Developing Fish Passage and Protection at Hydropower Dams: Carl R. Schilt

Applied Animal Behaviour Science 104 (2007) 295–325
www.elsevier.com/locate/applanim
Review
Developing fish passage and protection

at hydropower dams
Carl R. Schilt *
LGL Ltd., Environmental Research Associates, P.O. Box 225, North Bonneville, WA 98639, USA
Available online 13 November 2006
Abstract
The development of waterways, for hydropower and other industrial uses, has substantially altered many of
the freshwater habitats of the planet and this has had considerable impact upon aquatic organisms. Industrial
changes in aquatic ecosystems, including hydropower development, can restrict or delay fish migration,
increase predation, affect water quantity and quality, and subject fish to direct damage and stress. This review
will focus on the consequences for fish welfare and the progress towards developing the means to pass and
protect fish at hydropower dams, at water withdrawal facilities, and in other engineered aquatic environments.
It primarily concerns the large mainstem hydropower dams in the Columbia-Snake River Basin in the
northwestern United States. Some methods for improving fish passage and protection at hydropower dams
involve modifications and additions to engineered structures and occasionally use sensory stimuli such as light,
sound, turbulence, or electric fields to influence fish distributions. Measures to improve fish survival, like
spilling water at a dam to provide non-turbine passage, can cause other problems for fish, for example higher
dissolved gas concentrations downstream. Reducing losses of fish in industrial environments is desirable in
both the industrialized world, where many fish-related problems currently exist, and in the developing world,
where lessons already learned may make future development more cost-effective and benign.
# 2006 Elsevier B.V. All rights reserved.
Keywords: Fish passage; Fish conservation; Hydropower; Dams; Environmental impact
Contents
1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 296
1.1. Background . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 296
1.2. Statement of problem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 299
* Tel.: +1 509 427 4840; fax: +1 509 427 4846.

E-mail address: cschilt@lgl.com.
0168-1591/$ – see front matter # 2006 Elsevier B.V. All rights reserved.
doi:10.1016/j.applanim.2006.09.004
296 C.R. Schilt / Applied Animal Behaviour Science 104 (2007) 295–325
1.3. Anatomy of a hydropower dam . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 300

