DFO Libra y MPO Bibliotheque
11 1 1 11
08014483
9236 G 1014
EVALUATION OF SWIMMING CAPABILITY
AND POTENTIAL VELOCITY BARRIER
PROBLEMS FOR FISH
Part B: New Telemetric Approaches to the
Assessment of Fish Swimming Performance
Prepared by:
FISHERIES AND OCEANS
St. John's, NF
and
UNIVERSITY OF WATERLOO
Waterloo, ON
1+ 1
QL
Fisheries and Oceans Peches et Oceans
Canada
Canada
Canadian Electricity Association
Association canadienne de l'electricite
639.4
E92
September 1998
CEA RESEARCH & DEVELOPMENT DIVISION
The CEA R&D Program, created in 1974,
coordinates and complements the research
requirements of the Canadian electric utility industry.
The Program is jointly funded by member
utilities in Canada and the federal government through Energy, Mines and Resources
Canada and the Panel on Energy Research
and Development.
The objectives of the Program, as defined
within the Constitution of the Canadian
Electrical Association, are:
. Develop technologies to improve safety,
reliability, performance, and predictable
service life of electrical systems equipment and processes so as to reduce utility
capital and operating costs.
Develop and investigate new and advanced technologies for more efficient electricity generation, transmission, distribution and use so as to conserve energy and
natural resources and reduce utility capital
and operating costs.
• Through research, establish facts and
develop technologies with the aim of reducing adverse environmental and socioeconomic impact of electricity generation,
transmission, distribution and use.
. Provide data, analysis, methodology and
overall R&D perspective to aid the utility
industry, the federal government and other
policy setting bodies in present and future
energy decision making.
. Maximize the benefits of resources allocated to research and development by CEA
through the provision of joint R&D programs and liaisons with national and international organizations engaged in similar
work.
. Promote and facilitate the transfer of
information gained and technologies developed through the R&D Program to
CEA member organizations and to government and the scientific, academic and
industrial communities.
REPORT FOR THE
CANADIAN ELECTRICITY ASSOCIATION
Research & Development
1155, rue Metcalfe, bureau 1120
Montreal QC H3B 2V6
CEA No. 9236 G 1014
EVALUATION OF SWIMMING CAPABILITY AND
POTENTIAL VELOCITY BARRIER PROBLEMS FOR FISH
PART 8:
New Telemetric Approaches to the Assessment of Fish
Swimming Performance
PREPARED BY:
FISHERIES AND OCEANS
Science Branch
P.O. box 5667
St. John's NF A1C 5X1
UNIVERSITY OF WATERLOO (2)
Waterloo Biotelemetry Institute
Waterloo ON N2L 3G1
(1)
Project Leaders:
D .A. Scruton1
R.S. McKinley2
Principal Investigators
R.K. Booth 2
M. Colavecchia2
R.G. Goosney1
SEPTEMBER 1998
NOTICE
This report was prepared by the CONTRACTOR and sponsored by the Canadian
Electricity Association (CEA) which does not necessarily agree with the opinions
expressed herein.
Neither CEA (including its members), nor the CONTRACTOR, nor any other person
acting on their behalf makes any warranty, expressed or implied, or assumes any legal
responsibility for the accuracy of any information or for the completeness or usefulness
of any apparatus, product or process disclosed, or accept liability for the use, or damages
resulting from the use, thereof.
Neither do they represent that their use would not
infringe upon privately owned rights.
Any reference in this report to any specific commercial product, process or service by
tradename, trademark, manufacturer or otherwise does not necessarily constitute or imply
its endorsement or recommendation by the CONTRACTOR, CEA or any of its members.
The correct citation for this report is as follows:
Scruton, D.A., R.S. McKinley, R.K. Booth, S.J. Peake, and R.F. Goosney. 1998.
Evaluation of Swimming Capability and Potential Velocity Barrier Problems for
Fish Part A. Swimming Performance of Selected Warm and Cold Water Fish
Species Relative to Fish Passage and Fishway Design. CEA Project No. 9236 G
1014, Montreal, Quebec. xiv + 62 pp., 2 appendices.
Copyright© 1998 - Canadian Electricity Association. All rights reserved.
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1155, rue Metcalfe, bureau 1120
Montreal, Quebec H3B 2V6
Tel: (514) 866-6121 Telecopieur: (514) 866-1880
courr.el.: [email protected]
ABSTRACT
This study was undertaken to develop a set of information or criteria related to
swimming capability for several important fish species to provide biological design
criteria to mitigate potential velocity barrier problems associated with hydroelectric
facilities. A major objective of this project report included the development and
evaluation of innovative approaches to assessing locomotory activity, swimming
performance, and energetic costs to fish under naturally occurring conditions and in
relation to potential barrier problems. Physiological telemetry was used and involved
implantation of a bio-sensitive radio transmitter (EMG or electromyogram tag) in
swimming muscle of fish, calibration to locomotory ability and energetic scope, and
subsequent use of radio transmitted EMG signals to assess swimming performance
and metabolic costs in situ. Studies were also conducted to assess the effects of sex,
maturity, location and timing with respect to migratory distance, and body morphology
on muscle activity. Additionally, 'state-of-the-art' telemetry systems (DSP or digital
signal processing with antennae switching) were used to study high speed swimming
performance, behaviour, and migratory strategy in relation to ascent of an experimental
flume. Finally, these techniques were applied to assessing swimming performance,
behavioural strategy, and energetics associated with passage of an existing fishway at
Grand Falls, Newfoundland. Collectively, these studies have demonstrated the
capabilities of these technologies and techniques in addressing fish passage issues
and furthermore have indicated the complexity of factors that regulate fish swimming
energy expenditure that need be considered in the design and operation of fish
bypasses.
Keywords: swimming performance, swimming speed, burst swimming, physiological
telemetry, electromyograms, fish passage, fishways, Atlantic salmon, water velocity;
digital telemetry
111
lV
ACKNOWLEDGMENTS
This project was primarily co-funded by the Canadian Electricity Association in
partnership with the Canadian Department of Fisheries and Oceans (DFO). Additional
funding and in kind support was provided by the following: DFO, Marine Environment
and Habitat Management Division (Newfoundland Region)- burst swimming research
and fabrication of Blaska swim speed tube; the University of Waterloo - NSERC grant to
R.S. McKinley- principle investigator; DFO (Central and Arctic Region)- travel costs for
participation of C. Katopodis; Manitoba Hydro -funding support for walleye and lake
whitefish research in Manitoba; Manitoba Department of Fish and Wildlife - use of
Whiteshell Provincial Fish Hatchery for walleye and lake whitefish research; U.S. Fish
and Wildlife Service (Region 3), the S.O. Conte Anadromous Fish Research Centre of
the U.S. National Biological Survey, and the Wisconsin and Michigan Departments of
Natural Resources -funding for research on lake sturgeon; Lotek Engineering Inc. - in
kind support for telemetry studies; Alma Fish Hatchery - support for sturgeon swimming
performance research; Abitibi-Consolidated (Grand Falls Division)- funding of
swimming performance research on Atlantic salmon smolt and kelt; and Newfoundland
and Labrador Hydro- in kind support.
The work reported here is primarily research completed by a number of M.Sc. and
Ph.D. students at the University of Waterloo under the supervision of the principal
investigators R.S. McKinley (chief supervisor) and D.A. Scruton (committee member).
In addition to the students identified as authors, technical assistance was also provided
by. J. Mitchell, E. Bombardier, and K. Conners. Other individuals and agencies have
played an important role in support of this research including: Mr. Ed Hill (B.C. Hydro
formerly of Newfoundland and Labrador Hydro)- advice and support in development of
the study proposal; R. Bukowski (Manitoba Hydro)- coordination of, and securing
funding for, studies in Manitoba; C. Katopodis (DFO - Central and Arctic Region)scientific advice; S. Richter (Acres International Limited)- coordination for smolt and
kelt research at Grand Falls, Newfoundland; and the Environmental Resources
Management Association (ERMA), in particular F. Parsons- fish capture and
transportation in Newfoundland.
A number of Fisheries and Oceans (DFO) staff have also supported this project
including L. Cole, A. Bowdring, P. Rose, C. Kelley, C. Pennell, L. Fudge, G. Clarke, T.
Nichols, K. Smith, N. Hefford, and many others. Support from DFO Engineering field
staff including G. Higgins, C. Armstrong, W. Butler , and S. Maynard was also greatly
appreciated. The authors would also like to acknowledge the advice and direction
provided by the Canadian Electricity Association's (CEA) Technical Monitors for the
project; Mr. Roy Bukowski (Manitoba Hydro), Mr. Marcell LaPerle and Jean-Maurice
Gauthier of Hydro Quebec and Mr. Ed Hill, B.C. Hydro. Ms. Tayna Galvicic-Theberge
and Ms. Cecile Gragner, CEA Project Managers, have also shown much support for the
project and have demonstrated considerable patience for the project Final Reports.
v
Vl
EXECUTIVE SUMMARY
This study was conceived to address a problem of mutual concern to the Canadian
Electricity Association, representing Canada's hydroelectric utilities, and the
Department of Fisheries and Oceans, as a government regulatory body responsible for
the management and conservation of freshwater and anadromous fish and their
habitats. Specifically, this study was undertaken to develop a set of information or
criteria related to the swimming capability for several widely distributed and important,
both from a recreational and economic perspective, fish species. This information is to
be used to provide design criteria to mitigate potential velocity barrier problems
associated with hydroelectric facilities and other developments. The study also
developed new and innovative methods for assessing swimming performance in situ,
primarily employing recent advances in conventional and physiological telemetry. Data
from this study provides an information base to assist industry and government assess
the design, construction, and evaluation of fish passage systems.
The initial focus of this study was to investigate the swimming capability of anadromous
(pre-spawning adults, post-spawning adults, juveniles, smolts) and landlocked
Uuveniles, adults) Atlantic salmon (Sa/mo sa/ar); brook trout (Salve/inus fontina/is;
juveniles and adults), brown trout (Sa/mo trutta; juveniles and adults), lake sturgeon
(Acipenser fulvesens; juveniles and adults), and walleye (Stizostedion vitreum; adults),
collected from various locations throughout Canada. Criteria were developed related to
sustained, prolonged, burst swimming performance characteristics of the study
species/life stages and included investigation of the effect of environmental variables
(e.g. temperature), fish physiology (e.g. sex, maturity, body morphology), life history,
and migration distance on swimming performance (companion report; Part A.
Swimming Performance of Selected Warm And Cold Water Fish Species Relative to
Fish Passage And Fishway Design).
The second major emphasis of this research was to develop and evaluate innovative
approaches to assessing locomotory activity, swimming performance, and energetic
costs to fish under naturally occurring conditions in relation to potential barrier
problems. This involved surgical implantation of a bio-sensitive radio transmitter (EMG
or electromyogram tag) in individual fish, calibration to locomotory ability and energetic
scope (calibration of EMGs to swimming speed and oxygen consumption), and
subsequent use of radio transmitted EMG signals to assess swimming performance
and metabolic costs in situ. Studies were also conducted to assess the effects of sex,
maturity, location and timing with respect to migratory distance, and body morphology
on muscle activity as determined from EMG telemetry. Additionally, 'state-of-the-art'
telemetry systems (DSP or digital signal processing with antennae switching) were
used to study high speed swimming performance, behaviour, and migratory strategy in
relation to ascent of a an experimental flume. Finally, these techniques were applied to
assessing swimming performance, behavioural strategy, and energetics associated with
passage of an existing fishway at Grand Falls, Newfoundland. Collectively, these
studies have demonstrated the capabilities of these technologies and techniques in
vii
addressing fish passage issues and furthermore have indicated the complexity of
factors that regulate fish swimming energy expenditure that need be considered in the
design and operation of fish bypasses (this report; Part B. New telemetric Approaches
to the Assessment of Fish Swimming Performance). Further, results from the
physiological telemetry would allow validation and extrapolation of laboratory based
bioenergetic models and basic swimming performance criteria as developed in Part A of
this study.
Swimming capabilities and in situ measurement of muscle activity from adult Atlantic
salmon (Salmo salar), at two seasonal temperatures, were measured using radio
transmitted electromyogram (EMG) signals. Critical swimming speeds were determined
and correlated to radio transmitted EMG signals in a modified Blazka swim speed
chamber. At 18 °C, sustained activity and critical swimming speeds were approximately
70% and 20% higher than at 12 °C, respectively. No differences in burst activity were
observed at these temperatures. EMGs recorded from salmon during ascent of an
artificial flume at cold temperatures revealed that overall muscle activity was greater
than that observed for critical swimming speeds, implying that white muscle was being
recruited at these temperatures. Data demonstrated that salmon may recruit white
muscle fibres and incur an oxygen debt at colder temperatures as a strategy for
ascending velocity obstructions at a quicker rate.
The influence of environmental temperature and changes in body morphology on the
swimming capabilities and muscle activity patterns in migrating Atlantic salmon were
also studied. Significant increases in girth and cross-sectional area were observed
among females but not males. No differences in muscle activity indices were observed
until the onset of spawning when mean muscle activity indices increased significantly
for both sexes. Sex dependent differences in muscle activity indices were most
pronounced during prolonged swimming (i.e. 2 body length per second) and were
significantly higher for females. Muscle activity was correlated to both temperature and
cross-sectional area for females, but only to temperature for males. Results indicated
that environmental temperature was an important determinant of swimming
performance in anadromous Atlantic salmon. Changes in the body morphology of
females placed additional demands on locomotory muscle and may be responsible for
the significantly lower aerobic swimming capabilities observed.
High speed (burst) swimming performance of wild Atlantic salmon was investigated in
an experimental flume using coded radio signals, with a digital spectrum processor
using near real-time spectrum analysis, to measure distance moved and time elapsed.
Fish voluntarily swam against water velocities, ranging from 1.32 to 2.85 m s· 1, in an 18
m long flume. At water velocities of 1.32 to 1.55 m·s· 1, individuals successfully
ascended the flume at swimming speeds of 1.61 to 2.55 m·s·1 , or 3.30 to 4. 79 body
lengths per second (bl·s- 1 ), respectively. At high water velocities ranging from 1.92 to
2.85 m·s·1, individual swimming speeds increased from 2.55 to 3.60 m·s· 1, or 4.94 to
7.27 bl·s· 1 , respectively. However, above a threshold value of 1.92 m·s·1, individuals
traversed shorter distances and were unable to ascend the flume. Results
Vlll
demonstrated that digital telemetry would be an excellent tool in design and evaluation
of future fishways and culvert installations.
Swimming performance, including movement patterns and passage times, of wild
Atlantic salmon was then investigated in an existing fishway again using coded radiotransmitted signals. Radio-tagged adults swam voluntarily through a 116 m long
vertical slot fishway with passage occurring primarily during late morning (57 .1%) and
late afternoon (35.7%), with night passage (7.1 %) of secondary importance. As
velocities increased from 1.69 ± 0.07 m·s-1 to 1.82 ± 0.03 m·s-1 , ascent times
significantly increased from 3.33 ± 0.72 to 27.95 ± 8.86 h whereas, the number of
entrance attempts significantly declined from 18.70 ± 3.68 to 7.75 ± 1.71 per day.
Tracking data provided fine resolution of movements and position of fish within the main
sections, resting pools and entrance of the fishway. EMG signals from fish moving
through the fishway indicated excessive energy requirements for movement between
pools of the fishway, for holding position in pools, and in use of the designed resting
areas. Results indicated that the DSP and EMG telemetry technology is highly
applicable for providing design criteria for new fishways or for modifying existing bypass structures.
This information is of importance to industry and government in responding in a
scientifically responsible manner to fish passage concerns. Collectively the information
contained in these two reports (Part A and B) will be applicable to assessing the
impacts and specifying design criteria for existing and proposed hydroelectric
developments, bridge and culvert installations, fish passage facilities, etc., throughout
the range of the study species. This information will be of benefit in the development of
mitigative strategies for structures or conditions that may potentially impede fish
passage or alienate habitats.
IX
X
TABLE OF CONTENTS
1.0
INTRODUCTION
1-1
1.1
1.2
1-1
1-2
1.5
Background
Measurement of Locomotory Performance Using Physiological
Telemetry
High Speed (Burst) Swimming
Use of Telemetry to Assess Fish Swimming Performance Within an
Existing Fishway
Study Objectives
1-5
2.0
MATERIAL AND METHODS
2-1
2.1
2.2
Study Locations
Physiological Telemetry Studies
2.2.1 Experimental Animals
2.2.2 Electromyogram (EMG) Transmitter and Telemetry
Equipment
2.2.3 Surgical Procedures
2.2.4 Calibration of EMG Signals with Swimming Speed
2.2.5 In Situ Measurement of Swimming (Muscle) Activity
2.2.6 Statistical Analyzes
Burst Swimming Studies
2.3.1 Experimental Flume
2.3.2 Telemetry Equipment
2.3.3 Experimental Animals
2.3.3.1
Physiology and Blood Collection
2.3.4 Flume Hydraulics
2.3.5 Data Analysis
2.3.6 Statistical Analyzes
Assessment of Swimming Performance in a Fishway
2.4.1 Fishway Description
2.4.2 Experimental Animals and Transmitter Attachment
2.4.3 Digital Spectrum Processing (DSP) Telemetry Studies
2.4.4 Electromyogram (EMG) Telemetry Studies
2.4.5 Statistical Analysis
2-1
2-2
2-2
2-2
1.3
1.4
2.3
2.4
1-3
1-5
2-3
2-4
2-5
2-5
2-6
2-6
2-6
2-8
2-9
2-9
2-10
2-10
2-11
2-11
2-12
2-13
2-13
2-13
3.0
RESULTS
3-1
3.1
Physiological Telemetry Studies
3.1.1 Swimming Performance and Temperature
3.1.2 Relationship of Muscle Activity to Swimming Speed
3.1.3 Correlation of Oxygen Consumption and Swimming Speed
3.1.4 Effect of Body Morphology
3-1
3-1
3-1
3-2
3-2
Xl
3.2
3.3
3.1.5 Muscle Activity During Spawning Migration
3.1.6 In Situ Muscle Activity (Experimental Flume Study)
Burst Swimming
3.2.1 Chemical, Temperature and Flow Conditions
3.2.2 Timing of Ascent of Flume
3.2.3 Success Rates
3.2.4 Swimming Performance Kinetics in Relation to Velocity
3.2.5 Blood Lactate
Assessment of Swimming Performance in a Fishway
3.3.1 Timing of Ascent of Fishway
3.3.2 Rate of Passage
3.3.3 Muscle Activity (EMGs) From Fish Moving Through Fishway
3-2
3-3
3-4
3-4
3-5
3-5
3-5
3-9
3-9
3-9
3-10
3-11
4.0
DISCUSSION
4-1
5.0
CONCLUSIONS
5-1
6.0
REFERENCES
6-1
7.0
APPENDICES
APPENDIX A:
APPENDIX B:
Report Illustrations
Annotated Bibliography of Publications Arising from
the Study
Xll
A-1
B-1
LIST OF TABLES
2-7
Table 2-1.
DSP_500 and SRX_400 Specifications
Table 2-2:
Physical and hydraulic characteristics of the Grand Falls
fishway.
Table 3-1.
Multiple squared regression coefficients and significance
levels from comparisons of girth and temperature with
muscle activity measured during swimming at 1 and 2 body
lengths per second from wild Atlantic salmon collected
during their spawning migration.
3-3
Table 3-2.
