Morphology and ultrastructure of the gills of terrestrial crabs (Crustacea,Gecarcinidae and Grapsidae): Adaptations for air-breathing

Summary The terrestrial crabsGeograpsus grayi, Geograpsus crinipes, Cardisoma hirtipes andGecarcoidea natalis have a reduced number of gills and show a reduced planar gill surface (SA) compared to aquatic species. Gill lamellae are stiffened and thickened (increasing blood/gas (BG) diffusion distances) and nodules maintain wide spacing between lamellae. Haemolymph is directed through the gill lamellae by rows of pillar cells and in the afferent region an intralamellar septum splits the haemolymph into two parallel networks. Gaps in the lines of pillar cells allow movement of haemolymph between adjacent channels. The afferent vessel distributes haemolymph to the lamella via a number of direct channels including the marginal canal and in large gills with the aid of a long, forked sinus which supplies the ventral and central regions of the lamellae. The marginal canal functions in both distribution and collection of haemolymph; the role varies with species. Potential flow-control sites were identified at the junctions between afferent and efferent areas and where the efferent channels enter the efferent branchial vessel. Each gill receives a branch from the sternal artery which supplies all the lamellae. Transport epithelia is the principal cell type in the gills of all species examined though its location varies between species, either being confined to certain gills or specific parts of the lamellae.The gill lamellae of air-breathing crabs are clearly modified to breathe air (stiffening and presence of nodules), though the overall contribution of the gills to gas exchange has been reduced (smaller SA and longer BG diffusion distances). The role of the gills in air-breathing crabs thus appears to have switched from one of an efficient aquatic gas-exchanger (thin with large surface area) and transport tissue, to one that is predominantly set up for ion-regulation.Abbreviations a afferent branchial vessel - ac afferent channels - art arteriole - ass artifactual subcuticular space - bl basal lamina - c cuticle - col collagen - ct connective tissue - e efferent branchial vessel - ec efferent channels - epi epithelium - f folds - g Glycogen - h haemolymph - hc haemocyte - is intralamellar septum - m marginal canal - mi mitochondria - mt microtubules - n nucleus - p pillar cell - s shaft of efferent vessel - sd septate desmosome     

The ecophysiological significance of lung-air retention during submersion by the air-breathing crabs Cardisoma carnifex and Cardisoma hirtipes

The air-breathing crabs Cardisoma hirtipes and C. carnifex trap 18.8 and 15.5 ml of air respectively within the lumen of the lung (branchial chamber) when submerging, rather than breathe water, and suffer a hypoxic hypercapnia. The role, effect and advantage of trapping air within the lung during submergence was assessed for both species. Cardisomaretained at least 85% of the lung air during a 30-min submergence. The mass-specific O₂uptake rate (MO₂) of air-breathing C. hirtipes and C. carnifexwas reduced by at least 60% during submersion and the crabs additionally consumed 80% or more of the O₂stored in the trapped air. After 30 min of submergence the large arterial-venous difference in haemolymph partial pressure of O₂ (PO₂) and O₂ content (CO₂) of air-breathingCardisoma was completely removed, consistent with near-zero O₂ transport. Removing the air from the lung lumen did not restore MO₂ but rather deprived the crab of an O₂ store and, in C. hirtipes, promoted anaerobiosis. Submergence reduced the haemolymph CO₂ by 50% or more, regardless of the presence of trapped air in their lungs. C. hirtipes and C. carnifex with a retained air bubble lost Na at only 2.6 and 2.3 μmol g^-1 h^-1, respectively, but at 3.9 and 5.0 μmol g^-1 h^-1 without the air bubble. Unidirectional Na uptake in C. hirtipes was only 0.90 μmol g^-1 h^-1 when air was trapped in the lung but 1.74 μmol g^-1 h^-1 when the bubble was removed. In C. carnifex these rates were 1.88 and 2.79 μmol g^-1 h^-1 respectively. C. hirtipes and C. carnifex both trap air within the lung and avoid exposing exchange surfaces to water. There is no large respiratory advantage to expelling the air but there are significant ion regulation cost savings in retaining it.     