1.3.1. Turbine passage . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 300
1.3.2. Juvenile bypass system passage . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 300
1.3.3. Spillway passage . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 301
1.3.4. Surface passage . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 301
2. Routes of passage . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 301
2.1. Upstream fish passage by lorries, ladders, and lifts . . . . . . . . . . . . . . . . . . . . . . . . 301
2.2. Downstream passage options . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 302
2.2.1. Turbines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 303
2.2.2. Juvenile bypass systems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 303
2.2.3. Spillways. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 303
2.2.4. Sluiceways and other surface routes . . . . . . . . . . . . . . . . . . . . . . . . . . . . 303
2.2.5. Transportation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 303
3. Sources of passage-related delay, stress, injury, and mortality . . . . . . . . . . . . . . . . . . . . . 304
3.1. The importance of run timing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 304
3.2. Turbine passage . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 305
3.3. Juvenile bypass system passage . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 306
3.4. Spillway passage . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 306
3.5. Surface passage . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 306
3.6. Transportation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 306
4. Improving fish passage at hydropower dams . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 307
4.1. Changing project structures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 307
4.2. Changing project operations. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 308
4.3. Using sensory stimuli to control local fish distributions. . . . . . . . . . . . . . . . . . . . . 310
4.3.1. Light. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 310
4.3.2. Low-frequency sound . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 311
4.3.3. High-frequency sound and alosine herrings . . . . . . . . . . . . . . . . . . . . . . . 312
4.3.4. Flows and turbulence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 313
4.3.5. Electric fields. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 313
4.4. Conclusions on reducing turbine passage. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 314
5. Improving fish passage at hydropower dams by improving turbine passage. . . . . . . . . . . . 314
6. Special challenges . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 315
6.1. Scale. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 315
6.2. Perspectives . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 315
6.3. Variation among species, stocks, and origins . . . . . . . . . . . . . . . . . . . . . . . . . . . . 316
7. Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 316
Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 317
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 318
1. Introduction
1.1. Background
Aquatic ecosystems have provided us sustenance and transportation since prehistoric times
and we have incrementally developed methods and technologies that enable us to use them to
irrigate our crops, to remove our waste products, to supply mechanical energy, to make and clean
things, to cool thermal power plants and other heat sources, and to generate a substantial fraction
of our electric power (about one fifth worldwide, see World Energy Council, 2001).
C.R. Schilt / Applied Animal Behaviour Science 104 (2007) 295–325 297
Unfortunately, human modification of natural waterways has often been damaging and even
devastating to the ecosystems. It is likely that the scale of future hydropower and other
anthropogenic impacts on freshwater systems, especially in the developing world, will
sometimes be on a sub-continental scale, interconnecting several currently separate river
drainages (Pearse, 2005). Even in currently developed systems such as the Columbia-Snake
River Basin (CSRB), where human population and power demand are rapidly growing, the
prognosis for wild salmon survival seems bleak (Lackey, 2003). In that and similar systems,
where snowmelt is an important source of water for summer passage, a systematic increase in
temperature does not bode well (Mote et al., 2003).
Hydropower dams are but one of many kinds of modifications that human beings make to
natural waterways, but they can have especially profound ramifications for riverine ecosystems
and the animals that live in them (Petts, 1989). In temperate and tropical latitudes water loss
through evaporation can be substantial and reservoirs often undergo seasonal stratification that,
especially when coupled with turbine withdrawal from oxygen-poor strata, can lead to fish kills
and other unfortunate consequences downstream. Conversely generation withdrawal from cold,
oxygen-rich strata also can attract fish to a dam’s tailrace from a downstream reservoir (Nestler
et al., 2002).
Dams inevitably make profound ecosystem-wide changes in rivers, altering everything from
silt transportation, water clarity, and temperature to species diversity and abundance within the
freshwater system itself, from headwaters to estuary, and even into the marine environment.
Dams are increasingly important factors in freshwater systems around the world (Dynesius and
Nilsson, 1994; McAllister et al., 2001; Nilsson et al., 2005).
Unfortunately many indigenous fishes, such as those like salmon, suckers, and darters, that
require fast, cold, and shallow water and clean, well aerated gravel substrates, usually do not fare
well in reservoirs and may be eliminated (Bowen et al., 2003). Unregulated rivers are open
systems (Ward and Stanford, 1989) and by interrupting river interconnectivity dams profoundly
affect, for better or worse depending on species and circumstances, fish populations. However
impoundment is not necessarily or even usually a net loss for the ichthyofauna in general. Some
native fishes, including many popular sport and commercial species and their prey, do quite well
in reservoirs and downstream of dams (Dudley, 1974). Fish of non-native species often have been
introduced by human intervention, and impoundments may increase the abundance and range of
non-native fishes (Martinez et al., 1994). Poe et al. (1994) make the case that introduced channel
catfish (Ictalurus punctatus), smallmouth bass (Micropterus dolomieu) and walleye (Stizostedion
vitreum) have prospered in the CSRB so as to become a major component of juvenile salmonid
mortality themselves and, by competing for its preferred prey of sculpins, sandrollers, and
crayfishes, driving a native fish predator, the northern pikeminnow (Ptychocheilus oregonensis)
from its preferred prey to juvenile salmonids.
Dams inherently present impediments to migration. This is especially problematic for
diadromous fishes (including anadromous salmons and steelhead, shads, lampreys, sturgeons,
some whitefishes, and others which migrate from freshwater spawning to sea and back and
catadromous eels, mullets, and some others which spawn at sea and mature in rivers, reviewed in
Dadswell et al., 1987) or potadromous fishes (which make extensive in-river migrations) to
complete their life cycles (Freeman et al., 2003). Dam-induced changes in distributions of fishes
that do not make long migrations can also occur for ecological reasons, passage reasons, or both
(Schmetterling and McEvoy, 2000; Winston et al., 1991).
Anthropocentric concerns, which inspire many government-mandated rules and protections,
are still much of the motivation behind fish passage and protection efforts and regulations. In
general the fisheries community is concerned with population rather than with individual animal
welfare. Those fish populations in many cases represent perceived human resources, from being
the basis of commercial fisheries to providing recreational fishing opportunities or forage for
such fish populations. It is much easier to protect populations and habitats of fishes that support
economically important sport and commercial fisheries than of those that do not (Ono et al.,
1983; Wissmar and Bisson, 2003).
Conservation concerns increasingly involve water-management plans that may be needed for
the economically important fishes’ survival and transport but also serve more ecosystem-level
concerns, so that an ecosystem’s integrity is becoming more the focus than the welfare of just one
species or population. Still the needs of valued sport and commercial fish populations, which are
often non-natives (e.g. McKinney et al., 2001), and those of other less-valued indigenous fishes
are sometimes at odds.
Although the effects of dams and their operations extend to all aspects of riverine ecosystems
and delayed mortality is an important factor (Ferguson et al., 2006), this paper primarily
addresses the immediate welfare, survival, and timely passage of fish in the vicinity of a
hydropower dam. Though problems are increasing world wide (Gleick, 2004), and the scale of
waterway manipulation in the developing world may become even more immense, this
discussion will be biased towards problems and attempted solutions in the U.S. and Canada in
general and in the U.S. Pacific Northwest’s CSRB in particular.
This system, which is often called the ‘‘Federal Columbia River Power System’’ (FCRPS), is
very heavily dammed with 31 large dams built for the U.S. Federal Government by the U.S.
Army Corps of Engineers (USACE) and the U.S. Bureau of Reclamation (USBR). See Williams
et al. (2005) for a recent and comprehensive discussion of the effects of the FCRPS on system
salmonid populations. As compared with the free-flowing rivers that they replace, impoundments
are deeper, slower-running, generally warmer and often clearer, and support different suites of
populations from phytoplankton to top predators. There are also several large mainstem
nonfederal dams that were built and are owned and operated by local utility districts and other
operators.
Since prehistoric times the CSRB has been the spawning ground of very abundant and
valuable native migratory fish populations, especially those of the anadromous Pacific salmonids
(Oncorhynchus spp.) including Pacific salmon and sea-run steelhead (O. mykiss), as well as
Pacific lamprey (Lampreta tridentata). Some populations of salmonids now have special legal
protection in the form of the U.S. Federal Endangered Species Act (ESA). Pacific lamprey
numbers are much reduced and they, along with three other Lampreta species, have been the
subjects of petitions for ESA listing as well (see the Columbia River Basin Lamprey Technical
Workgroup website http://www.fws.gov/columbiariver/lamprey.htm).
Conflict is inevitable. The dams provide flood control, enable large-scale commercial barge
navigation, supply irrigation for naturally arid agricultural landscapes, and provide public
recreation and substantial related economy for much of the U.S. Pacific Northwest. The dams of
the FCRPS, along with their nonfederal counterparts, provide electric power for much of the
western US. On the other hand the salmonid populations of the Pacific Northwest, besides being
important ecosystem components, are both popular with the general public and support very
valuable sport and commercial fisheries, both at sea and in-river. The native peoples of the Pacific
Northwest are, by law, entitled to and do take an important segment of the commercial in-river
salmonid harvest. Those same salmonid fish populations, as well as the Pacific lamprey, also are
important in the subsistence, religions, and other aspects of the traditions and daily lives of those
native peoples.
The scale of the conflict of utility versus conservation here has meant that considerable effort
has been expended on fish passage and protection over more than a half century. Many
improvements have been made but substantial challenges remain, especially in the downstream
passage of juvenile salmonids and the upstream passage of adult Pacific lamprey. This short
discussion of some of the challenges that have been encountered here, as well as attempts at solving
them, may be helpful in industrial and other large-scale water management conflicts worldwide.
1.2. Statement of problem
Hydropower dams impede the flows of rivers. At a thermal electrical power plant there may be
a need to reduce the numbers of fish being drawn (‘‘entrained’’) into the cooling system. Such
facilities are typically sited alongside water bodies and withdraw some quantity of water for their
operation and return much of it (as with cooling water) to the original water body.
In water withdrawal applications keeping fish from being attracted to or otherwise being
entrained into an intake (fish protection) is often the objective. In such applications water must be
drawn through screens of some sort so as to exclude fish, perhaps even including their planktonic
eggs and larvae (Weisberg et al., 1987; Čada, 1990), without the screened animals becoming
trapped and held (‘‘impinged’’) against the screens (Mathur et al., 1997). Some dams, which do
not interfere with migratory fish populations, are similar to water withdrawals; just excluding fish
from the immediate vicinity of turbine intakes would suffice.
But for most dams on rivers that are obligate routes for migratory fishes, passing the project
successfully in terms of stressors, predation threat, and run timing are as important for subsequent
long-term survival and reproduction, the pertinent currency of ‘‘success’’, as is simply avoiding
being killed by a turbine. It is important to recognize that in many systems, certainly including
the CSRB where fish must pass many dams and reservoirs, that there may be cumulative effects
(Mesa, 1994; Budy et al., 2002).
Anytime we build a water-management structure of any kind we are likely to have important
effects on the distribution, abundance, and well being of fish and their populations. Just building a
structure in water can attract fish of many kinds, both in fresh water (Helfman, 1979) and in the
sea (Klima and Wickham, 1971; Wickham et al., 1973; Love et al., 2005) and rigid structures in
moving water invariably create shadows, eddies, and other anomalies that can attract, retain,
fatigue, or repel migrating fish. Sometimes a structure such as a wing wall or a corner will
produce eddies that can attract or retain migrants. This is the case with a corner of the tailrace at a
small hydropower dam on the lower Susquehanna River in the eastern US where upstream
migrating anadromous American shad (Alosa sapidissima) congregate and are delayed in their
migration. Currents into turbine intakes can attract downstream migrants, which swim with flow.
Since dams delay and concentrate fish they are often attractive to fish predators. Piscivorous
mammals, birds, and fish often congregate near dams and feed on fish that are delayed or injured
there. Recently Federally protected California sea lions (Zalophus californianus) have been
aggregating at Bonneville Dam on the lower Columbia River. Non-lethal attempts at discouraging
birds, which are also protected, include wires hung with shining ribbons stretched across the
tailrace and high-power water cannons. There is a bounty predator removal fishery on the northern
pikeminnow (P. oregonensis, Friesen and Ward, 1999), an important native fish predator that may
have increased in number and ecological importance through human intervention (Poe et al.,
1994). Like the management of dams and reservoirs for the enhancement of nonnative fish
populations (as in McKinney et al., 2001), a government-sponsored predator removal fishery on a
native fish points out the complexity of a notion like ‘‘fish welfare’’ in fisheries management.
The dams of the post-colonial US (Hart, 2004) were tiny by modern standards, but for some
species, for example the economically important American shad (McPhee, 2002), even small
dams can have profound effects. Upstream-migrating adult American shad can be blocked in
their spawning run by a dam just a meter high (Beasley and Hightower, 2000) and upstream
passage problems at some dams continue to limit the return of American shad in parts of their
native range in the eastern U.S. and Canada. Some other fish species may be similarly range-
limited by small dams (examples include Cooke and Leach, 2004; Zigler et al., 2004), including
non-native or otherwise undesirable fishes (Bullow et al., 1988; Thompson and Rahel, 1998;
Porto et al., 1999; Baxter et al., 2003.).
Conversely, even large mainstem dams can benefit populations of some fishes. One dramatic
example involves the growth of the population of non-native (introduced in the late 1870s)
American shad that spawn in the Columbia River (Petersen et al., 2003). In 2004 Bonneville Dam
passed over five million upstream-migrating (pre-spawning) adult American shad through its
fishways. This population growth was in great part made possible by the lower and middle
Columbia River dams which have expanded potential shad spawning habitat and turned a cold,
rapid river to a slower, warmer series of reservoirs (the shad are pelagic spawners, unlike gravel
nesting salmonids). A dam also enabled the upstream passage of shad over a very large waterfall
(Celilo Falls) that, in 1957, was inundated by the closure of Federal (USACE) The Dalles Dam at
The Dalles, Oregon (http://www.cqs.washington.edu/hinrich/shad/JOURNAL3/vol3n2.pdf).
1.3. Anatomy of a hydropower dam
Besides size and configuration there are several different sorts of hydropower dams according
to how they are operated. Three of the most important are ‘‘run-of-river’’ projects which maintain
relatively constant reservoir pool levels so that operators are rather limited as to how much water
they can discharge from a reservoir. ‘‘Peaking’’ dams respond to changes in power demand (air
conditioning in the southern U.S. is a primary factor). Peaking dams, accordingly, have periodic
and often daily fluctuations of reservoir pool level. ‘‘Pumped-storage’’ hydropower projects are
dams that can run some or all of their turbines in reverse, as pumps. These use electric power to
pump water upstream at low-demand times, when electricity is cheaply available, and then
generate power using the pump-turbine units as turbines during high-demand times.
Hydropower dams come in many sizes and take many forms, often depending on the geology
and bathymetry of the site, but there are usually several rather typical components that each can
pass a proportion of downstream flow and migrants (Fig. 2). Pictures and descriptions of a
number of mainstem CSRB dams can be seen at a University of Washington School of Aquatic
and Fishery Science website at http://www.cbr.washington.edu/hinrich/shad/JOURNAL3/
vol3n2.pdf and should give some idea of the scale and structural diversity involved just in one
system.
1.3.1. Turbine passage

