! 1!A Scientific Review of the Physiology of Pacific Salmo

! 1!
A Scientific Review of the Physiology of Pacific Salmon
Migration
B. C. McKinney1
1 Department of Natural Sciences, University of South Carolina
Beaufort, One University
Boulevard, Bluffton, South Carolina 29909, USA
Abstract For many generations, humans have altered practically
every
ecosystem in the entire world. The footprint humans leave
behind on ecosystems
on Earth has continuously matted the ecosystems and critical
habitat in which all
species on Earth depend on for survival. When considering
Pacific and Atlantic
salmon populations, the array of human caused stressors is
responsible for the
population depletions across the United States and Canada.
This review will
coordinate the impacts of river impoundments (i.e., hydropower
systems) on

upstream and downstream migration as well as visit the impacts
of natural and
human caused change on the quality of habitat in which
salmonids inhabit through
all life stages.
Introduction
A variety of teleost species are classified within the Family
Salmonidae under the Order
Salmoniformes. Salmonidae is comprised of a variety of trouts
(Salmo spp.), chars (Salvelinus
spp.), graylings (Thymallus spp.), taimen (Parahucho spp.), and
salmons (Salmo &
Oncorhynchus spp.). The anatomy of this family is similar to
other ray-finned fish having
dorsal, pelvic, pectoral, anal, and dorsal fins, however they
possess an additional fin posterior to
the dorsal called the adipose fin.
Salmonid lifecycles are very complex and have been a topic of
research for many
generations (Briggs, 1953; Holmes & Stainer 1966; Vronskiy,
1972; Thompson & Sargent, 1977;
Healy, 1980; McCormick &Saunders, 1987; Murray & Rosenau,
1989; Nehlson et al., 1991). In

recent findings, the introduction of telemetry techniques and
field sampling routines have given
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researchers insight about the duration, timing, and patterns of
homing and staying (Healy, 1980;
Giorgi et al., 1997; Walker et al., 2016). Through the
protection of the Endangered Species Act
(ESA) select Pacific salmon populations have been granted
protection by federal regulations in
relation to the habitat that is essential to their survival
(USNMFS 1995). In this review, relevant
available published literature will be compiled to discuss a
variety of explanations towards the
physiology and morphological complexities associated with
Pacific salmon.!
Overview of Salmon Biology
In this section, emphasis will focus on the evolutionary history
of Salmon (see Groot &
Margolis, 1991, Hendry et al., 2000, and Waples et al., 2007 for
more details). North America’s
populations of Pacific Salmon consist of five distinct species:
chinook salmon (Onchorhynchus

tshawytscha), pink salmon (O. gorbusha), chum salmon (O.
keta), coho salmon (O. kisutch), and
sockeye salmon (O. nerka). Pacific salmon are uniquely
characterized as anadromous
(migratory) and semelaparous (i.e., die after spawning) species.
This life strategy is opposite of
their cousin, steelhead are the anadromous form of rainbow
trout (O. mykiss), which are
iteroparus (i.e., spawn more than once).
Spawning adults (spawn and die within 2 weeks) must migrate
far enough upriver to an
area there has good water quality and low predation rates,
seeking this habitat is essential to the
survival of their offspring as eggs (i.e., 0-3 months), alevin
(i.e., feed off yolk sac for 1-2 weeks),
fry (i.e., 5-10 weeks), and parr (i.e., 1-2 months old). Yearling
smolt will reside in the brackish
estuary for weeks, months, or even years before moving
offshore depending on the species and
their ability to osmoregulate to increasing salinities. Their
voyage into the ocean occurs next,
migrating to more productive water for growth will last for
multiple years (i.e., 1-3 depending on

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the species) until sexual maturity (i.e., 3-8 years old) when the
homing phase will be initiated.
Shoji et al. (2003) and Hara (1992) investigated an essential
mechanism for successful homing to
the natal grounds; explaining the dependency on olfactory
organs to electro-physically respond
to dissolved chemicals (i.e., amino acids, steroids, bile acids,
and prostaglandins) queues in
rivers and streams which they were born. Salmon face a var iety
of physical and morphological
obstacles along the way, for example, sockeye salmon morph
from their silver (i.e., offshore)
coloration into a vibrant reddish/pink color and the formation of
a hooked snout (i.e., kype)
(Groot and Margolis, 1991; McCormick & Saunders, 1987).
Effects of Hydropower Systems
The implication of hydropower systems in the United States and
Canada has significantly
affected the passage of spawning salmon. Due to their vast
size, salmon are often times limited
in terms of upstream distance traveled (Anderson & Salinger,

