North American Journal of Fisheries Management
ISSN: 0275-5947 (Print) 1548-8675 (Online) Journal homepage: http://afs.tandfonline.com/loi/ujfm20
Swimming Performance of Rock Carp Procypris rabaudi and Prenant’s Schizothoracin Schizothorax prenanti Acclimated to Total Dissolved Gas Supersaturated Water Yuanming Wang, Ruidong An, Yong Li & Li Kefeng To cite this article: Yuanming Wang, Ruidong An, Yong Li & Li Kefeng (2017): Swimming Performance of Rock Carp Procypris rabaudi and Prenant’s Schizothoracin Schizothorax prenanti Acclimated to Total Dissolved Gas Supersaturated Water, North American Journal of Fisheries Management, DOI: 10.1080/02755947.2017.1353558 To link to this article: http://dx.doi.org/10.1080/02755947.2017.1353558
Accepted author version posted online: 18 Jul 2017.
Submit your article to this journal
Article views: 1
View Crossmark data
Full Terms & Conditions of access and use can be found at http://afs.tandfonline.com/action/journalInformation?journalCode=ujfm20 Download by: [Sichuan University]
Date: 20 July 2017, At: 22:00
Swimming Performance of Rock Carp Procypris rabaudi and Prenant’s Schizothoracin Schizothorax prenanti Acclimated to Total Dissolved Gas Supersaturated Water *
Yuanming Wang, Ruidong An , Yong Li, and Kefeng Li
ip t
State Key Laboratory of Hydraulics and Mountain River Engineering, Sichuan University, Chengdu 610065, China
cr
Abstract: Total dissolved gas (TDG) supersaturation is becoming an increasingly
us
serious problem in the upper Yangtze River Basin and has been identified as a
an
potentially negative environmental effect of hydropower development and operations. Juvenile Rock Carp Procypris rabaudi and Prenant’s Schizothoracin Schizothorax
M
prenanti dwell in the upper reaches of the Yangtze River and were selected as
ed
experimental fish to evaluate the impact of TDG supersaturation on swimming performance. The results showed that the swimming speeds of Rock Carp and
ce pt
Prenant’s Schizothoracin differed significantly under different exposure times to TDG supersaturated water (129 – 131%). Exposure of fish to supersaturated TDG for 2 h and 4 h with no recovery resulted in significantly lower critical swimming (Ucrit) and
Ac
burst swimming (Uburst) speeds compared to those of fish that were not exposed to supersaturated TDG. Ucrit and Uburst values for Rock Carp not exposed to supersaturated TDG were 1.4 to 1.6 times higher than those of fish exposed to
*
Corresponding author. E-mail:
[email protected]
1
supersaturated TDG for 2 h and 1.9 to 2.0 times higher than those of fish exposed for 4 h. Similarly, the Ucrit and Uburst values for Prenant’s Schizothoracin not exposed to supersaturated TDG were 1.4 to 1.8 times higher than those of fish exposed to supersaturated TDG for 2 h and 2.5 to 3.1 times higher than those of fish exposed for
ip t
4 h. Fish that were exposed to supersaturated TDG for 2 h and 4 h and then allowed to
cr
recover for 2 d swam better than fish exposed to supersaturated TDG for the same
us
amount of time without recovery. Although a 2-d recovery improved swimming performance compared to that of fish not allowed to recover, only the Rock Carp (and
an
not the Prenant’s Schizothoracin) returned to the swimming speeds observed in the
M
control fish that were not exposed to supersaturated TDG. The results of our research can be applied to the operation of hydropower stations and can contribute
Ac
ce pt
ed
fundamental data for establishing water-related environmental standards in China.