Respiration and Circulation in Land Crabs: Novel Variations on the Marine Design

Both the "true" crabs (Brachyura) and hermit crabs (Anomura)include species that show numerous behavioral, morphological,and physiological specializations permitting terrestrial life.This paper examines respiratory and circulatory adaptationsfor air breathing in these land crabs. Respiratory specializationsinclude modification of gas exchange structures for air breathing(gills and elaborated branchial chamber linings), ventilatorymechanisms permitting effective air pumping, an elevated hemolymphoxygen capacity, and a primarily CO₂- rather than O₂- sensitiveventilatory control system. The qualitative aspects of hemolymphoxygen transport and metabolic rate are apparently unchangedfrom that of marine crabs. While the basic cardiovascular morphologyof land crabs appears similar to that of marine forms, thereis considerable elaboration of the vasculature of the branchialchamber lining, which in some species includes a unique doubleportal system. Cardiac output is lower in land crabs (probablyrelated to their higher hemolymph O₂ capacity), but insufficientdata on hemolymph pressures prevent comparisons with marineforms. In general, land crabs have modified (sometimes extensively)existing structures and processes found in their marine relativesrather than evolving structures for terrestrial life de novo.Accordingly, land crabs present a useful model for the evolutionof terrestriality, showing that even subtle anatomical changescan result in the large changes in physiological function necessaryfor the terrestrial invasion.     

Acid-base and electrolyte regulation,and haemolymph gas transport in crayfish,Astacus astacus,exposed to soft,acid water with and without aluminium

Summary Intermoult crayfish (Astacus astacus) were exposed to acid (pH 4), soft water ([Ca⁺⁺]=100 mol·l^–1) in the absence and presence of aluminium (25 mol·l^–1) for variable time periods (up to 21 days) in order to assess the consequences for acid-base and electrolyte balance and haemolymph gas transport. Haemolymph osmolality and concentration of major ions decreased drastically and to a similar extent in acid and acid-aluminium water. Muscle tissue ion concentrations were, however, regulated at an almost constant level. A severe metabolic acidosis was gradually developed, attaining a haemolymph metabolic acid load of 6–7 mequiv·l^–1 after 12–21 days. The acidosis was partially compensated by ventilatory means, with the postbranchial haemolymph PCO₂ decreasing earlier in acidaluminium-exposed than in acid-exposed specimens. Hyperventilation seemed to be a direct acid-base regulatory response, since the rise in postbranchial PCO₂ had only minimal influence on haemolymph O₂ transport. The Bohr effect of Astacus astacus haemocyanin was low (log P₅₀/GdpH=-0.24), and the mean P₅₀ only increased from 15 to 19 mmHg after 21 days of acid exposure. The decrease in O₂ affinity with decreasing pH was accompanied by a decrease in the cooperativity of O₂ binding. The haemolymph haemocyanin concentration was not affected by acid and acid-aluminium exposure, but decreased after 21 days due to starvation. Muscle tissue aluminium concentrations were unaffected, whereas gill tissue concentrations increased in acid-aluminium exposed crayfish, most likely due to accumulation of aluminium on the gill surface. Mortality was low, and an internal hypoxia and lactacidosis was not developed in either of the experimental groups. This suggests that the gas transfer qualities of the chitincovered gills of crayfish are much less sensitive to acid and acid-aluminium stress than the gills of teleost fish.Abbreviations Hc haemocyanin - SO₂ saturation of Hc with O₂ - P ₅₀ oxygen tension of haemolymph at 50% SO₂ - n ₅₀ Hills coelficient around 50% SO₂     

Histopathological changes in gills of the estuarine crab Chasmagnathus granulata (Crustacea-Decapoda) following acute exposure to ammonia

Histopathological effects of ammonia on the gills of the estuarine crab Chasmagnathus granulata (Dana, 1851) were evaluated after acute exposure to ammonia concentrations around LC₅₀ value (17.85 Mm). Disruption of pilaster cells and a subsequent collapse of gill lamellae were the main effects observed. Ephitelial necrosis and hyperplasia were also detected. Significant (P<0.05) increases in pCO₂ and lactate, and significant decreases of pO₂ were detected in the haemolymph of ammonia-exposed crabs. These changes suggest that the observed histopathological damage affected gas exchange, possibly leading to death.     

