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1.
Cardiac activity and anaerobic metabolism were analyzed in zebrafish larvae raised under normoxia (PO(2) = 20 kPa) and under chronic hypoxia (PO(2) = 10 kPa) at three different temperatures (25, 28, and 31 degrees C). Heart rate increased with development and with temperature. Under normoxia, cardiac output increased significantly at high temperature (31 degrees C), but not at 28 or at 25 degrees C. Under chronic hypoxia, however, heart rate as well as cardiac output increased at all temperatures in larvae at about hatching time or shortly thereafter. Cardiac activity of larvae raised for 2 wk after fertilization with a reduced hemoglobin oxygen-carrying capacity in their blood (hypoxemia; due to the presence of CO or of phenylhydrazine in the incubation water) was not different from control animals. Whole body lactate content of these animals did not increase. Thus there was no indication of a stimulated anaerobic energy metabolism. The increase in cardiac activity observed during hypoxia suggests that at about hatching time receptors are present that sense hypoxic conditions, and this information can be used to induce a stimulation of convective oxygen transport to compensate for a reduction in bulk oxygen diffusion in the face of a reduced oxygen gradient between environmental water and tissues. Under normoxia, however, the PO(2) gradient between environmental water and tissues and diffusional oxygen transport assure sufficient oxygen supply even if hemoglobin oxygen transport in the blood is severely impaired. Thus, under normoxic conditions and with a normal metabolic rate of the tissues, convective oxygen transport is not required until approximately 2 wk after fertilization.  相似文献   

2.
River impoundments are characterized by low oxygen levels as a result of reduced water velocity and increased water depth. Bimodally respiring turtle species are likely to be highly sensitive to changes in aquatic PO2 with decreases in oxygen levels impacting upon their diving ability. The acute and long-term effects of aquatic hypoxia on dive duration, oxygen consumption and blood respiratory properties were examined in hatchlings of the endangered Mary River turtle Elusor macrurus . It was hypothesized that acute exposure to aquatic hypoxia would cause a decrease in dive duration as a consequence of a decrease in reliance on aquatic respiration. With long-term exposure to hypoxia, we predicted that Elu. macrurus would have the capacity to compensate for the acute effect of hypoxia and that dive duration would increase due to an increase in aquatic respiration, haemoglobin concentration and oxygen affinity (P50). When exposed to hypoxic conditions, aquatic respiration in Elu. macrurus was substantially reduced resulting in a 51% decrease in dive duration. Contrary to our predictions, Elu. macrurus hatchlings did not acclimate, and long-term exposure to hypoxic conditions caused Elu. macrurus to lose significantly more oxygen to the hypoxic water than the normoxic acclimated turtles. The exacerbation of long-term hypoxia on the respiratory physiology and diving ecology of Elu. macrurus raises concerns about the impacts of long-term environmental change as a result of habitat alteration on the survival of freshwater turtle populations.  相似文献   

3.
Effect of salinity on oxygen consumption in fishes: a review   总被引:1,自引:0,他引:1  
The effect of salinity on resting oxygen uptake was measured in the perch Perca fluviatilis and available information on oxygen uptake in teleost species at a variety of salinities was reviewed. Trans‐epithelial ion transport against a concentration gradient requires energy and exposure to salinities osmotically different from the body fluids therefore imposes an energetic demand that is expected to be lowest in brackish water compared to fresh and sea water. Across species, there is no clear trend between oxygen uptake and salinity, and estimates of cost of osmotic and ionic regulation vary from a few per cent to >30% of standard metabolism.  相似文献   

4.
Toxic “sulphide eruptions” sporadically occur in the highly productive inshore regions of the central Namibian Benguela upwelling system. The surf clam Donax serra (Röding, 1798) dominates the intertidal and upper subtidal of large exposed sandy beaches of southern Africa and its recruitment seems to be affected by sulphide events. The reaction of juvenile surf clams to low oxygen concentrations and sulphide occurrence (0.1 mmol l−1) was examined by in vitro exposure experiments in a gas-tight continuous flow system. After 2 h of hypoxic- and hypoxic/sulphidic conditions, clams moved to the sediment surface, aiding their passive transport to areas with more favourable conditions. The clams showed a high sulphide detoxification capacity by oxidising the penetrating hydrogen sulphide to non-toxic thiosulphate. Moreover, juvenile D. serra switched to anaerobic energy production, indicated by the significant accumulation of succinate and, to some extent, alanine. Test animals were not able to reduce their energy requirements enough to withstand long periods of exposure, leading to a median survival time (LT50) of 80 h under hypoxic sulphide incubation. In conclusion, natural “sulphide eruptions”, especially those with a large spatial and temporal extension, have to be considered as an important factor for D. serra recruitment failures. Hydrogen sulphide is assumed to be a potential community structuring factor.  相似文献   