Summarized swimming data for experimental (summer)
1996 fish grouped by water velocity (m·s- 1). VF, VFW,
Vfmax, and Vfwmax are reported in m·s- 1 and bl·s-1 and refer
to average ground speed, average total speed, maximum
ground speed and maximum total speed, respectively.
3-7
Table 3-3.
Summarized swimming data for experimental (fall)1996 fish
grouped by water velocity (m·s- 1 ). VF, VFW, Vfmax. and
Vfwmax, are reported in m·s-1 and bl·s-1 and refer to average
ground speed, average total speed, maximum ground speed
and maximum total speed, respectively.
3-8
Table 3-4.
Summarized plasma lactate values (m·moles·L- 1 ) for controls
and salmon ascending the flume under high water velocities.
3-9
Table 3-5.
Time periods of salmon ascent up the Grand Falls fishway.
3-10
Table 3-6.
Biological characteristics, passage times, entrance attempts,
and behavioral patterns of adult anadromous Atlantic
salmon monitored at Grand Falls fishway.
3-11
Xlll
2-11
xiv
LIST OF FIGURES
Figure 2-1.
Study locations along the migratory route of Atlantic salmon
in the Exploits River, Newfoundland. Mean water
temperature and sampling dates are shown.
A1-1
Figure 2-2.
A schematic showing the location the electromyogram
(EMG) transmitter in the body cavity of fish. Gold tipped
electrodes are inserted into the red swimming muscle.
A1-2
Figure 2-3.
A schematic diagram showing 7 fixed antennae stations
connected to the receiver/coprocessor system in the
experimental flume used for this study.
A 1-3
Figure 2-4.
Hydraulics of the experimental flume, Noel Paul's Brook. A
stoplog section in the first sluiceway of the dam controls
head elevation (~H) and subsequent water velocities (V)
downstream.
A 1-4
Figure 2-5
A schematic diagram of Grand Falls fishway with
corresponding antennae locations (denoted 1 to 7) used in
this study. Resting facilities are located at antennae five and
seven.
A 1-5
Figure 2-6
A schematic diagram of Grand Falls fishway showing
locations of EMG telemetry monitoring. The EMG signals
from traversing the lower 11 pools and from spending time in
the resting pool (pool 12) are shown.
A1-6
Figure 3-1.
Fatigue tests of swimming performance of wild Atlantic
salmon (n=5) conducted at 12 and 18 °C. Transition to
exhaustion has been sub-divided into the following
components: sustained, prolonged and burst swimming
speeds. Dotted lined represent swimming endurance (i.e.
time to fatigue) beyond 120 minutes.
A1-7
Figure 3-2.
Calibration of muscle activity to swimming performance in
wild Atlantic salmon (n=5), at 12 and 18 °C.
A1-8
Figure 3-3.
Correlation of oxygen consumption with muscle activity in
wild Atlantic salmon (n=5), at 12 and 18 °C.
A1-9
Figure 3-4.
Relationships between muscle activity and swimming
speeds for male (solid) and female (white) salmon at various
stages of their spawning migration. Site 1 represents
XV
A1-10
freshwater entry and site 5 represents the pre-spawning
period. Sample sizes are as follows sites 2-4 N=4 males
and 4 females, site 1 and 5 N=3 males and 3 females.
Figure 3-5.
The muscle activity indices of male (solid circle) and female
(white circle) Atlantic salmon at rest (i) and swum at 1 (ii)
and 2 (iii) body lengths per second. Hatched bar indicates
the pre-spawning period. Sampling periods with similar
letters are not significantly different. Significant differences
between sexes are indicated by an asterix (*). In all cases
the accepted level of significance was P<0.05.
A1-11
Figure 3-6.
Muscle activity recorded in wild Atlantic salmon (n=4) during
ascent of 20 m long experimental flume recorded at 12 °C
(fall) and 18 °C (late summer) in relation to critical swimming
speed (Ucrit).
A1-12
Figure 3-7.
Water temperature in Noel Paul Brook during the 1996
experimental period as obtained by hourly thermograph
readings.
A1-13
Figure 3-8.
Depth profiles in the flume with corresponding pool
elevations. Elevations of 0.25, 0.34, and 0.43 represent
mean water velocities of 1.89, 2.79, and 3.09 m·s·\
respectively.
A1-14
Figure 3-9.
Time periods and fish activity in the flume. Time periods
(hours: minutes) of 00:01-04:00, 04:01-08:00, 08:01-12:00,
12:01-16:00,16:01-20:00, and 20:01-24:00 refer to night,
early morning, late morning, early afternoon, late afternoon,
and evening periods, respectively.
A1-15
Figure 3-10. Success rates of passage of the flume in relation to water
velocity (m·s- 1).
A1-16
Figure 3-11. Maximum distances attained by salmon ascending the flume
at various water velocities (m·s- 1).
A1-17
Figure 3-12. Total time required for salmon to ascend the flume at various
water velocities (m·s- 1).
A1-18
Figure 3-13. Average ground speeds (m·s- 1) for varying water velocities
(m·s- 1).
A1-19
Figure 3-14. Average total speeds (m·s-1) for varying water velocities (m·s·
A1-20
XVI
1 ).
Figure 3-15. Average total speeds, in bl·s- 1 , for varying water velocities
(m·s- 1 ).
A1-21
Figure 3-16. Maximum total speeds, in bl·s- 1 , in relation to increasing
water velocities (m·s-1 ).
A1-22
Figure 3-17. Maximum ground speeds (m·s- 1 ) in relation to increasing
water velocities (m·s- 1 ).
A1-23
Figure 3-18. Representative tracks of fish ascent of the flume under
moderate water flow conditions. Panel (A) shows an
unsuccessful attempt with panel (C) showing corresponding
speeds attained. Panel (B) shows a successful attempt with
panel (D) showing corresponding speeds attained.
A1-24
Figure 3-19. Time versus position profiles for several tagged salmon at
two water flows. Flow period 1 refers to 1.69 ± 0.07 m·s- 1
and flow period 2 refers to 1.82 ± 0.03 m·s- 1 • The y-axis
represents the seven sections of the fishway as illustrated in
Figure 2-5.
A1-25
Figure 3-20. The time spent (min) at various sections of the fishway
during two water flows. Results are expressed as a mean±
standard error(** P ~ 0.01 ).
A1-26
Figure 3-21. The number of unsuccessful attempts (total) at the fishway
entrance at two flow periods. Unsuccessful refers to
individuals who did not ascend the fishway during the study.
Successful refers to fish who traversed the fishway. Results
are expressed as a mean ± standard error(* P ~ 0.05).
A1-27
Figure 3-22. A schematic of Grand Falls fishway with corresponding
pools (1-11) and resting pool (12). EMG signals for passage
through pools 1-11 and for remaining in resting pool12 are
depicted.
A1-28
xvn
New Telemetric Approaches To The Assessment OfFjsh Swjmmjng Peifarmance
1.0
INTRODUCTION
1.1
Background
The design and operation of fish bypass systems generally fail to integrate the
swimming ability and activity pattern of the fish they were constructed to serve. As a
result, the water velocity at bypasses, culverts, fishways, bridges, etc. may be beyond
the scope of the fish and therefore act as a velocity barrier to upstream passage.
Positioning of structures may also prevent or hinder fish from reaching or locating a
bypass. Information is also required to ensure that the design of fish passage
structures themselves do not represent behaviourial barriers to fish.
A possible negative impact of hydroelectric development is the creation of a 'velocity
barrier', a situation where the velocity of water prevents fish from accessing habitat that
is critical to the maintenance of the population (e.g. spawning habitat). Velocity barriers
can be created by increasing flows in a stream above natural conditions by diversion,
channelization, flow augmentation, etc. or by restricting channel width as in construction
of power canals, bridges and culverts. The requirement for fish to maintain position or
traverse areas of high velocity may have a physiological cost beyond which energy
budgeted for migration and/or reproduction may be utilized. The consequences of
imposing excessive energy expenditure on migrating fish could severely influence
reproductive success through failure to reach spawning habitats and reduced or
impaired production of gonadal products. Fish swimming ability is therefor a critical
consideration in providing access (i.e. fishway construction) above an impoundment or
in gaining access to habitat.
Proposed actions or construction activities associated with hydroelectric development
may potentially constitute a velocity barrier, therefore it is necessary to compare water
velocity conditions associated with the undertaking with the swimming ability of fish
(species and size) in question. Water velocities can generally be measured or
modeled, however, information on the swimming abilities of several important fish
species is generally lacking. Consequently this study was conceived to develop
swimming performance criteria for wild (non-hatchery) for important fish species within
Canada. The study was also to develop new and innovative methods for assessing
swimming performance in situ, primarily employing recent advances in conventional
and physiological telemetry.
1- 1
New
1.2
Telemetrjc Approaches To The Assessment OfFish Swjmmiag Pedormance
Measurement of Locomotory Performance Using Physiological Telemetry
Direct measurement of intensity of locomotory activity displayed by wild fish has
previously been difficult (Fry 1947; Beamish 1978; Brett and Groves 1979). Estimates
of fish activity in the field are difficult to quantify and previous attempts have involved
measurements of movement between fixed points using mark and recapture techniques
(Ellis 1966), conventional radio telemetry (McCleave et al. 1978) and ultrasonic
telemetry (Quinn 1988). These methods provide estimates of minimum swimming
speed, however they are unable to distinguish between swimming mode (i.e. aerobic
vs. anaerobic) or relative swimming intensity.
Biotelemetry has played an important role in furthering understanding fish migration and
identification of critical habitats. Knowledge of the energetics costs of migration to
access these critical habitats requires direct measures of fish activity in the field. This
will allow assessment of fish swimming performance and pattern of energy expenditure
relative to a fishes physiological limitations. Physiological telemetry, using new devices
and equipment, is considered a major advancement that will permit measure of
physiological attributes in fish in situ and will allow validation and extrapolation of
laboratory based bioenergetic models and basic swimming performance criteria (e.g.
sustained, prolonged, and burst swimming). Developments in physiological telemetry
has produced several promising techniques for directly estimating the activity and energetics of free living fish under field conditions including correlations of heart rate (Priede
and Young 1977; Armstrong et al. 1989; Lucas et al. 1993), opercular rate (Rogers and
Weatherly 1983; McKinley and Power 1992; Demers et al. 1996; Weatherly et al. 1996)
and locomotor muscle activity using electromyograms (EMGs) (Rogers and Weatherly
1984; Kaseloo et al. 1992; McKinley and Power 1992). EMGs are best deployed in fish
where the metabolic rate is largely determined by locomotory activity and consequently
is ideally suited to measurement of physiological correlates of fish swimming
performance. While radio transmitted measurements of muscle activity (i.e. EMG) have
been used to assess the activity patterns and estimate energy expenditures of fish
(McKinley and Power 1992; Demers et al. 1996; Hinch et al 1996; Weatherley et al.
1996), application to assessment of swimming efficiency has not yet been
demonstrated.
Much of a fish's routine activity involves sustained and prolonged swimming typically
supported by the red muscle (Beamish 1978). Energy required for this activity is
obtained through the metabolism of energy-rich substrates such as fats and proteins
acquired during feeding. In migratory species such as the Atlantic salmon (Salmo
salar) feeding is suspended during their spawning period, consequently energy required
for locomotion must be obtained solely from endogenous sources. This situation is
further complicated by a period of sexual development which also occurs during the
migratory period. Gonadal development requires enormous amounts of energy and
may leave little for other needs such as body maintenance and migration. Changes to
1 -2
New Telemetr;c Approaches To The Assessment OfFish Swjroroing Periarmance
either the intensity or duration of migration could significantly accelerate the rate at
which energy is used and could result in unsuccessful spawning, low egg viability, poor
recruitment and even death of spawning adults.
Considerable research has been conducted towards understanding the energetics
(Jonsson et al. 1991; 1997), migratory patterns (McCleave et al. 1978; Power and
McCleave 1980) and reproductive behavior (Heggberget 1988; Bagliniere et al. 1990)
of Atlantic salmon. There is a paucity of information concerning the swimming
capabilities and efficiencies of Atlantic salmon during their spawning migration. During
spawning migration, salmon experience significant fluctuations in temperature as well
as pronounced changes in body morphology due to starvation and morphogenic
changes associated with sexual development. The influence of such factors on
swimming performance suggest decreases in swimming performance may be expected
during migration. Migration is an important component in the energy budget of
anadromous fish (Lambert and Dodson 1990) and changes in the intensity of the
migration or decreases in the swimming capabilities of migratory fish can have
consequences on energy available for other purposes (e.g. spawning).
A number of factors have can influence the swimming capabilities of fish including water
temperature (Quinn et al. 1996), body size/morphology (Taylor and McPhail1985;
Goolish 1991; Hawkins and Quinn 1996), and reproductive status (Thorstad et al.
1997). Information obtained from studies of muscle activity patterns in swimming fish
suggest that swimming speeds can be maintained independent of the influence of
external factors by increasing the number of active muscle fibers (Rome et al.
1984, 1990; Jayne and Lauder 1994 ). Increased muscle activity can therefor mask the
influence of environmental and physiological factors on swimming capabilities. Some
previous studies have investigated the swimming capabilities of Atlantic salmon (e.g.
Booth et al. 1997), however, no studies have described the sustained, critical and burst
swimming capabilities of Atlantic salmon during spawning migration. Swimming
performance studies have typically not involved assessment of swimming intensity,
consequently a major purpose of this research was to examine the influence of body
morphology and temperature on the sustained and burst swimming capabilities and
muscle activity of male and female Atlantic salmon during spawning migration.
1.3
High Speed (Burst) Swimming
Swimming of fish can be classified into three major categories: sustained, prolonged
and burst swimming speeds. Sustained swimming occurs at relatively low water
velocities and represents speeds which can be maintained for longer than 200 minutes
using energy derived exclusively from aerobic processes (Beamish 1978). Prolonged
swimming covers a spectrum of speeds between burst and sustained and is often
categorized by st~ady swimming interspersed with periods of vigorous efforts. The
1 -3
New Telemetric Approaches To The Assessment Of Bsh $wjmmjng Peiformance
highest speeds of which fish are capable are classified as burst swimming. In fish, as in
all vertebrates, the highest levels of exercise performance are achieved anaerobically
(Jones 1982). These high speeds can be maintained only for brief periods (less than
20-30 seconds) and are terminated by the exhaustion of extracellular energy supplies
or by accumulation of waste products.
The capacity for high speed swimming for short periods of time is important for many
fish species. Acceleration involved in fast-starts from rest, high speed maneuvers, or
speed changes from one steady speed to another is an integral part of swimming. The
former two propulsive patterns involve high rates of acceleration and are important in
prey capture, escape from predators (Hunter 1972), and in the successful negotiation of
such obstacles as waterfalls and fishways (Katopodis 1994 ). A few species like salmon
use high speed bursts to swim up or leap otherwise impassable water. Fish can greatly
reduce energy expenditure by alternating periods of fast swimming and gliding (Weihs
1974; Videler and Weihs 1982). Burst swimming is therefor a vital components of a
fish's locomotory repertoire and of ecological importance.
There are several biological and environmental constraints on fish swimming which
deserve consideration. Early studies suggested fish up to 1 m in length could swim up
to 10 times their body length for a short time (about 1 second), beyond which swimming
speed would decrease exponentially (Bainbridge 1958). Further research suggested
variability in burst swimming among species and, for some, the relative performance
maximum of 10 bl·s· 1 was conservative (Webb and Corolla 1981; Wardle and He 1988).
The maximum speed which fish can achieve is influenced by fish size (Webb 1975)
and body form (Webb 1978; Taylor and McPhail 1985). Anaerobic metabolism at burst
activity levels is relatively independent of water temperatures (Webb 1978), however,
recent studies indicate that temperature acclimation can effect muscle properties
(Johnston et al. 1990; Beddow et al. 1995). Therefore, biological factors such as size
and species, as well as environmental factors, may affect burst swimming performance.
Dams have been widely constructed on waterways to provide water supply, control
floods and generate hydroelectric power and fishways have been constructed to
facilitate upstream passage for fishes. High flow rates are often associated with
fishway entrances (attraction) and for a fish to progress through a fishway they must be
able to swim at velocities greater than the opposing water currents and generally,
successful passage requires a high swimming speed. Few studies have addressed
high speed (burst) swimming performance in salmonids, particularly in relation to
fishway design. A major component of this research was therefor to assess high
speed swimming performance in migrating anadromous Atlantic salmon (Sa/mo sa far
L.) in an experimental passage structure. In particular, this study was to focus on the
influence of flume velocity on the burst swimming speeds, and behavioural strategy for
passage, of Atlantic salmon during spawning migration.
1-4
New
1.4
Tefemetcjc Approaches To The Assessment OfFish Swimming Pedqrmance
Use of Telemetry to Assess Fish Swimming Performance Within an
Existing Fishway
Numerous waterways throughout North America have employed fish passage
structures and the efficiency of these facilities for successfully passing fish is often
unknown. Data on the rate of ascent of various species of fish through different types
of fishways under varying conditions are lacking (Clay 1995). Typically assessment has
involved annual enumeration of adult migrants, while detailed information on the rate of
movement through the fishways and behavioral patterns within these structures is not
available. Consequently, little published data on the movements and behavior of adult
salmonids through fishways is available for use in fishway design and operation
(Weaver 1963). Previous studies of fishway performance have frequently been
conducted under laboratory conditions (e.g. Slatick 1975; Mallen-Cooper 1992).
Extensive research has also examined the physiology and behavior of fish during
migration and fishway passage (e.g. Schwalme et al. 1985; Slatick and Basham 1985;
Blackett 1987; Monk et al. 1989) but these have not been conducted in relation to a
specific fish passage structure. Other studies have utilized conventional mark-andrecapture estimates or timed survey counts which provide no detailed information on
behavior or the movement patterns of fish within a structure. The utility of these
methods depends on species, their habits and habitat, hydraulic conditions (water
turbulence), water clarity, and diel pattern of movements. Timed entry and exit counts
provide relative comparisons of entry and passage at a point in time but not actual
passage rates and energetics associated with traversing a fish by-pass.
Recent advances in telemetry equipment (e.g. Gray and Haynes 1979; Heggberget et
al. 1988; Webb 1990; BagliniE3re et al. 1990; Arnekleiv and Kraab0l 1996) have
provided detailed analyzes of fish movements. Many radio tracking studies of salmonid
spawning migration have been limited to areas in estuaries or specific areas of rivers
(Trefether and Sutherland 1968; Gray and Haynes 1977; Couturier et al. 1986;
Solomon and Potter 1988). Consequently, an additional component of this study was
directed at applying 'state-of-the-art' telemetry techniques to assessing swimming
performance, behavioural strategy, and energetics for adult Atlantic salmon associated
with passage of an existing fishway at Grand Falls, Newfoundland, during spawning
migration.
1.5
Study Objectives
This research project had two major objectives with the results of each being described
in separate reports (Part A and B).
The primary objective of this study was to determine the empirical swimming capability
(sustained, prolonged, and burst) of juvenile and adult anadromous Atlantic salmon,
1-5
New Te/emefric Approaches To The Assessment OfBsh ,Swjmming Pedarmance
landlocked Atlantic salmon or ouananiche, brook trout, brown trout, lake sturgeon and
walleye. The intention was to use wild (non-hatchery) fish and conduct studies at
ambient temperatures (in some instances controlled temperatures) using common
apparatus (e.g. Blaska swim speed respirometers) and similar experimental protocols.