Respiratory gas transport, metabolic status, and locomotor capacity of the Christmas Island red crab Gecarcoidea natalis assessed in the field with respect to dichotomous seasonal activity levels

Red crabs, Gecarcoidea natalis, exhibit seasonal activity patterns: low activity during the dry season when they shelter in burrows to avoid dehydration, and high activity during the wet season. Red crabs were examined in situ in the rainforest of Christmas Island to determine if there were underlying seasonal differences in the capacity for exercise and associated metabolism. During both seasons, free-ranging (FR) crabs engaged in their normal activities and, together with crabs induced to exercise for 5 min, were sampled for haemolymph and muscle tissue. Respiratory gases in the haemolymph and key metabolites were measured to assess differences in metabolic status of FR and exercised crabs. Actively foraging FR crabs during the wet season exhibited a relative haemolymph hypoxia (2.9 kPa) and accumulated an extra 3 mmol. litre(-1) of CO(2) compared to the relatively inactive FR crabs during the dry season. Wet-season crabs appeared to be in a state of relative respiratory acidosis compared to dry-season animals. This hypercapnia may arise as a consequence of a relative hypoventilation in animals with a relatively higher metabolic rate during the wet season. Oxygenation of pulmonary and arterial haemolymph was similar and remained high after 5 min of exercise, indicating that the gills and lungs functioned similarly in gas exchange in both FR and exercised crabs. During exercise, venous O(2) reserves decreased and red crabs experienced a mixed respiratory/metabolic acidosis. Similar changes, after 5 min of enforced exercise, in metabolite concentrations, pH and respiratory gas status in the haemolymph during both sampling seasons suggest that the crabs maintain similar capacity to increase exercise during the wet and the dry seasons, despite the differences in underlying physiological status. This is important since after prolonged inactivity during the dry season, with the arrival of moonsoonal rains, red crabs must engage in their annual breeding migration.     

Aerial and Aquatic Respiration in the River Crab Potamonautes Warreni Calman with Notes on Gill Structure

Summary

The oxygen consumption rate (?O₂) for Potamonauteus warreni Calman (= Potamon warreni (Calman) kept in 25 °C water was 34,4 μmol 1^?1 O₂ kg^?1 and after 72 hours in 98% R.H. air the rate was 31,9 μmol 1^?1 O₂ kg^?1 min^?1. The ?O₂ values for each of the two groups are not significantly different (P > 0,05). The partial oxygen tension of pre-branchial (v = venous) haemolymph (P_vCO₂) is 15,3 mm Hg in water and 13,0 mm Hg in air); partial carbon dioxide tension of pre-branchial (v) haemolymph (P_vCO₂) is 13,2 mm Hg in water and 13,0 mm Hg in air); the total carbon dioxide concentration in pre-branchial (v) haemolymph (C_vCO₂) tot. is 12,3 mmol 1^?1 in air and 13,9 mmol 1^?1 in water) are not significantly different for the two groups (P > 0,05). The haemolymph pH and the lactate concentration for crabs in water was found to be 7,51 and 0,38 mmol 1^?1 respectively. No significant differences were found in pre-branchial haemolymph oxygen tension, carbon dioxide tension, total carbon dioxide content, haemolymph pH, lactate level, chloride concentration, P₅₀ and haemocyanin-oxygen cooperativity in control crabs kept in water, and experimental crabs held in air for 72 hours. The chloride concentration, (327,0 mmol 1^?1) for crabs kept in water does not differ from that of crabs held in air for 72 hours but is at least 15% higher than the sodium concentration (255 mmol 1^?1) for crabs kept in water. The gill surface area is 520 mm² g^?1 wet body mass; on average 9,2 gill platelets (lamellae) can be found on a gill length of one millimetre. Each lamella is spaced 60–70 μm apart, each with a thickness of 30–40 μm. It is concluded that P. warreni may be described as a truly amphibious fresh-water crab.     