2006). Brett (1995) further
explains that because salmonids do not feed during upstream
migration, the flow velocity of the
river is essential. Brett (1995); Anderson & Salinger (2006);
and Tillotson & Quinn (2017) all
contribute reasons why hydropower systems are responsible for
pre-natal spawning and pre-
spawn death.
Water Quality:
Ever since the introduction of hydropower systems, (Waples et
al., 2007; Regetz, 2003)
strongly suggest that the redirection of water for agricultural
use is the major reason for pour
water quality (i.e., lower dissolved oxygen, higher water
temperatures, salinity, sedimentation,
and contaminants from pesticides and fertilizers) in the lakes
and streams of the Pacific
Northwest.
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Physical Conditions:
Brett (1995) explains that the ocean phase before upstream

migration is essential to
migration because this is the period in which the salmon
consume their last bit of energy.
Semaloparous salmon begin to deteriorate and transform during
this migration, their mouth
structure changes significantly, which limits their ability to
consume food during migration.
Evolutionary Significance
Egg Phase:
During a spawning event, gravid females will create a
spawning bed (i.e., redd); this
structure is often variable in size depending on the surface area
and flow velocity of the river
(Vronskiy, 1972). In Healy’s (1980) review of Utilization of
the Naniamo River Estuary By
Figure 1. Sockeye Salmon (O. nerka) Spawning Phase can be
observed by the change in coloration from silver (ocean phase)
to reddish/pink during spawning. Males (top) show significant

morphological changes: the hump in front of the dorsal fin and
kype of the jaw structure. (Photo credit: Washington
Department of Fish & Wildlife)
! 5!
Juvenile Chinook Salmon, (Briggs, 1953) observed that chinook
salmon tended to deposit eggs in
gravel substrate at a mean depth of 20-36 cm in a small steam
located in California. In contrast,
Vronskiy (1972) observed deposition in the same substrate
characteristics but at a more variable
depth (i.e., 10-80 cm).
The incubation (1-3 months) of eggs in the spawning bed is
variable among Pacific
salmonids; however, a factor that affects development into
alevin and fry is stream quality.
Severe flooding in rivers during the time of incubation is noted
as a major cause of mortality; in
addition, the flow of water (percolation) through the spawning
bed is another factor that
promotes mortality of eggs and larvae (Gangmark & Bakkala,
1960)
Juvenile Phase:

The emergence of fry from the spawning beds occurs mostly at
night; from here they will
be dispersed downstream into the estuary or may occupy a slack
pool (less current) or stream for
a short period of time (Reimers, 1971). The progeny of fry that
emerge from spawning beds are
faced with a variety of challenges which determine their
survival, in today’s era, scientists are
developing a better understanding about survival rates,
abundance, and growth rates of juvenile
salmon through telemetry research as well as intensive sampling
of river and estuary habitats
(see Downstream Migration).
Downstream Migration:
Healy (1980) established a sampling routine in the Nanaimo
River, Canada to
establishing an understanding about duration of residency of
juvenile chinook salmon in
freshwater before migrating to the estuary and the duration of
residency in the estuary before
migrating offshore. Healy’s sampling protocol focused on
sampling fry in freshwater stream

! 6!
channels (i.e., incline plane fry traps), estuarine intertidal mud
flats (i.e., beach seine) and marine
rivers/creeks (i.e., beach and purse seine). Healy (1980)
recorded fork length (FL), weight of
total catch, and weight of stomach content from a subset (i.e.,
10-15 individuals) of the total
collection. In conclusion, his findings from sampling fresh and
saline habitats revealed that
juveniles reside in the freshwater and the estuary of Nanaimo
River for a year before migrating
offshore, respectively (Healy, 1980).
Giorgi et al. (1997) estimated the downstream migration rate of
juvenile chinook salmon
throughout 259-km of river between the Rock Island Dam (RI)
and McNary Dam (McN) of the
Columbia River. In 1997, Giorgi implanted 14,723 chinook
smolts (47-171 mm) with a Passive
intergraded transponder (PIT) tag which allowed him to collect
telemetry data about the timing
of downstream migration by an electronic scanner attached to
various smolt by-passes.
.