Although obtaining survivorship estimates for all fish passage routes presents challenges to be
discussed presently, turbine passage is generally considered the passage route most likely to
injure or kill fish (Fig. 2, bottom centre).
1.3.2. Juvenile bypass system passage

In the turbine intakes of most CRFPS dams there are screens, usually rotating conveyor-belt
like mesh screens (but sometimes bar screens), that are designed to deflect flow and fish in the
upper portion of the turbine intake (juvenile salmonids are mostly fairly shallow in distribution)
upward and into a passage called a ‘‘gatewell slot’’, a vertical passageway. From there they pass
through an orifice and into a juvenile bypass system. The actual bypass is often built as a
modification of the ‘‘ice and trash sluiceways’’ that were designed to pass debris downstream
(Fig. 2, lower centre and right inset).
1.3.3. Spillway passage

Many small dams ‘‘spill’’ water over the top, as do some very large dams without spillway fish
passage, but at most large mainstem dams the spill gates open from the bottom up so that water
and fish ‘‘spill’’ underneath. Spill is currently considered the most benign passage route for
juvenile salmonids here. Disadvantages of spill include elevated dissolved gas downstream,
which can stress or kill fish (Beiningen and Ebel, 1970; Bouck, 1980; Crunkilton et al., 1980;
Lutz, 1995; Backman and Evans, 2002; Backman et al., 2002) as well as reducing swimming
performance (Sciewe, 1974) and resistance to pathogens (Weiland et al., 1999). Spillway passage
also concentrates downstream juvenile passage, thereby increasing predation, and damage fish by
causing turbulence or pressure stresses in high spillway discharge (Fig. 2, upper left).
1.3.4. Surface passage

Surface passage is simply a route that collects fish near the surface of the near-dam forebay
and passes them, sometimes after considerable dewatering, into the tailrace (see Ferguson et al.,
1998). In terms of the proportion of fish passed to the proportion of total water discharge passed
surface passage is by far the most cost-effective route but it passes a relatively small proportion of
the total downstream migration (Fig. 2, upper right).
Potential problems with any fish passage route include trauma and stressors encountered when
passing through the dam and disorientation and injury upon reaching the tailrace downstream.
Any passage route that concentrates fish, especially potentially stressed, disoriented, and
damaged juvenile fish, can increase predation losses since predators can and do congregate at
juvenile bypass outfalls and spillway or powerhouse tailraces (Ruggerone, 1986; Reiman et al.,
1991; Petersen et al., 1994; Isaak and Bjornn, 1996; Blackwell and Juanes, 1998). That, along
with the sometimes violently turbulent and gas-supersaturated conditions there, is why a prompt
departure from the tailrace (or ‘‘tailrace egress’’) is an important consideration for downstream
fish passage.
2. Routes of passage
These four primary passage routes (turbine, juvenile bypass, spillway, and surface passage)
are the only ways, save for some passage through navigation locks, by which any downstream
migrating fish can pass through one of these dams. Most upstream fish passage in the CSRB is via
fishways (called ‘‘fish ladders’’ here). The several primary downstream passage routes, as well as
upstream passage via collection and hauling, fish ladders, and fish lifts (sometimes called fish
elevators) will be introduced independently.
2.1. Upstream fish passage by lorries, ladders, and lifts
There are three means that have been developed for upstream passage of fish around dams that
otherwise block upstream fish passage. A primitive but direct and still sometimes useful method
is capturing upstream migrants below a dam and trucking or otherwise hauling them above the
dam for release. Capture and hauling are costly and difficult in human terms and stressful to the
fish, but can succeed in the short term to re-establish an upstream population (Schmetterling,
2003). Trucking and barging are also an important part of downstream passage of juvenile
salmonids in the CSRB, to be discussed briefly below.
Fish ladders (or ‘‘fishways’’) have been very successful in the CSRB and some designs that are
used worldwide were developed here (Monk et al., 1989). A fish ladder is a sloping, baffled
raceway or sometimes a more naturalistic slope with boulders and other structures that provide
hydrodynamic refuges for resting and function as uphill stairways for fish. There are many
designs of baffles, pools, and weirs that dissipate energy as water descends from the forebay
upstream of the dam to the downstream tailrace. Like stairways, fish ladders are typically rather
long so that the uphill slope can be gradual.
Fish ladders attract upstream migrants into their downstream ends and then on upstream by
providing appropriate attracting flow to guide the upstream migrating fish. Although fishways
here and elsewhere have mostly been designed to pass anadromous adult upstream migrants,
especially shads and salmonids, they can also pass fish of other taxa (Bunt et al., 1999). At
projects where there are fishway entrances at the ends of powerhouses and spillways it is
sometimes helpful to use spillway or turbine discharge in those areas for attraction flow as well.
There are still upstream passage problems remaining in the CSRB (see Moser et al., 2002 for
the special challenges presented by providing upstream passage for Pacific lamprey). But the
fishways in place in the Pacific Northwest, the development of which goes back nearly half a
century (Monk et al., 1989), largely have been successful at passing the large, strongly
swimming, and highly motivated pre-spawning adult salmonids and the non-native American
shad. Diagrams of fishways can be seen in Clay, 1995.
Fish lifts (or fish elevators) are similar to fish ladders in that they use attracting flow
(Barry and Kynard, 1986) but, rather than being constantly open passageways like fish
ladders, passage through them is more complicated. Fish enter a downstream hopper and
are held until, triggered by a timer, a sonar system, or a human operator, the hopper is
closed and then raised from the tailrace level to the forebay level (diagrams of fish lifts
can be seen in Clay, 1995). The hopper then is opened and the fish are released into the forebay
on the upstream side of the dam. Fish lifts can be very effective and have to a considerable
extent replaced ladders at many eastern U.S. dams where upstream passage for shads is an
issue.
Although fish ladders and lifts have been evolving for a long time and work well for many
species, important challenges remain. Unlike salmonids, the lampreys (family Petromyzontidae)
and the eels (family Anguillidae), have bodies and swimming modes that are not very good
against strong currents. The catadromous eels (which reproduce at sea and grow upstream, the
inverse of anadromy in salmonids, shads, sea lampreys, and others) also must get upstream but at
a very small body size. Verdon and Desrochers (2003) suggest that most upstream migrating
juvenile American eels (Anguilla rostrata) do not have enough time to cross two dams, about
80 km apart, in two passage seasons.
2.2. Downstream passage options
Whereas there must usually be some engineered route, such as a fishway or fish lift, to allow
upstream passage at a dam, downstream passage, in one form or another, is hard to stop. As a rule,
downstream migrants don’t use fishways. Neither juvenile nor adult (many post-spawning shad
and some steelhead adults, called ‘‘kelts’’ return to sea and catadromous eels migrate downstream
to spawning grounds at sea) downstream migrants use fishways for passage in appreciable
numbers.
Concerns with safer downstream passage of juveniles began to be addressed as soon as the
means (e.g. primarily biotelemetry and fisheries sonar) were available and downstream fish
passage and survival could be studied. The development of upstream passage routes for large,
powerful, and strongly motivated upstream-migrating adults to and through fishways is a much
simpler proposition than is collecting downstream migrating juveniles which are small, young,
and at various stages of development out of a deep and wide hydropower forebay and passing
them safely and quickly downstream. Understanding of and engineering for downstream passage
is a much more difficult and more recently addressed set of problems that are exacerbated by the
dearth of knowledge of the motivation, orientation, sensory capacities, and hydrodynamic
preferences of downstream migrants.
2.2.1. Turbines
Generally turbine passage is thought to be the most likely of all of the downstream passage
routes to harm or kill fish. A good deal of work has gone and is going into improving turbines to
reduce damage and increase survival and will be briefly discussed below. Water speeds are often
very high.
2.2.2. Juvenile bypass systems

At a dam there is a downstream (juvenile) bypass route. Fish are deflected upward by intake
screens they pass upwards through a vertical passage called a ‘‘gatewell slot’’ and then through a
horizontal orifice (see Fig. 2) and into the juvenile fish bypass.
2.2.3. Spillways
When the spill gates (either curved ‘‘Tainter’’ gates as in Fig. 2 or flat gates) are raised the
water flows underneath them and down into a ‘‘stilling basin’’ (Fig. 2) before continuing
downstream through the spillway tailrace. In the stilling basin water and air are forced deep
underwater and the pressure there can cause the water to become supersaturated with air.
2.2.4. Sluiceways and other surface routes