This was to ensure applicability of research results to practical field applications and to
ensure comparability of results. From this empirical data, criteria were to be developed
relating swimming ability to fish size (length/weight) for each species and for the
temperature ranges studied. Models were to be derived to describe the swimming
ability of each species and/or life stage so that fish passage structures could be
designed to optimize fish passage. Criteria were also to be developed relating
swimming ability to required length of fish passage (i.e. fishway length, culvert length) to
further facilitate practical application of results. The effect of temperature and other
environmental variables, sex, maturity, and influence of migration distance and difficulty
on swimming performance was to be investigated, primarily for upstream migrating
adult Atlantic salmon. Comparison of the swimming ability of sympatric salmon parr
and brook trout were to be conducted to determine if velocity is important in segregating
species. Comparison of swimming performance of anadromous and landlocked salmon
Atlantic salmon were to be conducted to determine if the populations had diverged with
respect to swimming ability. The results of this work are contained in a companion
report (Part A. Swimming Performance of Selected Warm And Cold Water Fish Species
Relative to Fish Passage And Fishway Design).
The second objective of the study was to develop innovative approaches to the
evaluation of locomotory activity, swimming performance, and energetic costs to fish
under naturally occurring conditions in relation to potential barrier problems at natural
barriers (e.g. falls, rapids), hydroelectric installations, culverts, fishways, etc. This was
to involve employing new physiological telemetry techniques, specifically the surgical
implantation of a bio-sensitive radio transmitter (EMG or electromyogram tag) in
individual fish, calibration to locomotory ability and energetic scope (calibration of EMGs
to swimming speed and oxygen consumption), and subsequent use of radio transmitted
EMG signals to assess swimming performance and metabolic costs in situ. New
'state-of-the-art' telemetry systems (DSP or digital signal processing with antennae
switching) were to be used to assess high speed (burst) swimming performance,
behaviour, and migratory strategy in relation to ascent of a fishway and experimental
flume to demonstrate application of the technology to resolution of fish passage issues.
The results of this work are contained in this report (Part B. New telemetric Approaches
to the Assessment of Fish Swimming Performance).
1 -6
New Tetemefric Approaches To The Assessment OfEjsh Swimming Pedocrnance
2.0
MATERIALS AND METHODS
2.1
Study Locations
All study components related to development and application of new methods
employing radio telemetry to assess fish swimming performance in situ were conducted
on upstream migrating adult (bright) anadromous Atlantic salmon (Salmo salar) on the
Exploits River in the Province of Newfoundland and Labrador from 1994 to 1997. The
Exploits River is the longest river and largest drainage basin on the island of
Newfoundland. The river is approximately 267 km long, has a drainage area of 11,272
km 2 , a mean annual discharge of approximately 120 m3s· 1, and drains in a
southwesterly direction to the Bay of Exploits. Other fish species in the Exploits River,
in addition to anadromous Atlantic salmon, include ouananiche (landlocked Atlantic
salmon), brook trout (Salvelinus fontinalis), Arctic charr (Salvelinus a/pinus), American
eel (Anguilla rostrata), and threespine stickleback (Gasterosteus aculeatus). Figure 2-1
identifies the key study locations, including sites for fish capture.
The site of investigation for initial development and application of electromyogram
(EMG) telemetry in 1994 to 1996 was the Fisheries and Oceans' facility at Noel Paul
Brook (49°N latitude, 57°W longitude), in the middle portion of the Exploits River (Figure
2-1 ). This site was ideally situated with respect to accessing experimental animals, had
excellent fish holding facilities and accommodations for the investigators, and was close
to hydroelectric and other facilities where in situ experiments will be conducted. Adult
Atlantic salmon were collected from the Exploits river and transported back to Noel Paul
Brook at regular intervals during the season. Adult salmon were collected from 5
locations on the Exploits River, corresponding to various stages in their upstream
migration. These sites included (1) at the salt/fresh water confluence (1 0 km north of
Jumper's Brook (approximately 49°07.38' N latitude, 55° 18.61' W longitude), (2) above
the estuary (approximately 49°01.90' N latitude, 55° 24.35' W longitude), (3) the Bishop
Falls fishway (approximately 49° 00.85' N latitude, 55° 28.30' W longitude), (4) the
Grand Falls fishway (approximately 48° 55.60' N latitude, 55° 40.22' W longitude), and
(5) the Red Indian Lake fish elevator (approximately 48° 45.76' N latitude, 56° 35.96' W
longitude). These sites were located approximately 0, 2, 10, 25, and 110 km from salt
water, respectively.
The development and application of telemetric methods for assessment of fish
swimming performance within fishways was conducted at a vertical slot fishway located
at Grand Falls (48° 55.60' N latitude, 55° 40.22' W longitude) on the lower Exploits River
(Figure 2-1 ). Salmon ascending the river must negotiate a stretch of turbulent "white
water'' and a passable lower falls before reaching this fishway.
2- 1
New Telemetric Approaches To The Assessment OfRsh Swimming Periormance
2.2
Physiological Telemetry Studies
2.2.1 Experimental Animals
Developmental work on EMG signals from adult Atlantic salmon were conducted on fish
collected from the Exploits river and transported back to Noel Paul Brook at regular
intervals during the summer/fall season. Wild adult Atlantic salmon (55-60 em) were
collected between 15 July and 10 October, 1995. Collection occurred between 21 and
27 July when ambient water temperatures averaged 18 °C and 27 September to 3
October when water temperatures averaged 12 °C. Animals were transported using a
1000 liter transport tank supplied continuously with oxygen (1-3 psi) delivered through 1
meter of Micropore (TM) tubing. At Noel Paul Brook, fish were held in large
rectangular outdoor pens (8 m wide, 15 m long and 80 em deep) and indoor cement
upwelling incubation boxes. All animals were allowed to recover from transportation for
three days prior to swim speed trials and/or surgery. Water temperature was
measured daily but was not controlled such that all fish were exposed to ambient river
temperatures.
Studies on muscle activity of wild Atlantic salmon were conducted on fish ranging from
1.0-2.6 kg weight collected from the Exploits River during their spawning migration (May
261h to October 1]lh ) in 1996. Collection of salmon was based on the peak migration at
five sampling locations ranging from the point of freshwater entry (estuary) to spawning
(Figure 2-1 ). Salmon were collected from the estuary using a tended small mesh gill
net. The net was monitored constantly and entangled fish were typically removed
within 4 minutes and immediately placed in fresh aerated river water. Collection of fish
from upstream sites involved dip-netting individuals from traps located at fishways
(Bishop falls, Grand falls) and elevator (Red Indian Lake), located 5 km, 20 km and 100
km, respectively from the river mouth. Pre-spawning individuals were collected from
site 4 approximately 2 weeks prior to spawning and taken to Noel Paul's Brook (site 5)
where they were held in an artificial spawning channel until the onset of spawning. Prespawning behavior was signified by the initiation of redd making activity by female
salmon and later confirmed using hormone analyzes and measurements of gonad
weight.
2.2.2 Electromyogram (EMG) Transmitter and Telemetry Equipment
Activity of the muscle was measured using radio-transmitted electromyogram or EMG
signals at various swimming speeds in preparation for in situ studies (Figure 2-2).
Transmitters measured 50 mm in length and 13 mm in diameter and weighed 8.5 g in
water (less than 2% of the experimental animal's body weight). The EMG from
contracting muscle were detected via 18 carat gold tip sensors and transmitted along
2-2
New Telemetric Approaches To The Assessment OfBsh Swimming Pedarcnance
insulated stainless steel electrodes. A precision half wave rectifier and integrator
processed the input EMG signals within the bandwidth 30-350 MHZ. EMG signals were
processed through an integrator and a radio pulse corresponding to the pulse interval in
milliseconds (ms) was transmitted when the integrated EMG (i.e. EMGi) equaled the
predetermined threshold value of 150 J.A-V. Increasing muscle activity resulted in a
corresponding decrease in the interval between successive radio pulses. Transmitters
were designed to broadcast at frequency intervals of 10 kHz within an operating band
of 148 to 150 MHZ. Lifespan (battery life) of the transmitters was approximately two to
three months. The receiver and transmitter were programmed to continuously monitor
in situ muscle activity on 1 second intervals. Transmitted signals were detected and
recorded automatically using a SRX_400 radio receiver/ data logger (Lotek Engineering
Inc., Newmarket, Ontario) and downloaded to a laptop computer via an RS232 serial
communication port. The receiver was programmed to record all EMGi signals from
each tagged individual.
2.2.3 Surgical Procedures
Prior to experimentation, a protocol for the implant and measurement of EMG signals
was developed for work on Atlantic salmon. This protocol was designed for salmonid
species and is based on the technique used by McKinley and Power (1992) for lake
sturgeon. Individual Atlantic salmon were removed from a common enclosure and
anaesthetized in an aerated and buffered solution of MS-222 (50-75 mg·l-1, pH 7.0).
When a slow irregular operculum rate was observed, surgery was initiated. This stage
of anaesthesia was generally reached within four minutes. Anaesthetized individuals
were placed ventral side up onto a non-abrasive V-shaped surgical table and the gills
were irrigated with fresh oxygenated water.
The transmitter was implanted into the body cavity of fully anaesthetized adult salmon
via a 3 em incision and located such that it would rest above the pelvic girdle (Figure 22). The transmitter was gently inserted through the incision and pushed anteriorly into
the body cavity. Electrodes were positioned approximately 5 mm apart in the lateral
muscle using 21 gauge rods. Once the electrodes were secure, the rods were
removed. The antenna was then placed through the body wall and was allowed to trail
outside the animal. The incision was closed using three independent sutures (2/0
Ethicon silk) and, prior to the last suture, an antibiotic (Liquamycin LP 1 ml·kg-1} was
injected intraperitoneally. Surgical time averaged 4-5 minutes. In the present study, 4
male and 4 female salmon were collected from each site and used for assessment of
muscle activity. The fish was allowed to recover for a minimum of two weeks prior to
experimentation.
2-3
New Telemetric A,apmaches To The Assessment OfFish Swimming Pec(ormance
2.2.4 Calibration of EMG Signals with Swimming Speed
Muscle activity obtained from Atlantic salmon was correlated against sustained and
prolonged swimming speeds. Muscle activity of salmon was expressed using an
activity index (pulse interval in ms) based on previous studies of muscle activity in lake
sturgeon (McKinley and Power, 1992).
Swimming performance trials were conducted (at least one week after surgery) using a
Blaska-type swim speed chamber/respirometer (Biaska et al. 1960). This type of swim
speed chamber is characterized by a tube within a tube design which permits water to
be circulated within a smaller volume compared to Brett-type tunnel swim speed
chambers (Brett, 1964 ). Ambient, untreated river water was pumped into the chamber.
Water velocity within the chamber is rheostatically controlled and the maximum water
velocity could be obtained within 5 seconds. Maximum cross sectional area of salmon
used in swim trials was measured to ensure that adjustments could be made for the
influence of the fish's body on water flow within the chamber, if necessary (Smit et al.
1971 ).
Endurance was measured at velocities just above the maximum sustainable swim
speeds at 12 °C and 18 °C. Tests involved placing individuals into the chamber for an
acclamation period of one hour after which the speed was immediately adjusted to the
test velocity and time to fatigue noted. Five individuals were tested at each test velocity
and temperature. Swimming endurance includes three distinct phases of swimming
which Beamish (1978) has previously described as sustained, prolonged and burst.
Sustained swimming endurance was defined as the speed at which salmon could
maintain swimming for 200 minutes. Burst swimming were those velocities maintained
for less than 30 seconds, and the prolonged swimming phase were those speeds
between the sustained and burst velocities. Endurance trials were performed by
swimming individual fish at a known water velocity until fatigue (Beamish 1978).
The swimming performance of both tagged and untagged salmon was evaluated by
measuring critical swimming speeds. The critical swimming speed is a special category
of prolonged swimming and is typically used to indicate the upper limit of a fishes
aerobic swimming capacity (Beamish 1978). The procedure used to determine critical
swimming speeds is described in Beamish (1978). Critical swimming speeds were
investigated using 5 male and 5 female salmon from each site. Before being swum,
salmon were allowed to acclimate to the swim chamber for 3 hours at a velocity of 0.5
bl sec- 1• Depending on the nature of the study, and fish being tested, different criteria
can be used to determine critical swimming speeds. In the present study, salmon were
subjected to water velocities which were increased by 0.5 body lengths per second at
10 minutes intervals, until fatigue. Critical swimming speeds (Ucrit) were calculated
using the equation presented in Brett (1964 ):
2-4
New Telemetric Approaches To The Assessment OfRsh Swimming Perlormance
equation 2-1
where: Vis the highest velocity maintained for the prescribed period (em s· 1 ), ti is the
time elapsed at final velocity (min.), tii is the time increment (min.) and ui is the velocity
increment (em sec- 1).
Measurements of critical speed and endurance depend on accurate and consistent
recognition and definition of fish fatigue. The procedure used to determine metabolic
fatigue involved stimulating individuals to continue swimming using rapid changes in
water velocity. In all cases, fish were considered fatigued when they failed to leave the
downstream screen despite two to three attempts to stimulate them.
Correlation of salmon muscle activity of Atlantic salmon (N = 4 at each temperature) to
swim performance was determined under forced swim conditions in the swim chamber
described previously. Salmon fitted with transmitters were placed individually into the
swim chamber and swum over the following range of water velocities: 0.8, 1.0, 1.2, 1.4,
1.6, 1.8, 2.0 and 2.2 m.s-1. The relationship between muscle activity and swimming
speed was determined using linear regression analyzes.
2.2.5 In Situ Measurement of Swimming (Muscle) Activity
Experiments were conducted to determine the applicability of measuring in situ
swimming performance of EMG tagged adult Atlantic salmon in an experimental flume.
The flume consisted of a head pond and a tail pond connected by a 20 m long sluice
(30 em wide and 30 em high). The water velocity in the flume could be adjusted
between 1 m sec· 1 and 3 m sec·1 by changing the slope of the sluice and the depth of
water in the head pond. The water velocity in the flume at which muscle activity was
measured was 2.0 m sec· 1• The flume provided information on swimming performance
at high water velocities over a 20 m length under natural conditions.
Atlantic salmon (n=4 at each temperature) were placed into the tail pond and allowed to
acclimate to the enclosure for at least 48 hours before experiments were allowed to
begin. The activity of salmon was measured continuously during the evenings as
salmon were found to most active during this period. Salmon were allowed to ascend
the flume by their own volition, without providing stimulus. Muscle activity in these
salmon was measured at 1 second intervals and continuously recorded.
2.2.6 Statistical Analyzes
All data values were represented as means ± standard error. Linear regression
analysis was used to correlate swimming speed and muscle activity. Unpaired t-tests
2-5
New Te[emetcjc A.a.acoaches To The Assessment OfBsh Swjmming Pedormance
were used to compare swimming endurance of 12 °C fish with that of 18 °C fish. In all
cases, P < 0.05 was the accepted level of significance.
Comparisons between sexes and among sites were accomplished using two-way
ANOVAs. Significance differences were identified using LSMEANS (Steeles and Torrie
1980). The accepted level of significance in all cases was P<0.05. Linear regressions
were used to correlate muscle activity with swimming performance. Linear regressions
were also used to investigate relationships between changes in temperature and crosssectional area with swimming performance and muscle activity. Multiple linear
regressions were used to investigate changes in swimming capabilities and muscle
activity with cross-sectional area, temperature and water velocity (swimming speed).
2.3
Burst Swimming Studies
2.3.1 Experimental Flume
An experimental stream flume, constructed at Noel Paul's Brook, consisted of an
upstream head pond (upper pool, area 6 m2 ) created in the existing sluiceway of a dam
and associated small reservoir and a downstream pond (lower pool, area 17 m2 ) (Figure
2-3). A wooden flume, measuring 18m long by 0.50 m wide and 0.61 m deep, was
situated between these two pools. Water was diverted into the flume through a vertical
sluice gate located in the dam. Discharge into the upper pool was controlled by varying
the number of stoplogs in the sluiceway. Two gates in the stoplog section provided
additional regulation to adjust and stabilize the water level in the head pond. The fish
exit (lower pool) was modified to reduce turbulence, remove standing waves from the
water surface and develop a laminar flow. The flume had a slope of 2% and where it
entered the upper pool its bottom was flush with the floor of the upper pool.
2.3.2 Telemetry Equipment
Radio telemetry equipment manufactured by Lotek Engineering Ltd. (Newmarket,
Ontario) was used to monitor the movements of fish in the flume. Radio transmitters
were cylindrical (tapered to a 0.5 em diameter at one end), 5 em in length and 1 em in
diameter, weighed 8.6 grams in air, and had a 24 em insulated trailing antennae.
Transmitters were encapsulated in waterproof epoxy resin and activated by a magnetic
reed switch. Transmitters were uniquely coded and were designed to broadcast at
frequency intervals of 20kHz within an operating band of 149.54 to 149.74 MHZ with a
battery life of 15 days. The receiving system was programmed to record coded signals
from each fish at 1.25 second intervals.
2-6
New Telemetric A,nprnaches To The Assessment Offish Swimming Perlorroance
Transmitted signals were detected by seven fixed antennae connected to a receiver
(model SRX_400) and digital spectrum processor (model DSP_500). The digital
spectrum processor (DSP) is a receiver/coprocessor capable of providing frequency
discrimination using near real-time spectrum analysis (Table 2-1 ). The DSP was
optimized to simultaneously detect many frequencies and transfer information
concerning pulse arrival times, antenna position and frequencies to the receiver which
performed code discrimination, code error correction and data logging/storage. In
addition, the receiving system was equipped with multiple antennae switching capability
which determined the location of a transmitter relative to the 7 antennae (Figure 2-3).
This was achieved via pulse position code discrimination, in which each radio
transmission is assigned a unique coded time signature. Antennae were simultaneously
scanned every 7.5 msec. The underwater antennae array was placed at equally spaced
intervals along the bottom of the flume ( 0, 2.53, 5.06, 7 .59, 10.12, 12.65, 15.18 m).
Calibration of signal strength permitted determination of distance between transmitter
and antennae, thereby eliminating non-quantitative visual monitoring. This procedure
involved mapping individual antenna reception areas and areas of cell connection
zones down the length of the flume prior to experimentation. The DSP continually
recorded events in real time, provided measurements of transmitter position, thereby
permitting precise spatial and temporal monitoring of fish passage through the flume.
Table 2-1.
DSP_500 and SRX_400 Specifications
General
Size:
Weight:
Operating voltage range:
Operating current:
Operating Temperature Range:
22.0 x 20.4 x 8.8 em
2.5 kg
12VDC
1.5A
-3o•c to +5o•c (LCD:-2o•c)
Electrical
Detection Bandwidth:
Detection Sensitivity:
Dynamic Range:
Dynamic Gain Control Range:
25 discrete frequencies in 20 kHz steps
-115dBm
<100 dB
90dB
Memo
Program Memory:
Data Memory:
128 K
512 K
Controls and 110
Interface:
Controls:
50 ohms BNC input for RFRS-232 (9-pin male) serial
communication port and antenna switch control port
2 external power sockets, 2 RS-232 ports, LCD display, BNC
(50 ohm) antenna jack, 16 key weatherproof keypad
2-7
New Telemetric Approaches To The Assessment OfFish Swimming Pedorroance
2.3.3 Experimental Animals
Wild adult Atlantic salmon (N=75, 1.22 ± 0.42 kg, fork length=51.9 ± 5.2 em, girth=23.7
± 4.8 em) were collected from fishway traps located on the Exploits River watershed
during late August 1995 and July to October 1996. Fish were transported to Noel
Paul's Brook and were held in large rectangular outdoor pens, measuring 8 m long and
12 m wide and with depth ranging from 0.6 to 1 m. The bottom was composed of
gravel substrate similar to that of the riverbed and overhead cover was provided. Pens
were located in a 100 m long channel and river water was diverted through the raceway
by upstream intake valves at a rate of 40 em s· 1 • All animals were allowed to recover
from transport for seven days prior to swim speed trials in the flume.