2
Hydropower development is inevitable in China due to the country’s energy requirements, and many high-head dams have been built in the upper Yangtze River Basin, which is the richest hydropower resource in China (Huang and Zheng 2009; Fang and Deng 2011; Tang and Zhou 2012). During the flood season, water is allowed
ip t
to spill over dams through discharge structures, and air is trapped in the plunge pools
cr
downstream of the dams, thus leading to increased total dissolved gases (TDGs) in the
us
downstream river (Hibbs and Gulliver 1997; Qu et al. 2012). TDG supersaturation can affect fish and other aquatic resources downstream of dams, and a high level of TDG
an
supersaturation can easily cause gas bubble disease (GBD) and mortality in fish
M
(Weitkamp and Katz 1980; Weitkamp 2008; Chen et al. 2012; Wang et al. 2015a; Wang et al. 2015b). A serious supersaturation problem existed in the Columbia River
ed
Basin (USA), and GBD in fish resulting from TDG supersaturation has long been a cause of concern (Ebel 1969; Meekin 1971; McGrath et al. 2006; Arntzen et al. 2008;
ce pt
Geist et al. 2013). Fish downstream of Three Gorges Dam Gezhouba Dam also show signs of GBD during the flood season (Water Resources Protection Bureau of Yangtze
Ac
River 1983; Tan 2006). After Xiluodu Dam discharged its first flood waters in 2014, more than 40 tons of fish died and floated on the Jinsha River, which attracted widespread attention from the government and the public (CCTV 2014). Management agencies around the world (e.g., in the Columbia River Basin in the USA) have set water quality criteria for dissolved gas to protect fish. Waivers are granted to the water quality criteria when evidence suggests operations won’t have an 3
impact on aquatic biota (Weitkamp 2008). However, information that can be used to establish water quality criteria for TDG has not been collected for Chinese rivers that are impounded by hydroelectric dams; therefore, more research is needed. Water quality criteria (and waivers) must consider fish behavior because if they are not
ip t
severely affected by dissolved gas, fish can swim to avoid high TDG, but high TDG
cr
could affect upstream migration abilities (Gray and Haynes 1977; Lund and
us
Heggberget 1985; Johnson et al. 2005; Johnson et al. 2010).
Swimming is an important physiological activity in fish that is closely related to
an
foraging behavior, evasion capacity and propagation (Webb 1984). Fish swimming
M
behavior can be classified into three major categories: sustained, prolonged, and burst swimming. Critical swimming (Ucrit) is a special category of prolonged swimming
ed
that reflects the aerobic swimming performance of fish. Burst swimming (Uburst) is generally considered to be performed anaerobically (Beamish 1979; Hammer 1995).
ce pt
With the construction of high-head dams, the swimming performance of downstream fish is inevitably affected by TDG supersaturation (Schiewe 1974; Jia 2016). However,
Ac
while previous studies have primarily concentrated on the tolerance of fish to TDG supersaturation (Jensen 1988; Smiley et al. 2011; Chen et al. 2012; Geist et al. 2013; Wang et al. 2015a), some researchers have assessed fish detection and avoidance to supersaturated water (Stevens et al. 1980; Lund and Heggberget 1985; Wang et al. 2015a). While the swimming capability of juvenile Chinook Salmon Oncorhynchus tshawytscha and Steelhead Trout Oncorhynchus mykiss exposed to TDG 4
supersaturated water (100 – 125%) has been studied (Schiewe 1974; Dawley and Ebel 1975), the impact of TDG supersaturation on the swimming performance of fish in the upper Yangtze River has not been properly investigated. Both Rock Carp Procypris rabaudi and Prenant's Schizothoracin Schizothorax
ip t
prenanti are benthic species that are endemic the upper Yangtze River, where the water
cr
temperature is between 5 – 27 °C and the dissolved oxygen (DO) concentration is
us
above 4 mg/L. These species grow slowly and are favored by the Chinese people for their flavor and nutritional value (Ding 1994; Yang et al. 2009). However, the
an
populations of these fish have greatly declined in recent years, and this decline is
M
associated with dam construction (Liu 2004). The flood duration in the upper Yangtze River is 1-2 d, and endemic species have always been seriously threatened by TDG
ed
supersaturation during the flood season (Yu et al. 2009). However, the duration of continuous TDG supersaturation may be shorter due to dam regulation.
ce pt
To study the impact of TDG supersaturation on the swimming performance of
fish living in the upper Yangtze River, we selected Rock Carp and Prenant's
Ac
Schizothoracin as our experimental species and then assessed the Ucrit and Uburst values
of fish exposed to TDG supersaturated water. Based on previous studies, we proposed two hypotheses: (1) TDG exposure (129 – 131%) for a short time (2 to 4 h) would result in lower swimming speeds and even mortality (Wang et al. 2015a and b), and (2)
swimming speeds would return to normal levels after sufficient recovery time (2 d) (Schiewe 1974; Dawley and Ebel 1975; Wang et al. 2015b). The results of this study 5
can be used to inform the operation of hydropower stations and contribute fundamental data for establishing water-related environmental standards in China. Materials and Methods Experimental fish. – Healthy juvenile Rock Carp and Prenant’s Schizothoracin were
ip t
obtained from the Sichuan Fisheries Research Institute of China for use in the
cr
experiment. Prior to experimentation, the fish were kept in a saturated-equilibrium
us
(100% TDG) water pool (water depth × length × width: 35 cm × 400 cm × 100 cm) for 7 d to acclimate to the environment. The water temperature ranged from 23 -
an
25 °C, the DO level ranged from 7.0 - 8.1 mg/L, and the pH was 7.5-8.0. Experimental
M
fish were fed Tubificidae twice daily, and to avoid the impact of food on swimming performance, feeding was stopped 24 h before the experiment.
