Leverage and muscle type in crab chelae (Crustacea: Brachyura)

The chelae of Cancer pagurus and Macropipus depurator were examined with respect to mechanical advantage. The closer muscles were investigated with respect to sarcomere length in the constituent fibres and to the force developed by the whole muscle during isometric contraction. Cancer chelae have a relatively high mechanical advantage, 0.329 ± 001. Cancer closer muscles contain a high proportion of fibres with long sarcomeres, mean lengths mostly falling between 12 and 15 μm, and develop a maximum stress of about 496 kN.m⁻² during contraction. These figures are typical for "slow" crustacean muscle. The chelae of M. depurator are dimorphic. In one, the strong chela, the mechanical advantage is 0.248 ± 0.066 while in the other, the fast chela, the mechanical advantage is 0.177 ± 0.006. M. depurator closer muscles contain fibres with mean sarcomere lengths mostly falling between 6 and 10 μm. The muscle develops a maximum stress of about 145 kN.m⁻² during contraction. These figures are typical of "intermediate" crustacean muscles. "Fast" muscle fibres with short sarcomeres (about 30 um) were found in the chelae of both Cancer and M. depurator but were much commoner in the latter. Thus in Cancer a high mechanical advantage is correlated with slow muscle while in M. depurator lower mechanical advantages are broadly correlated with faster muscle. Consistent correlation between mechanical advantage and muscle type in the dimorphic chelae of M. depurator , however, is lacking. No consistent regionation of fibres with similar properties was found in the muscles.     

A comparison of the strengths of gastropod shells with forces generated by potential crab predators

Carcinus manenas, Liocarcinus puber and Cancer pagurs are thought to be three likely crab predators of the gastropod Calliostoma Zizyphinum. In order to compare the strenghts of predators and their prey, the whole shell and aperture lip strengh of white and pink Calliostoma morphotypes and the maximum forces exerted by the chelipeds of three crab species were measured. Although white shells were thicker than pink shells, Calliostoma colour morphotyes did not differ significantly in either the force required to break the shell lip or the whole shell. Both Liocarcinus puber and Carcinus maenas have dimorphic chelipeds and their "crusher" chelipeds deliver almost double the forces generated by the'cutter'chelipeds. In constrast, Cancer pagurus has monomorphic chelipeds both delivering similar forces.
When compared with Calliostoma shell strenght, the forces generated by the'crusher'chelipeds of most L. puber tested were, in general, sufficient to break the shell lip of Calliostoma shells, whereas forces generated by the'cutter'chelipeds of only the larger individuals were sufficient to break the shell lip. In C. manenas , forces generated by both the'cutter'and'crusher'chelipeds often exceeded the minimum recorded force required to break the shell lip and the'crusher'cheliped reached the minimum force required to break whole Calliostoma shells. Both chelipeds of all C. pagurus tested generated forces in excess of the minimum required to break the shell lip, and the proportion of individuals capable of generating the minimum force required to break the whole shell increased with the size of the size of the crab. Carcinus maenas and Cancer pagurus were capable of breaking both the shell lips and the whole shells of a wider range of shell sizes than L. puber.     

Hole boring of crustacean prey by the octopus Eledone cirrhosa (Mollusca, Cephalopoda)

Eledone cirrhosa has been found to make a borehole in the carapace of a high proportion of its crustacean prey. This is the first account of drilling in crustaceans by octopus. The frequency of incidence of the boring behaviour varied between prey species from 17 to 93% of those killed. The incidence of boreholes was higher in crabs killed by small octopuses. Using crabs mainly of the genera Cancer, Carcinus, Corystes and Macropipus , the distribution and orientation of the boreholes was recorded. The boreholes occurred in any part of the carapace but the great majority were found close to the mid-line and in the posterior half. The long axis of the oval penetration was usually aligned with the anteroposterior axis of the crab. The Mediterranean species Eledone moschata was also found to bore crabs, and newly hatched juveniles of this species are capable of boring as early as their second killing of a live crab. No evidence was found that Octopus vulgaris normally bores the carapace of crabs.     