Chlorophyll a Fluorescence Emission,Xanthophyll Cycle Activity,and Net Photosynthetic Rate Responses to Ozone in Some Foliose and Fruticose Lichen Species

The lichens Parmelia quercina, Parmelia sulcata, Evernia prunastri, Hypogymnia physodes, and Anaptychia ciliaris were exposed to ozone (O₃) in controlled environment cuvettes designed to maintain the lichens at optimal physiological activity during exposure. Measurements of gas exchange, modulated chlorophyll (Chl) fluorescence, and pigment analysis were conducted before and after exposure to 300 mm³ (O₃) m^–3, 4 h per d for 14 d. No changes in the efficiency of photosystem 2 (PS2) photochemistry, the reduction state of Q_A, or the electron flow through PS2, measured by Chl fluorescence, were detected in any of the five lichen species studied. Additionally, neither photosynthetic CO₂ assimilation nor xanthophyll cycle activity or photosynthetic pigment concentration were affected by high O₃ concentrations. Thus the studied lichen species have significant capacities to withstand oxidative stresses induced by high concentration of O₃.     

Gas exchange and habitat selection in the aquatic salamanders Necturus maculosus and Cryptobranchus alleganiensis

Summary The standard metabolic rate (SMR) and critical O₂ tension (P_c) of water-breathing mudpuppies and hellbenders were determined at 20° C using open-system respirometry. Both species are metabolic O₂ regulators, although the P_c of hellbenders (90 mmHg) is much higher than that of mudpuppies (40 mmHg). The SMR of the two species in water saturated with air was similar (19.5 and 20.0 l O₂/g·h for Cryptobranchus and Necturus, respectively) and not different from that of salamanders in general. Both species were able to survive for at least 5–11 days in severely hypoxic water (9–10 mmHg) by breathing air, indicating that the lungs are functional accessory respiratory structures.We conclude that hellbenders are restricted to relatively cool and flowing waters because of their limited gas exchange capabilities, particularly with regard to their limited aerobic scope for activity and slow recovery from exercise. Necturus maculosus is much more tolerant of hypoxia, but it is not known if they can inhabit areas were hypoxia is combined with hypercarbia.     

Aerial respiration in the semaphore crab,Heloecius cordiformis,with or without branchial water

• 1.1. Semaphore crabs (Heloecius cordiformis) are active in air at low tide. Their branchial chambers are lined with a vascular epithelium and are expanded above the gills (five pairs) to form air cavities which could function as lungs. Water is continuously circulated over the gills.
• 2.2. The relative contribution made by the gills and lungs to gas exchange in semaphore crabs active in air and circulating branchial water, was determined by measuring oxygen consumption (at 25°C) in crabs with and without branchial water, and in crabs with their lungs subsequently occluded.
• 3.3. Activity levels and VO₂ were unaffected by the absence of branchial water.
• 4.4. With their lungs occluded, VO₂ dropped (on average) by 61% in crabs with branchial water (i.e. gills still functional) and by 81% in crabs without branchial water (gill function impaired).
• 5.5. It is concluded that semaphore crabs are obligate air breathers while active on land, despite carrying water within their branchial chambers. Lung development and gill reduction in land crabs is discussed briefly in relation to “terrestriality”.

     

Ventilatory responses to temperature variation in the fresh water turtle,Mauremys caspica leprosa

