Elasmobranch pericardial function. 1. Pericardial pressures are not always negative

Acute studies have led to the generalization that negative pericardial pressure is necessary for optimal cardiac function in elasmobranchs. We chronically instrumented horn sharks with pericardial catheters to test the hypothesis that ejection of pericardial fluid through the pericardioperitoneal canal (PPC) during routine handling could have accounted in part for previous measurements of exclusively negative pressures (–0.3 to –9.1 cm H₂O) in elasmobranchs. Maximum and minimum pericardial pressures measured immediately following routine handling (acute pressures) were more negative than those measured in resting horn sharks at intervals from 1 to 27 days following handling (chronic pressures). Chronic pericardial pulse pressure was less than acute. Entirely positive pericardial pressures were observed on occasion. Handling of chronically catheterized horn sharks resulted in ejection of 21 per cent (range=10–26, n=5) of the initial pericardial fluid volume through the PPC and reduced pericardial pressure. Operating pericardial fluid volume of horn sharks averaged 2.0 ml.kg^–1 (range=1.6–2.6, n=9). The PPC opened after 4.3±0.2 ml.kg^–1 (x±S.E.) of elasmobranch saline had been slowly infused into the pericardium, corresponding to an average pressure of 1.3±0.2 cm H₂O (n=10). The presence of the PPC plus a comparatively large pericardial fluid volume allows horn sharks to regulate pericardial pressure. Our analysis of pericardial pulse pressure, which can be an index of cardiac activity, suggests in contrast to previous studies that the elasmobranch heart can have relatively high stroke volumes at pericardial pressures near ambient. Thus, for venous return in resting or even moderately active elasmobranchs, it is more important that pericardial pressure be pulsatile than at a mean level which is negative.     

Elasmobranch pericardial function. 3. The pericardioperitoneal canal in the horn sharkHeterodontus francisci

The pericardial and peritoneal spaces of elasmobranch fishes are connected by the pericardioperitoneal canal (PPC), which allows pericardial fluid to escape when pressures exceed 0.1–0.3 kPA. Using the horn shark (Heterodontus francisci), we tested the hypothesis that the PPC functions to increase cardiac stroke volume by lowering pericardial pressure during activity. We also assessed the role of the PPC during coughing, feeding, or burst swimming and examined the effects of PPC occlusion. Increases in heart size were not prevented following augmented venous return in sharks with undisturbed or occluded PCP, evidence that argues that pericardial fluid loss through the PPC is a cause of increased cardiac stroke volume and not the result. Coughs, feeding, and burst swimming led to discharge of pericardial fluid. Chronic PPC occlusion resulted in an increased pericardial pressure, fluid volume, and frequency of coughing, and a decreased survival time compared to shams. Thus, in the horn shark the PPC likely compensates for constraints that may be imposed by the pericardium, provides a route for pericardial drainage, and regulates cardiac stroke volume during periods of activity.     

Regulation of cardiac function in the horn shark by changes in pericardial fluid volume mediated through the pericardioperitoneal canal

The California horn shark (Heterodontus francisci), instrumented with a pericardial catheter and a ventral-aortic flow probe, was studied to determine the effect of complete pericardial chamber evacuation on the time course for restitution of pericardial fluid-volume and pressure, and the effects of both fluid removal and its restitution on cardiac output. Prior to evacuation, pericardial pressure was –0.02±0.02 kPa, and cardiac output was 18±2 ml min^–1 kg^–1. Evacuation reduced pericardial pressure to –0.73±0.14 kPa, and increased cardiac output to 23±4 ml min^–1 kg^–1. The time course for restoration of post-evacuation pressure is described by a non-linear asymptotic function. A large percentage of the pericardial pressure and volume recovery occurred within the first hour, while, complete restoration of pre-withdrawal conditions required about 11 h. Pericardial pressure-volume relationships, determined by incremental infusion of small volumes of elasmobranch saline into the pericardium, confirm previous findings that the operating pericardial pressure in the horn shark is at or near ambient pressure and that both pericardial fluid volume and cardiac stroke volume influence horn shark pericardial pressure.     