ed
Experimental design. – After the 7 d of acclimation, a total of 160 Rock Carp and 120 Prenant’s Schizothoracin were introduced into TDG supersaturated water tanks with a
ce pt
depth of 35 cm. The supersaturation level was between 129 and 131% TDG which is representative of conditions downstream from the dams on the upper Yangtze River
Ac
Basin (Li et al. 2009; Qu et al. 2011). The TDG supersaturated water was generated using a pump and a compressor (Wang et al. 2015a,b), and the dissolved gas levels were monitored by a TDG measuring instrument (Tracker, Point Four Systems Inc., Canada) at the surface of the water tank. At the second hour of exposure (the time point when Rock Carp began to die), 18 Rock Carp and 12 Prenant’s Schizothoracin were immediately subjected to either Ucrit 6
or Uburst tests. Similarly, 12 Rock Carp and 10 Prenant’s Schizothoracin were removed from the TDG supersaturated water tanks and subjected to the tests at the fourth h of exposure (the time point when Prenant’s Schizothoracin began to die). At the second hour of exposure, 40 Rock Carp and 30 Prenant’s Schizothoracin were transferred to
ip t
the saturated-equilibrium water pool for 2 d to recover, and at the fourth hour of
cr
exposure, 63 Rock Carp and 62 Prenant’s Schizothoracin were also transferred for 2 d
us
of recovery. The fish were not fed during the 2-d recovery process, after which their swimming performance was assessed via Ucrit and Uburst tests. During the exposure to
an
TDG supersaturation and the recovery process, dead experimental fish were
M
immediately removed, and the behavior of the living fish was recorded. As a control, we also tested the Ucrit and Uburst of healthy Rock Carp and Prenant’s Schizothoracin
ed
(0 h of TDG supersaturation exposure) prior to TDG supersaturation exposure. The body masses and lengths of the fish used in the Ucrit and Uburst tests are shown in Table
ce pt
1, and each fish was used only once.
Measurement of Ucrit and Uburst. – A Brett-type swim tunnel respirometer (Loligo
Ac
Systems SW10200, Denmark) was used to measure the Ucrit and Uburst of the fish. The swim tunnel respirometer had a total water volume of 90 L, and the swimming chamber was 70 × 20 ×20 cm (length × width × height). A honeycomb screen was fixed at the upstream point of the swimming chamber to reduce turbulence and ensure a uniform water velocity across the swimming chamber (for details see Cai et al. 2013). Water flow was driven by a single propeller powered by an electric motor with 7
a variable frequency drive, and the water flow velocity was measured by a propeller flow meter and ranged from 5 - 150 cm s-1. The water temperature in the swimming chamber ranged from 23 - 25 °C, and the DO ranged from 7.0-8.1 mg/L. The Ucrit and Uburst values were determined using the increasing velocity method
ip t
(Brett 1964). Fish were placed in the swimming chamber under a 5 cm s-1 flow for 20
cr
min to eliminate the stress of the transfer process, and the Ucrit test began with an
us
increase in the water velocity to 50% of the estimated Ucrit from the 5 cm s-1 acclimation velocity (Jain et al. 1997; Penghan et al. 2014). The water velocity in the
an
swimming chamber was then steadily increased in 10 cm s-1 increments every 20 min
=
)∆
(1)
is the highest speed at which the fish swam during the entire time
ed
where
+(
M
until the fish were exhausted. The Ucrit was calculated as follows:
period (cm s-1); ∆
is the velocity increment (10 cm s-1, which is equal to 1/4 to is the constant swimming period at each
ce pt
1/5th of the Ucrit of the control fish); speed (20 min); and
is the time (min) for which the fish swam at the final speed.
Ac
The solid blocking effect was considered to be negligible because the volume of the fish was < 10% of the swim chamber (Webb, 1971). All absolute Ucrit (cm s-1) values were converted to body length per second, which denotes the relative Ucrit (BL s-1). The duration of the test period typically did not exceed 2 h (Schiewe 1974). The Uburst test was similar to the Ucrit test. Fish were held in the swimming
chamber for 20 min at a water speed of 5 cm s-1 for acclimation. Then, the Uburst test 8
began with an increase in the water velocity to 50% of the Uburst estimated from the 5 cm s-1 acclimation velocity. The water velocity in the swimming chamber was then steadily raised in 10 cm s-1 increments every 1 min until the fish were exhausted. Statistical analysis. – The impact of TDG supersaturation exposure time on swimming
ip t
speed was determined using one-way analysis of variance (ANOVA), which was
cr
followed by a post hoc multiple comparison test (least significant difference) to
us
determine the difference between the values of the different treatment groups. The effect of the 2 d of recovery on the swimming speed was determined by an
an
independent samples t-test. The level of significant difference was set at P < 0.05.