A study of lung gas exchange in the fresh water turtle Mauremys caspica leprosa at normal physiological body temperatures (15, 25 and 35 °C) was extended to extreme temperatures (5 and 40 °C) to determine whether the direct relationship between body temperature and ventilatory response found in many lung-breathing ectotherms including other chelonian species was maintained. From 5 to 35 °C the lung ventilation per unit of O₂ uptake and CO₂ removed declined with temperature. Consequently, lung CO₂ partial pressure increased with temperature. Its value was maintained within narrow limits at each thermal constant, suggesting a suitable control throughout the complete ventilatory cycle. At 40 °C the ventilatory response showed the opposite trend. The ratios of ventilation to lung gas exchange increased compared to their values at 35 °C. The impact of this increased breathing-lowering the estimated mean alveolar CO₂ partial pressure-was nevertheless less than expected due to an increase in calculated physiological dead space. This suggests that the relative hyperventilation in response to hyperthermia found in Mauremys caspica leprosa is related to evaporative heat loss.Abbreviations BTPS body temperature, ambient pressure, saturated with water vapour - CTM critical thermal maximum - F_N2 fractional concentration of nitrogen - PA _CO2or PL _CO2 alveolar or lung CO₂ pressure - PA_O2or PL_O2 alveolar or lung O₂ pressure - PI_O2 inspired O₂ pressure - R respiratory exchange ratio - STPD standard temperature, standard pressure, dry - T _a ambient temperature - T _b body temperature - VA alveolar ventilation - VA/VCO₂ relative alveolar ventilation (alveolar ventilation per unit of CO₂ removed) - VO₂ O₂ uptake - VCO₂ CO₂ output - V _D anatomical dead space volume - V _D physiological dead space volume - VE/VO₂ ventilatory equivalent for O₂ - VE pulmonary ventilation or expiratory minute volume - VE/VCO₂ ventilatory equivalent for CO₂ - V _T tidal volume     

Dietary assimilation and the digestive strategy of the omnivorous anomuran land crab<Emphasis Type="Italic"> Birgus latro</Emphasis> (Coenobitidae)

On Christmas Island, Indian Ocean, the diet of robber crabs, Birgus latro (Linnaeus) was generally high in fat, storage polysaccharides or protein and largely comprised fruits, seeds, nuts and animal material. The plant items also contained significant amounts of hemicellulose and cellulose. In laboratory feeding trials, crabs had similar intakes of dry matter when fed artificial diets high in either fat or storage polysaccharide, but intake was lower on a high protein diet. Assimilation coefficients of dry matter (69–74%), carbon (72–81%), nitrogen (76–100%), lipid (71–96%) and storage polysaccharide (89–99%) were high on all three diets. B. latro also assimilated significant amounts of the chitin ingested in the high protein diet ( 93%) and hemicellulose (49.6–65%) and cellulose (16–53%) from the high carbohydrate and high fat diets. This is consistent with the presence of chitinase, hemicellulase and cellulase enzymes in the digestive tract of B. latro. The mean retention time (27.2 h) for a dietary particle marker (⁵⁷Co-labelled microspheres) was longer than measured in leaf-eating land crabs. The feeding strategy of B. latro involves the selection of highly digestible and nutrient-rich plant and animal material and retention of the digesta for a period long enough to allow extensive exploitation of storage carbohydrates, lipids, protein and significant amounts of structural carbohydrates (hemicellulose, cellulose and chitin).Communicated by I.D. Hume     

Haemocyanin oxygen binding and the physiological ecology of a range of talitroidean amphipods (Crustacea)

Summary Haemolymph PO₂ and pH of two amphipod species, Apohyale pugettensis (aquatic) and Megalorchestia californiana (semi-terrestrial) in vivo were examined during immersion and emersion at 15 and 25°C, and also after activity in air at 15°C. For M. californiana arterial O₂ tensions were higher in air than in water. This situation was reversed in A. pugettensis, although all O₂ tensions measured for both species were comparatively high. No arterial-venous PO₂ difference was apparent in the haemolymph of quiescent M. californiana. Haemocyanin (Hc) was 100% saturated in vivo only in the following; A. pugettensis in water (15 and 25°C) and air (15°C), and M. californiana in air (15°C). The Hc of both species becomes important in O₂ transport during activity; under such circumstances the haemolymph of M. californiana delivered more O₂ to the tissues than did that of A. pugettensis, despite the greater O₂ content of the latter. The animals studied here may exhibit a stage (size class?) where cutaneous gas exchange is sufficient for resting aerobic metabolism while specialized respiratory carriers (and respiratory structures) are important in meeting the increased aerobic demands of activity or environmental stress.Abbreviations Hc haemocyanin - PO₂ partial pressure of oxygen     