Plasticity in the diel vertical movement of two pelagic predators (Prionace glauca and Alopias vulpinus) in the southeastern Indian Ocean

Management and conservation of marine predator species relies on a fundamental knowledge of their movements and behaviour. Pop‐up satellite archival tags were used to investigate the vertical movement patterns of five blue sharks (Prionace glauca) and one thresher shark (Alopias vulpinus) within the southeastern Indian Ocean. Sections of similar depth distribution, identified using a split moving window analysis, were investigated in relation to the thermal structure of the water column and activity rates. Minimum horizontal displacement of between 66 and 5,187 km for blue sharks and 16 km for the thresher shark were recorded over 863 tracking days. Maximum depths ranged from 540 to 807 m for blue sharks and 144 m for the thresher shark. All sharks displayed plasticity in their depth distribution, with diel vertical movements and surface‐oriented movements the two most common patterns. Diel movement of prey is the most likely explanation for diel vertical movements of thresher and blue sharks. This study has improved our understanding of the vertical movement patterns of these pelagic predators and the relationship between their depth distribution, temperature, and activity.     

Molecular markers of cancer in cartilaginous fish: immunocytochemical study of PCNA, p-53, myc and ras expression in neoplastic and hyperplastic tissues from free ranging blue sharks, Prionace glauca (L.)

Archival formalin-fixed tissues from wild-caught adult blue sharks, Prionace glauca (L.), were used for immunocytochemical detection of proliferating cell nuclear antigen (PCNA), two oncoproteins from the oncogenes c-myc and pan-ras, and a protein product from the tumour suppressor gene p-53. All sharks were caught during summer months between 2000 and 2006 by recreational fishermen off the USA coast in the northwestern Atlantic. The sharks were necropsied on landing and selected organ samples were collected into elasmobranch formalin and processed for paraffin embedding and light microscopy. Paraffin-embedded sections from collected tissue were both stained with haematoxylin and eosin and processed by immunocytochemical techniques using antibodies raised against the PCNA, p-ras, c-myc and p-53 proteins. The lesions examined in this study included two well differentiated adenomatous gastric polyps, a testicular capsular mesothelioma, a gingival fibropapilloma with elements of ameloblastoma, three liver tumours, two pericardial fibropapillomas and six cases of proliferative serositis (pericarditis and peritonitis). Normal and hyperplastic tissues from blue sharks, and human neoplastic tissues served as negative and positive controls, respectively. We detected upregulation of PCNA in many neoplastic, one dysplastic and in some hyperplastic lesions, and positive p-ras and c-myc signals in some of the neoplastic lesions. None of the examined tissues showed positive p-53 signalling. This is the first literature report on immunocytochemical detection of molecular markers of cancer in sharks and in fish of the class Chondrichthyes.     

Tissue‐specific stable isotope ratios of shortfin mako (Isurus oxyrinchus) and white (Carcharodon carcharias) sharks as indicators of size‐based differences in foraging habitat and trophic level

We evaluated whether the trophic level and feeding grounds (coastal versus offshore) of white and shortfin mako sharks captured in the northern Mexican Pacific varied as a function of size using carbon (δ¹³C values) and nitrogen (δ¹⁵N values) stable isotope analysis of soft tissues. A step increase in δ¹⁵N values was observed at approximately 85 cm TL and 150–170 cm TL for shortfin mako and white sharks, respectively, consistent with the dilution of the maternal isotopic signal following the incorporation of exogenous food. Shortfin mako and white shark muscle tissue and whole blood were enriched in ¹⁵N as a function of size, which is consistent with increasing trophic level, but this pattern may be confounded by differences in habitat. To infer differences in foraging habitat throughout life, we compared isotope ratios of liver and blood fractions with those of white muscle tissue; these tissues have relatively faster and slower isotope turnover rates, respectively. Whole blood and muscle δ¹³C values of small white sharks (<200 cm total length) caught near the coast suggested benthic foraging. Very limited differences between δ¹³C values of muscle tissue and whole blood from small and medium mako (<196 cm TL) and small white sharks indicated equilibrium to local feeding grounds. In contrast, differences in the δ¹³C values of liver tissue from small and medium mako and small white sharks indicated changes in diet or feeding grounds. The δ¹³C values of plasma and muscle tissue of large white sharks (>350 cm total length) caught offshore indicate feeding in coastal and oceanic areas and are consistent with recent migration to pelagic feeding grounds. Our study provides insight into size‐based habitat use and migration patterns of two species of migratory sharks for which there is limited information, particularly in their nursery grounds in Mexican waters.     