Surface passage is, in terms of the cost (in water diverted from power generation) vs. benefit
(in terms of number of fish bypassing turbines), very desirable. Juvenile and other downstream
migrating fish have been passing over waterfalls, including the once spectacular Celilo Falls
(http://www.critfc.org/text/celilo.html) on the lower Columbia River, although there are no
estimates of what the mortality rate of that passage route may have been.
Challenges in surface bypass development, as with other bypass routes, include attracting fish
to the entrance, conveying them quickly and safely to the tailrace or further downstream, and
delivering them there in such a condition and in such a place so that their survival is likely. The
physical nature of the bypass route and outfall are important (Den Bleyker et al., 1997; Johnson
et al., 2003).
2.2.5. Transportation
Since 1981 (Ward et al., 1997) the USACE has been collecting juvenile salmonids from
bypass systems at several dams (McNary Dam on the middle Columbia River and three of the
four USACE dams on the lower Snake River) and barging or trucking them downstream (Fig. 1).
The bypass is partially dewatered and a rather elaborate system of gates enable the fish to be
Fig. 1. Schematic drawing showing the main waterways and mainstem dams in the Columbia-Snake River Basin. Note
the four dams, marked in white, at which some proportion of the juvenile salmonids that are screened into the juvenile
bypass systems, can be collected for transport. Originally published in Wertheimer and Evans (2005) and used by
permission of Robert Wertheimer, U.S. Army Corps of Engineers District, Portland, OR, USA.
bypassed directly to release into the dam’s tailrace or diverted and held for separation,
examination and tagging, and sometimes for transportation by barge or truck downstream past
the last dam, which is Bonneville Dam.
3. Sources of passage-related delay, stress, injury, and mortality
3.1. The importance of run timing
To improve fish migration success, delay in hydropower-effected rivers occurs at two levels:
overall migration timing and timing with regard to encountering and passing through a
hydropower dam, including the tailrace. The passage of the immediate forebay-dam-tailrace (or
tailrace-fishway, forebay for upstream migrants) is of primary interest here. Delay of upstream
migrants can occur from difficulty finding a fishway entrance or refusing to enter and remain
inside for lifting or to continue upstream through and from ladders.
There is an especially pernicious form of upstream delay called ‘‘fallback’’ (Reischel and
Bjornn, 2003; Boggs et al., 2004) which is just what the word suggests. Sometimes fish leave
fishways or lifts just to be drawn back downstream through some other passage route such as a
spillway or turbine intake. That is the explanation for the otherwise disturbing fact that upstream
(ladder) counts of upstream migrating American shad are often higher at The Dalles Dam than at
Bonneville Dam. Bonneville Dam is about 50 miles downstream of The Dalles and so every
upstream migrant of any species must pass through Bonneville Dam before reaching The Dalles
Dam. Many shad seem to fall back at The Dalles Dam and so are counted there more than once.
Rather little is known about the sensory and behavioral mechanisms by which downstream-
migrating juvenile salmon migrate but the wide and deep waters of a hydropower forebay can
cause substantial delays, up to several days, as fish seem to seek out sensory stimuli that direct
them downstream. We do know that juvenile salmonid passage, at least in this system, follows
fairly clear diel trends with ‘‘deep’’ passage through turbines and spill bays occurring in late
evenings and early mornings (Thorne and Johnson, 1993) and ‘‘shallow’’ surface passage through
some engineered surface route occurring more in daytime. For juvenile salmonid dam passage at
dams in the CSRB that means that, since most passage occurs from an hour or so before twilight
until an hour or so after dawn, a great many fish pass through turbines and spill bays. One
advantage of surface passage, besides its dramatic water economy, is that it provides for daytime
passage and thereby reduces delay.
The tailrace environment can also be the site of delay for downstream migrants, perhaps due to
disorientation and stress from passage or water circulation patterns that hold migrants and delay
tailrace egress. Even inside the dam itself there can be substantial delay (Liscom, 1971; Beeman
and Maule, 2001). At the project (forebay-dam-tailrace) scale delay is especially important
because it increases both the level and duration of predation pressure by concentrating
downstream migrants, allowing predators to congregate. In general delay, of either downstream
or upstream migrants, should be reduced as much as possible (Castro-Santos and Haro, 2003).
The longer time scale of the full downstream migration is somewhat more complex. Long-
distance animal migrations are often timed to take advantage of or avoid temporally and spatially
varying conditions such as temperature, flood stage, trophic resources, or other environmental
variables. For a reproductive migration the appropriately timed arrival of potential mates and
conditions for reproduction are important. For diadromous fish that are moving from fresh to salt
water the body must undergo profound physiological changes to make that transition (called
‘‘smoltification’’, see Tiffan et al., 2000 on Pacific salmonids, the juvenile downstream migrating
juveniles of which are called ‘‘smolts’’) and being either delayed or accelerated can reduce
survival in the new environment. In general, impounded rivers pass fish downstream more slowly
than do free-flowing rivers and so juveniles arrive in the estuary and near-shore marine
environment later and with lower energy reserves than they would have in a free-flowing river
system (Williams et al., 2005).
3.2. Turbine passage
Not all fish that pass dams by turbine die as a result of doing so. The proportions of fish killed,
injured, or stressed by turbine passage can vary with species, age, time of year, water
temperature, turbine type and operations, and other factors. For example Taylor and Kynard
(1985) found mortality rates over 40% for juvenile American shad and blueback herring (Alosa
aestivalis), which are notoriously fragile animals, but Mathur et al. (1994) reported immediate
(within 1 h of passage) mortality from zero to about 15% for juvenile American shad. Such
discrepancies may be due to many things including time of year and water temperature, differing
experimental and analytical methods, and different turbine types and operating conditions.
Bickford and Skalski (2000) reanalyzed 25 years of Columbia-Snake River System survival
studies and found juvenile salmonid (Oncorhynchus spp.,) mortality rates between about 7 and
13%. Even though some estimates of turbine mortality rates may seem fairly low, it should be
recalled that mortality studies are mostly of immediate and fairly short-term experiments, often
48 h, and that there may be many dams and reservoirs that must be passed both ways by upstream
spawning populations. Most Snake River in-river migrating juveniles must pass eight dams.
There has been a good deal of thought and work done on the mechanisms by which fish are
damaged and killed in turbine passage (Wittinger et al., 1995) and they include pressure changes,
shearing between differently moving water masses, impact with turbine or dam structures,
cavitations (very low pressure zones), grinding, and abrasion. Coutant and Whitney (2000)
review fish behavior with regard to turbine passage. They discuss dam approach, forebay
disorientation, buoyancy, fish condition, and perhaps most important, the vast gaps in our
understanding of fish behavior in dam passage.
3.3. Juvenile bypass system passage
Besides delay (above and see Liscom, 1971; Beeman and Maule, 2001), juvenile bypasses
present other problems. These include impact with and descaling and other damage from screens
and other bypass system hardware, in-bypass predation, and predation at the point that bypassed
fish are returned to the river (potentially disoriented and stressed) to a place where predators can
and typically do congregate.
3.4. Spillway passage
Spillway passage is, in the Columbia-Snake River Basin, generally considered the most
benign way to pass large numbers of fish. From the hydropower perspective, the primary
drawback of spill is foregone power generation represented by the spilled water. There is also the
previously mentioned increase in dissolved gas that can cause stress and death in fish downstream
of the spillway. Other potential problems include many of those involved in turbine passage
(striking structures, descaling, damage in violently turbulent flows). In spillway tailraces
potential fish predators, including piscivorous fishes, birds, and sometimes mammals congregate.
3.5. Surface passage
The only negative aspect of surface passage seems to be that too few fish in the CSRB use
those routes. It is by far the most cost-effective in terms of number of fish per bypassed water.
Fish mostly avoid the lethal aspects of turbine and spill passage including pressure changes,
abrasion, cavitations, etc. The structure of the bypass outfall must be carefully considered (Den
Bleyker et al., 1997; Johnson et al., 2003). There can be unknown problems and properly
designed and conducted survival studies are needed but it seems likely that surface passage is, if
the system works well and the outfall is sited and constructed properly, the most benign passage
route and the one that diverts the least water from turbines.
3.6. Transportation
The barging or trucking of downstream migrants is an important tool in fish passage and
protection in this system (Williams et al., 2005). Return rates (determined by methods to be
described briefly below) compared between transported and in-reservoir migrant salmonids are
variable, with one route doing better in some years and the reverse in others. The reasons are
unclear but important variables seem to be in-system environmental conditions, especially water
temperature and flow rates and feeding and predation conditions in the estuary and just offshore
at the time the smolts are delivered and released below the most downstream dam as well as the
developmental, energetic, and other physiological condition of the fish (Williams et al., 2005).
Whereas the entire downstream passage time, from either natural or hatchery production
to estuary and near-shore environment is presumably slower than would be the case in a
free-flowing system, trucking or barging delivers fish downstream faster than would a free-
flowing system.
It seems reasonable that, even though there are stresses and mortality attributable to fish
collection and transportation (Maule et al., 1988), in a drought year when spill rates may be low
and in-system conditions poor (reservoir flow rates and spill levels low and water temperatures
high) that transportation might be helpful. Transportation is also not an option for any of the
millions of juvenile fish that enter the system downstream of the lowest collection dams so that
many juveniles in this system cannot be transported (Fig. 1). Columbia-Snake System-wide
downstream passage strategies for dams, as well as in-reservoir passage and survival
enhancement through flow augmentation is reviewed by Giorgi et al. (2003).
4. Improving fish passage at hydropower dams
The existence and operation of many water withdrawal facilities and hydropower dams
produce unfortunate effects for river and other freshwater systems and the animals that live and
move through them. Dam removal, at least the removal of small dams, can lead to dramatic
improvements in habitat quality and population dynamics (Kanehl et al., 1997; Smith, 2000), and
even some larger dams are and surely will be considered for decommissioning and effective
removal (‘‘breaching’’, or opening to restore relatively unimpeded flow). But it is certain that
most of those facilities will persist for the foreseeable future.
Hydropower dams were not designed to pass fish safely downstream. They were designed and
built as power-generating plants and so juvenile (downstream) fish-passage improvements, both
structural and operational, are obliged to be retrofit to the structures and operations that already
exist. Dams will always have ecosystem-wide effects but it may be that, as new dams are
inevitably built in the developing world, what has been and will be learned can be applied in the
original construction. New dams may start service as better passage structures and be more
readily monitored and improved than are those of the current generation.
4.1. Changing project structures
Dams and water withdrawal facilities are structures and so structural changes, such as the
development of screening systems (as described in Gessel et al., 1991), louvers (as in Amaral
et al., 2002), or even a riprap groin to alter a fish-entraining circulation pattern (Nestler et al.,
1995; Ploskey et al., 1995), are possible. Recently the USACE has developed both a prototype
surface collector for Bonneville Dam’s first powerhouse and a fully operational expansion of its
second powerhouse sluiceway. This second passage route, called a ‘‘Corner Collector’’ because it
is on one end of the powerhouse, collects juvenile downstream migrants and then passes them
down a conveyance channel, which delivers them into the spillway discharge plume so that they
move downstream quickly and spend little time in the tailrace.
Besides surface passage, spill bays have been modified to reduce dissolved gas saturation and
turbulence (Muir et al., 2001), fishways and downstream bypasses are being improved (Kynard
and Buerkett, 1997; Kynard and Horgan, 2001), and new forms of fishways have been developed
for especially challenging species such as Pacific lamprey. Lampreys are jawless fishes and
upstream-migrating adults passing rapids must rest frequently by holding on to some rigid
structure with their oral discs. Lamprey passage at mainstem dams has been problematic (Moser
et al., 2002). An innovative new upstream passage route design for adult Pacific lamprey is now
being tested.
The new lamprey passage structure consists of a series of inclined, shallow troughs
interspersed with resting boxes with water cascading down them. The adult lamprey grasp onto
the smooth, wet, metal surfaces with their oral discs to rest and to move upstream and uphill,
which they accomplish by both swimming and by literally throwing their upper bodies ahead of
their oral discs and then reattaching quickly before they fall back. The Pacific lamprey’s climbing
ability is considerable; they once quite successfully climbed and passed upstream at Celilo Falls
and they can scale a vertical concrete wall in air as long as it is wet, but they have trouble passing
around corners, which are ubiquitous in the fishways, without losing their grip and being washed
back downstream. Besides the ongoing effort to provide adult lamprey with their own passage
structure those corners are being rounded as possible during seasonal dewatering for maintenance
in order to facilitate lamprey fishway passage.
The USACE Walla Walla (Washington) District has installed and tested a large-scale
behavioral guidance structure, essentially a steel curtain about 330 m long and ranging from 17 to
24 m deep (Adams et al., 2001) to guide fish to a surface bypass collector (Johnson et al., 2000,
2005a,b). Over the last several years the Portland District of the USACE has modified and tested
turbine intakes at Bonneville Dam’s second powerhouse, continuing the effort described in
Gessel et al. (1991).
4.2. Changing project operations
Many of these additions and other modifications have reduced delay and increased survival
and those efforts should and will continue. However, modifying structures on the scale of a
mainstem hydroelectric dam can be very expensive. In some cases it is possible to achieve fish
passage goals (decreasing delay and increasing survival) by changing project operations (the
scheduling and proportions of water passed through various routes. Project operators may have
considerable scope for adjusting discharge locations and rates so as to pass fish more effectively
and safely, within the limits of their available water, required generation schedule, and equipment
availability.
For example both fish passage distributions and electric energy demand may be cyclic on a
diel schedule. Spilling more water at night, when more juvenile salmon pass the dams than during
daytime (Thorne and Johnson, 1993) but conserving spill during the day when fewer pass deep is
a standard strategy in this system. Fewer juvenile fish pass by spill during daytime and lower
daytime spill, besides conserving water for power generation or spill at another time, it is less
likely to induce adult fallback (see Section 3.1 above) than is higher daytime spill. If years of data
indicate that more fish pass downstream by spill during nighttime then, in drought or whenever
spill is limited by water availability, it may be prudent to reduce daytime spill so as to conserve
water for power generation or for fish passage later.
On the other hand some minimum spill level or spill to generation ratio may be needed to
maintain spillway attraction currents for upstream migrants (such as returning adult salmonids) and
to keep the spillway forebay water quality from degenerating. If there is a threshold of spillway
discharge or proportion above which many more fish are not passing, then spilling much more may
be counterproductive. Very low spill levels may retain spill-passed fish in the very turbulent
‘‘stilling basin’’ just downstream of the spill gates (See Fig. 2) where predation rates can be high.
Retaining or passing water from colder upstream reservoirs can help reduce and delay summer
warming and flush juveniles down the reservoir system. It may be possible to make operational
changes such as limiting power generation or spilling water for fish passage in line with the
presence or absence of large numbers of fish and the need either to exclude them from a project
C.R. Schilt / Applied Animal Behaviour Science 104 (2007) 295–325
Fig. 2. Study area showing locations of Federal Columbia River Power System dams on the lower Columbia River in Washington and Oregon and the lower Snake River,
Washington, including Lower Granite, McNary, and John Day dams, where steelhead kelts were radio-tagged and released in 2001 and 2002.
309
area or to provide them with passage. Whenever it is possible the natural history of the
populations of interest should be studied and should inform the development of fish protection
and passage measures. In these profoundly altered ecosystems a new natural history, that of
human-created environments like dam forebays and tailraces, is needed to inform construction,
operation, and policy decisions (Schilt, 2002).
4.3. Using sensory stimuli to control local fish distributions
Fish are actively behaving and mobile animals with a wide range of sensory systems with
which they perceive changes in the environment (Atema et al., 1988; Kapoor and Hara, 2001;
Collin and Marshall, 2003). Fish of all sizes can be surprisingly powerful swimmers. It is
reasonable to consider using engineered stimuli to direct fish movement for their own protection
or passage at an industrial site. In the Pacific Northwest the search for such tools goes at least
back to the early 1950s (Burner and Moore, 1953) and some fairly well developed structural
guidance systems, such as screens and louvers, have inherent sensory components. Much more
recently Popper and Carlson (1997) have reviewed problems, progress, and opportunities. Of all
the many sensory modalities that fish have and use, most save for the chemical senses and touch-
based senses have been explored. What follows is a brief catalogue of efforts to use sensory
stimuli to accomplish or improve fish protection and passage arranged by sensory system.
4.3.1. Light
Light stimuli are relatively easily and cheaply produced and fish of most kinds are visually
sensitive animals (Fernald, 1988). Aquatic animals of many kinds carry on diel vertical
migrations (Kelso, 1978; Levy, 1990a,b; TeWinkle and Fleischer, 1999; Scheuerell and
Schindler, 2003), which are probably mediated by some change in light level, direction, polarity,
or other quality (Ringelberg, 1995).
Light has been used both as an attractor and as a deterrent. For centuries it has been know that
light can attract fish and other aquatic animals (Wickham, 1973) and attracting lights are part of
subsistence, commercial, and sport fisheries world wide. Continuous light has been used as an
attractor to draw fish away from hydropower turbines including those at Richard B. Russell Dam,
a USACE Savannah District pumped-storage project (Nestler et al., 1995; Ploskey et al., 1995) on
the Savannah River in the southeastern U.S. Strobe lights have been found to be effective in
laboratory and cage (usually tank or net pen) tests (Nemeth and Anderson, 1992; Amaral et al.,
2001; Mueller et al., 2001; Patrick et al., 2001; Ploskey and Johnson, 2001; Taft et al., 2001) and
at some field sites (Johnson et al., 2001a,b; Mueller et al., 2001; Taft et al., 2001; Johnson et al.,
2005a,b). Although strobe lights have been useful in some places with some populations the
efficacy of any light-based behavioral system can be limited by turbidity, which is often high at
mainstem dams. Sites like the Hiram H. Chittendon Locks in Seattle (Johnson and Ploskey, 2001;
Johnson et al., 2001a,b, 2005a,b) where water is usually fairly clear and pertinent distances are
short or in clear mountain lakes in Idaho (Maiolie et al., 2001) where water clarity is very high are
likely places for lighting systems. As water clarity declines or distances increase light’s
effectiveness is liable to decline. The same may be true under daylight conditions (Amaral et al.,
2001); any sensory stimulus occurs in the presence and in the context of a background noise
regime that potentially can mask it from an animal’s perception, recognition, or source direction
determination.
Light, both constant and flashing, has promise for fish protection and passage. The visibility to
migrants of intake screens and other bypass-related hardware and how that might relate to
passage and survival are little explored. We know that more salmonid juveniles pass these large
Columbia and Snake River dams deep (via turbines or spill bays) at night and that surface passage
is generally higher during daytime but we do not know the relationship, if any, of ambient light
level to surface fish passage rate. As with any behavioral system, continuous and strobe lights
may be subject to habituation, which is an issue in any sensory stimulus application. Spectrum-
specific differences in fish response, the effects of background and contrast, light polarity, and
many other aspects of fish response to light remain to be explored.
4.3.2. Low-frequency sound