Attachment and penetration of Centrocestus armatus (Digenea: Heterophyidae) cercariae to gills of secondary intermediate fish hosts

The attachment and penetration of Centrocestus armatus cercariae into the fish host Zacco temmincki are described in this study. Light and scanning electron microscopy (SEM) were used to examine the topographical features and behavior of cercariae. Histochemistry and transmission electron microscopy (TEM) were employed to trace glandular products and secretions released by cercariae during penetration. Cercariae are first carried into the fish gill chambers via the respiratory currents. The frequency of respiratory-current reversals of fish increased when infected with cercariae. The behavior of cercariae, during breaks in current flow that preceded each current reversal, was observed using a specially devised apparatus. Cercariae produce a mucus-like secretion upon attachment, shed their tail, and employ a brief period of leech-like creeping behavior before penetration. In all cases, the site of penetration was via the surface of the primary gill lamellae. SEM revealed a well-developed anterior penetration apparatus, and a highly contractile body region, that created a driving force for penetration. TEM and histochemistry showed that the mucus observed on the surface tegument of cercariae during attachment were glandular secretions from the parasite. The significance of fish respiratory current reversals to the success of cercariae penetration nicely illustrates the exploitation by the parasite of a host response to a stimulus.     

Structural adaptations for ram ventilation: Gill fusions in scombrids and billfishes

For ram‐gill ventilators such as tunas and mackerels (family Scombridae) and billfishes (families Istiophoridae, Xiphiidae), fusions binding the gill lamellae and filaments prevent gill deformation by a fast and continuous ventilatory stream. This study examines the gills from 28 scombrid and seven billfish species in order to determine how factors such as body size, swimming speed, and the degree of dependence upon ram ventilation influence the site of occurrence and type of fusions. In the family Scombridae there is a progressive increase in the reliance on ram ventilation that correlates with the elaboration of gill fusions. This ranges from mackerels (tribe Scombrini), which only utilize ram ventilation at fast cruising speeds and lack gill fusions, to tunas (tribe Thunnini) of the genus Thunnus, which are obligate ram ventilators and have two distinct fusion types (one binding the gill lamellae and a second connecting the gill filaments). The billfishes appear to have independently evolved gill fusions that rival those of tunas in terms of structural complexity. Examination of a wide range of body sizes for some scombrids and billfishes shows that gill fusions begin to develop at lengths as small as 2.0 cm fork length. In addition to securing the spatial configuration of the gill sieve, gill fusions also appear to increase branchial resistance to slow the high‐speed current produced by ram ventilation to distribute flow evenly and optimally to the respiratory exchange surfaces. J. Morphol. 2012. © 2012 Wiley Periodicals, Inc.     

Antifouling adaptations of marine shrimp (Decapoda: Caridea): gill cleaning mechanisms and grooming of brooded embryos

Gills in the branchial chambers of caridean shrimps, as well as the brooded embryos in females, are subject to fouling by particulate debris and epizoites. Important mechanisms for cleaning the gills are brushing of the gills by the grooming or cleaning chelipeds in some species, while in others, setae from the bases of the thoracic legs brush up among the gills during movement of the limbs (epipodsetobranch complexes). Setae of cleaning chelipeds and of epipod-setobranch complexes show similar ultrastructural adaptions for scraping gill surfaces. Ablation of the cleaning chelipeds of the shrimp Heptacarpus pictus results in severe fouling of the gills in experimentals, while those of controls remain clean. Embryos brooded by female carideans are often brushed and jostled by the grooming chelipeds. In H. pictus , removal of the cleaning chelae results in heavier microbial and sediment fouling than in controls.     