Years later in 2016, Walker et al. investigated the size thr eshold
which juvenile chinook
salmon were capable of bearing a passive integrated transponder
tag (PIT). This study focused
on minimalizing the size of the tag so that the implantation of
this telemetry transmitter would
Figure 2. Passive Internal Transponder
(PIT) illustrating the size comparison of
old technology (A-B) vs. new technology
(C-D) (Walker et al., 2016).!
! 7!
have no negative effect on juveniles during development and
migration. Walker et al. (2016)
conducted swimming performance and predator avoidance
simulations to gain insight about the
effects of adding additional weight (i.e., 0.22g in air; 0.11g in
water) to the fish during
development. Through these experiments, he was able to
conclude that implantation of internal
tags for telemetry research in fact has no negative impacts on

the fish during his study.
Reluctantly, Walker et al. (2016) concluded these results
because the use of telemetry had
previously been used for centuries before his experiments (e.g.
Giorgi et al., 1997).
Early Ocean Phase:
McCormick & Saunders (1987) describes the transformation of
stream-dwelling parr to
the seaward-migrating smolt (i.e., smoltification) as a
significant stage of life, which is essential
for the survivorship of various salmon species. Morphological
changes of parr transformation is
described (Folmar & Dickhoff, 1980) to have dark bars that are
perpendicular to the lateral line
and as the parr grow into smolts, these bars tend to disappear
through the accumulation of purine
amino acids (i.e., guanine and hypoxanthine), which change the
coloration into a reflective silver
appearance (Folmar & Dickhoff, 1980). Physiological changes
are also important to migrating
smolts; changes in ionic composition by the gills and the
excretory and endocrine systems. (see
Osmoregulation)

Adult migration phase:
Quinn et al. (2001) believed that populations of migrant and
spawning salmon were
repeatedly differed in that the time of spawning is a product of
adaptation and spatial isolation
of populations to facilitate and accelerate divergence in traits.
The timing of spawning migrations
! 8!
is temporally and spatially different among salmonids, however,
there is a common link in
phenotypic size and growth of the expected progeny (Quinn et
al., 2001).
Shoji et al. (2003) investigated the ability of salmon to
recognize stream-specific
chemical composition (i.e., amino acids, steroids, bile acids,
and prostaglandins) during the
homing phase. During this study Shoji used mature chum
salmon from the Osaru River, Japan
where he then simulated an environment that had two inputs of
water: on one side of the test tank
had inflow of artificial stream water which was a mixture of
Osaru River water and artificial

freshwater (0.5 mM NaCl, 0.05 mM KCl, 0.4 mM CaCl2 and 0.2
mM MaHCO3, pH 6.9) and on
the opposite side was natural lake water. Both water types were
added to the test tank over a
nine-hour period, fish movement was monitored by a remote
camera system (T-water 2000C,
Tukamoto Musen Co., Mie) and then the selection to each arm
was recorded. During his
experiment, of the forty-four total chum salmon, twenty-eight
(64%) showed upstream
movement (i.e., selecting the left of right arm), 24% of fish
were recorded selecting the arm
which had an inflow of artificial water and 15% of fish chose
the natural lake water. From these
results, it is plausible to implicate the olfactory capabilities of
salmon to smell the chemical
composition of water for successful migration to their natal
grounds (Shoji et al., 2003).!
Physiological Adaptations
Osmoregulation:
Spawned salmonids reside in freshwater for a short period of
time (i.e., 1-2 weeks) before
they migrate into the estuary where the water becomes more
saline (Healy, 1980; McCormick &