M
Results
Effect of TDG supersaturation on fish. – When the fish were initially exposed to the
ed
TDG supersaturated water, most gathered at the bottom of the tanks. After 0.5 h of exposure, the fish exhibited elevated activity. Some fish swam more slowly with
ce pt
increased exposure time; some lost equilibrium and fell to the bottom of the experimental tanks; and some jumped and hit the nets in the upper part of the tanks. Gas
Ac
bubbles were evident on the fins of the fish after 2 h of exposure. Two Rock Carp (1.25%) died during the second hour of exposure and 25 (15.6%) died during the fourth hour. In addition, six Prenant’s Schizothoracin (5.0%) died within 4 h of exposure to TDG supersaturation. During the first day of recovery, most fish, particularly those that experienced 4 h of exposure, either swam slowly or were stationary at the bottom of the pool, and a number of the fish swam to the surface and 9
released gas bubbles from their mouth. Three Rock Carp (7.5%) that experienced 2 h of exposure and 9 Rock Carp (14.3%) that experienced 4 h of exposure died in the recovery pool within 12 h in the recovery pool. Five Prenant’s Schizothoracin (8.1%) that experienced 4 h of exposure also died during the recovery process. Among the
ip t
other fish, the gas bubbles on the fins gradually disappeared, and all fish exhibited
us
Prenant’s Schizothoracin exposed to 2 h of TDG died.
cr
normal behavior after 2 d of recovery. During the recovery process, none of the
Swimming performance of fish under TDG supersaturation stress. – The swimming
an
ability of the fish in the experiment under exposure to TDG supersaturation is shown
M
in Figure 1. The ANOVA revealed significant differences were observed in the swimming speed of the Rock Carp between the different TDG supersaturation
ed
exposure times (Ucrit: F(2, 20) = 13.10, P < 0.001; and Uburst: F(2, 23) = 56.77, P < 0.001). Exposure of Rock Carp and Prenant’s Schizothoracin to supersaturated TDG for 2 h
ce pt
and 4 h with no recovery resulted in significantly lower Ucrit and Uburst swimming speeds than those of fish not exposed to supersaturated TDG. The Ucrit and Uburst
Ac
values for unexposed Rock Carp were 1.4 to 1.6 times higher than fish exposed to supersaturated TDG for 2 h and 1.9 to 2.0 times higher than those of fish exposed for 4 h. Similarly, the swimming speed of Prenant’s Schizothoracin also differed significantly between the different exposure times according to the ANOVA (Ucrit: F(2,
13)
= 20.76, P < 0.001; and Uburst: F(2, 13) = 35.86, P < 0.001). The Ucrit and Uburst values
for unexposed Prenant’s Schizothoracin were 1.4 to 1.8 times higher than those of the 10
fish exposed to supersaturated TDG for 2 h and 2.5 to 3.1 times higher than fish exposed for 4 h. Swimming performance of fish after recovery. – The Ucrit values of the Rock Carp after 2 h and 4 h of exposure and 2 d of recovery were 4.33 and 4.59 BL s-1, respectively,
ip t
and the Uburst values were 6.11 and 5.85 BL s-1, respectively (Figure 2). Significant
cr
differences in the Ucrit or Uburst values between the Rock Carp after 2 h and 4 h of
us
exposure were not detected by the independent t-test analysis (Ucrit: F18 = 2.98, P = 0.71; and Uburst: F18 = 0.15, P = 0.69). The Ucrit of the Prenant’s Schizothoracin after 2
an
h of exposure was 7.64 BL s-1 after 2 d of recovery, and this value was significantly
M
higher than that after 4 h of exposure (6.14 BL s-1) according to the independent t-test (F8 = 0.023, P = 0.04). Significant differences were not observed between the Uburst
ed
values of between the Prenant’s Schizothoracin after 2 and 4 h of exposure according to the independent T-test (F9 = 3.10, P = 0.20), although the Uburst after 2 h of exposure
ce pt
was 9.4% higher than that after 4 h. The swimming speeds of the Rock Carp after 2 d of recovery were not significantly different from those of the control Rock Carp (P >
Ac
0.05), except for the Ucrit of the Rock Carp after 2 h of exposure, but the speeds of the Prenant’s Schizothoracin exposed to TDG were significantly lower than those of the control group (P < 0.05) except for the Ucrit after 2 h of exposure. Compared with the
swimming ability of fish after 2 or 4 h of exposure, the swimming ability of the corresponding fish improved after 2 d of recovery (Figure 3). The swimming speed of the Rock Carp after 2 days of recovery was significantly higher than that of the fish 11
without recovery (P < 0.05), except for the Ucrit after 2 h of exposure. The swimming speed of the Prenant’s Schizothoracin after 2 d of recovery was more than 35% higher than that of the fish without recovery (P < 0.05). Discussion
ip t
Effect of TDG supersaturation on fish. – TDG supersaturation can cause GBD in fish,
cr
thereby threatening their survival (Weitkamp and Katz 1980; Weitkamp 2008). Gas
us
bubbles were visible on the fins of the two species in this study, and these fish also suffered mortality within 4 h of TDG supersaturation (15.6% for the Rock Carp and
an
5.0% for the Prenant’s Schizothoracin). The results indicated that Prenant’s
M
Schizothoracin was more tolerant of TDG supersaturated water than Rock Carp during the 4 h exposure treatment, which was consistent with the results of previous
ed
studies (Wang et al. 2015a; Wang et al. 2015b).