THE STRUCTURE OF FISH GILLS IN RELATION TO THEIR RESPIRATORY FUNCTION

1. The general structure of the gills of different fishes is compared and it is concluded that, though essentially the same, there are certain differences by which they can be recognized. Possible ways in which they may have evolved from one another are considered. 2. A detailed account is given of the structure of the secondary lamellae, where gaseous exchange takes place, and it is shown that two epithelial sheets are separated by a vascular axis mainly composed of pillar cells overlain by a basement membrane on each side. Blood pathways through the gills are discussed in relation to their respiratory function. 3. The embryonic development of gills is described and evidence regarding homo-logies of different structures, particularly the pillar cells, is reviewed. 4. The gills of fish having different modes of life show variations in (a) the number of arches, (b) the number and length of the gill filaments, and (c) the size and frequency of the secondary lamellae. Ways in which measurements of gill area may be carried out and some of the complications involved are reviewed and a summary given of measurements made for a wide variety of species. Measurements of the thickness of the water-blood barrier are also discussed; the more active fish generally have thinner water-blood barriers and larger gill areas. 5. The different mechanisms of gill ventilation are summarized and characteristics of gill resistance in elasmobranchs and teleosts are compared. Gas exchange is discussed in relation to available techniques and the current terminology and symbols, and to indicate the value of analogies between gill exchangers and systems studied by engineers. 6. It is outlined how studies of the functioning of gills during coughing, parasitic infection, and in polluted waters add to knowledge of their role in respiration.     

Sensory and central nervous control of gill ventilation in Limulus.

The gills of Limulus are ventilated by a metachronal rhythm of movements of five pairs of gill plates. A gill plate is promoted and remoted by action of alternating nerve impulse bursts to antagonist promotor and remotor muscles. The motor output pattern is centrally generated, requiring no sensory feedback. Intracellularly recorded rhythmic activity of respiratory motoneurons consists of cyclic depolarization and spiking, and repolarization. The repolarizations have reversal potentials that indicate that motoneuron burst terminations result from synaptic inhibition. Intracellular and antidromic stimulation of motoneurons has little effect on other motoneurons. This apparent lack of interaction between motoneurons indicates that the central respiratory pattern is generated at interneuronal levels. Proprioceptive reflexes are present; they play little role in modulating the centrally generated motor pattern, but they are capable of partially entraining the rhythm when all gill plates are cycled at frequencies near the respiratory rate. Respiratory rate in intact animals is proportional to the ambient oxygen content, respiration ceasing in an anoxic environment. This oxygen dependence may result from sensory input from external oxygen receptors located in the cuticle between the coxae of the walking legs and within the lamellas of the book gills. The intercoxal units are inhibited by anoxia. Three classes of units are recorded from the gills: units excited by oxygen, units inhibited by oxygen, and units whose mechanosensitivity is oxygen dependent. These external oxygen receptors may modulate ventilation via command fibers present in the ventral nerve cord.     

Ventilation in rainbow trout (Salmo gairdneri, Richardson) with damaged gills

An account is given of damage to the gills of trout which involved reduction in total filament length and consequently reduced respiratory surface area. It was noted that fish with the greatest gill damage tended to ventilate the gills by active swimming (ram ventilation) to a greater extent than those with more normal gills.     

Fine structural survey of old cuticle degradation during pre-ecdysis in two European Atlantic crabs

Light and transmission electron microscopy were used to monitor changes due to the degradation of the old exoskeleton and related events in the sclerites, articular membranes, and gills of two decapod crustaceans (Carcinus maenas and Macropipus puber) during pre-ecdysis. In both sclerites and articular membranes, degradation follows a similar general pattern in both crab species, while the gill cuticle appears unaltered. In early pre-ecdysis (D(0)), the degradation of the old cuticle starts with the secretion of ecdysial droplets by the epidermis. Apolysis, occurring at stage D(1)', is re-defined as an event, not necessarily morphologically observable, consisting in the loss of adherence between the epidermis and the old cuticle during early pre-ecdysis of arthropods. At the stage D(1)', the moulding of the epidermal cell surface occurs in preparation to the deposition of the new cuticle and causes the opening of the ecdysial cleft. In the principal layer of sclerites, degradation of the chitin-protein microfibres should precede mineral dissolution. In contrast to the other degraded cuticle layers, the membranous layer of sclerites and the innermost endocuticular lamellae of articular membranes are transformed into a digestion-resistant fibrous network resembling the ecdysial membrane of insects.     