5.
There is little information regarding the effect of hypoxia on alveolar fluid clearance capacity. We measured alveolar fluid clearance, lung water volume, plasma catecholamine concentrations, and serum osmolality in rats exposed to 10% oxygen for up to 120 h and explored the mechanisms responsible for the increase in alveolar fluid clearance. The principal results were 1) alveolar fluid clearance did not change for 48 h and then increased between 72 and 120 h of exposure to hypoxia; 2) although nutritional impairment during hypoxia decreased basal alveolar fluid clearance, endogenous norepinephrine increased net alveolar fluid clearance; 3) the changes of lung water volume and serum osmolality were not associated with those of alveolar fluid clearance; 4) an administration of beta-adrenergic agonists further increased alveolar fluid clearance; and 5) alveolar fluid clearance returned to normal within 24 h of reoxygenation after hypoxia. In conclusion, alveolar epithelial fluid transport capacity increases in rats exposed to hypoxia. It is likely that a combination of endogenous norepinephrine and nutritional impairment regulates alveolar fluid clearance under hypoxic conditions.  相似文献   

6.
The rate of biosynthesis of pyruvate kinase (ATP:pyruvate 2-O-phosphotransferase, EC 2.7.1.40) was compared in cells maintained under normoxic or hypoxic conditions. L8 cells (a myoblast cell line) were pulse-labeled with [3H]leucine and incorporation of radioactivity into pyruvate kinase was measured after quantitative affinity separation with anti-pyruvate kinase monoclonal antibody. During chronic hypoxia there is an increased rate of biosynthesis of pyruvate kinase leading to an increase in enzyme content and augmented glycolytic capacity. An inhibitor of the electron transport chain, antimycin A, was used to determine whether changes in pyruvate kinase content occurring during hypoxia are a result of reduction in molecular oxygen directly or an indirect consequence of oxygen depletion. Pyruvate kinase activity increased during chronic antimycin A exposure under normoxic conditions. The increase was quantitatively accounted for by an increase in cellular pyruvate kinase enzyme content. This suggested that decreases in the levels of molecular O2 are not the direct stimulus for the increased content of pyruvate kinase. It is more likely that the increased pyruvate kinase content results from depressed rates of electron transport through the mitochondrial electron transport chain.  相似文献   

7.
The article deals with studies of serotonergic system activity in different tissues of semimigratory fish—the Caspian roach (Rutilus rutilus caspicus) and the carpbream (Abramis brama orientalis)—and of migratory fish—shemaya (Chalcalburnus chalcoides) caught in fresh and brackish waters, as well as in the common carp (Cyprinus carpio L.) tissues under effect of brackish water in model experiments. Using the indirect solid-phase ELISA-test, the serotonergic system activity was evaluated by measuring in the tissues of the studied fish the serotonin-modulated anticonsolidation protein (SMAP) which is in linear relationship with the serotonin level. There was found a significant elevation of SMAP levels in brains of the Caspian roach, carpbream, shemaya, and the common carp under action of increased water salinity. A revealed increase of the SMAP content in brain of the Caspian roach, carpbream, shemaya and the common carp under action of increased water salinity reflects the corresponding elevated activity of the serotoninergic system and indicates involvement of adaptive readjustments in the animals’ body.  相似文献   