Swimming performance of fish under TDG supersaturation stress. – Previous studies
ce pt
have indicated that the swimming performance of fish may be affected by many factors. For example, Fu et al. (2011) exposed Goldfish Carassius auratus to hypoxic
Ac
conditions (0.3 mg O2 L-1) for 48 h and then determined the Ucrit of the fish in normoxic conditions (10 mg O2 L-1) and hypoxic conditions (1 mg O2 L-1), their results showed that the Goldfish exposed to hypoxic conditions (0.3 mg O2 L-1) for 48
h had a significantly higher Ucrit than the fish that were not exposed. Melzner et al. (2009) studied the swimming performance of Atlantic Cod Gadus morhua acclimated to elevated seawater PCO2 and found that the Ucrit of fish incubated for 1 year at a PCO2 12
of 0.6 kPa or for 4 months at a PCO2 of 0.3 kPa did not show differences in Ucrit relative to control fish. Our results indicated that TDG supersaturation severely impacted the swimming performance of Rock Carp and Prenant’s Schizothoracin. Prenant’s Schizothoracin
ip t
swam better than Rock Carp, and under TDG supersaturation, the swimming speed of
cr
both species significantly decreased. Schiewe (1974) found that exposure to 117 and
us
120% TDG supersaturation levels (in tanks with depths of 25 cm) for more than 10 h or exposure to 104, 106 and 112% TDG supersaturated water for 35 d decreased the
an
swimming capability of juvenile Chinook Salmon. However, the swimming
M
performance of control fish was only significantly better than that of the test fish at 120% TDG supersaturation. Dawley and Ebel (1975) also found that the swimming
ed
performance of the control fish was not significantly different from that of Steelhead Trout exposed to 105 - 125% TDG supersaturated water for 35 d or until 10 or 50%
ce pt
mortality was reached. Compared with the treatments for the juvenile Chinook Salmon and Steelhead Trout, the Rock Carp and Prenant’s Schizothoracin in this
Ac
study were exposed to a higher level of TDG supersaturation (129 - 131% TDG) over a shorter time (2 or 4 h), and their swimming speeds differed significantly between the different exposure times. However, the swimming performance of Chinook Salmon and Steelhead Trout were only measured by distance gained and swimming duration against a constant water current; their Ucrit and Uburst were not tested.
13
The present results showed that the Uburst values of healthy Rock Carp and Prenant’s Schizothoracin was 123 and 135% of the Ucrit, respectively, which is consistent with the findings of previous reports that showed that the Uburst of Largemouth bass Micropterus salmoides, Atlantic cod and Rainbow trout
ip t
Onchorhynchus mykiss were 120 to 180% of their Ucrit (Farlinger and Beamish 1977;
cr
Reidy et al. 2000; Farrell 2008). After 2 or 4 h of TDG supersaturation exposure, the
us
Uburst values of the two species in this study were slightly higher than their Ucrit values, although the difference was not significant. This finding indicated that the anaerobic
M
their aerobic swimming performance.