Nitric oxide changes its role as a modulator of respiratory motor activity during development in the bullfrog (Rana catesbeiana)

Nitric oxide (NO) is a unique chemical messenger that has been shown to play a role in the modulation of breathing in amphibians and other vertebrates. In the post-metamorphic tadpole and adult amphibian brainstem, NO, acting via the neuronal isoform of nitric oxide synthase (nNOS), is excitatory to the generation of lung burst activity. In this study, we examine the modulation of breathing by NO during development of the amphibian brainstem. Isolated brainstem preparations from pre-metamorphic and late-stage post-metamorphic tadpoles (Rana catesbeiana) were used to determine the role of NO in modulating central respiratory neural activity. Respiratory neural activity was monitored with suction electrodes recording extracellular activity of cranial nerve rootlets that innervate respiratory musculature. Brainstems were superfused with an artificial cerebrospinal fluid (aCSF) at 20-22 degrees C containing l-nitroarginine (l-NA; 1-10 mM), a non-selective NOS inhibitor. In pre-metamorphic tadpoles, l-NA increased fictive gill ventilation frequency and amplitude, and increased lung burst frequency. By contrast, l-NA applied to the post-metamorphic tadpole brainstem had little effect on fictive buccal activity, but significantly decreased lung burst frequency and the frequency of lung burst episodes. These data indicate that early in development, NO provides a tonic inhibitory input to gill and lung burst activity, but as development progresses, NO provides an excitatory input to lung ventilation. This changing role for NO coincides with the shift in importance in the different respiratory modes during development in amphibians; that is, pre-metamorphic tadpoles rely predominantly on gill ventilation whereas post-metamorphic tadpoles have lost the gills and are obligate air-breathers primarily using lungs for gas exchange. We hypothesize that NO provides a tonic input to the respiratory CPG during development and this changing role reflects the modulatory influence of NO on inhibitory or excitatory modulators or neurotransmitters involved in the generation of respiratory rhythm.     

Uptake of cadmium through isolated perfused gills of the Chinese mitten crab, Eriocheir sinensis

Using the perfusion method, we compared cadmium accumulation and influx across the gills of the euryhaline Chinese mitten crab Eriocheir sinensis, exposed to 4.8 microM cadmium in the incubation medium (OUT). Cadmium influx was not observed across posterior gills while it ranged from 0.15 to 6.82 nmol Cd g(-1) gill w.w. h(-1) across anterior ones. For these respiratory gills, a strong increase (40 times) was observed when calcium was removed in both incubation and perfusion media while the lack of sodium in the perfusion medium resulted in a 46 times decrease. For crabs acclimated 15 days to artificial seawater, cadmium influx across anterior gills showed a 21 times decrease when compared with freshwater acclimated ones. On the other hand, after 3 h of perfusion, we detected cadmium accumulation in both types of gills, ranging from 3.8 to 68 nmol Cd g(-1) gill w.w. in anterior gills and from 2.1 to 39 nmol Cd g(-1) gill w.w. in posterior ones. Such accumulations represent between 61.3 and 100% of the total uptake of cadmium through the gills. From these results, we suggest that cadmium can penetrate more easily into the hemolymph space through the 'respiratory' type epithelium present in the anterior gills but absent in the posterior ones. This metal uptake is likely to occur at least in part through the same pathways as calcium. On the contrary, cadmium seems to be sequestered inside the posterior gills, perhaps in the cuticle of the salt-transporting type epithelium.     