Serum steroid concentrations and development of reproductive organs during puberty in male bonnethead sharks, Sphyrna tiburo

Puberty is a critical, hormone-mediated event during which an animal acquires the ability to breed and propagate. Despite the importance of this stage in animal reproduction, little is known regarding the physiological factors that regulate and/or accompany puberty in several vertebrate groups including elasmobranchs. To address the need for such information, the present study investigated morphological and hormonal changes that occur during puberty in male bonnethead sharks (Sphyrna tiburo). Serial changes in development of claspers, paired copulatory organs in male elasmobranchs, and serum steroid concentrations during puberty were evaluated in captive-held male S. tiburo. Captive-animal studies were supplemented by observations on gonadal development, gonaduct morphology, and serum steroid concentrations in feral, peripubertal male S. tiburo. Changes in size and histological architecture of testes and gonaducts of peripubertal sharks mirrored the seasonal progression of events that occur in these structures in mature males. Claspers grew in length continuously during puberty, but sharks did not reach functional maturity until a short period before mating activity commences in the mature population. Clasper growth appeared to be strictly regulated in S. tiburo, perhaps to ensure growth of these organs to sizes deemed critical for reproductive success. Serum concentrations of testosterone, dihydrotestosterone, progesterone, and 17β-estradiol increased in both captive and feral sharks during pubertal development, and may be associated with development of the gonads and gonaducts. Differences in hormone profiles of captive and feral sharks were observed at certain periods during puberty, but their origin remains unclear. This revised version was published online in July 2006 with corrections to the Cover Date.     

热带大西洋金枪鱼延绳钓兼捕鲨鱼种类组成和渔获率及其与表温的关系

根据2007年12月~2008年3月采集的热带大西洋(05°37′~12°01′N、29°00′~36°51′W)金枪鱼延绳钓渔获物数据,分析了金枪鱼延绳钓兼捕鲨鱼的种类组成、渔获量、渔获率及其与表温的关系。本次调查共捕获鲨鱼8种,隶属3目7科7属,总渔获尾数为633 ind,总渔获量达26 837.4 kg,其中大青鲨为主要兼捕种类。各种鲨鱼渔获率平均值在0.003~1.524 ind/1 000 hooks之间,其中大青鲨最高,其值为1.524 ind/1 000 hooks,大眼砂锥齿鲨最低,其值为0.003 ind/1 000 hooks。各种鲨鱼渔获率月变化不明显(ANOVA,P=0.901)。鲨鱼总渔获率和大青鲨渔获率与表温都呈显著性负相关。大青鲨主要出现渔场的表温范围为24.6~25.8℃。     

You can swim but you can't hide: the global status and conservation of oceanic pelagic sharks and rays

• 1. Fishing spans all oceans and the impact on ocean predators such as sharks and rays is largely unknown. A lack of data and complicated jurisdictional issues present particular challenges for assessing and conserving high seas biodiversity. It is clear, however, that pelagic sharks and rays of the open ocean are subject to high and often unrestricted levels of mortality from bycatch and targeted fisheries for their meat and valuable fins.
• 2. These species exhibit a wide range of life‐history characteristics, but many have relatively low productivity and consequently relatively high intrinsic vulnerability to over‐exploitation. The IUCN — World Conservation Union Red List criteria were used to assess the global status of 21 oceanic pelagic shark and ray species.
• 3. Three‐quarters (16) of these species are classified as Threatened or Near Threatened. Eleven species are globally threatened with higher risk of extinction: the giant devilray is Endangered, ten sharks are Vulnerable and a further five species are Near Threatened. Threat status depends on the interaction between the demographic resilience of the species and intensity of fisheries exploitation.
• 4. 4. Most threatened species, like the shortfin mako shark, have low population increase rates and suffer high fishing mortality throughout their range. Species with a lower risk of extinction have either fast, resilient life histories (e.g. pelagic stingray) or are species with slow, less resilient life histories but subject to fisheries management (e.g. salmon shark).
• 5. 5. Recommendations, including implementing and enforcing finning bans and catch limits, are made to guide effective conservation and management of these sharks and rays.

Copyright © 2008 John Wiley & Sons, Ltd.     