8.
Little is known of the cardiorespiratory control mechanisms utilized by hypoxia-tolerant teleost fish to tolerate prolonged periods (h) of near anoxic exposure. Here, we report on the cardiorespiratory control mechanisms of the common carp Cyprinus carpio L. during normoxia and prolonged, severe hypoxic (<0.3 mg O(2) L(-1)) exposure at acclimation temperatures of 5 degrees C, 10 degrees C, and 15 degrees C. Through serial intra-arterial injections of alpha - and beta -adrenergic, cholinergic, and purinergic antagonists while measuring cardiac output (Q), heart rate (f(H)), ventral aortic blood pressure, and respiration rate, we established that autonomic cardiovascular and respiratory control was preserved during severe hypoxia at all three acclimation temperatures and contributed to downregulation of cardiorespiratory activity. Specifically, inhibitory cholinergic tone mediated up to 76% reductions in f(H) and Q during hypoxia, whereas the accompanying arterial hypotension was attenuated by an upregulation of an alpha -adrenergically mediated peripheral vasoconstriction. Despite the overall cardiac downregulation, a large, stimulatory cardiac beta -adrenergic tone was present during prolonged, severe hypoxia, possibly to protect the heart from attendant acidotic conditions. Purinergic blockade, following alpha -adrenergic and cholinergic antagonists, showed that the hypoxic ventilatory depression, which reversed the 2.3- to 7.7-fold increases in respiration rate that occurred with the onset of hypoxia, was a result of purinergic inhibition at all three acclimation temperatures. In contrast, purinergic inhibition of cardiac activity during hypoxia might be important only at 5 degrees C. Finally, given that cardiac power output was reduced 72%-87% during prolonged, severe hypoxia and that glycolysis yields approximately 94% less ATP per mole glucose than oxidative phosphorylation, it seems unlikely that the common carp sufficiently reduces its cardiac energy demand to a level to preclude activation of a partial Pasteur effect. This means that glycogen stores will be used and waste products will accumulate at faster rates, a finding that may help explain why the common carp cannot tolerate such extended periods of severe hypoxia (weeks to months) at cold acclimation temperatures as the freshwater turtle, which is able to reduce its cardiac energy demand to a level that does not require a Pasteur effect and also blunts autonomic cardiovascular control.  相似文献   

9.
10.
综合养鱼高产池塘的溶氧变化周期   总被引:13,自引:1,他引:12  
根据1979—1984年对主养鲢、鳙、非鲫,主养青、草鱼,主养青鱼3种养殖结构类型高产鱼池溶氧变化周期的系统研究,揭示了高产养鱼池塘溶氧的昼夜、垂直、水平变化及季节变化周期,分析了光合作用、呼吸作用、扩散作用在高产养鱼池塘溶氧动力学上的地位和作用。对高产养鱼池塘的溶氧收入、支出及平衡情况进行了定量研究,在主要饲养季节,测得晴到多云天、晴天的溶氧来源分别为:浮游植物光合产氧占86.0和95.3%,大气扩散溶入占14.0和4.7%;氧的消耗分别为:“水呼吸”消耗氧占72.0和72.0%,鱼类呼吸消耗氧占22.0和13.1%,淤泥中生物呼吸消耗氧占2.9、5.5%,扩散逸出占3.1、8.8%。文中还对溶氧变化周期与养鱼池塘管理的环境控制、结构控制之间的关系进行了分析。    相似文献   

11.
Hypoxia affects mammalian mitochondrial function, as well as mitochondria-based energy metabolism. The detail mechanism has not been fully understood. In this study, we detected protein expression levels in mitochondrial fractions of Wistar rats exposed to hypobaric hypoxia by use of proteomic methods. Adult male Wistar rats were randomized into an hypoxic (4,500?m, 30 days) group and a normoxic control group (sea level). Gastrocnemius muscles mitochondria were extracted and purified. Mitochondrial oxygen consumption was measured with a Clark oxygen electrode; mitochondrial transmembrane potential was detected with Rhodamine 123 as a fluoresce probe. Using 2-DE and MALDI-TOF MS analysis, we identified eight mitochondrial protein spots that were differentially expressed in the hypoxic group compared with the normoxic control. These proteins included Chain A of F1-ATPase, voltage dependent anion channel 1 (VDAC), hydroxyacyl Coenzyme A dehydrogenase α-subunit, mitochondrial F1 complex γ-subunit, androgen-regulated protein and tripartite motif protein 50. Two of the spots, VDAC and ATP synthase α-subunit, were confirmed by Western blotting analysis. Oxygen consumption during State 3 respiration, as well as the respiratory control ratio (RCR) was significantly higher in the control than that in the hypoxic group; mitochondrial transmembrane potential was significantly higher in hypoxic group than that in the control. With successful use of multiple proteomic analysis techniques, we demonstrates that 30 days hypoxia exposure has effects on the expression of mitochondrial proteins involved in ATP production and lipid metabolism, decrease the stability of mitochondrial membrane, and affect the mitochondrial electron transport chain.  相似文献   