Functional respiratory anatomy of a deep-sea orbiniid polychaete from the Brine Pool NR-1 in the Gulf of Mexico

Abstract. An undescribed species of Orbiniidae (Annelida; Polychaeta) is found in large numbers associated with communities of the mussel, Bathymodiolus childressi at hydrocarbon seeps on the Louisiana slope (Gulf of Mexico). Their microhabitat is often hypoxic and sulfidic, which poses serious respiratory challenges for an aerobic metazoan. They display several anatomical features that are quite unusual for this family, which likely allow them to live in their food-rich, but oxygen-limited, habitat. The anterior gills are hypertrophied whereas the posterior gills are not. These anterior gills provide the worms with a large gill surface area (≤9.9 cm²/g wet weight), which represents 90–95% of the total gill surface area. The gills contain two blood vessels: a central blood vessel, delimited by a coelomic epithelium, and an intra-epidermal vessel. The diffusion distance between this latter and the environment is only 3 μ-m in the anterior gills, which facilitates gas diffusion. Only the anterior gills are ciliated, which may also facilitate gas exchange across this respiratory surface. The gill cells also contain numerous mitochondria and other electron-dense organelles that might be involved in sulfide detoxification.     

Gross structure of the respiratory organs and dimensions of the gill in the mud–skipper, Periophthalmodon schlosseri (Bleeker)

In Periopkrlialnwdon scldosseri the respiratory organs consist of the gills, the suprabranchial and opercular chambers. The gills are more suited for aerial than aquatic respiration as is shown by the presence of the vascular papillae, blood sinusesand dilated blood vessels in their lamellae. The gill lamellae possess a surface coat of sulphated mucopolysaccharides that prevents water loss during exposure to the air. The filaments of the outer hemibranchs in the first gill arch are reduced to nearly one quarter of those of its posterior hemibranch. The gill area in relation to body weight shows a high slope value ( b =0·93).     

Functional morphology of the gills of the shortfin mako,Isurus oxyrinchus,a lamnid shark

This study examines the functional gill morphology of the shortfin mako, Isurus oxyrinchus, to determine the extent to which its gill structure is convergent with that of tunas for specializations required to increase gas exchange and withstand the forceful branchial flow induced by ram ventilation. Mako gill structure is also compared to that of the blue shark, Prionace glauca, an epipelagic species with lower metabolic requirements and a reduced dependence on fast, continuous swimming to ventilate the gills. The gill surface area of the mako is about one‐half that of a comparably sized tuna, but more than twice that of the blue shark and other nonlamnid shark species. Mako gills are also distinguished from those of other sharks by shorter diffusion distances and a more fully developed diagonal blood‐flow pattern through the gill lamellae, which is similar to that found in tunas. Although the mako lacks the filament and lamellar fusions of tunas and other ram‐ventilating teleosts, its gill filaments are stiffened by the elasmobranch interbranchial septum, and the lamellae appear to be stabilized by one to two vascular sacs that protrude from the lamellar surface and abut sacs of adjacent lamellae. Vasoactive agents and changes in vascular pressure potentially influence sac size, consequently effecting lamellar rigidity and both the volume and speed of water through the interlamellar channels. However, vascular sacs also occur in the blue shark, and no other structural elements of the mako gill appear specialized for ram ventilation. Rather, the basic elasmobranch gill design and pattern of branchial circulation are both conserved. Despite specializations that increase mako gill area and efficacy relative to other sharks, the basic features of the elasmobranch gill design appear to have limited selection for a larger gill surface area, and this may ultimately constrain mako aerobic performance in comparison to tunas. J. Morphol. 271:937–948, 2010. © 2010 Wiley‐Liss, Inc.     

Effects of osmotic shock on the ultrastructure of cell nuclei in euryhaline and stenohaline crustaceans

Changes in cell ultrastructure in relation to volume regulation following application of osmotic challenges have been but poorly documented up to now. In a previous study, we demonstrated that large and immediate changes in the organization of nuclear chromatin occur in mammalian cells under such conditions. Similar chromatin modifications are shown in the present study in the cells of the stenohaline crab Maia verrucosa. They are not seen in the cells of the euryhaline species tested, hyperosmoregulating (Eriocheir sinensis) or osmoconforming (Cancer pagurus). Nor are they seen in cells of Portunus puber, another stenohaline species. These results are discussed in the context of the disrupting effects that inorganic ions have on macromolecular structures and of the protection which may be conferred by organic osmolytes used by various crustacean species.     