an
swimming performance of the fish was more susceptible to the effects of TDG than
Swimming performance of fish after recovery. – Our results showed that most of the
ed
Rock Carp and Prenant’s Schizothoracin recovered within 2 d in the saturated-equilibrium water, which is consistent with the results of Dawley and Ebel
ce pt
(1975) and Wang et al. (2015b). Wang et al. (2015b) placed Rock Carp into 35 cm deep tanks at 130% supersaturation for 2.33 h and then held them in
Ac
saturated-equilibrium (100% TDG) water for 5 d. Their results indicated that these fish had a poorer tolerance to TDG supersaturated water than the control fish, although their horizontal and vertical behaviors were not significantly different. Our experiment showed that the swimming speed of Prenant’s Schizothoracin after 2 d of recovery was significantly lower than that of the control fish, whereas that of the Rock Carp under the same treatment reached baseline levels. This result indicates that the 14
Rock Carp recovered their swimming ability more easily than the Prenant’s Schizothoracin. Schiewe (1974) exposed juvenile Chinook Salmon to 120% TDG supersaturated water until 50% mortality was reached (mean value of 13.4 h) and found that the swimming capabilities of the surviving fish could completely recover
ip t
after 2 h of exposure in saturated-equilibrium (100% TDG) water. The swimming
us
of the Chinook Salmon examined by Schiewe (1974).
cr
capabilities of the two species in this study took longer to recover compared with that
Wang et al. (2015a and b) found that Rock Carp and Prenant’s Schizothoracin
an
could exploit the compensation depth to reduce the effect of TDG supersaturation
M
under laboratory conditions. In our study, the fish were exposed to extremely acute TDG supersaturation because of the absence of a compensatory depth (35 cm test
ed
depth). Because Rock Carp and Prenant’s Schizothoracin are both benthic species in the upper Yangtze River, they can use the compensation depth in natural river
ce pt
conditions to reduce the effects of TDG supersaturation and may exhibit better swimming performance than under laboratory conditions.
Ac
Management Implications. – Resident fish in the upper Yangtze River (such as Rock Carp and Prenant’s Schizothoracin) stressed by TDG supersaturation may become more vulnerable to predators; their foraging behavior may become inefficient; and their ability to evade threats (such as TDG supersaturation) may be compromised due to decreased swimming capabilities. However, our results showed that fish could recover their swimming ability with sufficient time in un-saturated water, and under 15
natural conditions, fish could migrate to a branch of the river (with only small dams or without dams) to avoid the impact of TDG supersaturation. Thus, artificial tributaries without TDG supersaturation could be created for fish, which would be especially important because of the many rare fish that live in the upper Yangtze River (Dudgeon
ip t
2010). The results presented here provide important information that can be used to
cr
develop methods to protect fish threatened by TDG supersaturation and to guide
us
operation of hydropower stations.
an
Acknowledgments
This work was sponsored by the National Key Research Project of China under
M
permission number 2016YFC0401710 and the National Natural Science Foundation
REFERENCES
ed
of China under permission number 51379136.
ce pt
Arntzen, E. V., K. J. Murray, E. M. Dawley, K. D. Hand, J. L. Panther, V. I. Cullinan, and D. R. Geist. 2008. Effects of total dissolved gas on Chum salmon fry
Ac
incubating in the lower Columbia River. Pacific Northwest National Laboratory.
Brett, J. R. 1964. The respiratory metabolism and swimming performance of young sockeye salmon. Journal of the Fisheries Research Board of Canada 21: 1183-1226. Beamish, F. W. H. 1979. Swimming Capacity. Fish physiology 7: 101-187. Chen, S. C., X. Q. Liu, W. Jiang, K. F. Li, J. Du, D. Z. Shen, and Q. Gong. 2012. Effects of total dissolved gas supersaturated water on lethality and catalase activity 16
of Chinese sucker (Myxocyprinus asiaticus Bleeker). Journal of Zhejiang University-SCIENCE B (Biomedicine & Biotechnology) 13: 791-796. Cai, L., R. Taupier, D. Johnson, Z. Tu, G. Liu, and Y. Huang. 2013. Swimming capability and swimming behavior of juvenile Acipenser schrenckii. Journal of
ip t
Experimental Zoology Part A: Ecological Genetics and Physiology 319:
cr
149-155.
us
CCTV. 2014. Shichuan Jinsha salvage 40 tons of dead fish poisoning may exclude official. http://news.cntv.cn/2014/07/19/ARTI1405750580177886.shtml.
an
Dawley, E. M., and W. J. Ebel. 1975. Effects of various concentrations of dissolved
M
atmospheric gas on juvenile Chinook salmon and steelhead trout. United States National Marine Fisheries Service Fishery Bulletin 73: 787-796.
ed
Ding, R. H. 1994. Fishes of the Sichuan. Sichuan Publishing of Science and Technology, Chengdu, China.
ce pt
David, D. 2010. Requiem for a river: extinctions,climate change and the last of the Yangtze. Aquatic Conservation: Marine and Freshwater 20: 127–131.
