The current study used portable swimming respirometers to compare exercise performance and respiratory metabolism of fish exposed in cages for three weeks to either clean or polluted sites on three urban European river systems: the river Lambro, Milan, Italy; the rivers Blythe, Cole and Tame, Birmingham, UK; and the river Amstel, Amsterdam, The Netherlands. The UK and Italian rivers were variously polluted with high levels of both bioavailable heavy metals and organics, and the Amstel by mixtures of bioavailable organics at high concentrations. In both the UK and Italy, indigenous chub (Leuciscus cephalus) exposed to clean or polluted sites swam equally well in an initial performance test, but the chub from polluted sites could not repeat this performance after a brief recovery interval. These animals were unable to raise the metabolic rate and allocate oxygen towards exercise in the second trial, an effect confirmed in successive campaigns in Italy. Swimming performance was therefore a biomarker indicator of pollutant exposure in chub exposed at these sites. Exposure to polluted sites on the river Amstel did not affect the repeat swimming performance of cultured cloned carp (Cyprinus carpio), indicating either a species-specific tolerance or relative absence of heavy metals. However, measurements of oxygen uptake during swimming revealed increased rates of routine aerobic metabolism in both chub and carp at polluted sites in all of the rivers studied, indicating a sub-lethal metabolic loading effect. Therefore, the physiological traits of exercise performance and metabolic rate have potential as biomarkers of the overall sub-lethal toxic effects of exposure to complex mixtures of pollutants in rivers, and may also provide insight into why fish do not colonize some polluted environments.
The jeju is a teleost fish with bimodal respiration that utilizes a modified swim bladder as an air-breathing organ (ABO). Like all air-breathing fish studied to date, jeju exhibit pronounced changes in heart rate (fH) during air-breathing events, and it is believed that these may facilitate oxygen uptake (MO2) from the ABO. The current study employed power spectral analysis (PSA) of fH patterns, coupled with instantaneous respirometry, to investigate the autonomic control of these phenomena and their functional significance for the efficacy of air breathing. The jeju obtained less than 5% of total MO2 (MtO2) from air breathing in normoxia at 26 degrees C, and PSA of beat-to-beat variability in fH revealed a pattern similar to that of unimodal water-breathing fish. In deep aquatic hypoxia (water PO2)=1 kPa) the jeju increased the frequency of air breathing (fAB) tenfold and maintained MtO2 unchanged from normoxia. This was associated with a significant increase in heart rate variability (HRV), each air breath (AB) being preceded by a brief bradycardia and then followed by a brief tachycardia. These fH changes are qualitatively similar to those associated with breathing in unimodal air-breathing vertebrates. Within 20 heartbeats after the AB, however, a beat-to-beat variability in fH typical of water-breathing fish was re-established. Pharmacological blockade revealed that both adrenergic and cholinergic tone increased simultaneously prior to each AB, and then decreased after it. However, modulation of inhibitory cholinergic tone was responsible for the major proportion of HRV, including the precise beat-to-beat modulation of fH around each AB. Pharmacological blockade of all variations in fH associated with air breathing in deep hypoxia did not, however, have a significant effect upon fAB or the regulation of MtO2. Thus, the functional significance of the profound HRV during air breathing remains a mystery.