Like light, sound is relatively easy to produce in water, especially at fairly high frequencies.
Fishes are diverse and so are their various capacities for sound transduction, processing, and
response. What we usually call ‘‘sound’’ is a propagating wave of pressure changes that causes,
either directly or indirectly, the bending of hair cells in the inner ears of fish and other animals.
Fishes that have their inner ears stimulated directly by the sound and the concomitant whole-body
and inner ear motions are called ‘‘hearing generalists’’ and they hear mostly below 1 kHz. The
hearing generalists include most fishes including all of those without gas bladders (also called
swim bladders in some literature) as well as salmonids and eels.
The hearing specialists all have gas pockets of one form or another, including the gas bladder,
and some means for transmitting pressure changes to the inner ears or gas pockets in or very near
the ears themselves. A gas bubble or pocket in water will convert scalar pressure changes (sound)
to vector motion that can stimulate hair cells. The hearing specialists include the catfishes, the
herrings and their allies, the minnows, carps, and others. Some hearing specialists can, because of
the influence of the gas pockets and their coupling to the ears, hear up to several kHz.
The mechanosensory systems of fishes are, like those of all vertebrates, built upon the sensory
transduction, sound to nerve impulse, accomplished by vertebrate hair cells (Howard et al., 1988;
Hudspeth and Markin, 1994). Hair cells function as accelerometers, both in the inner ears (which
mediate both hearing, Schellart and Popper, 1992; Fay and Popper, 1999; Fay and Megela-
Simmons, 1999) and the vestibular system, which senses whole-body and head linear and
rotational velocities (Kalmijn, 1988, 1989). The skin-borne ‘‘lateral line’’ transduces differential
pressures and water motions across the skin (Coombs et al., 1988, 1989; Kalmijn, 1988, 1989;
Coombs and Montgomery, 1999).
As compared to mammals most fish have hearing frequency sensitivities that are rather low,
limited to frequencies below just a few kHz. Despite being rather frequency-limited, fish hear
well and use hearing to assay their environments (Fay and Popper, 2000). The biomechanics of
fish hearing, and of directional hearing especially, is complex and beyond this discussion but
there are recent reviews, especially Fay and Popper (1999). Schilt and Nestler (1997) have
provided a primer of fish bioacoustics that was written with hydropower applications in mind.
Fish of several taxa are thought to respond to subsonic or ‘‘infrasonic’’ (below the human
lower sensitivity range limit of about 20 Hz) sound (Karlsen, 1992a,b; Enger et al., 1993; Sand
and Karlsen, 2000), and even extending down to lower than 1 Hz (Sand and Karlsen, 1986). Very
low (infrasound) frequencies have been tested with regard to fish passage and protection. Sand
et al. (2000, 2001), Amaral et al. (2001), Ploskey and Johnson (2001), Mueller et al. (2001) and
Taft et al. (2001) have found strong and predictable responses in a variety of species and life
stages. Unfortunately, perhaps due to the engineering difficulties involved with producing a
reliable stimulus source or the difficulty with predicting sound propagation and response in an
actual dam environment there have been no proven long-term applications for infrasound in fish
passage or protection.
Sound audible to humans also has been tested as attractant (Patrick et al., 2001 adult Angilla
eels) and as deterrent (Schwartz and Greer, 1984 Pacific herring (Clupea harengus), Smith and
Anderson, 1984 various reservoir fishes; Haymes and Patrick, 1986 alewife Alosa
pseudoharengus; Ploskey et al., 2000 juvenile salmonids at a hydropower dam). A wide
variety of sounds including impulsive bangs and recorded predator sounds have been tried but,
although there have sometimes been initially promising results (as in Popper et al., 2002) there
has yet to be a robust and effective fish protection system based on audible (to humans) sound, at
least in this river system. Reasons may be many and complex and may include the often very
noisy near-dam environment, difficulty for the fish in distinguishing the stimulus and its direction
in the noise, acoustic boundary conditions that were not properly considered, or habituation.
4.3.3. High-frequency sound and alosine herrings