Ac
Ebel, W. J. 1969. Supersaturation of nitrogen in the Columbia River and its effect on salmon and steelhead trout. United States National Marine Fisheries Service Fishery Bulletin 68: 1-11.
Farlinger, S., and F. W. H. Beamish. 1977. Effects of time and velocity increments on the critical swimming speed of largemouth bass (Micropterus salmoides). Transactions of the American Fisheries Society 106: 436-439. 17
Farrell, A. P. 2008. Comparisons of swimming performance in rainbow trout using constant acceleration and critical swimming speed tests. Journal of Fish Biology 72: 693-710. Fang, Y., and W. Deng. 2011. The critical scale and section management of cascade
ip t
hydropower exploitation in Southwestern China. Energy 36: 5944-5953.
cr
Fu, S. J., C. J. Brauner, Z. D. Cao, J. G. Richards, J. L. Peng, R. Dhillon, and Y.
us
X.Wang, 2011. The effect of acclimation to hypoxia and sustained exercise on subsequent hypoxia tolerance and swimming performance in goldfish (Carassius
an
auratus).The Journal of experimental biology 214: 2080-2088.
M
Gray, R. H., and J. M. Haynes. 1977. Depth distribution of adult chinook salmon (Oncorhynchus tshawytscha) in relation to season and gas-supersaturated water.
ed
Transactions of the American Fisheries Society 106: 617-620. Geist, D. R., T. J. Linley, V. Cullinan, and Z. Q. Deng. 2013. The Effects of Total
ce pt
Dissolved Gas on Chum Salmon Fry Survival, Growth, Gas Bubble Disease, and
Seawater Tolerance. North American Journal of Fisheries Management 33:
Ac
200-215.
Hammer, C. 1995. Fatigue and exercise tests with fish. Comparative Biochemistry and Physiology Part A: Physiology 112: 1-20.
Hibbs, D. E., and J. S. Gulliver. 1997. Prediction of effective saturation concentration at spillway plunge pools. Journal of Hydraulic Engineering 132: 940-949.
18
Huang, H. L., and Y. Zheng. 2009. Present situation and future prospect of hydropower in China. Renewable and Sustainable Energy Reviews 13: 1652-1656. Jensen, J. O. T. 1988. Combined effects of gas supersaturation and dissolved oxygen levels on steelhead (Salmo gairdneri) eggs, larvae, and fry. Aquaculture 68:
ip t
131-139.
cr
Jain, K. E., J. C. Hamilton, and A. P. Farrell. 1997. Use of a ramp velocity test to
us
measure critical swimming speed in rainbow trout (Onchorhynchus mykiss). Comparative Biochemistry and Physiology Part A: Physiology 117: 441-444.
an
Johnson, E. L., T. S. Clabough, D. H. Bennett, T. C. Bjornn, C. A. Peery, C. C. Caudill,
M
and L. C. Stuehrenberg. 2005. Migration depths of adult spring and summer Chinook salmon in the lower Columbia and Snake rivers in relation to dissolved
1213-1227.
ed
gas supersaturation. Transactions of the American Fisheries Society 134:
ce pt
Johnson, E. L., T. S. Clabough, C. C. Caudill, M. L. Keefer, C. A. Peery, and M. C. Richmond. 2010. Migration depths of adult steelhead Oncorhynchus mykiss in
Ac
relation to dissolved gas supersaturation in a regulated river system. Journal of
fish biology 76: 1520-1528.
Jia, J. 2016. A technical review of hydro-project development in China. Engineering 2:302-312.
19
Lund, M., and T. G. Heggberget. 1985. Avoidance response of two-year-old rainbow trout, Salmo gairdneri Richardson, to air-supersaturated water: hydrostatic compensation. Journal of fish biology 26: 193-200. Li, R., J. Li, K. F. Li, Y. Deng, and J. J. Feng. 2009. Prediction for supersaturated total
ip t
dissolved gas in high-dam hydropower projects. Science in China Series E:
cr
Technological Sciences 52: 3661−3667.
us
Liu, J. 2004. A quantitative analysis on threat and priority of conservation order of the endemic fishes in upper reaches of the Yangtze River. China Environmental
an
Science 24: 395-399.