Serotonergic modulation of respiratory motor output during tadpole development.

To test the hypothesis that serotonin (5-hydroxytryptamine; 5-HT)-receptor activation elicits age-dependent changes in respiratory motor output, we compared the effects of 5-HT bath application (5-HT concentration = 0.5-25 microM) onto in vitro brain stem preparations from pre- and postmetamorphic bullfrog tadpoles. Recording of motor output related to gill and lung ventilation showed that 5-HT elicits a dose-dependent depression of gill burst frequency in both groups. In contrast, the lung burst frequency response was stage dependent; an increase in lung burst frequency at low 5-HT concentration (< or =0.5 microM) was observed only in the postmetamorphic group. Higher 5-HT concentrations decreased lung burst frequency in all preparations. Gill burst frequency attenuation is mediated (at least in part) by 5-HT(1A)-receptor activation in an age-dependent fashion. We conclude that serotonergic modulation of respiratory motor output 1) changes during tadpole development and 2) is distinct for gill and lung ventilation.     

Neuroendocrine regulation of carbonic anhydrase expression in the gills of the euryhaline green crab, Carcinus maenas

The relationship between branchial carbonic anhydrase (CA) activity, CA gene expression and salinity, and potential mechanisms of regulation, was investigated in the euryhaline green crab, Carcinus maenas, acclimated to 33 ppt and transferred to 10 ppt, and the stenohaline rock crab, Cancer irroratus, acclimated to 32 ppt and transferred to 18 ppt. CA activity in green crabs acclimated to high and low salinity was a function of CA mRNA expression, with low salinity exposure resulting in an increase in both CA expression and activity. Eyestalk ablation (ESA) in green crabs acclimated to high salinity resulted in an increase in CA expression in the posterior, ion-transporting gills, in the absence of the low salinity stimulus. There were no changes in CA activity or expression in the anterior, respiratory gills. ESA also potentiated low salinity-stimulated CA induction, again, only in posterior gills. There were no changes in CA activity in any gills of Cancer irroratus, in response to either ESA or low salinity. These results suggest that CA expression in euryhaline, osmoregulating species, is under inhibitory regulation by a putative repressor found in the eyestalk, and that this mechanism is absent in stenohaline, osmoconforming species. CA expression is maintained at low, baseline levels in crabs acclimated to high salinity by the presence and action of this compound. The effects of the repressor appear to be reduced upon exposure to low salinity, allowing CA induction to occur.     

Male sea lampreys,Petromyzon marinus L., excrete a sex pheromone from gill epithelia

During the period when they are producing sperm, male sea lampreys (Petromyzon marinus L.) release a sex pheromone 7alpha, 12alpha, 24-trihydroxy-5alpha-cholan-3-one-24-sulfate (3 keto-petromyzonol sulfate, 3ketoPZS) that induces search and preference behaviors in ovulating females. In this study, we conducted a series of experiments to demonstrate that release of this pheromone into water takes place exclusively through the gills. In a behavioral maze, water conditioned with the anterior region of spermiating males induced an increase of search and preference behaviors in ovulating females. Similar behavior was not elicited by water conditioned by the posterior region. The anterior region washings and whole-body washings from spermiating males also elicited large and virtually identical electro-olfactogram responses from female sea lampreys, while the posterior washings produced negligible responses. Further, mass spectrometry and immunoassay confirmed that virtually all the 3ketoPZS released into water was through the gills. Immunocytochemistry revealed some gill epithelial cells and hepatocytes from spermiating males contained dense immunoreactive 3ketoPZS, but not those from prespermiating males. These results demonstrate that 3ketoPZS is released through the gill epithelia and suggest that this pheromone or its precursor may be produced in the liver.