Enzymes of urate synthesis and catabolism in the Gecarcinid land crab Gecarcoidea natalis

This study investigated the sites of urate synthesis and catabolism in the gecarcinid land crab Gecarcoidea natalis by assaying spongy connective tissue, midgut gland, muscle and gill for xanthine oxidoreductase, the last enzyme involved in urate synthesis, and uricase and urease, the first and last enzymes involved in urate catabolism. The spongy connective tissue and midgut gland of the G. natalis contained activities of xanthine oxidoreductase and were considered to be sites of urate synthesis. The midgut gland had a high activity of xanthine oxidoreductase [(58.87±4.6 (SE) nmol urate produced g^-1 wet wt. tissue min^-1], 2.7 times the xanthine oxidoreductase activity contained within the spongy connective tissue, and was thought to be the main site of urate synthesis. Xanthine dehydrogenase (EC 1.1.1.204) was the only form of xanthine oxidoreductase detected within the tissues. Its presence means that the cost of synthesising urate de novo is relatively small (between 1 and 3 ATP). Uricase (EC 1.7.3.3) and urease (EC 3.5.1.5) activities were present in the tissues of G. natalis. Spongy connective tissue contained the highest activities of uricase [48.44±4.29 (SE) nmol urate consumed g^-1 wet wt. tissue min^-1] while the highest activities of urease [365.31±37.21 (SE) nmol urate consumed g^-1 wet wt tissue min^-1] were contained within the gills. From this evidence it is clear that G. natalis possesses the uricolytic pathway and hence the ability to catabolise urate, and urate catabolism is begun at the site of urate storage, the spongy connective tissue, and is completed at the gills. As the gills are the site of ammonia excretion in this species the ammonia produced from the catabolism of urate is probably excreted. The urate deposits within the body of G. natalis may be involved in temporary storage of nitrogenous wastes.     

Changes in Na+/K+-ATPase activity, unsaturated fatty acids and metallothioneins in gills of the shore crab Carcinus aestuarii after dilute seawater acclimation

We have assessed the activity of Na⁺/K⁺-ATPase, cAMP, free fatty acids (FFA) and metallothionein (MT) in the posterior gills of the brackish water shore crab Carcinus aestuarii during acclimation to 10 ppt dilute seawater (DSW). Following 3–18 days acclimation in DSW specific activity of Na⁺/K⁺-ATPase in native gill homogenates and partially purified membrane vesicles was progressively increased, from 1.7- to 3.9-fold. After short-term acclimation of crabs in DSW with added sucrose to make media isosmotic with the haemolymph the specific Na⁺/K⁺-ATPase activity in homogenates was not increased, relative to SW enzyme activity. Moreover, hyposmotic conditions led to depletion of cAMP in gills.In partially purified membrane vesicles isolated from posterior gills, fatty acids with compositions 16:0, 18:0, 18:1, 20:4 and 20:5 dominated in both SW- and DSW-acclimated Carcinus. During a year in which the metabolic activity of crabs was increased, the arachidonic/linoleic acids ratio (ARA/LA) for DSW-acclimated crabs was markedly increased relative to that in SW. Increased Na⁺ + K⁺-ATPase activity under hyposmotic stress may be modulated at least partially by the changed proportion of fatty acids in the purified membranes of posterior gills. Long-term acclimation of shore crabs to DSW resulted in a 2.6-fold increase in cytosolic metallothionein (MT) content in posterior gills over those in SW crabs. Assuming an antioxidant role of MT associated with intracellular zinc partitioning, the observed MT induction in posterior gills may be considered an adaptive response of C. aestuarii to hyposmotic stress.     