12.
Hypoxia inhibits fish spawning via LH-dependent final oocyte maturation   总被引:1,自引:0,他引:1  
To evaluate the effects of long term hypoxia exposure on fish spawning, mature common carp, Cyprinus carpio carpio (Linnaeus) were subjected to either normoxia (7.4+/-0.2 mgO(2)mg O(2) L(-1)) or hypoxia (1.0+/-0.2 mgO(2)O(2) L(-1)) for more than two months. Gonadosomatic index (GSI), and concentrations of serum luteinizing hormone (LH), testosterone (T), and estroldiol (E2) were measured and gonad histology examined. Hypoxia inhibits fish spawning even though the gonad and oocytes developed under hypoxia exposure. LH levels of female carp were significantly decreased upon chronic exposure to hypoxia, and the final oocyte maturation in hypoxic females was significantly retarded. The results indicated that hypoxia may inhibit fish spawning through LH-dependent final oocyte maturation. In addition, no courtship was observed in hypoxic males. In conclusion, hypoxia impairs fish ovulation and, therefore, spawning and reproduction. LH levels were reduced leading to a failure of oocyte maturation. This, along with a lack of courtship by males may be the major mechanisms involved in hypoxic inhibition of reproduction in carp.  相似文献   

13.
1. Previous studies in a variety of ecosystems have shown that ecologically and economically important benthic and bentho‐pelagic fishes avoid hypoxic (<2 mg O2 L?1) habitats by moving vertically or horizontally to more oxygenated areas. While avoidance of hypoxic conditions generally leads to a complete shift away from preferred benthic prey, some individual fish continue to consume benthic prey items in spite of bottom hypoxia, suggesting complex habitat utilisation and foraging patterns. For example, Lake Erie yellow perch (Perca flavescens) continue to consume benthic prey, despite being displaced vertically and horizontally by hypolimnetic hypoxia. 2. We hypothesised that hypolimnetic hypoxia can negatively affect yellow perch by altering their distribution and inducing energetically expensive foraging behaviour. To test this hypothesis, we used drifting hydroacoustics and trawl sampling to quantify water column distribution, sub‐daily vertical movement and foraging behaviour of yellow perch within hypoxic and normoxic habitats of Lake Erie’s central basin during August‐September 2007. We also investigated the effects of rapid changes in ambient oxygen conditions on yellow perch consumption potential by exposing yellow perch to various static and fluctuating oxygen conditions in a controlled laboratory experiment. 3. Our results indicate that, while yellow perch in general avoid hypoxic conditions, some individuals undertake foraging forays into hypoxic habitats where they experience greater fluctuations in abiotic conditions (pressure, temperature and oxygen concentration) than at normoxic sites. However, laboratory results suggest short‐term exposure to low oxygen conditions did not negatively impact consumption potential of yellow perch. 4. Detailed understanding of sub‐daily individual behaviours may be crucial for determining interactive individual‐ and ecosystem‐level effects of stressors such as hypoxia.  相似文献   

14.
Flood discharge results in total dissolved gas (TDG) supersaturation downstream of a dam during the flood period. Fish suffer death from gas bubble disease (GBD) caused by TDG supersaturation. Nonetheless, current studies mainly attach importance to the survival of benthic fish affected by TDG supersaturation in the Yangtze River in China. Few studies have attempted to investigate the survival of pelagic fish influenced by TDG supersaturated water and compare the tolerance characteristics to TDG supersaturation between benthic and pelagic fish. To identify the survival of fish species that inhabit the various water layers affected by TDG supersaturation, silver carp (Hypophthalmichthys molitrix) (pelagic fish) and common carp (Cyprinus carpio) (benthic fish) were chosen to conduct an acute exposure experiment of four different TDG supersaturation levels (125%, 130%, 135% and 140%). The findings illustrated that the two fish species both exhibited evident aberrant behaviours of maladjustment in TDG supersaturated water. Obvious GBD symptoms were also found in the test fish. The survival probability of silver carp and common carp decreased with increasing levels of TDG supersaturation. The median survival time (ST50) values of the silver carp exposed to four levels of TDG supersaturated water (125%, 130%, 135% and 140%) were 26.84, 7.96, 5.56 and 3.62 h, respectively, whereas the ST50 values of common carp were 53.50, 26.00, 16.50 and 11.70 h, respectively. When compared with common carp, silver carp had a weaker tolerance to TDG-supersaturated water and were vulnerable to GBD. It shows that levels above 125% are not safe for common carp survival. In terms of the tolerance threshold value, silver carp merits further investigation because it showed lower tolerance to TDG than did common carp.  相似文献   