Salmon were removed from their holding pen 24 h prior to surgery. Prior to transmitter
implantation, fish were immobilized by immersion in an aerated and buffered solution of
clove oil (30 ppm). Anaesthetized fish were placed ventral-side up in a non-abrasive Vshaped surgical table with heads submerged in freshwater. Fish's head and trunk were
kept moist by a cover of pre-soaked towels. Implantation involved inserting the tapered
end of the tag into the urogenital papilla, and gently pushing it anteriorly into the body
cavity of the fish (Peake et al. 1996). The transmitter's antenna was allowed to trail
externally from the oviduct and the implantation procedure averaged one minute.
When size and volume of transmitter were compared to that of the fish (tag to fish
weight ratio was less than 1% ), stress following insertion was considered negligible
(Solomon and Storeton-West 1983; Heggberget et al. 1988).
After sufficient recovery in freshwater, fish were introduced into the lower pool of the
experimental flume. They were allowed a 24 h acclimation period at which point the
gate to the flume entrance was opened and the head elevation in the upper pool was
randomly raised to one of five water velocities (subgroups). Water velocities used in the
summer (1.61, 1.89, 2.30, 2.79, and 3.09 m·s- 1 ) were slightly lower than those
measured in the fall (1.70, 1.99, 2.33, 2.84, and 3.20 m·s- 1) owing to low water levels
experienced in August 1996. The number of fish that successfully negotiated the flume
depended on the volitional swimming of individuals as there was no attempt to force
fish to swim. Wildlife problems (otter predation) complicated one swimming trial while
technical problems (power/recording failures) affected three other trials. These fish
were subsequently removed from analyzes. After 72 hours, tracking data from the
digital spectrum processor was downloaded to a laptop computer via a RS232 serial
communication port. Tags were removed from individuals, cleaned, sterilized with
alcohol and reused. Fish were then measured for fork length, girth and wet-weight and
transferred to separate holding enclosures, grouped by treatment (water velocity), and
observed for abnormal behaviour and mortalities for 14 d post recovery period.
2-8
New Tetemetric Approaches To The Assessment QfBsh .Swimmjag Pedocmance
2.3.3.1
Physiology and Blood Collection
The level of anabolic glycolytic potential experienced by fish ascending the flume was
determined by measuring the level of plasma lactate enzymatically using the L-lactate
dehydrogenase (LDH) method (Lowry and Passonneau 1972). Since fish struggled
when removed from the flume for blood collection, lactate measurements obtained this
way may not truly reflect the levels of exercise and stress associated with traversing
high velocities. To overcome this concern, individuals were anaesthetized prior to
sampling as this approach has been shown to reduce any metabolic and acid-base
changes associated with handling (Tang and Boutillier 1991 ). Salmon that had
ascended the flume at high water velocities (n=1 0, fall fish only) were confined in the
upper pool for 2 h. Previous studies had shown that blood lactate levels are
substantially elevated within 5 min of strenuous exercise and that acidosis is most
severe during the 2 h immediately following exercise (Wood et al. 1983, Tufts et al.
1991 ). After 2 h, the sluiceway was closed and fish in the upper pool were immobilized
by adding a buffered solution of MS-222 (0.25 g·L- 1 ) to the enclosure.
Fish were then placed ventral side-up in a foam trough and bled via a caudal puncture
within 1 min of handling. Blood was immediately transferred to a heparinized
Eppendorf tube and then centrifuged for 2 min. Plasma (500 ,uL) collected after
centrifugation was quickly pipetted into another 1.5 ml tube with an equal volume of
perchloric acid (8% PCA). Samples were subsequently frozen in liquid nitrogen for
transport and then stored at -20 °C until analysis. Plasma samples were analyzed in
neutral PCA extracts and appropriate metabolic standards (Sigma Diagnostics) were
used to complete assays (Lactate Procedure 826-UV).
Control fish (n=5) were sampled in a similar manner as described above to obtain
resting values for lactate. Individual control fish were maintained quiescent in well
aerated, covered enclosures for 12 to 24 h prior to sampling. Prior to blood collection,
anaesthetic was introduced into the enclosures. A blood sample was then collected
from individuals as described above.
2.3.4 Flume Hydraulics
The approach used to quantify flow characteristics in the experimental flume was to use
the head pond elevation upstream of the flume. The hydraulics could then be described
by rating curves and equations relating upstream headpond elevations with the
discharge through the flume at a constant slope (Figure 2-4 ). Upstream head elevation
was fairly stable and was continually monitored during swimming tests. Exceptions to
this were noted on two occasions in which rapidly changing weather conditions altered
both the water level in the brook and lower pool. Water velocities and depths were
measured at sixt~en transects equally spaced along the flume's length and at the
2-9
New Te[emetric Approaches To The Assessment OfRsh Swimming Pedormance
Centre and sides of each transect. Velocity measurements were taken with a twodirectional electromagnetic current meter (Marsh-McBirney, Model 2000) positioned 3
em above the flume floor, 3 em below the water surface and at mid depth (0.6 times the
water depth below the surface). Mean mid depth measurements at the centre transects
were taken to represent the average velocity through the flume. Depth measurements
were recorded to the nearest mm at every transect in order to determine flow rates and
develop surface profiles in the flume.
2.3.5 Data Analyses
Fish movement through the flume was monitored continuously for the duration of the
study excepting a seven day period when heavy rains resulted in excessive discharges
beyond the scope of the study (3.6 m3·s- 1). As individuals entered the flume, their rate
of ascent against the corresponding water velocity was recorded as the distance
traveled over time, or ground speed ((Vf, ms- 1 ). Distance was determined from power
strength signals as obtained from the DSP and SRX receiver. Using velocity profile
equations, (Vw), the fish's swim speed with respect to water velocity (Vfw, in m·s-1) was
calculated as the mean of the ground speed and water velocity at each transect using
the following equation:
equation 2-2
2.3.6 Statistical Analyses
The effects of varying water velocities on kinematic variables (distance, time, average
and maximum ground speed, average and maximum total speed) were analysed in a
model-1, randomized design, one-way analysis of variance (ANOVA) with individuals as
a random factor and five test water velocities. AN OVA tests incorporated contrasts for
polynomial trends, specifically linear and non-linear trends. Prior to analyses, residuals
for each dependent variable were examined graphically and tested for normality and
homogeneity of variance and were subsequently log transformed within comparison
groups if these assumptions were not met. One way cross tabulations, standardized as
percentages with 95% confidence intervals, were completed on success rates of
passage and time periods for ascents at each water velocity. Seasonal water
temperatures (daily minimum, maximum, and mean), collected from hourly thermograph
recordings, were entered as covariates and an ANCOVA analyses was performed.
Plasma lactate values of 10 salmon who had ascended the flume at high water
velocities were compared to controls (n=5) using a Mann-Whitney U test (one-way). All
values were represented as means± standard error. Statistical analyses were
performed by SYSTAT 6.0 and tests were considered significant at an alpha level of
0.05.
2- 10
New Telemetric Approaches To The Assessment OfEjsh Swimming Pedarmance
2.4
Assessment of Swimming Performance in a Fishway
2.4.1 Fishway Description
This aspect of the study was conducted at a fish by-pass structure (vertical slot fishway)
on the lower Exploits River at Grand Falls (48°85'N, 55°90'W), about 40 km upstream
from the estuary. Salmon ascending the river must negotiate a stretch of turbulent
"white water" and a passable lower falls before reaching the fishway. Additionally fish
are exposed to secondary treated process industrial and municipal effluents from a
thermo-mechanical pulp mill (Abitibi-Consolidated) and sanitary sewer outfalls. At this
location a power dam and a natural waterfalls, with a combined height of 27 m, had
created a complete blockage to upstream salmon migration.
The fishway consists of a 116 m long x 2 m wide x 2 m deep reinforced concrete
channel (Figure 2-5). The channel in the fishway extends downstream, from the bottom
of the falls and top of the dam, and doubles back so that its entrance is close to the
face of the waterfall where fish congregate. The fishway consists of a series of 33
baffles each containing a tall narrow slot, 35 em wide by 1. 7 m long beginning 30 em
above the channel bottom. The lower section of the fishway consists of 19 pools and
ends in a collection facility. The calculated operational flows of the fishway range from
0.5 m3s·1, with a pool depth of 0.91 m, to 0.9 m3s·1, with a pool depth of 1.7 m. The
upper section of the fishway consisted of 10 pools and ends in a deep exit tank. The
calculated operational flows for this section range from 0.5 m3s·1 , with a pool depth of
0.91 m, to 1.2 m3s·1 , with a pool depth of 1.2 m. After ascending the fishway and
passing through an underwater viewing tank located near fishway exit, fish exited along
the north-shore through a removable barrier screen at the final slot opening. Some
descriptive characteristics of the fishway are given in Table 2-2
Table 2-2. Physical and hydraulic characteristics of Grand Falls fishway.
Parameters
Lower section
Upper section
Slope (m/m)
1:8
1:8.8
Inside pool length (m)
2.74
3.10
inside pool width (m)
1.83
2.40
1.52- 1.83
2.16
pool spacing (m)
3.05
3.30
drop per pool (em)
30.5
30
sill depth (em)
31
30
slot width (em)
31
38
maximum pool depth (m)
2- 11
New Telemetric Appcaaches To The Assessment Of Bsh .Swimming Performance
To traverse the fishway, fish must swim a distance of about 15 m against high water
velocities. The total distance of high velocity flow through each of the vertical slots is
about 45 em. Water velocities in the fishway were measured with an electromagnetic
digital flowmeter (Marsh McBirney, Model 201 D). The maximum water velocity that was
measured in the fishway occurred about 20 em downstream of each vertical slot, where
water flow is constricted to form a jet (vena contracta) into the next pool. During the
study period, daily water velocities (m·s· 1 at 0.6 depth) at several slots within the fishway
were recorded as the average of at least three readings per day. The median of the
daily water velocities and the upper and lower limits which contain 75% of the daily
water velocities between them (37.5% above and below the median) were calculated.
The fishway was studied under natural operating conditions and the water depth varied
between 55 and 85 em.
The water supply was from the natural river so temperature and dissolved oxygen were
not controlled and were monitored several times daily. Over the period September 7 to
24, 1996, water temperatures were fairly constant at 11.5 ± 0.48 °C and dissolved
oxygen was between 77 and 89% saturation.
2.4.2 Experimental Animals and Transmitter Attachment
Fish used were actively migrating adults (55.3 ± 3.8 em; mean weight 1.51 ± 0.24 kg)
collected from traps and pools below the Grand Falls dam during September, 1996.
Twelve Atlantic salmon (5 males, 7 females) were equipped with external coded radio
transmitters to enable their movements to be tracked. Prior to transmitter attachment,
fish were quickly anaesthetized in 30 ppm clove oil and ethanol solution following the
procedure outlined by Anderson et al. (1997). Once anaesthetized, fish were placed
into a V-shaped surgical table filled with oxygenated water. Oxygenated water was
directed across the gills through a tube inserted in the mouth and the individual's head
and trunk was kept moist by a cover of pre-soaked towels throughout the procedure.
External transmitters were attached anterior to the dorsal fin using two steel wires and
the transmitter's antenna wire was left hanging externally. After tagging, the fish were
kept in oxygenated water for 0.5 to 2 h to allow recovery before release. Once fish
regained equilibrium and resumed swimming, they were released. Fish were
transported downstream of the fishway entrance (approx. 15 m) by submerging the
recovery tank on the river bottom's substrate and carefully releasing the fish without
taking them out of the water. Salmon subsequently moved their way up the fishway on
their own volition.
2- 12
New Telemetric A,a,aroaches To The Assessment Offish .Swimming Pedorcnance
2.4.3 Digital Spectrum Processing (DSP) Telemetry Studies
Radio telemetry equipment, manufactured by Lotek Engineering Inc. (Newmarket,
Ontario), was used to monitor the movements of fish in the fishway. Radio transmitters
were cylindrical, 20.4 mm in length and 9.9 mm in diameter, weighed 2.1 grams in air,
had a 30 em insulated trailing antennae (model CFRT-3GM). Transmitters were
encapsulated in waterproof epoxy resin and activated by a magnetic reed switch.
Transmitters were uniquely coded and were designed to broadcast at frequency
intervals of 20kHz within an operating band of 149.56 to 149.66 MHz with a battery life
of about 20 days. The tag weight to fish weight ratio was less than 0.01 %. The
receiving system was programmed to record transmitted signals from each fish at 1.25
second intervals.
Transmitted signals were detected by seven fixed antennae stations connected to a
receiver (model SRX_400) and digital spectrum processor (model DSP_500). Similar
to burst swimming research, the receiving system was equipped with multiple antennae
switching capability which determined the location of a transmitter relative to seven
antennae. Antennae were simultaneously scanned every 7.5 msec which allowed all
fish and their corresponding codes to be tracked simultaneously. The antennae array
(coaxial cable with appropriate lengths stripped of shielding) was placed at equally
spaced intervals along the bottom of the fishway (Figure X). The receiving and
processing system were checked for malfunctions and settings against test transmitters
in early morning, mid afternoon, and late evening each day. Calibration of signal
strength permitted determination of distance between transmitter and antennae,
thereby precisely positioning fish in the pools of the fishway. Thus the DSP continually
recorded events in real time, and by providing estimates of transmitter position
monitored fish passage through the by-pass structure.
2.4.4 Electromyogram (EMG) Telemetry Studies
Additionally, migrating adult salmon (n=1 0) had EMG transmitters surgically implanted
as per the procedure previously described (Section 2.2.3). After sufficient recovery, fish
were placed downstream of the fishway entrance and allowed to volitionally enter the
fishway. EMG signals were recorded continuously in the lower 11 pools and the first
holding pool (Figure 2-6).
2.4.5 Statistical Analyzes
Data for response variables (time in antennae areas, total time in fishway, number of
entrance attempts) were tested for normality and homogeneity-of-variance by graphical
examination of the residuals. Log transformation of the raw data did not remove
2- 13
NeW Telemetric Approaches To The Assessment OfBsh Swimming Pedarmance
violations of the above assumptions, therefore the Friedman's test, a non-parametric
analog of a repeated measures analyzes of variance (one-way) was used. All data
filtering and statistical analyzes were conducted with SYSTAT 6.0, and significance was
determined at P <0.05.
2-14
New Telemetric A,a.ornaches To The Assessment OfRsh Swimming Perlorroance
3.0
RESULTS
3.1
Physiological Telemetry Studies
3.1.1 Swimming Performance and Temperature
Swim performance of wild Atlantic salmon was lower at 12 °C than at 18 °C (Figure 31). Maximum sustained swimming speed decreased by 1 m·s· 1 as temperature
decreased from 18 °C to 12 °C. Sustained swimming speed of salmon at 12 °C was
also significantly lower than at 18 °C. At burst velocities, swimming performance was
independent of temperature, as indicated by the inability of salmon to maintain swimming speeds in excess of 2.40 m·s· 1 for periods greater than 10 s at either temperature
(Figure 3-1 ).
Critical swimming speed (Ucrit) of salmon at 18 °C and 12 °C was 2.16 ± 0.18 m·s·1 and
1.76 ± 0.06 m·s·1, respectively (P < 0.05). Critical swimming velocities of untagged
individuals were found to be 2.16 ± 0.18 m·s·1 at 18 °C and 1. 76 ± 0.06 m·s· 1 at 12 °C,
while those of tagged salmon were 2.10 ± 0.05 m·s· 1 at 18 °C and 1.80 ± 0.03 m·s·1 at
12 °C. There were no statistical differences between tagged and untagged salmon at
either temperature (P < 0.05).
3.1.2 Relationship of Muscle Activity to Swimming Speed
Regression analyzes indicate that muscle activity was positively correlated to swimming
performance at both temperatures (Figure 3-2). The relationships between muscle
activity and swimming speed are described by the following equations:
muscle activity= 2,521.1 - 7.51 x swimming speed, R2 = 0.85, at 18 °C,
equation 3-1
muscle activity = 2,330.8 - 4.29 x swimming speed, R2 = 0.91, at 12 °C
equation 3-2
where, muscle activity is in milliseconds and swimming speed is in m·s· 1• Temperature
dependent differences in the muscle activity relative to swimming speed were only
apparent beyond the critical swimming speeds indicated by the divergence of the
regression lines. The relationship between swimming speed and muscle activity was
significantly different (slope 7.51) at 18 °C than at 12 °C (slope 4.51 ). The greater slope
of the relationship at 18 °C indicates that, at warmer temperatures, greater muscle
activity results in higher overall swimming speeds.
3-1
New Telemetric Approaches To The Assessment OfBsh .Swimming Pedarroance
3.1.3 Correlation of Oxygen Consumption and Swimming Speed
Oxygen consumption was highly correlated with muscle activity at both 18
as described by the following equations:
oc and 12 °C,
oxygen consumption = 1195.7 - (0.582*muscle activity), r-2=0.96 at 18° C
equation 3-3
oxygen consumption = 950.5 - (0.425*muscle activity), r-2=0.91, at 12° C
equation 3-4
oc
The oxygen consumption of salmon at 12
was markedly lower than at 18 °C, and
may reflect the lower aerobic muscle activity of Atlantic salmon at the colder
temperatures (Figure 3-3).
3.1.4 Effect of Body Morphology
Salmon possess an elliptical body shape with a dorsal ventral height ranging between
10 and 14 em and a lateral width ranging between 9 and 12 em, corresponding to cross
sectional areas ranging from 58 to 66 cm 2 • According to Smit et al. (1971), fish with
cross sectional areas greater than 10% of the swim chamber must have swimming
speeds corrected for influence of body size. Using the following equation from Smit et
al. (1971 ), 'blocking effect' of salmon in this study would increase the actual swimming
velocity by no more than 0.04 for the smallest salmon and 0. 12 m·s- 1 for the largest.
Therefore, conservative estimates of swimming performance were used and no
correction was made for cross sectional area.
equation 3-5
Ucorrected = Umeasured (1 + Areatish/Areachamber)
where, velocity is measured in m·s- 1, and U is the critical swimming speed (m·s- 1) of the
corrected and actual measurement, respectively.
3.1.5 Muscle Activity During Spawning Migration
Figure 2-1 shows the sampling sites and times of salmon collection from the Exploits
river. Temperature increased during the initial 10 weeks of sampling after which time it
declines until spawning. The highest temperature was observed during July and was
22 °C while the lowest temperature observed was 8.8
just prior to spawning.
oc
The relationship between muscle activity and water velocity (swimming speed) changes
significantly during the migratory period (Figure 3-4 ). Mean slopes did not differ
between individual male or female salmon at any site during the initial 16 weeks of
3-2
New Tetemetrjc Anrrnaches To The Assessment Offish Swjmmjng Peiformance
migration (site 1- 4 ). However, prior to spawning, a significant increase in the slope of
the relationship between muscle activity and velocity was observed in both males and
females (Figure 3-4.) The slope of the relationship was significantly greater for prespawning salmon than for salmon sampled 4 weeks prior to spawning, and was
significantly higher for females than males (site 4: females: -5.744 ± 0.39 male -4.28 ±
0.27; site 5: females -13.73 ± 1.05 males -9.62 ± 0.51 ).