Characterization of red sternum syndrome in mud crab farms from Thailand

Samples of abnormal mud crabs, Scylla serrata (Forsk?l, 1755) (Decapoda: Portunidae), were collected from crab farms in Samutsongkhram Province, Thailand. These crabs had hard carapaces, red chelipeds and joints, pale hepatopancreas, gills, and soft muscles. They were almost immobile and finally died. The haemolymph revealed three stages of the syndrome, namely orange, orange-white, and milky-white in colors. The haemolymph, integument, hepatopancreas, gills, abdominal and claw muscle, stomach, and heart were dissected and histologically examined using transmission and scanning electron microscopy. Closer examinations found infection with rod-, curve rod-, or coccus-shape bacteria with thin and thick cell walls in all investigated organs and haemolymph. Isolation of the microorganisms from the infected tissues of red sternum syndrome crabs resulted in five types of bacteria. No microorganism growth was observed in normal crabs. Interestingly, the types of isolated bacteria can be classified according to the severity of the disease. Additionally, the degree of bacterial infection found was consistent with the stages of the disease. It was postulated that the bacteria entered the crabs via the gills, and then migrated through circulating haemocytes, before reaching the internal organs.     

Bimodal respiration, water balance and acid-base regulation in a high-shore crab, Cyclograpsus lavauxi H. Milne Edwards

Cyclograpsus lavauxi H. Milne Edwards, 1853 occurs under boulders near the littoral fringe, where it is wetted only briefly, if at all, in each tide. Mean water content of sea-water equilibrated crabs was 62.3 % of body weight. On warm, windy days crabs on the shore lose > 17.5% of body water. Water loss for crabs of 1.6 g mean body weight was 0.29 % water content · h ⁻¹ · mm Hg saturation deficit ⁻¹, which is similar to that of species occupying positions lower on the shore. C. lavauxi is, however, able to tolerate comparatively greater losses of body water (up to 36%) compared with species from lower shore levels.

As the gills dry, the lamellae separate into regular clumps which may help to maintain gas exchange. Resting oxygen consumption (V_o2) after 3–5 h settlement was similar in water and air. During enforced activity, V_o2 increased by factors of 5.3 in water and 2.6 in air at a standardized body weight of 1.5 g, indicating an appreciable aerobic scope in both media. Loss of up to 16% of body water did not depress resting aerial V_o2 or aerobic scope.

In resting crabs there was no change in haemolymph pH after 24 h of either immersion or emersion, but haemolymph [Ca²⁺], [HCO⁻₃] + [CO²⁻₃], and calculated P_co2 all rose in air relative to aquatic values. These results suggest that cuticular or other endogenous CaCO₃ is mobilized to compensate the respiratory acidosis in air. The implications of such physiological properties for aquatic and aerial activity on the shore are discussed.     

Physiological responses to emersion in the intertidal green crab,Carcinus maenas (L.)

The green shore crab, Carcinus maenas, undergoes on average 6?h periods of emersion during each low-tide cycle during the summer months. Under those conditions, the crab is cut off from its normal water environment and is exposed to potential stress from a suite of environmental and physiological changes: dehydration, compromised gas exchange and resultant internal hypoxia and hypercapnia, thermal stress, and ammonia toxicity. This study examined the comprehensive responses of the green crab in water and to a 6?h emersion period laboratory simulation of a tidal cycle followed by a 1?h re-immersion period, measuring indicators of dehydration, hemolymph osmolality, oxygen uptake, hemolymph acid–base status, heart and ventilatory rate, and hemolymph ammonia and ammonia excretion. Green crabs showed physiological responses of varying magnitude to 6?h of emersion. Individuals were found in the field exclusively under rocks and large clumps of seaweed where temperatures were approximately half those of exposed surfaces and relative humidity was about twice as high as ambient air. During emersion, crabs lost less than 5% of their wet weight, and hemolymph osmolality did not increase significantly. Oxygen uptake continued in air at about 50% of the control, aquatic values; and the gills continued to be ventilated by the scaphognathite, albeit at lower rates. Hemolymph lactate concentrations increased, indicating a shift to a greater reliance on anaerobic metabolism to support energetic needs. A slight acidosis developed in the hemolymph after 1?h of emersion, but it did not increase thereafter. Ammonia concentrations in the hemolymph were unchanged, as ammonia was volatilized by the gills and excreted into the air as NH₃ gas. These results show that the green crab copes with emersion by seeking refuge in microhabitats that mitigate the changes in the physical parameters of intertidal emersion. Physiologically, desiccation is avoided, cardio-respiratory processes are maintained at reduced levels, and hemolymph acid–base balance is minimally affected. Ammonia toxicity appears to be avoided by a shift to excreting NH₃ gas directly or indirectly to air.     