The effectiveness of Shark-Management-Alert-in-Real-Time (SMART) drumlines as a tool for catching white sharks,Carcharodon carcharias,off coastal New South Wales,Australia

White (Carcharodon carcharias L.), bull (Carcharhinus leucas, Müller & Henle) and tiger (Galeocerdo cuvier, Péron & Lesueur) sharks are the primary species responsible for unprovoked shark bites. Historically, management practices were based on culling “target” shark species (i.e. white, bull and tiger sharks), which resulted in high levels of bycatch and mortality. Shark-Management-Alert-in-Real-Time (SMART) drumlines were trialled in New South Wales, Australia, aiming to optimise the capture of target shark species while minimising bycatch and mortality. Target shark species accounted for 70% of the total catch, with white sharks contributing 298 of the 350 sharks that were caught. Four animals died, and bycatch consisted of 13 species including two threatened species. Generalised linear mixed models (GLMMs) revealed a significant spatial, temporal, environmental and gear influence on white shark catch rates. SMART drumlines are a useful tool for catching target shark species with low bycatch and mortality relative to historical bather protection methods.     

Fisheries and conservation assessment of sharks in Pacific Panama

1. In Panama, shark fisheries were initially developed in the 1980s and progressively increased in production in the 1990s mainly due to the high demand for shark fins and meat from the international Asian market. Since then, and despite the exploitation rate (average 3,514 t year^–1) and endangered status of some species, shark fisheries have seldom been studied, and official statistics are general or incomplete and not suitable for the development of appropriate conservation and management strategies.
2. To understand the dynamics of shark fisheries in Panama, field surveys were conducted between 2007 and 2009 at several landing ports of small‐scale and industrial fisheries, at fish processing plants and on‐board fishing vessels in Pacific Panama, where most of the fishing vessels of the country operate.
3. In general, it was found that the artisanal and industrial fisheries of the Pacific coast of Panama regularly exploit at least 18 species of sharks, which are also being exploited by neighbouring countries in the eastern Pacific, suggesting the importance of coordinated conservation initiatives across the multiple jurisdictions. A large number of the individuals caught were immature, implying a certain level of impact on recruitment rates. This pattern was particularly evident in species such as Sphyrna lewini, for which immature individuals represented at least 99% and 63% of the total catch by small‐scale and industrial fisheries, respectively. Catch per unit of effort analyses showed that Carcharhinus and Sphyrna species were the most exploited (representing ~80% of the catches) by industrial fisheries in Panama between 2006 and 2009, suggesting that fishery management should provide special attention to these groups.
4. It is expected that the information presented here provides a baseline to develop new regulations, including the implementation of annual quotas and fishing seasons and the protection of nursery areas, for the long‐term sustainability and conservation of sharks in Panama.

     

Temperature effects on the blood oxygen affinity in sharks

In fish, regional endothermy (i.e., the capacity to significantly elevate tissue temperatures above ambient via vascular heat exchangers) in the red swimming muscles (RM) has evolved only in a few marine groups (e.g., sharks: Lamnidae, Alopiidae, and teleosts Scombridae). Within these taxa, several species have also been shown to share similar physiological adaptations to enhance oxygen delivery to the working tissues. Although the hemoglobin (Hb) of most fish has a decreased affinity for oxygen with an increase in temperature, some regionally endothermic teleosts (e.g., tunas) have evolved Hbs that have a very low or even an increased affinity for oxygen with an increase in temperature. For sharks, however, blood oxygen affinities remain largely unknown. We examined the effects of temperature on the blood oxygen affinity in two pelagic species (the regionally endothermic shortfin mako shark and the ectothermic blue shark) at 15, 20, and 25 °C, and two coastal ectothermic species (the leopard shark and brown smooth-hound shark) at 10, 15, and 20 °C. Relative to the effects of temperature on the blood oxygen affinity of ectothermic sharks (e.g., blue shark), shortfin mako shark blood was less affected by an increase in temperature, a scenario similar to that documented in some of the tunas. In the shortfin mako shark, this may act to prevent premature oxygen dissociation from Hb as the blood is warmed during its passage through vascular heat exchangers. Even though the shortfin mako shark and blue shark occupy a similar niche, the effects of temperature on blood oxygen affinity in the latter more closely resembled that of the blood in the two coastal shark species examined in this study. The only exception was a small, reverse temperature effect (an increase in blood oxygen affinity with temperature) observed during the warming of the leopard shark blood under simulated arterial conditions, a finding that is likely related to the estuarine ecology of this species. Taken together, we found species-specific differences in how temperature affects blood oxygen affinity in sharks, with some similarities between the regionally endothermic sharks and several regionally endothermic teleost fishes.     