Muscle activity indices from resting salmon did not differ between sites and sexes
throughout the migratory period (Figure 3-5). In contrast, significant increases in
muscle activity indices were observed between males and females swum at 1 bl· sec·1
(Figure 3-5). Swimming at 2 bl· sec· 1 resulted in the most noticeable change in muscle
activity between sites as well as between sexes (Figure 3-5). Muscle activity indices
recorded from female salmon just prior to spawning (site 5) and swimming at 2 bl· sec·1
were significantly greater than those of females 4 weeks prior to spawning.
Relationships between muscle activity and temperature and cross-sectional area are
show in Table 3-1. Muscle activity of both males and females increased with swimming
speed and the effects of temperature on muscle activity and swimming speed became
more pronounced at higher velocities (females: 1 bl· sec· 1 , R2=0.32 vs. 3 bl· sec·1 ,
R2 =0.64; males: 1 bl· sec· 1 , R2 =0.19 vs. 3 bl· sec·1 , R2 =0.63). Cross-sectional area and
swimming speed are important determinants of muscle activity for female salmon
(R2 =0.53 P=0.027) but not male salmon (R2 =0.01, P=0.758). Among females, the
relationship between cross-sectional area and muscle activity is stronger than the
relationship between temperature and muscle activity for any given swimming speed
(Table X). Moreover, the relationship between cross-sectional area and muscle activity
is stronger when female salmon are swum at higher velocities (R2 =0.74, P<0.001) than
at lower velocities (R2 =0.50, P<0.01 ).
Table 3-1.
Multiple squared regression coefficients and significance levels from
comparisons of girth and temperature with muscle activity measured
during swimming at 1 and 2 body lengths per second from wild Atlantic
salmon collected during their spawning migration.
female
male
activity(1 bl}
temperature
R2 =0.62, P<0.001
cross-sectional area
R2=0.71, P<0.001
temperature
R2=0.49, P=0.002
cross-sectional
R2=0.02, P=0.563
activity(2bl}
R2 =0.64, P<0.001
R2=0.74, P<0.001
R2 =0.63, P<0.001
R2=0.14, P=0.020
3.1.6 In Situ Muscle Activity (Experimental Flume Study)
Adult salmon exhibited two distinct patterns of swimming behaviour during ascent of the
3-3
New Telemetric Approaches To The Assessment OfFish Swimming Periarcnance
flume. At 18 °C, swimming was continuous and characterized by a constant increase in
muscle activity throughout the ascent. In contrast, at 12 °C, the swimming behaviour of
salmon was characterized by a rapid increase in muscle activity (within 10 s) to above
Ucrit levels, which then remained elevated throughout the ascent (Figure 3-6). At 18 °C,
ascent of the experimental flume by salmon was achieved within the salmon's aerobic
limits and below its critical swimming speed, indicating that an oxygen debt may have
been incurred during ascent.
Overall, salmon traversed the fishway significantly faster at 12 °C than at 18 °C,
requiring 31 ± 7 sand 47 ± 12 s, respectively. Part of the reason for the quicker ascent
at 12 °C is that fish swim at higher velocities. From equation X and X, the respective
mean swimming speed of salmon at 18 °C is 1.4 m·s- 1 while the mean at 12 °C is 2.87
m·s- 1 • As we could not use discreet positional telemetry, some of the activity recorded
from salmon may be while fish were holding position. During these periods, swimming
speeds may be high, while ground speeds are low or zero.
3.2
Burst Swimming
All salmon survived the transmitter attachment and appeared to rapidly recover from the
tagging and handling procedures. Mortality during swimming trials and the 14 d post
experimentation period was nil. Two of four fish that did not attempt to traverse the
flume at 22 °C (see 3.2.3) were later observed ventral side up with slow opercular
movements. These fish were removed form the flume and sacrificed for necropsies.
Expulsion rates of the transmitters from the urogenital papilla was also very low at 6. 7
% ( 5 of 75 attachments).
3.2.1 Chemical, Temperature and Flow Conditions
Chemical analyzes of water samples from the collection and experimental sites were
similar (median pH=6.5, average conductivities ranged from 20 to 35 ~S·cm- 1 and total
hardness=?.? ± 1.1 mg·L- 1). Generally, average conductivity was low as were
concentrations of nutrients and major anions and cations.
In 1996, swimming speed trials were performed during two seasons; mid-summer and
late fall. Summer water temperatures ranged from 14.1 to 25.6 °C, with a mean of 19.2
± 2.0 °C. Diel variation ranged from 0.8 to 4.9 °C. A rapid decline in temperatures
occurred mid-September owing to a heavy snowfall event which also resulted in a rapid
increase in discharge in the brook. No swim trails were conducted during this period.
Fall water temperatures varied from 5.5 to 12.9 °C with a mean of 9.3 ± 3.0 °C. Diel
variation ranged from 0.5 to 3.0 °C during the fall period. The temperature regimen
during the study period is provided in Figure 3-7.
3-4
New Tetemetrjc Approaches To The 4 ssessment Of Fish .$wjmmjng Pedqrmance
Water surface profiles for flows during fish tests are shown in Figure 3-8. Small
standing waves were evident as flows were near critical (Froude number was estimated
at 0.8 to 1.2). At the greatest water velocity tested, flow entered the flume as a
turbulent jet concentrated around the central areas of the flume. However, within 1 m
of the upstream end, the flow had become laminar and the distribution of velocity
across the structure was fairly uniform. Several fish were observed swimming up the
flume with their bodies submerged in the water column.
3.2.2 Timing of Ascent of Flume
Using seasonal water temperatures (daily maximum, minimum and mean) as
covariates, there were no significant effects for temperature on any of the
aforementioned kinematic variables (P > 0.2). Salmon monitored during the summer
period traversed up the flume primarily during twp periods: early morning (36.7 %) and
late afternoon (20.7 %). Late morning (134.3 %) and early afternoon (16.7% periods
were of secondary importance. Similarly, fall fish ascended the flume during the late
morning (22.2 %) and late afternoon (40.0 %). Early morning (11.1 %), early afternoon
(13.3 %) and evenings (13.3 %) were of secondary importance. During both seasons,
night activity was negligible (Figure 3-9).
3.2.3 Success Rates
A significant inverse relationship existed between success rate (of passage) and water
flows in both summer and fall fish (F 1•25=56.24, P<0.0001; F 1•25 =31.34, P<0.0001,
respectively; Figure 3-10). Fall fish were highly successful (83.3 to 71.4 %) in
traversing the entire length of the flume at relatively low velocities ( 1. 70 to 1.99 m·s·1,
respectively). A dramatic reduction in success (22.2 % to nil) was apparent at higher
velocities (2.33 to 3.2 m·s·1, respectively). Summer fish displayed a similar pattern with
all fish successfully ascending the flume at 1.61 m·s·1• However, at a threshold velocity
of 1.89 m·s· 1 , success levels dropped to 57.1 %. At relatively higher velocities (2.3 to
3.1 m·s-1), many fish completed partial ascent of the flume, however no were able to
traverse the entire flume length. At these higher velocities, 4 salmon displayed
swimming activity at the flume entrance however they did not enter of traverse the
flume over the 72 hour test period. Water temperatures during this trial averaged 22
oc.
3.2.4 Swimming Performance Kinematics in Relation to Velocity
In addition to success rates and information on timing of passage, swimming data for a
number of kinematic variables were collected and are summarized for both summer and
3-5
New Telemetric Approaches To The Assessment Offish Swimming Pedormance
fall fish in Tables 3-3 and 3-4, respectively. Swimming performance measurements
collected included data regarding : (i) maximum distance (m) attained during ascent, (ii)
total time (s) required for partial or complete ascent of the flume, (iii) average ground
speed (m·s- 1 , bl·s- 1}, (iv) average fish speed (m·s- 1 , bl·s-1}, (v) maximum ground speed
(m·s-\ bl·s- 1 }, and (vi) maximum fish speed (m·s-1, bl·s- 1 }.
An inverse linear relationship was apparent between maximum distances obtained and
water velocity. As velocity increased, the distances attained declined significantly for
both fall (F 140 =15.4, P < 0.01) and summer (F 121 =8.9, P < 0.05) fish (Figure 3-11 ).
Similarly, th'e total time to traverse the flume was negatively related to water velocity for
both summer (F 1•21 =168.2, P < 0.0001) and fall (F 1•21 =31.8, P < 0.0001) fish. As water
velocity increased, summer fish ascended the flume in approximately 33.5 ±1.3 to 9.0 ±
0.9 s while fall fish required from 22.3 ± 1.9 to 10.0 ± 1.3 s (Figure 3-12).
Average and total ground speed (m·s- 1, bl·s- 1) were positively related to water flows
during both seasons (Figures 3-13, 3-14, and 3-15). Summer fish exhibited significant
increases in average ground speeds from 0.5 ± 0.02 m·s-1 to 1.4 ± 0.4 m·s-1 as water
velocities increased from 1.61 to 3.09 m·s- 1 (F 1.21 =17.4, P < 0.001 ). Fall fish displayed
similar significant increases in average ground speeds from 0.8 ± 0.08 m·s-1 to 4.5 ±
0.3 m·s- 1 as water velocities increased from 1.70 to 3.20 m·s- 1 (F 1•40 =4.5, P < 0.05). As
water velocity rose, average total speeds significantly increased from 4.7 ± 0.1 bl·s-1 to
8.0 ± 0.4 bl·s- 1 in fall fish (F 121 =140.1, P < 0.0001) and from 4.0 ± 0.1 bl·s-1 to 8.8 ±
0.8 bl·s- 1 (F 1 40 =36.5, P < 0.0001) in summer fish. The highest average total speed for
fall fish was'11.7 bl·s- 1 and 10.6 bl·s- 1 for summer · h. Similarly, the highest maximum
total speed for fall fish was 16.3 bl·s- 1 ~nd 10.4 b · -1 for summer fish (Tables 3-3 and 34).
Maximum total speeds (m·s- 1 , bl·s- 1} were positively related to water velocities during
both seasons (Figures 3-16 and 3-17). Fall fish demonstrated significant increases in
maximum total speed from 9.2 ± 0.3 bl·s- 1 to 13.9 ± 0.2 bl·s- 1 (F 1 40 =73.3, P < 0.0001)
as velocities increased. Summer fish also demonstrated significant increases in
maximum total speed from 4.7 ± 0.3 bl·s- 1 to 8.9 ± 0.6 bl·s- 1 (F 1•21 =64.7, P < 0.0001) as
velocities increased. Although maximum ground speeds were positively related to
water flow in summer fish (F 121 =4.37, P < 0.05), fall fish displayed more variation in
speeds attained.
·
Beyond the threshold water velocity of 1.89 m·s- 1, many fish displayed variable average
ground speed and total speeds as a result of taking periodic pauses, before bursting at
a maximum speed to complete successful passage of the flume (Figure 3-18). This
suggests there are also behavioural aspects to 'strategies' used to pass high water
velocity areas.
3-6
New Tefemetrjc A,aprnaches To The Assessment OfFjsh .Swimming Pedocmance
Table 3-3.
Vw= 1.61 (n=8)
Mean±
SE
Minimum
Maximum
Range
Vw=1.89 (n=7)
Mean±
SE
Minimum
Maximum
Range
Vw=2.30 (n=5)
Mean±
SE
Minimum
Maximum
Range
Vw=2.79 (n=5)
Mean±
SE
Minimum
Maximum
Range
Vw=3.09 (n=5)
Mean±
SE
Minimum
Maximum
Range
Summarized swimming data for experimental (summer) 1996 fish grouped
by water velocity (m·s-1). Vf, Vfw, Vfmax, and Vfwmax are reported in m·s·
1
and bl·s- 1 and refer to average ground speed, average total speed,
maximum ground speed and maximum total speed, respectively.
Time
(5)
Vf
(m·s-1 )
Vfw
(m·s- 1 )
Vfw
(bl·s- 1 )
17.55±
0.15
16.50
17.70
1.20
33.50±
1.29
29.00
41.00
12.00
0.53±
0.02
0.43
0.61
0.18
2.14±
0.02
2.04
2.22
0.18
4.02±
0.12
3.61
4.58
0.97
0.89±
0.16
0.35
1.92
1.57
2.50±
0.16
1.96
3.53
1.57
4.70±
0.32
3.59
6.39
2.80
17.08
0.46
14.40
17.70
3.30
20.71
1.37
17.00
26.00
9.00
0.84
0.06
0.66
1.04
0.38
2.73
0.06
2.55
2.93
0.38
5.17
0.16
4.69
5.91
1.22
1.49
0.15
1.08
2.02
0.94
3.38
0.15
2.97
3.91
0.94
6.40
0.31
5.45
7.32
1.87
13.40
1.39
9.50
16.00
6.50
14.00
0.71
12.00
15.00
3.00
0.98
0.14
0.63
1.33
0.70
3.28
0.14
2.93
3.63
0.70
6.11
0.36
5.12
6.88
1.76
1.55
0.14
1.10
1.98
0.88
3.85
0.14
3.40
4.28
0.88
7.30
0.28
6.61
8.27
1.66
14.73
1.27
13.00
17.20
4.20
12.33
0.67
11.00
13.00
2.00
1.20
0.10
1.00
1.32
0.32
3.99
0.10
3.79
4.11
0.32
7.83
0.08
7.66
7.92
0.26
1.48
0.16
1.05
1.98
0.93
4.27
0.16
3.84
4.77
0.93
8.40
0.42
7.58
9.94
2.35
12.44
3.32
3.20
17.70
14.50
9.00
0.91
7.00
11.00
4.00
1.40
0.42
0.40
2.41
2.01
4.49
0.42
3.49
5.50
2.01
8.84
0.79
7.14
10.59
3.45
1.47
0.29
0.52
2.10
1.57
4.56
0.29
3.61
5.19
1.57
8.90 +
0.58
7.39
10.42
3.02
3-7
Vfmax
(m·s-1 )
Vfwmax Vfwmax
(m·s-1 )
(bl·s-1 )
Distance
(M)
New Telemetric A,npcoaches To The
Table 3-4.
Vw=1.70 (n=7)
Mean±
SE
Minimum
Maximum
Range
Vw--1.99 (n=9)
Mean±
SE
Minimum
Maximum
Range
Vw=2.33 (n=9)
Mean±
SE
Minimum
Maximum
Range
Vw=2.84 (n--16)
Mean±
SE
Minimum
Maximum
Range
Vw=3.20 (n=7)
Mean±
SE
Minimum
Maximum
Range
Assessment Offish Swimming Performance
Summarized swimming data for experimental fall 1996 fish grouped
by water velocity (m·s- 1). Vf, Vfw, Vfmax. and Vfwmax, are reported
in m·s- 1 and bl·s-1 and refer to average ground speed, average total
speed, maximum ground speed and maximum total speed,
respectively.
Distance
(m)
Time
(5)
Vf
Vfw
(m·s- 1 ) (m·s' 1 )
Vfw
(bl·s- 1 )
Vfmax
(m·s-1 )
VfWmax
(m·s-1 )
VfWmax
(bl·s'1 )
16.68±
1.02
11.60
17.70
6.10
22.33±
1.87
16.00
27.00
11.00
0.77±
0.08
0.58
1.11
0.53
2.47±
0.08
2.28
2.81
0.53
4.73±
0.14
4.34
5.27
0.95
1.32±
0.16
0.87
1.98
1.11
4.81±
0.17
4.36
5.38
1.02
9.24±
0.30
8.23
10.12
1.84
16.31
0.96
11.25
17.70
6.45
20.57
2.37
11.00
29.00
18.00
0.88
0.14
0.49
1.61
1.12
2.87
0.14
2.48
3.60
1.12
5.28
0.27
4.39
6.37
1.98
1.48
0.25
0.93
2.72
1.79
5.46
0.25
4.91
6.70
1.79
10.03
0.43
8.70
11.87
3.17
15.91
0.55
13.35
17.70
4.35
13.11
1.18
8.00
21.00
13.00
1.28
0.12
0.84
2.06
1.22
3.61
0.12
3.17
4.40
1.22
6.85
0.23
5.96
8.26
2.29
1.42±
0.13
0.88
2.00
1.12
6.08
0.13
5.54
6.66
1.12
11.53
0.24
10.42
12.77
2.35
13.31±
0.82
5.65
12.05
11.75
1.38
5.00
21.00
16.00
1.46
0.23
0.45
3.37
2.92
4.30
0.23
3.29
6.21
2.92
7.78
0.45
6.33
11.67
5.34
1.59
0.15
0.79
2.80
2.01
7.27
0.15
6.47
8.48
2.01
13.15
0.40
10.57
16.31
5.74
11.70
1.29
5.30
14.05
8.75
10-00
1.27
3.00
13.00
10.00
1.33
0.26
0.59
2.82
2.23
4.53
0.26
3.79
6.02
2.23
8.00
0.38
6.79
9.94
3.15
1.44
0.05
1.22
1.67
0.44
7.84
0.05
7.62
8.07
0.44
13.88
0.23
13.04
14.80
1.76
3-8
New Telemetric A(lpcoaches To The Assessment OfFjsh Swimming Performance
3.2.5 Blood Lactate
The plasma lactate values for control (resting) fish and for salmon ascending flume under
high water velocities are provided in Table 3-5. Plasma lactate levels in exercised fish
were significantly higher than for controls (Mann Whitney U-test, P=0.002). Some minor
deviations in plasma lactate levels were evident in the exercised fish and values were not
normally distributed.
Table 3-5.
Summarized plasma lactate values (m·moles·L. 1) for controls and salmon
ascending the flume under high water velocities.
Exercised {in flume}
Controls
10
5
minimum
6.21
0.06
maximum
8.29
0.15
range
2.08
0.09
7.33±0.19
0.10±0.016
N
mean± SE
3.3
Assessment of Swimming Performance in a Fishway
3.3.1 Timing of Ascent of Fishway
Upstream movement through the fishway and entrance were characterized into six time
periods of the day: night (00:01-04:00), early morning (04:01-08:00), late morning (08:0112:00), midday (12:01-16:00), late afternoon (16:01-20:00) and evening (20:01-00:00).
Movements occurred primarily during two time periods, late morning (57.2%) and late
afternoon (35. 7% ). Night movements (7 .1%) were of secondary importance (Table 3-6).
Radio-tracking data showed that many salmon accumulated in the resting pools during
the day periods and into the night and completed passage the following morning. Downladder movements occurred throughout the day.
3-9
New Telemetric A(lpcoaches To The Assessment otBsh $wjmmjng Pedqrmance
Table 3-6:
Time periods of salmon ascent up the Grand Falls fishway.
Period
Night
Real time
00:01 - 04:00
%of fish (N=14)
7.1
Early morning
04:01 - 08:00
0
Late morning
08:01 - 12:00
57.2
Midday
12:01 - 16:00
0
Late afternoon
16:01 - 20:00
35.7
Evening
20:01 -00:00
0
3.3.2 Rate of Passage
The time of passage through each of our seven sections in the fishway was measured
for all fourteen fish. Differences between sexes in body size or shape may contribute to
variability in fishway average passage time. Therefore, we compared passage time
between sexes at each of the seven areas. As no significant differences between sexes
were found (P > 0.30), male and female data were pooled for subsequent analyzes.