The discovery of ultrasound (over about 20 kHz) hearing and response in a fish owes its initial
discovery to fish-passage science. The hearing sensitivity ranges of fishes have long been known
to be much lower than that of most mammals and so it must have been surprising when, in 1982
at a raceway at Holyoke Dam on the Connecticut River Boyd Kynard observed that the
downstream migrating post-spawning American shad that he was sampling using sonar were
avoiding this sound beam (161.9 kHz in this case, Kynard and O’Leary, 1990). Subsequent work
at the site indicated that the sound field was effective at temporarily concentrating down-running
adults but that they would finally pass through or perhaps under the beam. Upstream running
(pre-spawning) shad were more successfully concentrated by the sound (Kynard and O’Leary,
1991). Fish hearing has long been know to be limited to just a few kHz and very few animals of
any kind hear above several dozen kHz and so it is especially surprising that any fish would hear
so high. Since Kynard’s discovery a good bit of work, both laboratory and applied, has been done.
Although ultrasonic hearing has been shown in the laboratory for cod Gadus morhua (Alstrup
and Møhl, 1993, 1998), the all-time high-frequency hearing champions for the fishes are, so far at
least, several species of the alosine herrings (subfamily alosinae, family clupeidae). The alosine
herrings are a subfamily of the anadromous herrings (Alosa and allied genera). Like the other
herrings, these are ‘‘hearing specialists’’ and might be expected to hear sounds somewhat higher
than 1 kHz (Enger, 1967) but they turn out to be able to hear and to respond by swimming away to
short-duration sounds of up to 180 kHz (Mann et al., 1997, juvenile American shad) or 200 kHz
(Gregory and Clabburn, 2003, adult upstream migrating adult twaite shad A. fallax fallax). Not
only can fish of all of the alosine species and life stages studied so far hear these sounds (Plachta
and Popper, 2003; Higgs et al., 2004), but all those appropriately tested swim away from the
source of the clicks. The response may be adapted as a predation-avoidance response to the
echolocation clicks of some marine mammals (Nestler et al., 1992; Mann et al., 1998; Alstrup,
1999), although the fully marine herrings do not seem to possess ultrasonic hearing (Mann et al.,
2001, 2005).
The repulsive effect of ultrasonic pulses on at least some alosine herrings has provided the
most successful attempts, to date, for acoustic fish protection in industrial environments. In the
late 1980s workers affiliated with Waterways Experiment Station, the USACE research center in
Vicksburg, MS., led by John Nestler, began investigating the potential for ultrasound response in
the alosine blueback herring to be useful in fish protection in the southeastern US (Nestler et al.,
1992). Thereafter pulsed ultrasound has been used in fish protection at several dams and other
power plants (Nestler et al., 1995; Ploskey et al., 1995; Dunning et al., 1992; Ross et al., 1993,
1996). There are two different ultrasonic pulsed sounds currently in use. In the southeastern U.S.
studies have used use a very simple sinusoidal waveform about 5 ms long, similar to the sonar
clicks that both fisheries sonar and, to a much lesser extent dolphins, use (Nestler et al., 1992,
1995; Ploskey et al., 1995). In the northeastern U.S. a half-second long band-limited noise signal
was used (Dunning et al., 1992; Ross et al., 1993, 1996). Unfortunately the performance of these
two signal types or any other candidate sounds have not been compared in terms of fish response,
resistance to acoustic masking in noisy industrial environments, sound source localization, or
habituation.
4.3.3.1. Important questions remain unanswered. What are the most effective pulse widths,
repetition rates, effects of noise and masking, and habituation for different species and life stages
of alosine herrings? What is the upper limit of the ultrasound sensitivity is in Alosa spp. (at least
to 200 kHz in twaite shad A. fallax, Gregory and Clabburn, 2003) and is ultrasound response
imanifested across the other genera of the subfamily alosinae around the world? Hydropower and
other development in Africa, Asia, and South America may make such studies timely.
4.3.4. Flows and turbulence