M
Meekin, T. K. 1971. Levels of nitrogen supersaturation at Chief Joseph Dam under various spill conditions, phase 1. Report to United States Army Corps of
ed
Engineers, Contract NPSSU-71-796, Washington Department of Fisheries, Olympia, Washington, USA.
ce pt
McGrath, K. E., E. M. Dawley, and D. R. Geist. 2006. Total dissolved gas effects on fishes of the Lower Columbia River. Pacific Northwest National Laboratory.
Ac
Melzner, F., S. Göbel, M. Langenbuch, M. A. Gutowska, H. O. Pörtner, and M. Lucassen. 2009. Swimming performance in Atlantic Cod (Gadus morhua)
following long-term (4–12 months) acclimation to elevated seawater PCO2.
Aquatic toxicology 92: 30-37. Penghan, L. Y., D. Z. Cao, and S. J. Fu. 2014. Effect of starvation on swimming performance of juvenile carp. Chinese Journal of Ecology 33: 2756-2760. 20
Qu, L., R. Li, J. Li, K. F. Li, and Y. Deng. 2011. Field observation of total dissolved gas supersaturation of high-dams. Science in China Series E: Technological Sciences 54: 156-162. Reidy, S. P., S. R. Kerr, and J. A. Nelson. 2000. Aerobic and anaerobic swimming
ip t
performance of individual Atlantic cod. The Journal of Experimental Biology 203:
cr
347-357.
us
Schiewe, M. H. 1974. Influence of dissolved atmospheric gas on swimming performance of juvenile Chinook salmon. Transactions of the American Fisheries
an
Society 103: 717-721.
M
Tan, D. C. 2006. Research on the lethal effect of the dissolved gas supersaturation resulted from Three Gorges Project to fish. Master’s thesis. Southwest University,
ed
Chongqing, China.
Tang, X., and J. Zhou. 2012. A future role for cascade hydropower in the electricity
ce pt
system of China. Energy Policy 51: 358-363.
Webb, P. W. 1971. The swimming energetics of trout. I. Thrust and power output at
Ac
cruising speeds. The Journal of experimental biology 55: 489-520.
Weitkamp, D. E., and M. Katz. 1980. A review of dissolved gas supersaturation literature. Transactions of the American Fisheries Society 109: 659-702.
Water Resources Protection Bureau of Yangtze River. 1983. Investigation of gas bubble disease and spill of Gezhouba Dam.
21
Webb, P. W. 1984. Body form, locomotion and foraging in aquatic vertebrates. American Zoologist 24: 107-120. Weitkamp, D. E. 2008. Total dissolved gas supersaturation biological effects, Review of Literature 1980-2007. Bellevue: Parametrix Inc., 2008:1-65.
ip t
Wang, Y., K. Li, J. Li, R. Li, and Y. Deng. 2015a. Tolerance and avoidance
cr
characteristics of Prenant's schizothoracin Schizothorax prenanti to total dissolved
us
gas supersaturated water. North American Journal of Fisheries Management 35: 827-834.
an
Wang, Y., R. Liang, Y. Tuo, K. Li, and B. Hodges. 2015b. Tolerance and avoidance
M
behavior towards gas supersaturation in rock carp Procypris rabaudi with a history of previous exposure. North American Journal of Aquaculture 77:
ed
478-484.
Yang, L., R. L. Mayden, and Y. Z. Cai. 2009. Threatened fishes of the world: Procypris
ce pt
rabaudi (Tchang, 1930) (Cyprinidae). Environmental biology of fishes 84: 275-276.
Ac
Yu, F., Z. Chen, X. Ren, and G. Yang. 2009. Analysis of historical floods on the Yangtze River, China: Characteristics and explanations. Geomorphology 113: 210-216.
22
FIGURE 1. Swimming speed of experimental fish after 0, 2, and 4 h of exposure to TDG supersaturation (129-131% TDG). Values are shown as the mean ± S.E. (Rock Carp: n = 4 - 10 per treatment; Prenant’s Schizothoracin: n = 4 - 6 per treatment). Letters above the bars indicate the results from a post hoc multiple comparison test
ip t
(least significant difference test); mean values that do not share a common lower case
Ac
ce pt
ed
M
an
us
cr
letter are significantly different (P < 0.05).
23
FIGURE 2. Swimming speed of experimental fish exposed to elevated levels of dissolved gas (129-131% TDG) for either 2 or 4 h and then allowed to recover for 2 d before testing. Also shown for comparison are the swimming speeds of control fish tested before any exposure to elevated dissolved gas. Values are shown as the mean ±
ip t
S.E. (Rock Carp: n = 4-10 per treatment; Prenant’s Schizothoracin: n = 4-6 per
cr
treatment). * Significant difference in swimming speed between treatments (P