There was great variation in the time taken by salmon to travel completely through the
fishway. Upstream and downstream movements were observed throughout the study
period, but most activity occurred during two flow periods over one to two days each.
These two periods were characterized by different hydraulic conditions, primarily varying
water depths and velocities. The mean of the daily water velocities at the beginning of
the study was 1.69 ± 0.07 m·s· 1 (flow period 1), with a water depth of 56 ± 2 em and a
corresponding fish ascent time of 3.33 ± 0.72 h (n = 7, mean± SE., range= 5.17). A
heavy rainfall late into the study caused the river to go into spate and produced higher
daily water velocities and depths, 1.82 ± 0.03 m·s·1 and 79 ± 5 em (flow period 11 ),
respectively. The river was considered to be in spate because both water level and
turbidity showed a marked, and very obvious increase. The total time to traverse the
ladder during this latter period was 27.95 ± 8.86 h (n = 4, mean± SE., range= 38.22).
The total time to traverse the fishway during this flow period was significantly higher than
at the lower flows (Mann-Whitney U-test, P = 0.04 ).
The majority of salmon successfully traversed the entire length of the fishway (66.7%),
however, several fish experienced a 'fall back' pattern in the upper section of the fishway
(Figure 3-19). The time spent in pools was significantly greater at two locations of the
fishway, sections five and seven (Friedman test, P = 0.041, P = 0.03; Figure 3-20). Area
five consisted of two large pools with a 60 em jump at the lower end. Section seven
contained a large fish pool with a 30 em jump into an upstream observation chamber. At
these two locations we observed large areas of quiet water with minimal turbulence. The
3- 10
New Telemetrjc Approaches To The Assessment OfFish Swimmjng Pedacmance
times spent at these locations during high flows were significantly greater than at low
flows (Friedman test, P = 0.007; Figure 3-20).
Radio-tagged adults engaged in five types of behavior: continuous upstream movement,
interrupted upstream movement, continuous downstream movement, interrupted
downstream movement and localized (at or near their respective release site) movement
(Table 3-7). Fish were stationary in two large pools (area 5) of the fishway during all
interrupted movements. Of the twelve salmon tagged, three displayed rapid movement
patterns at the entrance of the fishway and did not ascend beyond the first pool. These
three fish displayed an average of 20.33 ± 5.13 unsuccessful attempts·d-1 during
primarily flow period one (Figure 3-21 ). Similarly, salmon ascending the fishway
displayed a mean of 18.0 ± 3.11 attempts·d-1 during flow period one. However, there
was a significant reduction in the number of attempts·d- 1 (7.75 ± 1.71; mean± SD) at the
fishway entrance during higher flows (Mann-Whitney U-test, P = 0.04; Figure 5).
Table 3-7.
Fish
code
1868
1337
13376
1867
1332
1333
1865
18656
1866
1335
1334
1863
1640
1242
1
4
Biological characteristics, passage times, entrance attempts, and
behavioral patterns of adult anadromous Atlantic salmon monitored at
Grand Falls fishway.
Sex
Length
{em}
M
56.5
52.2
52.2
57.5
55.1
56.5
51.2
51.2
51.5
58.2
64.8
53.5
53.4
53.0
F
F
M
M
F
F
F
F
F
F
M
F
M
Weight
{9}
1690
1304
1304
1730
1530
1652
1218
1218
1288
1798
1930
1348
1344
1325
Total time
{hour}
1.82
2.20
4.20
6.92
16.10
1.75
2.18
21.78
19.62
4.28
54.32
NA
NA
NA
Flow
m·s· 1
1.69
1.69
1.69
1.69
1.82
1.69
1.69
1.82
1.82
1.69
1.82
1.69
1.69
1.69
Entrance
CU 1
IU 2
CD 3
104
L05
Attem~ts
22
16
14
21
8
20
15
6
7
18
10
26
19
16
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
CU=Continuous Upstream, 2 IU=Interrupted Upstream, 3CD=Continuous Downstream,
1D=Interrupted Downstream, 5LO=Localized (at or near their respective release area)
3.3.3 Muscle Activity (EMG s) From Fish Moving Through the Fishway
Tests were conducted at the Grand Falls fishway, Exploits River, to measure in-situ
muscle activity from EMG tagged fish as a means of evaluating energy expenditure and
3- 11
New Telemetric AAnroaches To The Assessment OfFish Swimming Performance
the difficulty of fish passing various sections of this major fishway. A total of 10 trials
were completed and a full set of EMG signals were recorded in the lower section of the
fishway (11 pools, vertical slot design) and a resting pool. The signals, and the
corresponding parts of the fishway in which they were recorded, are provided in Figure 322. The results suggested, considering previous calibration work (Section 3.1 ), that fish
were using burst mode to move between pools and that there was considerable energy
expenditure within in the various pools due to turbulence. Fish were also very active in
the holding pool where they remained for a considerable period of time.
3- 12
New Telemetric A(lproaches To The Assessment OfRsb Swimming Perlormance
4.0
DISCUSSION
During the spawning migration of Atlantic salmon no food is consumed and all the energy
required to migrate, sexually mature and spawn is derived from reserves stored in the
body (Jonsson et al. 1997). In some species, energetic costs associated with migration
and spawning result in death, such as in Pacific salmon. In Atlantic salmon, however,
migration and spawning does not result in death of all individuals and some salmon may
return to sea and spawn several times during their lifetime. Consequently, additional
energetic obstacles such as those induced anthropogenically (i.e. fishways) may result in
a depletion of fishes' limited energy reserves which may reduce gonadal production,
reduce quality of reproductive products (egg, sperm), may limit further upstream
migration to preferred/optimum spawning habitats, and, in extreme cases, may contribute
to post spawning mortality.
Fishways represent a potential barrier to the upstream migration of Atlantic salmon. The
limited numbers of studies concerning the effects of fishway design and water velocities
on migratory fish, however, have resulted in controversy regarding the ability of salmon
to navigate fishways. Fish are ectothermic and the energetic cost of activity is
determined, to a large extent, by water temperature. Decreases in water temperature
have been shown to significantly reduce a fishes' aerobic swimming capacity (Brett and
Glass, 1973; Jayne and Lauder, 1994). Similarly, in this study the critical swimming
speeds of wild Atlantic salmon were significantly reduced by temperature, decreasing
from 2.16 ± 0.18 m·s- 1 to 1. 76 ± 0.06 m·s- 1 at 18 °C and 12 °C, respectively.
Low speed swimming, which includes sustained and prolonged activity, is supported by
the red muscle fibres under aerobic conditions. Burst activity, that can only be
maintained for less than 30 seconds, which is supported by the white musculature using
anaerobic pathways. The sustained and prolonged swimming velocities of Atlantic
salmon also decreased significantly with decreased temperature but burst activity
appeared unaffected. Therefor salmon required to swim at high water velocities when
water temperatures are cold may depend more heavily on the use of white muscle fibres,
fuelled through anaerobic metabolism, and a significant oxygen debt may be acquired
during anaerobic activity (Wood et al. 1983). The energetic costs of repaying this oxygen
debt may be greater than the aerobic scope of the fish and exhaustive exercise may
contribute to delayed mortality (Beamish 1978; Wood et al. 1983). Consequently, ascent
of fishways by Atlantic salmon requiring activity beyond their prolonged capabilities could
result in increased levels of mortality.
There have been few studies of swimming activity in wild fish and physiological telemetry
is one tool that has been used previously to monitor the activity of fish in the wild (Priede
and Young 1977; McKinley and Power 1992; Demers et al. 1996; Weatherly et al. 1996).
In this study, successful calibration of radio transmitted EMG signals with swimming
performance permitted in situ measurement of muscle activity to assess the activity of
4- 1
New Telemetric Approaches To The Assessment OfFish Swimming Pedormance
free swimming Atlantic salmon in an experimental fishway (flume). At 18 °C, the water
velocity within the flume (i.e. 2.1 m·s· 1 ) was below the critical swimming speed of salmon
and was reflected in the ability of salmon to ascend the flume with constant muscle
activity and within aerobic capacity. In contrast, the muscle activity measured in salmon
ascending the fishway at 12 °C was more vigorous than at 18 °C and reflected reduced
swimming capabilities of Atlantic salmon at 12 °C, as determined under controlled
laboratory conditions. Furthermore, the majority of muscle activity exhibited by salmon at
colder temperatures indicated that ascent of the flume may have involved activity beyond
its aerobic scope. Thus water temperatures become a critical consideration in fish
passage when individuals are required to exceed their aerobic scope for a portion or all
of the ascent.
Atlantic salmon migrate up rivers at various times and at can experience a variety of
temperatures at the onset or during their migration. In the fall, water temperatures
decrease considerably and late migrants may be subjected to additional stress when
ascending fishways and navigating natural velocity barriers (e.g. rapids). These barriers,
either mechanical or hydraulic and either natural or anthropogenic, routinely impede the
natural migratory progress of salmon. On rivers which support dams requiring fish
passage, the amount of energy used by salmon during migration may increase and limit
spawning success. More study is required to determine the influence barriers have on
energy expenditure of migrating salmon and ultimately the affect on allocation of energy
for reproduction. Physiological telemetry, utilizing radio-transmitted electromyograms
(EMGs) of red swimming muscle, has been demonstrated in this study as an excellent
technique to assess in situ muscle activity of freely swimming fish in controlled and
natural settings.
Another aspect investigated in this study was the influence of season, body morphology,
and sex on swimming performance and muscle activity (EMGs). The reduction in the
sustained, critical and burst swimming capabilities of Atlantic salmon, consistent with the
observed declines in water temperature, may have reflected the influence of temperature
on the physiological and biochemical properties of red muscle tissue (Wardle 1980).
Although swimming capabilities declined in both sexes, the reductions in sustained and
critical swimming performance were more pronounced among females than males. One
reason for the greater loss of swimming ability among females was likely related to the
additional influence of body morphology. Ovarian development in females resulted in
pronounced increases in girth and cross-sectional area relative to males. These
increases in body morphology in females greatly increase the drag associated with
moving in water and result in a pronounced increase in the power required to swim by
fish (Osborne 1961 ).
Similarities between the burst swimming speeds of male and female salmon, and the
independence of burst capabilities on both temperature and morphology suggested the
influence of these factors may be limited to aerobically supported activity. Activity indices
4-2
New Telemetric A.a.acoaches To The Assessment OfRsh .Swimming Pedarmance
(EMGs) of male and female salmon both demonstrated increased muscle activity during
burst swimming and previous laboratory studies demonstrated that increased muscle
activity during burst swimming reflected recruitment of white muscle fibres (Rome et al.
1992; Jayne and Lauder 1994 ). The significant increase in activity indices of male and
female salmon taken from rest through burst levels (ie. 3.0-4.0 body lengths per second)
strongly suggested recruitment of additional muscle fibres and since red muscle is only
capable of maintaining sustained swimming (Beamish 1978), the observed muscle
activity patterns at burst speeds are considered to have indicated white muscle activity.
The largest increases in muscle activity were observed in spawning females at their burst
capabilities. These results imply that spawning females may depend on white muscle
activity to a greater extent than males, and consequently, may exceed their aerobic
capacity more frequently than males. Priede (1983) suggested that fish which exceed
their aerobic scope increase their potential mortality, and the declines observed in the
aerobic swimming capabilities of females may be an important factor determining postspawning mortality. These observations indicate that this could be important factor in the
relative survival of males and females after spawning, which in turn could influence the
sex ratio of repeat spawners which could influence the reproductive potential of the
population.
This study has also employed a state-of-the-art DSP telemetry system to study high
speed (burst) swimming in adult Atlantic salmon. Early studies on high-speed swimming
qualitatively investigated kinematics (Gero 1952; Hertel 1966) and most studies
employed procedures such as laboratory film analysis, or accelerometry. High-speed
cinematography has been primarily employed to evaluate aspects of fast-starts in
relation to predator-prey interactions and effects of temperature and body form on
escape performance (Webb 1976 1977). In many studies fish were stimulated or
shocked to induce movement so performance would be sub-optimal. In this study, all
fish movements were evaluated under natural conditions and the fish swam volitionally.
Furthermore, since many fish were active during low visibility hours (46.7 %) precluding
the use of film techniques for data collection.
Webb (1975) suggested that, to properly evaluate fast-start performance, data for the
duration of the event, mean and maximum accelerations, mean and maximum velocities
and the distance covered must all be considered. Maximum distances attained by fish
declined as water velocity increased, which suggested a distance-velocity barrier
resulting in fish fatigue and beyond moderate water velocities (e.g. 1.92 m·s- 1) success
rates fell significantly. At these water velocities fish varied their average speed by taking
periodic pauses then bursting to maximum speed to complete passage of the flume,
thereby displaying a burst-and-coast swimming pattern to reduce energy expenditure
(Videler and Weihs 1982). Some individuals also made several passes at the flume
entrance before returning to rest in the lower pool. This behaviour continued for hours
and even days before an individual attempted to pass through the flume and could have
4-3
New Te{emetric Approaches To The Assessment OfRsh Swimming Pedarmance
resulted in considerable additional energy expenditure. Delayed salmon passage and
excessive energy expended in passage can tax energy reserves (Osborne 1961 ),
decrease the ability to reach spawning areas and ultimately reproductive success (Geen
1975).
The physiological and biochemical mechanisms associated with burst-type exercise in
fish have been extensively studied (Dobson and Hochachka 1987; Wood 1991 ). Several
factors may contribute to variability in the physiological responses to exhaustive exercise
between individual fish observed during this research. This study examined the
swimming speeds of fish collected along their migratory routes at seasonal temperatures.
During migration, Atlantic salmon experience significant depletions of energy reserves
and changes in body morphology such as increased girth owing to extensive gonadal
development in females. Physical condition, as influenced by age, maturity stage,
stress and fatigue, health and probably to some extent the inherent genetic capability for
upstream migration, affects migration rates and capability for ascent of difficult natural
and anthropogenic fish passage. Physiological disturbances in preparation for spawning
include significant depletions of body proteins and fats which result in great reductions of
a salmon's energy reserves. Thus, additional energetic costs associated with barrier
falls, rapids and fishways can further contribute to depletion of a salmon's limited energy
budget.
The timing of salmon migration can, in part, be dependent upon varying light intensities
(Banks 1969) and activity patterns reported in this study illustrated a diel distribution, with
major peaks at dawn and dusk. Electronic monitoring studies have shown that salmon
migrate at dusk and during the hours of darkness in the summer however later in the
migratory period (e.g. during late fall), salmon appear to migrate at any time (Hellawell
1973; Stewart 1973; Webb and Hawkins 1989). Thus, salmon are flexible and variable in
their behaviour and may move during the day when close to maturity (Mills 1989).
The use of DSP and EMG telemetry to assess fish swimming performance in passing a
existing fishway at Grand Falls, Newfoundland, also provided interesting results that
could potentially be used to improve future fishway design and operation. For instance,
the proper location of fishway entrances and provision of sufficient attractant water is
critical. DSP tracking data indicated that individuals made several passes at the fishway
entrance before returning to downstream pool areas. This behaviour, in some instances,
continued for hours or even days before an individual either attempted to pass the
fishway or moved downstream out of the immediate area. The passage of tagged fish
into the fishway entrance was therefor subject to a range of delays and two aspects of
the behaviour of the fish may have accounted for these delays. Firstly, fish tended to
remain in the tailrace area which emerges a short distance from the fishway entrance, as
a result of the greater discharge in that area. Secondly, fish entered the lowest pool of
the fishway without continuing to ascend, apparently reluctant or unable to pass through
the slot openings.
4-4
New Telemetric A,npcoaches To The Assessment Offish Swimming Pedqrmance
Water flow is an important factor controlling the initiation, onset, rate, and success of
upstream migration (Arnekleiv and Kraab0l 1986; Jensen et al. 1986; Bagliniere et al.
1990), often modified by a variety of additional environmental factors. These include air
temperature, cloud cover, light intensity, barometric pressure, wind velocity and direction
(Banks 1969). In this study, discharge through the fishway increased relative to river
discharge (during heavy rainfall) and providing stimulus for increased movement in the
tagged fish through the fishway. The influence of river flow as migratory stimulus and on
other aspects of fish passage requires further investigation in order to more fully
understand the complexity salmon movements in relation to changes in environmental
and physical conditions. Tools such as the DSP and EMG telemetry offer new and
innovative approaches investigating these influences on fish migration.
Tracking data (DSP) revealed rapid fish movements through the main sections of the
fishway while salmon spent a significantly greater amount of time at two holding 'facilities'
(pools) within the fishway. Resting pools can be advantageous in fish passage in that
they can increasing system capacity and provide holding areas for slow or exhausted fish
(Blackett 1987). These results, when considered in conjunction with the EMG results,
indicated that salmon moving through the lower and upper sections of Grand Falls
fishway may become fatigued and may be bursting through slot openings between pools,
and rest periods at other sections of the fishway (i.e. the middle holding areas and exit
pools) are required to recover from exertion. The physiological disturbance which results
from intense swimming activity in fish may require 12 to 24 h for complete recovery
(Milligan and Wood 1986).
Tests were conducted to measure in-situ muscle activity from EMG tagged fish as a
means of evaluating energy expenditure and the difficulty of fish passing various sections
of a major fishway at Grand Falls. A total of 10 trials were completed and a full set of
EMG signals were recorded in the lower section of the fishway (vertical slot) and in a
resting pool. The signals indicated that fish were using burst mode to move between
pools and that there was considerable energy expenditure in the various pools, likely due
to turbulence. Fish were also very active in the holding pool where they remained for a
considerable period of time. These results indicated excessive energy requirements for
movement between pools of the fishway, for holding position in pools, and interestingly in
use of the designed resting areas. This aspect of the study has demonstrated the
application of this technology to measurement of energy expenditure in use of existing
fish passage structures which can be further used to assess fishway performance and
design problems.
4-5
New Telemetric Approaches To The Assessment Offish Swimming Pedqcmance
5.0
CONCLUSIONS
A major objective of this report (Part B. New Telemetric Approaches To The Assessment
Of Fish Swimming Performance) was to develop and evaluate innovative approaches to
assessing locomotory activity, swimming performance, and energetic costs to fish under
naturally occurring conditions in relation to potential barrier problems at natural barriers
(e.g. falls, rapids), hydroelectric installations, culverts, fishways, etc. This involved
surgical implantation of a bio-sensitive radio transmitter (EMG or electromyogram tag) in
individual fish, calibration to locomotary ability and energetic scope (calibration of EMGs
to swimming speed and oxygen consumption), and subsequent use of radio transmitted
EMG signals to assess swimming performance and metabolic costs in situ. Studies were
also conducted to assess the effects of sex, maturity, location and timing with respect to
migratory distance, and body morphology on muscle activity as determined from EMG
telemetry. Additionally, 'state-of-the-art' telemetry systems (DSP or digital signal
processing with antennae switching) were used to study high speed swimming
performance, behaviour, and migratory strategy in relation to ascent of a an experimental
flume. Finally, these techniques were also applied to assessing swimming performance,
behavioural strategy, and energetics associated with passage of an existing fishway at
Grand Falls, Newfoundland. Collectively, these studies have demonstrated the
capabilities of these technologies and techniques in addressing fish passage issues and
furthermore have indicated the complexity of factors that regulate fish swimming energy
expenditure that need be considered in the design and operation of fish bypasses.