Aspects of the mechanosensory systems also mediate a fish’s sensing both the orientation and
change in position of its body in the water and of the water on its body. As is the case with all
vertebrates, fish have semicircular canals, which are the end organs of the vestibular system in
their ears. That system transduces and sends to the brain sensations of linear, including that due to
gravity, and rotational accelerations. With the vestibular system, along with vision, fish maintain
balance, differentiate up and down, and recognize when the animal is speeding up or slowing
down, going around a curve, or tumbling end over end. Much of the fish-passage process takes
place in darkness and then the vestibular system may be the primary or only available sense for
spatial orientation, save for the lateral line system. Important aspects of vestibular and hearing
functions in inner-ear structures have recently been reviewed by Popper et al. (2005).
The lateral line system, using hair cell based end organs called neuromasts, which are arrayed
either on the skin’s outer surface or in tubes in the skin or both, detect near-body water motions
and differential pressures across the body (Coombs et al., 1989). Lateral line systems of many
descriptions occur on and in the skins of fishes (Coombs et al., 1988; Coombs and Montgomery,
1999). The lateral line system mediates orientation to flow (‘‘rheotaxis’’, Arnold, 1974;
Montgomery et al., 1997) as well as prey location (Montgomery and Milton, 1993; Kanter and
Coombs, 2003), schooling (Pitcher et al., 1976), and obstacle avoidance, especially in darkness
(Blaxter and Batty, 1985).
For fish passage the vestibular and the lateral line systems are involved in the fish’s orientation
in both the gravitational and flow fields in which a fish finds itself. Although little is known about
juvenile salmonid motivation and preferences for passage in the hydrodynamic environment, it
seems reasonable that there are physical signals which in part regulate fish distribution and,
especially, whether they accept or reject a given route. There has recently been some interest in
using engineered turbulence to attract migrating juvenile salmonids to bypass entrances
(Coutant, 1998, 2001) and some very preliminary studies have been done, but the endeavor is
very much in its infancy.
4.3.5. Electric fields