Performance of the heat of the hagfish,Eptatretus cirrhatus

The maximum power output of isolated perfused ventricles of the hafish (Eptatretus cirrhatus) averaged 0.367±0.031 mW g^–1 (n=9), considerably high than estimates for the heart of the Atlantic hagfish (Myxine glutinosa). Maximal minute volumes averaged 21.55±1.28 ml min^–1kg^–1, with a mean stroke volume of 0.71±0.14 ml kg^–1 body weight, values which are similar to those reported for many teleost and elasmobranch hearts.Ventricular output showed the characteristic dependence upon atrial filling pressure up to an optimum filling pressure ofc. 4 mm Hg. At output pressures exceeding 14 mm Hg the stroke volume and power output fell sharply. At these afterloads, the ventral aorta remained distended following semilunar valve closure and so the volume of fluid ejected on ventricular systole was reduced. There was little change in the frequency of the heart as either input or output pressures were varied.     

Data-mining social media platforms highlights conservation action for the Mediterranean Critically Endangered blue shark Prionace glauca

1. The Mediterranean Sea represents an area of elevated risk of extinction for sharks, where data deficiency is a pervasive problem.
2. To compensate for such a paucity of information, this study investigated the use of social media content as a complementary approach to evaluate the distribution and habitat use of the Critically Endangered blue shark Prionace glauca in coastal waters, as well as public perceptions of the sharks.
3. Through social media data mining a total of 146 records, comprising 158 individual blue sharks approaching Italian coastal waters, have been recorded from 2011 to 2020.
4. This study revealed that, over the past decade, blue sharks regularly visited Italian coastal habitats for extended periods of time. Differences in the temporal distribution of blue sharks by sex and size appear to be linked to reproductive activity. The higher number of adult females approaching the shore in spring and the increase in young-of-the-year (YOY) sightings in the following months possibly indicate parturition in coastal waters. Spatial analyses also showed that certain Italian coastal areas, such as those in Calabria and Puglia, were preferred coastal habitats for this species.
5. Results also indicate that social media platforms can be considered an ever-growing source of data on wildlife, which can shed light on the occurrence and distribution of endangered shark species in poorly known habitats. Furthermore, social media platforms should be used for awareness campaigns to educate the public, as this study showed that negative reactions to shark encounters remain widespread.

     

Benefits of marine protected areas for tropical coastal sharks

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Salmon shark connections: North Pacific climate change,indirect fisheries effects,or just variability?

A variety of changes are occurring in the ecosystems of the North Pacific Ocean and Bering Sea, but information about the mechanisms of change has been relatively limited, due in part to the region’s remoteness and subarctic conditions. Any number of ecosystem components or indicators could be used to exemplify this dilemma, but here we point to the salmon shark (Lamna ditropis, Lamnidae) as an example of a species that can potentially mediate considerable ecosystem change due to its high trophic level, but for which some basic information is lacking despite attracting some interesting research and widespread rumours and anecdotal evidence of increased abundance. Increases in the abundance of sharks such as salmon sharks in this region during the 1990s, if true, may help explain other observed changes such as declines in ocean survival rates of Pacific salmon (Oncorhynchus spp., Salmonidae) in the region and declines in some wild salmon stocks. Mechanisms that could cause salmon shark increases in Alaskan coastal waters include: (i) increases in sea temperature since the 1970s allowing distributional shifts of this species and changes in the abundance or distribution of their prey; (ii) the 1992 banning of high seas drift gillnets; and (iii) indirect fisheries effects such as competitive release of salmon sharks in the North Pacific transition region and towards the more southern geographic extent of their annual migration as the result of fishery‐related reductions in blue sharks (Prionace glauca, Carcharhinidae) and other pelagic predators. The relative plausibility of these alternative explanations can be evaluated using combinations of existing ecosystem models and empirical research and monitoring programmes including local and indigenous observations.     

Blood pressure and heart rate during tonic immobility in the black tipped reef shark,Carcharhinus melanoptera

Tonic immobility was induced in black tipped reef sharks (Carcharhinus melanoptera) and heart rate and ventral aortic blood pressure recorded. Without branchial irrigation, tonic immobility was correlated with a significant depression in blood pressure and heart rate irrespective of the sharks being in air or in water. Tonic immobility with branchial irrigation resulted in a significant increase in blood pressure in sharks in air, but not in water. Heart rate was unchanged when the gills were irrigated. Intra-arterial injections of atropine abolished the bradycardia and blood pressure rise associated with tonic immobility. We conclude that, during tonic immobility, sharks are able to receive afferent information from the ventilatory system and make appropriate responses via the vagus nerve.     