Radio transmitted EMG signals were successfully correlated with swimming performance
in Atlantic salmon thereby permitting this procedure to be used to evaluate salmon
ascent of an experimental flume utilising high intensity, burst swimming. This study also
demonstrated the influence of environmental temperature and reproductive related
changes in body morphology on the aerobic swimming capabilities of anadromous
Atlantic salmon. Consequently, increased demands on swimming capabilities in the latter
stages of migration, or during periods of low temperature may be determinant factors in
the migratory scope of Atlantic salmon. The influence of body morphology is most
prevalent in females related to the substantial increase in ovary size during sexual
maturation. Measurements of in situ muscle activity obtained from salmon at different
stages of their migration indicate that both male and female recruit additional muscle
fibres in order compensate for the effects of temperature and cross-sectional area on
swimming capabilities. Activity indices suggested that prolonged swimming by spawning
females involves increased white muscle activity which could lead to differential effects
on swimming performance and survival related to sex.
Ideally, fishways should be able to successfully pass all migratory species inhabiting the
river in which they are constructed consequently design criteria should be based on the
poorest swimming species requiring fish passage. Many existing structures have been
designed for economically, socially, or recreationally important species such as Atlantic
5-1
New Telemetric Approaches To The Assessment OfBsh Swimming Pedormance
salmon and these structures may be beyond the scope of other species within the
watershed. Results form these studies have suggested that water velocities within fish
by-pass structures should be less than the maximum attainable speed of all upstream
migrating species. For Atlantic salmon, a water velocity equal to or less than 1.55 m s· 1
within fishways would be ideal In order to ensure high passage rates. This study also
suggested a distance-velocity barrier and corresponding low passage success rates by
fish traversing at moderate to high water velocities. Thus, to avoid fish fatigue and
exhaustive exercise, water velocities within fish-bypass structures should be maintained
within the prolonged or sustained swimming scope of the species in question.
The use of DSP telemetry has permitted quantification of burst swimming ability and may
assist in optimizing the design of future fishways and culvert installations. Digital
telemetry has provided data on kinematics and strategies employed by salmon during
high speed swimming which were previously not attainable using conventional mark-andrecapture procedures. Furthermore, digital telemetry has been demonstrated to be an
important tool in evaluating fishways by providing finer resolution of movements and
position of fish within fish by-pass structures.
The management and protection of an important species such as Atlantic salmon
depends on a thorough understanding of the interrelationships between fishway
hydraulics, fish behaviour, physiology and swimming abilities. Data on the rate of ascent
of various species of fish through different types of fishways under varying conditions are
lacking. Results collected in this study provided some of the only existing information on
the movements and behaviour patterns of adult Atlantic salmon moving through a
fishway at different flows and substantiated the use of telemetry for evaluation of
existing fishways and to assist in optimization of designs for future fishways.
Construction of fishways to avoid high energy expenditure and provision of adequate
resting structures are necessary to maximize passage success. Therefore, the energy
an adult salmon must expend to reach and ascend a fishway and then finally to
successfully reach spawning areas and spawn, is an important consideration in fishway
design. The physiological telemetry component of the fishway assessment also
demonstrated detailed activity patterns of muscle activity at specific points of passage
through the fishway which could be used to rectify or modify problems. This aspect of
the research also demonstrated that, from the perspective of fish energetics, fishway
components may not function as designed.
This information is of importance to industry and government in responding in a
scientifically responsible manner to fish passage concerns. Collectively the information
contained in this report, as well as the companion document (Part A), on swimming
speed and energy budgets (locomotory activity and associated metabolic cost) of native
species, under natural conditions, will be applicable to assessing the impacts and
specifying design criteria for existing and proposed hydroelectric developments, bridge
and culvert installations, fish passage facilities, etc., throughout the range of the study
5-2
New Telemetrjc Approaches To The Assessment Offish Swimming Pedarmaace
species. This information will be of benefit in the development of mitigative strategies for
structures or conditions that may potentially impede fish passage or alienate habitats.
5-3
New Telemetrjc A,o,ocoaches To The Assessment Offish .$wjmmjng Pedarroancfl
6.0
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6-7
New Telemetric Approaches To The Assessment OfEjsh Swimming Performance
APPENDIX A
REPORT
ILLUSTRATIONS
New Tetemetrjc Approaches To The
Assessment Offish Swjmmjng Peifarmance
60km
site
location
Site 1
Site 2
Site 3
Site 4
Site 5
Bay of Exploits
Bishop's Falls
Grand Falls
Red Indian Dam
Noel Paul's Brook
Figure 2-1.
sampling date
May 26 -27th
June 18- 24th
July 16 - 20th
September 11-14th
October 12-17th
temperture (OC)
14.6+0.3
19.3+0.4
22.2+1.1
14.0+0.3
8.8+0.4
Study locations along the migratory route of Atlantic salmon in the Exploits
River, Newfoundland, Canada. Mean water temperature and sampling
dates are shown.
A1- 1
New Telemetric Approaches To The Assessment OfRsh .Swimming Performance
J
white muscle
electrodes with
gold tips
r----
EMG transmitter
Figure 2-2.
red muscle
antenna
A schematic showing the location the electromyogram (EMG) transmitter
in the body cavity of the fish. Gold tipped electrodes are inserted into the
red swimming muscle.
A1-2
New Telemetric A(lpcoaches To The Assessment OfFish Swimming Pedormance
Water flow
Upper pool
Lower pool
D
DSP/
receiver 11111111111 Bl
Figure 2-3.
A schematic diagram showing 7 fixed antennae stations connected to the
receiver/coprocessor system in the experimental flume used for this study.
As fish traversed the flume, signals were transmitted and received by the
antennae, and information regarding pulse arrival times, antenna position
and frequency were processed by the DSP. The receiver performed code
discrimination, verification and data storage.
A1-3
New Telemetric Approaches To The Assessment OfFish .Swimming Pedqrmaace
Bulkhead radius
Flow depth
Raceway width
Raceway length
r= 0.3 m
d = 0.3-0.6 m
b = 0.5 m
I= 18m
r
8H
d
v •
t
Sluiceway
of dam
Figure 2-4.
Hydraulics of the experimental flume, Noel Paul's Brook. . A stoplog
section in the first sluiceway of the dam controls head elevation (b. H) and
subsequent water velocities (V) downstream.
A1-4
New Telemetric A1npcoaches To The Assessment OfFish Swimming Pedormance
-a-controls ~
xit
:
:
1
I
1
1
1
I
I
I
I
I
I
I
~Intake
Flow
!
Entrance
Figure 2-5
A schematic diagram of Grand Falls fishway with corresponding antennae
locations (denoted 1 to 7) used in this study. Resting facilities are located
at antennae five and seven.
A1-5
New Telemetric Approaches To The Assessment Offish Swimming Performance
flow
) t~
salmon
Figure 2-6
A schematic diagram of the Grand Falls fishway showing locations of
EMG telemetry monitoring. The EMG signals from traversing the lower
11 pools and from spending time in the resting pool (pool 11) are shown.
A1-6
New Telemetric A,a,acoaches To The Assessment OfRsh Swimming Performance
160 140 -
l2°C
l8°C
120 ,-....
d
"§
100 -
'-"
a.>
::s
·-
0.0
80 -
~
~
0
60-
a.>
·8
E-<
40 200
-1
0.5
1.0
1.5
2.0
2.5
Swimming Speed (m sec-t)
Figure 3-1.
Fatigue tests of swimming performance of wild Atlantic salmon (n=5)
conducted at 12 and 18 °C. Transition to exhaustion has been subdivided into the following components: sustained, prolonged and burst
swimming speeds. Dotted lined represent swimming endurance (i.e. time
to fatigue) beyond 120 minutes.
A1-7
New Telemetric Approaches To The Assessment OfRsh Swimming Pedarmance
18°C
600
rn
E
-
800
.....(J)c
1000
ca
~
12°C
·-rn::::s 1200
Q)
-·s;
I
Q.
~
I
1400
~
(.)
ca
-rn
(J)
1600
(.)
::::s
E
1800
"C
... 2000
(J)
60
100
140
180
220
swimming speed (em sec-
Figure 3-2.
260
1
)
Calibration of muscle activity to swimming performance in wild Atlantic
salmon (n=5), at 12 and 18 °C.
A1-8
New Telemetric A,nprnaches To The Assessment Of Ejsh Swimming Performance
-
.......
.c
N
0
C)
-5
E
~
c.
E
:::J
900
800
700
600
I
500
400
tn
5 300
(.)
c
Q)
C)
~
0
200
100
0
2000
1800
1600
1400
1200
red muscle activity (pulse interval) ms
Figure 3-3.
Correlation of oxygen consumption with muscle activity in wild Atlantic
salmon (n=5), at 12 and 18 °C.
A1-9
New Telemetric Approaches To The Assessment OfFish .Swimming Periormance
500
1000
-
1500
-
activity (f)=2322.67-4.31 velocity r2= 0.97
site 1
2500
-
1500
-
2000
-
2500
-
~
500
-
.5
1000
-
~
1500
-
500
1000
"Ui'
e
-;
u
~
><
u
2000
s::
2500
"0
activity(m)=2231.05-3.57velocity r=0.98
activity(f)=2170. 75-3.59velocity r. 0.90
site 2
activity(m)=2277.43-3.81 velocity r=0.97
activity(f)=2232.25-3.54velocity r= 0.94
site 3
-
activity(m)=2142.73-2.87velocity r=0.95
.?;>
">
·.::
<
1500
-
2000
-
500
1000
activity(f)=2548.1 0-S. 77velocity r=0.87
site 4
a
2500
500
activity(m)=2298.60-4.28velocity r=0.95
-
2000
-
2500
-
1000
1500
0
activity(f)=3099 .60-13.79velocity
r=0.9s
site 5
r=O.so
activity(m)=2603.73-9.62velocity
0.5
1.0
1.5
Swimming Speed (m
Figure 3-4.
2.0
2.5
sec-1)
Relationships between muscle activity and swimming speeds for male
(solid) and female (white) salmon at various stages of their spawning
migration. Site 1 represents freshwater entry and site 5 represents the
pre-spawning period. Sample sizes are as follows sites 2-4 N=4 males
and 4 females, site 1 and 5 N=3 males and 3 females.
A1 -10
New Telemetric Approaches To The Assessment OfFish Swimming Performance
150
900
(i)
,_,
,_:··
....
A
a
-;;-
s
"E£
.5
?A
"3
,e,
><
u
"t::
c
·t
·-a
<
750
900
1050
1200
1350
1500
1650
1800
1950
2100
60015090010501200135015001650180019502100-
A
A
a
a
,
"""
'"
,-.
'\ ...'.
A
.. A
\~>
(h)
a
A
A
a
a
b
a
. B*
(ih)
b
~
-~··
a
a
a
a
;-\'
'"~·
..
·,
-,'-.
....
-.·...
~.
0
5
10
15
20
25
Length of Freshwater Migration (weeks)
Figure 3-4.
Relationships between muscle activity and swimming speeds for male
(solid) and female (white) salmon at various stages of their spawning
migration. Site 1 represents freshwater entry and site 5 represents the
pre-spawning period. Sample sizes are as follows sites 2-4 N=4 males
and 4 females, site 1 and 5 N=3 males and 3 females.
A1 -11
New Telemetric Approaches To The Assessment Offish Swimming Performance
400~---------------------------.
E
600
800
1000
1200
1400
16oo
1800
2000
-+------,;---~--.---,---,--=----t
0
10
20
30
40
50
60
X
Q)
"0
c
~
>
uro
400 ~-------------------------------,
600
800
1000
1200
1400
1600
1800
2000~~~----~---.~--r----.--~
0
10
20
30
40
50
60
time (seconds)
Figure 3-6.
Muscle activity recorded in wild Atlantic salmon (n=4) during ascent of 20
m long experimental flume recorded at 12 °C (fall) and 18 °C (late
summer) in relation to critical swimming speed (Ucn1).
A1 -12
New Telemetric A,a,nroaches To The Assessment Offish Swimming Pedarcnance
---Maximum
25
· · ·---- Minimum
-u
20
e
.a
e
15
Q)
!
....
,. ' ,....,
:\
.e.,
.,
I
..
\
\
.
I I
~·I I
.,
'•
•
'\
10
.
..••
:..
5
I
••••
I
0
14- 23- 1- 10- 19- 28- 6- 15- 24- 3- 12- 21Jul Jul Aug Aug Aug Aug Sep Sep Sep Oct Oct Oct
Date
Figure 3-7.
Water temperature in Noel Paul Brook during the 1996 experimental
period as obtained by hourly thermograph readings.
A1 -13
New Telemetric Approaches To The Assessment Offish Swimming Performance
0.60
0.50
-i'
6
o.40
·- •••
~
~
X~
----. ---~::::=:=;.:i::.:.: - -===
x
0.20
0.10
O.OOT-~.-~.-~.-~.-~.-~.-~.-~.-~
0
2
4
6
8
10
12
14
16
18
Distance up flume (m)
Pool elevations:
Figure 3-8.
-x-0.25 --0.34 ....... 0.43
Depth profiles in the flume with corresponding pool elevations. Elevations
of 0.25, 0.34, and 0.43 represent mean water velocities of 1.89, 2.79, and
3.09 m·s·1 , respectively.
A1- 14
New Telemetric A{l{'coaches To The Assessment OfRsh .Swimming Pedarmance
40
....om
=
G)
Summer
35
30
e 25
>
0
20
e
....
G)
0
~
15
10
5
0
40
40
35
...ccu
30
-
20
rl.l
ecu
>
0
e
0
~
25
15
10
5
0
0
0
0
0
0
0
0
0
0
0
0
0
0
- - -- -- - ~I
00
. I
9.
0
0
Figure 3-9.
~
c:J
\(:)
I
0
00
I
0
c:J
Q
Q
N
0I
-
0
I
0
\(:)
Q
N
Time periods and fish activity in the flume. Time periods (hours:minutes)
of 00:01-04:00, 04:01-08:00, 08:01-12:00, 12:01-16:00, 16:01-20:00, and
20:01-24:00 refer to night, early morning, late morning, early afternoon,
late afternoon, and evening periods, respectively.
A1- 15
New Telemetric Approaches To The Assessment OfFish .Swimming Performance
100-
100
~
Summer
-...=
80-
~
57.1
60-
8
VJ
VJ
~
40-
G.>
(,)
(,)
::s
(/)
200
1.61
1.89
0
0
0
2.3
2.82
3.09
Water velocity (m/s)
100
Fall
-~
80
60
40
20
0
0
1.7
1.99
2.33
2.84
3.2
Figure 3-10. Success rates of passage of the flume in relation to water velocity (m·s-1).
A1 -16
New Telemetric AJ1Proaches To The Assessment Offish Swimming Pedqcmance
20
,........
s
•t
15
~I
!
I
v(.)
B
·-e
e::s
·rll
""0
~
10
Season
• Fall
• Summer
5
0~------~------~------~----~
1.5
2.0
2.5
3.0
3.5
Water velocity (rnls)
Figure 3-11. Maximum distances attained by salmon ascending the flume at various
water velocities (m·s- 1).
A1 -17
New Telemetcjc Af1Proaches To The Assessment Of Eish Swjmmjng Pedocmance
Season
• Fall
• Summer
36
27
18
9
04-----~------~~----~r-----•
1.5
2.0
2.5
3.0
3.5
Water velocity (m/s)
Figure 3-12. Total time required for salmon to ascend the flume at various water
velocities (m·s-1).
A1 -18
New Telemetric Approaches To
--
The Assessment OfFish Swimming Performance
3.5
Season
1'1.)
e
'-"'
"0
II Fall
D Summer
2.8
G)
G)
c.
1'1.)
"0
c:
2.1
=
0
""bO
G)
1.4
bO
ccs
"">
<
G)
0.7
0.0
1.5
2.0
2.5
3.0
3.5
Water velocity (m/s)
Figure 3-13. Average ground speeds (m·s- 1) for varying water velocities (m·s- 1).
A1- 19
New Telemetric Approaches To The Assessment OfFish Swimming Performance
--
7
Season
f ll
Fall
e
.._,.
"'0
6
Summer
u
u
Q.
-......=
fll
5
0
u
bO
=
u
'"'
>
<
4
3
2
1.5
2.0
2.5
3.0
3.5
Water velocity (m /s)
Figure 3-14. Average total speeds (m·s- 1) for varying water velocities (m·s-1).
A1- 20
New Telemetric A,aproaches To The Assessment Of fish Swimming Pecfacmance
---
15
Season
• Fall
Summer
fll
~
0
a:l
"0
4)
4)
10
Q.,
-fll
~
0
4)
bO
~
....
4)
>
5
<
1.5
2.0
2.5
3.0
3.5
Water velocity (m/s)
Figure 3-15. Average total speeds, in bl·s·1, for varying water velocities (m·s-1).
A1- 21
New Telemetric Approaches To The Assessment Offish Swimming Peifocmance
Season
20
• Fall
Summer
,........
0
Cll
'~
~
'-"
15
"'0
Q)
Q)
~
Cll
t'\S
......
0
......
10
e::s
e
·-~
~
5
0
-+-.&.....f----:
1.61 1. 7 1.89 1.99 2.3 2.33 2. 79 2.84 3.09 3.2
Water velocity (m/s)
Figure 3-16. Maximum total speeds, in bl·s·1, in relation to increasing water velocities
(m·s- 1}.
A1- 22
New Telemetric Approaches To The Assessment Offish Swimming Pedorcnance
3
Season
• Fall
Summer
,.........
0
g
ell
~
G)
G)
2
~
ell
~
=
::s
0
bh
e::s
e
·-
1
~
::E
1.5
2.0
2.5
3.0
3.5
Water velocity (m/s)
Figure 3-17. Maximum ground speeds (m·s- 1) in relation to increasing water velocities
(m·s- 1 ).
A1- 23
New Telemetric Approaches To The Assessment OfBsh Swimming Performance
MODERATE
A
18
18
16
16
--e
14
fish aborts
14
12
12
.s= 8
10
10
i:5 6
6
4
4
2
2
t
8
"'
0~~~~~~~~--~~~
0
4
8
o~~~~~~~~~~~
12 16 20 24 28 32 36 40
0 4 8 12 16 20 24 28 32 36 40 44 48
Time (s)
4.5
4.5
B
4.0
4.0
-r 3.5
3.5
"'5
3.o
3.0
"C
2.5
2.5
~ 2.0
"' 1.5
.c:
2.0
~
..."'
~
1.5
1.0
1.0
0.5
0.5
0.0
0.0
0
4
8
12 16 20 24 28 32 36 40
0 4 8 12 16 20 24 28 32 36 40 44 48
Time (s)
Figure 3-18. Representative tracks of fish ascent of the flume under moderate wat~~
flow conditions. Panel (A) shows an unsuccessful attempt with panel 19)
showing corresponding speeds attained. Panel (~ shows a successful
attempt with panel (D) showing corresponding speeds attained.
A1- 24
New Te(eroetric Approaches To The Assessment OfEjsh $wjmming Pedarmance
Flow 1
7
6
5
4
3
2
llj
II
II
~
~
~
tl.)
~
··-·s::
s::
0
tl.)
0
~
0
!! .......