Resistance of Island Rainforest to Invasion by Alien Plants: Influence of Microhabitat and Herbivory on Seedling Performance

Experiments have promise in determining mechanisms by which communities resist invasion. Growth and survivorship of transplanted seedlings of introduced tree species (Leucaena leucocephala, Muntingia calabura, Adenanthera pavonia, and Clausena excavata) were used to assess abiotic (light regime) and biotic resistance (herbivory) to invasion of rainforest on Christmas Island (Indian Ocean). At four sites, seedlings were transplanted into the forest edge along roadside verges and into adjacent intact forest; half were caged to prevent access by the dominant seedling consumer, the red land crab (Gecarcoidea natalis). Red crab densities did not differ between roadside and forest plots. Red crabs initially reduced survivorship of Leucaena in both edge and interior plots but virtually all seedlings in the forest interior were dead after 41 weeks. Survivorship of Muntingia was also initially reduced by crabs in the forest edge, but again, all but one seedling died by the end of the experiment. Seedlings of Adenanthera and Clausena fared much better overall, surviving well in both locations. Red crabs had no overall effect on Adenanthera survival but significantly reduced survival of Clausena in forest plots. For both species, seedling performance was greater in the forest edge than in the forest interior. Red crabs had no effect on height increment for either species; however, for Clausena, red crabs reduced seedling mass in the forest interior. Both Adenanthera and Clausena were able to persist in the intact forest. Clausena is now actively invading intact rainforest, but Adenanthera appears dispersal-limited. Resistance factors in intact forest appear hierarchical: biotic resistance afforded by land crabs can impede establishment of some plant invaders but seedling responses to abiotic factors (e.g., the light regime) largely overwhelm its effect. Together, these two community attributes are likely to restrict the range of plant invaders to a small suite of species that can successfully establish in intact rainforest on the island. However, increased propagule pressure from a variety of shade-tolerant species and further declines in the abundance of the dominant seedling consumer may lead to increasing invasion success in this island rainforest. This revised version was published online in July 2006 with corrections to the Cover Date.     

Physiological consequences of prolonged aerial exposure in the American eel,Anguilla rostrata: blood respiratory and acid-base status

Summary The physiological consequences of prolonged air-exposure on blood respiratory and acid-base properties were examined in the American eel (Anguilla rostrata). Eels displayed a low capacity for aerial gas transfer as indicated by pronounced increases and decreases in arterial CO₂ and O₂ tensions, respectively. The increase in arterial CO₂ tension contributed to severe extracellular acidosis. The decrease in arterial O₂ tension, combined with a marked reduction in red blood cell pH and concomitant Bohr and Root effects, caused arterial O₂ content to decline to levels that were insufficient to support metabolic requirements, aerobically. Consequently, the rate of anaerobic glycolysis increased during air-exposure as suggested by a gradual elevation of blood lactate levels after 12 h. Increased anaerobic glycolysis and associated ATP hydrolysis and/or degradation of internal ATP stores further depressed blood pH as metabolic acid, produced by these processes, entered the circulation. Unlike other fishes previously examined, red blood cell pH was not regulated preferentially during the extracellular acidosis but simply conformed to the in vitro relationship between red blood cell and whole blood pH. Although capable of surviving prolonged air-exposure, the results demonstrate nevertheless and perhaps not surprisingly that eels, unlike true amphibious fishes that utilize gills or buccal epithelia for gas transfer, are not particularly well-adapted for gas exchange in air but do display an unusual tolerance to hypoxemia.Symbols and abbreviations B buffer value - Hct hematocrit - RBC red blood cell