Dynamic use of coastal areas by bull sharks and the conciliation of conservation and management of negative human–wildlife interactions

1. Knowledge about spatial and temporal variability in the distribution and abundance of predators is necessary to adapt measures to mitigate human–wildlife interactions.
2. Acoustic telemetry and network analyses were used to investigate the spatial ecology of bull sharks, the species responsible for most shark bites in Reunion Island, one of the world's shark bite hotspots.
3. The west coast of the island was not used uniformly by every individual, with size predicting the movements of sharks along the coast.
4. Node-based metrics – closeness, node strength, and cumulated continuous residency times – derived from up to 181 monthly movement networks from 20 individuals, revealed that smaller sharks (<250 cm total length) primarily used the south-west coast while larger individuals spent most of their time in the northern region with regular visits to multiple areas along the coast.
5. This study provides essential knowledge on bull shark behaviour and central areas used at different periods of the year, which correlates well with the dynamics of observed shark bites. Our approach provides a non-invasive alternative to help predicting and anticipating human–shark conflicts and avoid shark culling programmes detrimental to the conservation of large predators such as sharks.

     

大西洋中上层鲨鱼资源状况的初步探讨

据1994—2001年4个航次在金枪鱼延绳钓渔船对热带大洋性中上层鲨鱼资源进行调查，并分析国际大西洋金枪鱼资源保护委员会提供的关于中上层鲨鱼的渔业数据。结果表明：延绳钓渔业共兼捕13种鲨鱼，其中，尖吻鲭鲨和大青鲨是优势种类。尖吻鲭鲨在第1、第2和第4航次的CPUE分别为每千钩0．3502、0．1754和0．0642尾，呈下降趋势。大青鲨在第1～3航次的的CPUE达到每千钩5～7尾，而在第4航次下降为每千钩0．8尾。研究报告了中国金枪鱼船队自1993年开始在大西洋兼捕尖吻鲭鲨和大青鲨的渔获量，并根据捕捞死亡系数和大西洋总渔获量数据，初步探讨了尖吻鲭鲨和大青鲨年平均资源量，影响CPUE的因素和大西洋中上层资源状况。     

盐度对瘤背石磺不同部位Na /K -ATP酶活性、围心腔液和腹腔液渗透压及离子含量的影响

为了研究盐度对瘤背石磺不同部位Na+/K+-ATP酶活性、围心腔液和腹腔液渗透压及离子含量(Na+、K+和Cl-)的影响,选取健康的瘤背石磺[湿重(10.24±2.16)g]600只,在不同盐度条件下(5、15、25、35和45)采用模拟生态环境的方式饲养30 d后测定各项指标.结果显示:(1)瘤背石磺腹腔液和围心腔液的盐度和渗透压均随着环境盐度的升高而增高,腹腔液的盐度和渗透压的变动幅度分别为25～ 43和514 ～ 796 mOsm/kg,其增高幅度均大于围心腔液(分别为23 ～ 28和502～ 726 mOsm/kg);(2)围心腔液中Na+、Cl-和K+含量随环境盐度的变化具有一致性,均呈先上升后下降的趋势,且各离子浓度均在盐度25时达到最大值,分别为295.50、322.50和5.15 mmol/L,但盐度对Na+和Cl-含量有显著影响,对K+含量无显著影响.(3)瘤背石磺不同部位的Na +/K+-ATP酶活性具有组织特异性,贲门胃、肠、表皮、肌肉和围心腔膜等部位的Na +/K+-ATP酶活性较高,同时,盐度激活不同部位的Na +/K+-ATP酶活性具有显著差异性,肌肉和贲门胃的Na +/K+-ATP酶在低盐度时活性较高,而肠、肾及围心腔膜等在高盐度时Na+/K+-ATP酶被激活,酶活性较高.研究表明:盐度能显著影响瘤背石磺腹腔液和围心腔液的渗透压和离子浓度以及各器官的Na+/K+-ATP酶活性,贲门胃、肠和肌肉等均是其离子转运和渗透调节的主要部位,具有较强的渗透压调节能力.