15.
Oxygen preference of the planktonic crustacean Daphnia magna was observed within an oxygen gradient. Long-term acclimation to specific oxygen conditions determined the preferred oxygen tension. Animals from hypoxic acclimation stayed in oxygen-poor water at first. They chose oxygen tensions above the critical partial pressure, thus maintaining aerobic energy supply, but they avoided higher oxygen concentrations. However, they gradually took more oxygen-rich positions within 2 d. Animals from severe hypoxic and hyperoxic acclimation showed stenoxic behavior. Specimens raised in moderately hypoxic or normoxic medium spread out more widely in the oxygen gradient (euryoxic behavior). The possible role of hemoglobin and antioxidant defense capacity limiting the animals' distribution at low or high oxygen concentrations, respectively, is discussed. In tests with mixed groups of D. magna from opposing oxygen acclimation conditions, the animals clustered at intermediate oxygen conditions. However, the acclimation condition of the major group was found to determine the position taken within the gradient to a greater extent. Selected oxygen tensions were closer to the preference values of the larger group, yet the influence of the minority prevented the majority in heterogeneous groups from taking their originally determined preference position. Thus, aggregation behavior interfered with the acclimation-dependent oxygen preference behavior.  相似文献   

16.
The physiological responses to hypoxic stress were studied in the common reed, Phragmites australis (Cav.) Trin. ex Steudel. Growth, leaf gas exchange, water (and ion) relations and osmotic adjustment were determined in hydroponically grown plants exposed to 10, 20 and 30 days of oxygen deficiency. The highest growth of reed seedlings was found in normoxic (aerobic) conditions. Treatment effects on biomass production were relatively consistent within each harvest. Leaf water potential and osmotic potential declined significantly as hypoxia periods increased. However, leaf turgor pressure showed a consistent pattern of increase, suggesting that reed plants adjusted their water status by osmotic adjustment in response to root hypoxia. After 20 and 30 days in the low oxygen treatment, net CO2 assimilation and stomatal conductance were positively associated and the former variable also had a strong positive relationship with transpiration. Short-term hypoxic stress had a slight effect on the ionic status (K+, Ca2+ and Mg2+) of reed plants. In contrast, soluble sugar concentrations increased more under hypoxic conditions as compared to normoxia. These findings indicate that hypoxia slightly affected the physiological behavior of reed plants.  相似文献   

17.
The physiological responses to hypoxic stress were studied in the common reed, Phragmites australis (Cav.) Trin. ex Steudel. Growth, leaf gas exchange, water (and ion) relations and osmotic adjustment were determined in hydroponically grown plants exposed to 10, 20 and 30 days of oxygen deficiency. The highest growth of reed seedlings was found in normoxic (aerobic) conditions. Treatment effects on biomass production were relatively consistent within each harvest. Leaf water potential and osmotic potential declined significantly as hypoxia periods increased. However, leaf turgor pressure showed a consistent pattern of increase, suggesting that reed plants adjusted their water status by osmotic adjustment in response to root hypoxia. After 20 and 30 days in the low oxygen treatment, net CO2 assimilation and stomatal conductance were positively associated and the former variable also had a strong positive relationship with transpiration. Short-term hypoxic stress had a slight effect on the ionic status (K+, Ca2+ and Mg2+) of reed plants. In contrast, soluble sugar concentrations increased more under hypoxic conditions as compared to normoxia. These findings indicate that hypoxia slightly affected the physiological behavior of reed plants.  相似文献   