The use of electric fields to control fish distributions is fundamentally different from the other
sensory methods discussed so far for two reasons. First, in order to function it serves as a strongly
aversive stimulus that potentially can be damaging to the fish. Second, fish, especially if they are
upstream of the field, can be stunned and debilitated by the field (Roth et al., 2004) and so be
drawn deeper into and through whatever route they were meant to be repelled from. Usually the
field must be downstream of whatever access is to be denied the fish so that a disabled animal will
be washed back downstream and out of the field. Other limitations include human and wildlife
safety and being restricted to fresh water environments. For these reasons electric barriers have
not held an important place in the toolbox of things that hydropower project managers and
researchers can use. Primary uses have included stopping or slowing the spread of invasive fishes
(Verril and Berry, 1995; Macaena et al., 1999; Swink, 1999; Savino et al., 2001; Clarkson, 2004)
but, since the electric field is often not fatal, the downstream limitation still applies.
4.4. Conclusions on reducing turbine passage
With all behavioral methods it is important to be aware that the visual system and hearing and
all of the other systems operate in concert within themselves and among each other and responses
to a sound might be affected by light level, current, or any number of other sensory factors. An
animal’s sensitivity to any physical phenomenon, as well its response to that stimulus, is
dependent on many things. Enhancing or limiting factors might range from the genetic makeup
and life stage of the subject animal through environmental factors including season, time of day,
light level, presence of predators, distance to cover, temperature, group size, noise regime,
current, and many other things (Schilt and Norris, 1997). Assuming similarity of any behavioral
response across taxa is always ill advised, an argument made especially earnestly and well by
Bullock and Heiligenberg (1986). There may even be important differences in sensitivity or
response at subspecies levels. Nemeth et al. (2003) report not just different but opposite
orientation and movement between two strains of landlocked Atlantic salmon (Salmo salar) in
response to the same hydrodynamic stimulus.
At hydropower dams, at least, fish passage phenomena seem to be surprisingly variable with
respect to site, season, time of day, and project operations. Given the complexity of industrial
waterscapes and the site specificity and transient nature of passage the evaluation of a behavioral
treatment’s efficacy is difficult. Often it is impossible to generate an experimental design with
experimental units sufficient to detect small differences in whatever passage metrics have been
chosen as response variables. On the other hand, in most cases of protection and passage the scale
and stakes are such that a substantial effect, something far more than statistical significance, is
required for a treatment to be useful.
For a project’s owners and operators or a regulatory resource agency to concede or to deny
efficacy, a concession that can have important ramifications for subsequent project licensing and
operations, conservation dictates caution. In general what we consider ‘‘efficacy’’ is not just
moving, excluding, or acceptably passing enough more (or entraining fewer) fish than under
control conditions to infer that the result was, within some confidence limits, not due to chance.
We are looking for substantial effects in attracting fish or chasing them away. Short migration
seasons, diel cycles of fish distribution and behavior, project operational constraints, and other
considerations often limit us to fairly small sample sizes. It is well that the validity of an
hypothesis test’s outcome of significance is not compromised by small sample size and low
experimental power (Chittendon, 2002).
5. Improving fish passage at hydropower dams by improving turbine passage
For many years an important and perhaps the primary measure of success in fish passage in the
Columbia-Snake River system has been fish passage efficiency (FPE), which is the estimated
ratio of all fish passing a project by non-turbine routes to the total number passing the entire
project during the same time period. An FPE of 80% and a project survival of 95% is the
mandated goal of the U.S. Government’s National Marine Fisheries Service (NMFS, 2000) for
the dams in the Columbia-Snake River Basin.
No downstream passage route is perfect and estimates of turbine survival rates are far from
100% so it is not unreasonable to try to improve the passage environment for whatever fish will
inevitably pass through turbines. While large-scale hydropower turbines are very expensive and
last a long time, those at many dams in the CSRB and elsewhere will soon need updating or
replacement. There has been a concerted effort by the U.S. Department of Energy to design, test,
and build turbines that will pass a greater proportion of animals relatively unharmed. Whereas the
popular term ‘‘fish friendly turbines’’ may be a bit optimistic it is should be possible to build
turbines that provide a substantially less hostile fish passage environment. A new design
involving a ‘‘minimum gap runner’’ (Čada, 2001) has been designed. Much of this work is done
at the U.S. Department of Energy’s Idaho National Engineering and Environmental Laboratory
Hydropower Program in Idaho and considerable current information is available for both the
Advanced Turbine http://hydropower.inel.gov/turbines/index.shtml and Environmental Research
http://hydropower.inel.gov/environmental/index.shtml programs.
6. Special challenges
Fish passage and protection is an essential part of any large-scale attempts to maintain the best
possible biotic integrity in the aquatic systems we have and will develop. Some progress has been
made but there are special challenges involved, some of which may be peculiar to the field.
Industrial environments are extensive and, from a fish perspective, can be very challenging.
Sensory and behavioral mechanisms of navigation that function well in natural systems may be
less effective in engineered waterways. If the payoff for making the wrong decision in passing a
dam is death then learning is unlikely to help. In the attempt to develop more successful
engineered passage routes we encounter several interesting and challenging peculiarities of
dams.
6.1. Scale
The spatial scale on which fish behavior experiments are conducted is important. Managers in
the CSRB are increasingly unlikely to fund laboratory, net pen, and flume studies. It is correct
that a fish’s response in such an environment might be very different than it would be in an actual
dam forebay with unpredictable currents, noise, and predators. On the other hand the full-scale
‘‘try it and see what happens’’ method takes years and costs millions and may exacerbate rather
than help the problem.
6.2. Perspectives
Over the last decade there has been an effort to bring together hydropower managers,
engineers, and fish mechanosensory biologists for meetings and workshops (Carlson, 1994;
Carlson and Popper, 1997) to exchange ideas and perspectives. In general, industry addresses
problems on a project-specific basis. Especially in the case of a private utility with one or just a
few dams the priority is, perhaps rightly, to try whatever might ‘‘work’’ at a specific project.
‘‘Work’’ here means to move the project towards whatever fish passage goals have been set by the
regulating agencies. That is quite reasonable in that hydropower dam managers, like others in
industry, are charged with producing a product (in this case electric power) in a consistent and
cost-effective way.
Very little systematic empirical work has been done to elucidate the motivational, sensory, and
behavioral systems and mechanisms that facilitate juvenile navigation through dam forebays.
Some efforts have been made to model juvenile passage through reservoirs (Nestler et al., 2002)
and northwestern dam forebays (Goodwin et al., 2004), but there are few really relevant empirical
data upon which to base such models.
6.3. Variation among species, stocks, and origins
In the CSRB there are many different fish species and stocks. Just among the salmonids there
are many different runs and life histories as well as hatchery vs. wild stock differences (Quinn,
2005). There may be important differences in passage behavior and success as a function of
species and life stage (age), and perhaps among fish stocks of different origins. Just one species,
chinook salmon (O. tshawytscha), in spring migration range from subyearling fish that are
sometimes less than nine cm long to yearlings that are over 25 cm long. There are probably
differences in any number of fish passage factors, including swimming performance, migration
depth, day and night-time behavior, and susceptibility to screens and surface passage, sensory
capacities and responses, hydrodynamic preferences, and countless others. One of the strengths
of biotelemetry over sonar is that it permits the tracking of known individuals and so species, age,
and origin, etc. are known. One of the strengths of sonar over biotelemetry is that it samples the
run at large, regardless of all of those things. In order to begin to understand the variability
inherent in the fish themselves both are needed.
7. Conclusion
Hydropower dams and other industrial water management sites will continue to change
waterways in the developed world and their effects worldwide will only grow. It is not only
ecologically wise but, in the long run, economically prudent to learn how to design and operate
these structures to do the least harm possible to the ecosystems that we modify. Innovations in
turbine design are promising but it may often be helpful to maximize the proportion of fish
migrants that pass through non-turbine routes. Since the conflict between water for human needs
and water for fish passage and ecosystem management will persist and probably increase it
would be well to use the water that we do invest in passage as strategically as possible so as to
maximize the number of fish passed per water diverted from power generation. It is reasonable to
suspect that at some threshold of spillway discharge or proportion of total project discharge
spilled there may be a point of diminishing returns. If fish are entering the spillway forebay and
then passing without undue delay then there may be little to gain and considerable to lose (lost
power generation and maybe higher turbulence and gas super saturation in the spillway tailrace
and on downstream) with spilling much more. You can’t pass fish that aren’t available for
passage.
As for surface passage, there may be some work to do. Already the sluiceways are proving to
be very good at passing a lot of fish in a little water. Surface passage is appealing in several ways.
Surface passage routes are reasonable analogues of waterfalls and rapids that have been passing
downstream migrants, no doubt with less than 100% survival but acceptably, for a long time.
Understanding what stimuli provide Steve Rainey’s ‘‘opportunity for discovery’’ (Rainey, 1997)
and makes an entrance most readily acceptable to migrants must be the key to increasing the
number of fish passing by surface routes in the same amount of water.
It may be instructive that the success of the fishways developed here in the U.S. Pacific
Northwest (Monk et al., 1989) came from a substantial dedicated effort to understand and design
for adult salmonid upstream attraction and passage. If downstream passage problems are more
difficult, which seems likely due to the small size of the migrants, the size and complexity of dam
forebays, and the dearth of data on motivation and sensory biology of the fish themselves, then it
may be that such a dedicated effort to understand the juvenile salmonid sensory capacities and
behavioral responses is needed. Blumstein and Fernández-Juicic (2004), in an essay on the
emergence of a new discipline they call ‘‘conservation behavior’’, caution against failure ‘‘to
appreciate the importance of proximate mechanisms’’. That’s the scientist’s ‘‘understanding
what’s going on’’ counterpoint to the engineer’s ‘‘fixing what’s wrong’’ paradigm.
Sometimes it is well to look outside one’s own cultural and professional home range for
inspiration. Innovations developed by Temple Grandin, a scientist and engineer at Colorado State
University at Fort Collins (Grandin, 1993; http://www.grandin.com/index.html) have
revolutionized the equipment and procedures for livestock handling and slaughter. That has
been accomplished, to a large extent, by her understanding the suites of sensory stimuli (light,
sound, air movements, etc.) that facilitate stock animals’ acceptance and cooperation in moving
through chutes and ramps, a process that is in some ways similar to fish passage. If we know that
animals balk or hold up at some place in the process (the increasingly meandering fish trajectories
approaching the Bonneville Dam first powerhouse prototype surface collector in Johnson, 2001
come to mind) then determining what the hold up is seems like it should be a high priority. The
trouble is that the fish are more different from us humans than are cattle, they have more and
different (as well as similar) sensory systems, and it is difficult to see what they are doing. The
tools are getting better.
New tools are useful but the ability to integrate information from many levels or organization
to explain and effectively manipulate the world is as important (Bartholomew, 1986). Fish in
dams are part of nature; it’s just a fairly new part and maybe we can improve the passage habitat
characteristics for the fish. Does turbulence attract or repel downstream migrating salmonids?
Right now we do not really know but it probably depends on many different things. The answer
may well be ‘‘Yes, sometimes.’’ to both. Knowing which, and when, could help us help fish to
help themselves past the dams in the safest and most cost-effective way possible.
Successes will mostly be partial and failures may be frequent but it is important that all efforts
be evaluated carefully and fairly and well reported whether success or failure. In the field of
sensory stimulus systems (‘‘behavioral technology’’) it should be recognized that it is the
stimulus that is producing any observed effect, not the device that produces the stimulus.
Acknowledgements
First I would thank Lynne Sneddon for the opportunity to contribute this paper and, especially,
for her relentless good spirit. Lynne and the several reviewers generously helped me to make a
very rough draft acceptable. Peter Johnson, my boss while most of this was written, has been very
tolerant. I would like to take this opportunity to thank John Nestler, of the U.S. Army Corps of
Engineers’ Research and Development Center in Vicksburg, MS, USA for introducing me to the
field of fish passage and protection so long ago. Discussions with some professional friends and
mentors, especially Tom Carlson, Chuck Coutant, Gene Ploskey, and Art Popper, have been both
formative and informative, although any mistakes and shortcomings that appear here are my own.
Chuck Coutant and Glen Čada have been especially helpful in assembling materials. I appreciate
the willingness of the many website operators whose links appear herein to permit their
publication. Special thanks go to Robert Wertheimer and the Portland District of the U.S. Army
Corps of Engineers for permission to use their figures.
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Developing fish passage and protection

Available online 13 November 2006

Keywords: Fish passage; Fish conservation; Hydropower; Dams; Environmental impact

* Tel.: +1 509 427 4840; fax: +1 509 427 4846.

1.3. Anatomy of a hydropower dam . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 300

1.2. Statement of problem

1.3. Anatomy of a hydropower dam

1.3.1. Turbine passage

1.3.2. Juvenile bypass system passage

1.3.3. Spillway passage

1.3.4. Surface passage

2.1. Upstream fish passage by lorries, ladders, and lifts

2.2. Downstream passage options

2.2.2. Juvenile bypass systems

2.2.4. Sluiceways and other surface routes

3. Sources of passage-related delay, stress, injury, and mortality

3.1. The importance of run timing

3.2. Turbine passage

3.3. Juvenile bypass system passage

3.4. Spillway passage

3.5. Surface passage

4. Improving fish passage at hydropower dams

4.1. Changing project structures

4.2. Changing project operations

4.3. Using sensory stimuli to control local fish distributions

4.3.2. Low-frequency sound

4.3.3. High-frequency sound and alosine herrings

4.3.4. Flows and turbulence

4.3.5. Electric fields

4.4. Conclusions on reducing turbine passage

5. Improving fish passage at hydropower dams by improving turbine passage

6.3. Variation among species, stocks, and origins

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