0
12
0 0 6
14Sep
19Sep
12 0 12 0 12 0 12 0 12
20 Sep 21 Sep 22 Sep 23 Sep 24 Sep
6
5
4
3
2
1
6
12
11 Sep
0
~
12
Flow2
18
0
6
12 Sep
12
18
6
12
20Sep
0
6
21 Sep
12
22
14
20Sep
7
6
5
4
3
2
1
6 7 8
11Sep
9
10 11 12 13 14 15
17 18 19
19Sep
22
9
21 Sep
20
21
23
5
16 18
11 Sep
20
0
2
12 Sep
4
6
10
11
13
Real Time (h)
Figure 3-19. Time versus position profiles for several tagged salmon at two water
flows. Flow period 1 refers to 1.69 ± 0.07 m·s·1 and flow period 2 refers to
1.82 ± 0.03 m·s· 1• The y-xis represents the seven sections of the fishway
as illustrated in Figure 2-5.
A1- 25
New Tetemetric Approaches To The Assessment Offish Swjroroing Pedormance
=1000
••
Water
velocity
(m/s)
a
'-'
.....
5 100
Q.c
f'l.l
~1.69
~
a
••
.1.89
::
10
Q
eJ)
Q
~
1
1 2 3 4 56 7
Position in fishway
Figure 3-20. The time spent (min) at various sections of the fishway during two water
flows. Results are expressed as a mean ± standard error(** P ~ 0.01 ).
A1- 26
New Telemetric AAaroaches To The Assessment OfFish Swimming Peiformance
30
3
~
~
"'0
..._
{1.2
.......
20
7
~
e
~
.......
.......
~
10
•
Q
z
0
Flowl
Flow2
Successful
fish
Flowl
Unsuccessful
fish
Figure 3-21. The number of unsuccessful attempts (total) at the fishway entrance at
two flow periods. Unsuccessful refers to individuals who did not ascend
the fishway during the study. Successful refers to fish who traversed the
fishway. Results are expressed as a mean ± standard error(* P ~ 0.05).
A1- 27
New Telemetric Approaches To The Assessment OfFish Swimming Performance
EMGs from Passage
through Pools 1 to 11
2200X
Q)
"'0
c
>.
.......
·:;:
u
180014001000-
<(
800400-600
600
0
1800
3000
4200
1200
2400
3600
4800
Time (seconds)
EMGs from Resting Pool
X
2200-
(1)
-c
c 1800-
~
> 1400-
+J
(.)
<(
10000
100
200
300
400
500
600
Time (seconds)
Figure 3-22. EMG signals for passage through pools 1-11 and for remaining in resting
pool 12 are depicted. Refer to Figure 2-6 for a schematic of Grand Falls
fishway with corresponding pools (1-11) and resting pool (12).
A1- 28
New
Tetemetrjc Approaches To The Assessment OfEjsh .Swjmmjng Pedarmance
APPENDIX B
ANNOTATED
BIBLIOGRAPHY OF
PUBLICATIONS ARISING
FROM THE STUDY
New Telemetric Approaches To The Assessment Offish Swimming Performance
Anderson, W. G., R. S. McKinley, and M. V. Colavecchia. 1997. The use of clove oil
as an anaesthetic for rainbow trout and its effects on swimming performance. N. Am. J.
Fish. Manage. 17:302-307.
Abstract: The only anesthetic registered for use in North American Fisheries Science
is 3-aminobenzoic acid ethyl ester methanosulfate (MS-222). MS-222 has been
shown to be a very effective anesthesia for several fish species but its application in
the field is limited because U.S. Food and Drug Administration guidelines demand a
21-day withdrawal period post exposure to MS-222 before fish can be released and
enter the food chain. As a consequence carbon dioxide (C02}, has been used as a
substitute anesthetic. However, induction and recovery times have been shown to be
long and anesthesia is somewhat shallow in comparison to MS-222. We compared
the efficacy of MS-222 to that of a naturally occurring substance, clove oil, as an
anesthetic on juvenile and adult rainbow trout Onchorhynchus mykiss. Clove oil was
as effective as MS-222 in inducing anesthesia in both age groups of fish.
Furthermore, exposure to either clove oil or MS-222 at the concentrations tested was
not detrimental to critical swimming speed of juvenile or adult rainbow trout. We
propose that clove oil be considered as an alternative to MS-222 for use as a fish
anesthetic.
Booth, R.K., D.A. Scruton, R.G. Goosney, and R.S. McKinley. 1995. Measurement
of red muscle activity and oxygen consumption in wild Atlantic salmon (Sa/mo sa/ar) in
relation to swimming speed using radio transmitted signals. p. 209-215, In C. Cristalli,
C. Amlaner, and M. Neuman (eds.), Biotelemetry XIII, Proceedings, Williamsburg,
Virginia.
No abstract.
Booth, R.K. 1998. Swimming performance of anadromous Atlantic salmon, Sa/mo
sa far L., during their spawning migration in the Exploits River. Newfoundland, Canada.
Ph.D. Thesis, University of Waterloo, Waterloo, ON.
Abstract: Swimming performance, muscle activity patterns and plasma non-esterified
fatty acid profiles were examined in wild Atlantic salmon (Sa/mo salar L.) during their
upstream spawning migration and downstream post-spawning migrations. These
studies were conducted on the Exploit's River, Newfoundland, Canada between June
of 1994 and October of 1996. Significant reductions in sustained and prolonged
swimming performance were observed during the upstream migration of adult Atlantic
salmon. Associated with the reductions in swimming performance, spawning Atlantic
salmon demonstrated higher muscle activity indices for any given swimming speed
than non-spawn_ing individuals. The greatest loss of swimming performance, and
B- 1
New Teleroefdc A,opcoaches To The A ssessmeat Of Fjsh ,Swjrorojng Perlocmance
change in muscle activity was observed for females just prior to spawning. Both
swimming performance and muscle activity indices were correlated with observed
changes in temperature and body cross sectional area. The change in cross section
area was more pronounced among females and was related to the final stages of
ovarian maturation. Sustained, prolonged and burst swimming performance of
Atlantic salmon kelts were significantly lower than those of upstream migrating
individuals. Smolts were also investigated. Smolts had were capable of swimming
significantly faster than adults, when swimming speed were expressed relative to body
length.
Total plasma non-esterified fatty acid (NEFA) levels declined significantly between
freshwater entrance and spawning, and continued to decline during the post-spawning
period. Plasma NEFA levels were significantly higher for females but declined to a
greater extent during the upstream migration. The rapid decline in plasma NEFAs
among females coincided with the largest increase in their gonadaosomatic indices.
Differences in the circulating levels of polyunsaturated and saturated fatty acids
became evident in males and females just prior to spawning. At spawning, males and
females possessed similar amounts of all plasma NEFAs and these did not change
during the post-spawning period.
Decreases in temperature, changes in body morphology and depletion of lipid (i.e.
plasma NEFAs) were observed and recorded during the freshwater migration of
Atlantic salmon. Collectively, these factors, many have resulted in the pronounced
changes in swimming capabilities and muscle activity patterns observed in migrating
salmon. The observed changes in the swimming performance the significant loss of
plasma NEFAs suggest that Atlantic salmon may become more susceptible to
disturbances in their migrations as they approach sexual maturity and prepare to
spawn.
Booth, R. K. R.S. McKinley, F. Okland and M.M. Sisak. 1997. In situ measurement
of swimming performance of wild Atlantic salmon (Salmo salar) using radio transmitted
electromyogram signals. Aquat. Living Resour. 10:213-219.
Keywords: Atlantic salmon, telemetry, temperature, electromyogram, swimming
performance.
Abstract: Swimming capabilities and in situ measurement of muscle activity from
adult Atlantic salmon (Salmo salar) at two seasonal temperatures were measured
using radio transmitted electromyogram (EMG) signals. Forced sustained levels of
activity and critical swimming speeds were determined and correlated to radio
transmitted EMG signals using a modified Blazka swim speed chamber. There were
no differences ir:t swimming performance levels between tagged and untagged
B-2
New Telemetric Approaches To The Assessment Offish Swimming Pedormance
individuals. At 18 °C, sustained activity and critical swimming speeds were
approximately 70% and 20% higher than at 12 °C, respectively. No differences in
burst activity were observed at these temperatures. EMGs recorded from salmon
during ascent of an artificial flume at cold temperatures revealed that overall muscle
activity is greater than that observed for critical swimming speeds. This implies that
white muscle may be recruited at this temperature. However, in contrast, most activity
at 18 °C is below that observed during critical swimming speed. Moreover, salmon
required almost twice as long to traverse the flume at 18 °C than at 12 °C. Together,
our data demonstrates that salmon may recruit white muscle fibres and incur an
oxygen debt at colder temperature as a strategy for ascending velocity obstructions at
a quicker rate.
Booth, R.K, R.S. McKinley, G. Power, and D.A. Scruton. 1998 (submitted). The
influence of body morphology and environmental temperature on the swimming
capabilities and muscle activity patterns of wild Atlantic salmon (Salmo sa far L. ). Trans.
Am. Fish. Soc.
Keywords: Atlantic salmon, swimming, electromyograms, reproduction, migration
Abstract: The present study describes the influence of environmental temperature
and changes in body morphology on the swimming capabilities and muscle activity
patterns in migrating Atlantic salmon. During the study period (May 23rd-Oct. 14th
1996), water temperature ranged from 8.8°C to 22.2°C. Morphological measurements
indicated that stomach weight declined by approximately 88% in both male and female
salmon while liver weight declined by approximately 59% in females and 26% in
males. Just prior to spawning, ovaries weighed 218±23.77 g and were heavier than
testes ( 54.41±2.26 g). Significant increases in girth and cross-sectional area were
observed among females but not males. Sustained, critical and burst swimming
capabilities were correlated with temperature and cross-sectional area for females, but
only with temperature for males. No differences in muscle activity indices were
observed until the onset of spawning. At this time, mean muscle activity indices
increased significantly for both sexes. Sex dependent differences in muscle activity
indices were most pronounced during prolonged swimming (ie. 2 body length per
second) and were significantly higher for females. Muscle activity was correlated to
both temperature {R2=0.64) and cross-sectional area (R2=0. 74) for females, but only to
temperature for males (R2=0.63). The results of the present study indicate that
environmental temperature is an important determinant of swimming performance in
anadromous Atlantic salmon. Changes in the body morphology of females places
additional demands on locomotory muscle and may be responsible for the significantly
lower aerobic swimming capabilities observed among pre-spawning females.
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New Telemetric Approaches To The Assessment OfFjsh Swimming Pedacmance
Booth,R.K., E.B. Bombardier, D.A. Scruton, and R.S. McKinley. 1998. (submitted)
Swimming ability of Atlantic salmon, Salmo sa far L. At three migratory phases. J.
Fish. Bioi. (submitted)
Abstract: Sustained, prolonged and burst swimming capabilities were investigated for
pre-spawning adult (bright), post-spawning adult (kelts) and juvenile (smolts) Atlantic
salmon (Sa/mo sa/ar) collected from the Exploits River, Newfoundland, Canada.
Significant differences in weight were observed among male and female bright
salmon. There were, however, no differences in fork length, girth or condition factors
between sexes. In male and female kelts, there were no differences in fork lengths,
girths, weights and condition factors. Bright salmon were similar in length to kelts but
weighed 1.7 times as much. Condition factors of bright salmon and kelts were
0.84±0.04 and 0.55±0.02, respectively (P<0.05). Although smaller than adults, smolts
possess the greatest swimming capabilities of any group relative to their fork length
(4.39±.0.09 body length (bl) sec- 1 ). Among adults, the sustained swimming
capabilities of bright salmon were found to be 2.51 ±0.08 bl sec·1 and were significantly
higher than those of kelts (0.97±0.04 bl sec-1). Smolts also had the greatest burst
swimming capabilities (10.24±0.08 bl sec- 1). Among adults, bright salmon also had
significantly higher burst swimming speeds than (bright salmon 4.24±0.14 bl sec·1,
kelts 2.067±0.10 bl sec- 1). Prolonged swimming speeds of bright salmon were also
significantly higher than those of kelts The present study demonstrates that life
history stage is an important factor determining the swimming characteristics of
migratory fish species such as the Atlantic salmon.
M. V. Colavecchia. 1997. The use of telemetry to assess high speed swimming and
ftshway performance in Atlantic salmon. M.Sc. Thesis, University of Waterloo,
Waterloo, ON. x + 106 pp.
Abstract: High speed swimming performance in wild Atlantic salmon, Sa/mo sa far L.,
was examined using volitional swim trials in a flume model that simulated varying
water velocities {>1.6 to 3.2 m3s-1 ) that might be present in fishways. To calculate
ground speeds, uniquely coded signals transmitted information regarding distance
moved and time elapsed to a digital spectrum processor (DSP) which thereby
processed the data using near real time spectrum analysis. This provided significant
information on numerous kinematic parameters (distance, time, ground speed, total
speeds, etc.) and strategies (burst and coast) employed by salmon during high speed
swimming ..
In addition, the monitoring of volitional movements of radio tagged salmon at an
existing fishway provided valuable information with respect to timing of movements.
Ascent times, success levels, and entrance activity in relation to water flow. Salmon
displayed a diel_pattern in movements within the fishway, primarily late morning and
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New Telemetric Approaches To The Assessment QfFjsh Swimming Pedarmance
late afternoon. As water velocities increased form 1.69 to 1.82 m3s-1, ascent times
significantly increased from 3 to 28 hours. As well, fish displayed significantly longer
amounts of time in resting structures in the fishway.
Last, we conducted a laboratory study in which we compared the efficacy of MS-222
to clove oil and later assessed post-anaesthetic swimming speeds of both these
substances to controls. Results indicated that clove oil was as effective as MS-222
and exposures to the substance was not detrimental to critical swimming speeds.
Therefore, clove oil was a reliable, safe, and effective anaesthetic we could utilize in
the field.
Colavecchia, M.V., C. Katopodis, R.G. Goosney, D.A. Scruton, and R.S. McKinley.
1998. Measurement of burst swimming performance in wild Atlantic salmon (Salmo
salar) using digital telemetry. Regulated Rivers 14:41-51.
Abstract: Swimming performance of wild Atlantic salmon (Salmo salar L.) was
investigated in an experimental flume using coded radio signals. To calculate
swimming speed, distance moved and time elapsed were measured with a digital
spectrum processor using near real-time spectrum analysis. This device was designed
to be used in a co-processing arrangement with a receiver, thereby providing pulse
position code discrimination, verification and continuous data storage. Radio-tagged
adults (48.3 to 54.8 em long) voluntarily swam against water velocities, ranging from
1.32 to 2.85 m s· 1, in an 18 m long flume at a mean water temperature of 10.1 ± 1.6
°C. At water velocities of 1.32 to 1.55 m s· 1, individuals successfully ascended the
flume at swimming speeds of 1.61 to 2.55 m s· 1 , or 3.30 to 4. 79 body lengths per
second (L s·1), respectively. At high water velocities ranging from 1.92 to 2.85 m s·1,
individual swimming speeds increased from 2.55 to 3.60 m s·1, or 4.94 to 7.27 L s·1,
respectively. However, above a threshold value of 1.92 m s·1, individuals traversed
shorter distances and were unable to ascend the flume. The highest swimming speed
observed was 4.13 m s·1 or 8.35 L s· 1• The results of this study indicate that in addition
to its applicability in the determination of burst swimming speeds, digital telemetry
could prove a useful tool in the design and evaluation of future fishways and culvert
installations.
key words: burst swimming; Salmo sa/ar, water velocity; digital telemetry
M. V. Colavecchia, R. S. McKinley, R. F. Goosney and D.A. Scruton. 1998.
(submitted). The use of telemetry to assess activity patterns of Atlantic salmon (Sa/mo
sa/ar) in a vertical slot fishway. Trans. Am. Fish. Soc. (submitted)
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New Telemetric Appcoaches To The Assessment OfFish Swjmming Perlocmance
Abstract: The swimming performance of wild Atlantic salmon (Salmo sa far L.) in an
existing fishway was investigated using coded radio-transmitted signals. Movement
patterns and passage times were recorded with a digital spectrum processor using
near real-time spectrum analysis. This device was designed to be utilized in a coprocessing arrangement with a receiver, thereby providing pulse position code
discrimination, code error detection and continuous data storage. Radio-tagged adults
(n= 12, 55.3 ± 3.8 em; mean fork length± SO) swam voluntarily through a 116m long
vertical slot fishway at a mean water temperature of 11.5 °C ± 0.5 (mean ± SE).
Passage at the fishway occurred primarily during late morning (57.1 %) and late
afternoon (35.7%); night passages (7.1 %) were of secondary importance. As
velocities increased from 1.69 ± 0.07 m/s to 1.82 ± 0.03 m/s, ascent times significantly
increased from 3.33 ± 0.72 to 27.95 ± 8.86 h (mean± SE) whereas, the number of
entrance attempts significantly declined from 18.70 ± 3.68 to 7.75 ± 1.71 per day
(mean ± SO). Analyzes of the tracking data have provided fine resolution of
movements and position of fish within the main sections, resting pools and entrance of
the fishway. The results from this study could be used to improve design criteria for
new fishways or for modifying existing by-pass structures for salmonid passage. It is
clear that digital telemetry can be a useful tool in the design and evaluation of future
fish passes, and studies dealing with fish migration in relation to water-flow
management.
F. Okland, B. Finstad, R. S. McKinley, E. B. Thorstad and R. K. Booth. 1997.
Radio-transmitted electromyogram signals as indicators of physical activity in Atlantic
salmon. J. Fish. Bioi. 51:476-488.
Abstract: Surgical methods developed to implant EMG (electromyogram)
transmitters in Atlantic salmon Sa/mo salarwere tested to calibrate electromyograms
from axial red musculature to swimming speed in a swim speed chamber, and to
compare electromyograms of fish from two stocks (Lone and lmsa). Ten Lone and
eight lmsa salmon were equipped with internal EMG transmitters. Surgical
procedures were acceptable, with 100% survival of all implanted fish during the study.
It was possible to calibrate EMG pulse intervals to swimming speed in 14 of the 18
salmon run in the swim speed chamber (r-2=0.35-0. 76 for individuals, 0.63 for pooled
data). Individuals differed in their EMG resting levels, and so higher correlations were
obtained between swimming speed and an activity index (EMG pulse intervals at
different speeds/EMG resting levels) (pooled data, r-2=0.75). The linear relationship
between swimming speed and EMG pulse intervals differed significantly between the
two stocks (P<0.05). This successful calibration of EMGs to swimming speed opens
the possibility of calibrating EMGs to oxygen consumption and the measurement of
the metabolic costs of activity in field experiments.
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New Telemetric Ap.proaches To The Assessment Of Ejsh .Swimming Pedorcnance
Key words: Salmo salar; radio telemetry; electromyograms; swim speed chamber.
Peake, S., McKinley, R.S., Beddow, T.A., and Marmula, G. 1996. A new procedure
for radio transmitter attachment: Oviduct insertion. North Am. J. Fish. Management
17:757-762.
Abstract: We looked the effects of internally tagging adult female Atlantic salmon
Salmo salar and rainbow trout Oncorhynchus mykiss via the urogenital tract. This
method takes advantage of the fact that the oviduct in salmonids and some nonsalmonid species is not connected to the ovary; therefor, transmitters can be inserted
into the body cavity of females through the urogenital opening. We found that dummy
tags inserted into adult Atlantic salmon prior to egg formation did not alter survival,
behaviour, or general egg development. Sixty-nine percent of transmitters were
retained for the entire study period (60 d). Transmitter retention (45 d) was 83% in
rainbow trout tagged approximately 6 weeks before ovulation. However, proper
insertion of transmitters in rainbow trout was prevented by the developeing ovary,
affecting egg expulsion and viability. It appears that oviduct tagging is a promising
non-surgical option in certian fish species if the transmitter is inserted prior to ovary
development or after eggs have been shed.
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