18.
The energetic consequences of fighting, which may depend on environmental conditions, can be an important factor shaping contest strategy and duration. Energy expenditure may be costly to fitness because it depletes reserves that could otherwise have been allocated to reproduction, and metabolites are produced that may constrain subsequent activities. We examined the variation in the metabolic consequences of fighting in relation to hypoxia. Contests were staged between pairs of size-matched male shore crabs Carcinus maenas L., under a range of water oxygen tensions (between 10 and 100% oxygen saturation) which crabs experience in their natural habitat. Fighting under normoxic and hypoxic conditions resulted in significantly elevated concentrations of haemolymph metabolites (L-lactate and glucose) compared with crabs at rest. However, these concentrations were much lower than in crabs that had been walking on a treadmill. Glycogen concentrations differed only under hypoxic conditions: glycogen stores were reduced in crabs after fighting and this reduction was similar to that after exercise on a treadmill. Contests were shorter when they were staged below a water P o2of 6.7 kPa ( approximately 30% normoxia). As water oxygen tensions were reduced, fighting crabs had greater concentrations of L-lactate and glucose in their blood and tissues whilst glycogen stores were reduced. Fights became shorter when crabs were exposed to severe hypoxia (P o2=2 kPa) for increasing lengths of time, and blood L-lactate concentrations increased. The results suggest that as fights progressed, crabs experienced an increasing metabolic debt, in the form of accumulation of L-lactate and a reduction in energy stores, which was amplified by hypoxic conditions. Copyright 1999 The Association for the Study of Animal Behaviour.  相似文献   

19.
All 20.000 different fish species vary greatly in their ability to tolerate and survive fluctuating oxygen concentrations in the water. Especially fish of the genus Carassius, e.g. the crucian carp and the goldfish, exhibit a remarkable tolerance to limited/absent oxygen concentrations. The metabolic changes of anoxia-tolerant crucian carp were recently studied and published. Contrary to crucian carp, the hypoxia-tolerant common carp cannot survive a complete lack of oxygen (anoxia). Therefore, we studied the 1H-NMR-based metabolomics of brain, heart, liver and white muscle extracts of common carp, subjected to anoxia (0 mg O2 l?1) and hypoxia (0.9 mg O2 l?1) at 5 °C. Specifically, fish were exposed to normoxia (i.e. 9 mg O2 l?1; controls 24 h, 1 week and 2 weeks), acute hypoxia (24 h), chronic hypoxia (1 week) and chronic hypoxia (1 week) with normoxic reoxygenation (1 week). Additionally, we also investigated the metabolic responses of fish to anoxia for 2 h. Both anoxia and hypoxia significantly changed the tissue levels of standard energy metabolites as lactate, glycogen, ATP/ADP and phosphocreatine. Remarkably, anoxia induced increased lactate levels in all tissues except for the heart whereas hypoxia resulted in decreased lactate concentrations in all tissues except for brains. Furthermore, hypoxia and anoxia influenced amino acids (alanine, valine/(iso)leucine) and neurotransmitters levels (GABA, glutamate). Lastly, we also detected ‘other’ i.e. previously not reported compounds to play a role in the present context. Scyllo-inositol levels changed significantly in heart, liver and muscle, providing novel insights into the anoxia/hypoxic responses of the common carp.  相似文献   

20.
间歇性低氧适应的心脏保护   总被引:9,自引:0,他引:9  
Zhang Y  Yang HT  Zhou ZN 《生理学报》2007,59(5):601-613
间歇性低氧(intermittent hypoxia,IH)是指一定时间间断地暴露于低氧环境,而其余时间处于常氧环境。IH是机体某种生理和病理状态下的低氧形式。研究表明:间歇性低氧适应(IHadaptation),类似缺血预适应(ischemic preconditioning,IPC)和长期高原低氧适应(long-termhigh-altitude hypoxic adaptation,LHA),具有明显的心脏保护作用,表现为增强心肌对缺血/再灌注损伤的耐受性、限制心肌梗死面积和形态学改变、抗细胞凋亡、促进缺血/再灌注心脏舒缩功能的恢复,以及抗心律失常。尽管IH对心脏的保护作用不容质疑,但其作用机制远未阐明。IH心脏保护作用可能涉及氧的运输、能量代谢、神经体液调节、抗氧化酶、应激蛋白、腺苷系统、ATP敏感钾通道、线粒体及其钙调控、一氧化氮和蛋白激酶等多方面机制,并受低氧处理方式、动物年龄和性别等因素影响。IH心脏保护持续时间明显长于IPC,而对机体的不良影响远小于LHA,具有潜在的应用价值。  相似文献   

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