Saunders, 1987). The gill structure in teleost fish is the primary
organ that physiologically
! 9!
transforms during migration; gill Na+, K+-ATPase activity
increases as the salt concentration
increases from fresh into salt water systems (Epstein et al.,
1967).
Loretz et al. (1982) concluded that during the downstream
migration of coho smolts,
chloride cell abundance and size increase proportionally to the
size of the fish. In addition,
during this osmoregulation process, the internal medium of
ionic salt concentration must be
regulated, (Holmes & Stainer, 1966) found that the excretory
system is responsible for this
change. Kidney Na+, K+-ATPase activity of the Atlantic
salmon is highly variable in the
estuary in regards to seasonal fluctuations of freshwater input
from the streams during the rainy
season and visa versa during the dry seasons (Loretz et al.,
1982).
In conclusion, salmonids have a “strategy” for osmoregulation,

which entails a
continuous migration event from fresh to saltwater (i.e.,
juvenile migration) and salt to
freshwater (i.e., adult migration) (Thompson & Sargent, 1977),
whereas euryhaline species (e.g.,
fundulus spp.) have a “strategy” for osmoregulation, which
entails multiple migration events
from fresh to saltwater or salt to freshwater.
Growth:
The Nanaimo estuary is an important habitat for the growth of
chinook salmon, (Healy,
1980) found that the length of juvenile in the estuary was
slightly greater than those found in the
stream surveys. Healy (1980) concluded that the average length
of juvenile chinook in the
estuary was 70 mm fork length at the end of the downstream fry
run. During 1975-1977, Healy
(1980) concluded that there were differences in size class in the
different years however there
was not a significant difference, in conclusion the average size
threshold, which leaves the
estuary in migration to the ocean, was 70 mm fork length,
respectively.

! 10!
Metabolism:
Oxygen consumption changes in proportion to the rate of
growth in juveniles in the
smoltification phase, here, coho salmon tend to decrease blood
glucose levels were as Atlantic
salmon increase blood glucose levels during development
(McCormick & Saunders, 1987).
Creatine composition in juveniles during the smoltification
phase is significantly correlated with
downstream migration to the estuary and also during seaward
migration; Cowey & Parry (1963)
concluded that this 30% increase is due to the increased muscle
mass needed the future
migration.
Competition and Predation:
Salmon are an important step in the food web complex for a
variety of reasons. Predation
on adult salmon by brown bears (Ursus arctos) in Karluk Lake,
Alaska was significant in 1964
(Gard, 1971). The timing of upstream migration by sockeye
salmon in relation to predation by

bears was experimented by visual surveys; this evaluation
concluded that the bears killed up to
79 % of the migrant sockeye population in 1964 (Gard, 1971).
This predation is a natural
behavior that has been occurring for many generations,
conversely, Gard (1971) also estimated
bear predation was only a part of the loss of eggs during a
spawn, the estimates made in this
experiment explained that bears killed 1000 adults which is not
many considering the total death
of adults was estimated to be 8000, respectively.
Tillotson & Quinn (2017) evaluated the conspecific density
trigger for pre-spawning
mortality of sockeye salmon in the Fraser River, Canada was a
combination of predation by
terrestrial animals, anthropogenic impacts, fishing pressure, and
competition for habitable
freshwater.
! 11!
Collateral Damage / Sustainability:
Since the late 1970s, the economic value of wild salmon has

been decreasing due to the
introductioun of fish farms in Canada (Noakes et al., 2000;
Carr!&!Whoriskey, 2006) and years
later in the U.S. Noakes (2000) explained the escape of farmed
salmon by human error and
natural events is a factor that decreased wild salmon stocks in
British Columbia. Hybrid species
of salmon have impacted the migration and developmental life
stages of wild coho and chinook
salmon stocks, however, these are only a small piece of the
puzzle which further is completed
with the impacts of climate change, overfishing, and the loss of
critical freshwater habitat
(Noakes et al., 2000).!
Climate Change:
Climate fluctuation has been a relevant topic of research for
many generations, Perry et
al., (2005) investigated Climate Change and Distribution Shifts
in Marine Fishes in the North
Sea, his findings promote that water temperatures are increasing
with increased latitude. Perry et
al. (2005) concluded that shifts in distribution would eventually
result in a loss of species

through habitat loss. His concept can be applied to many other
water bodies around the world.
Beamish et al. (1997; 1999) used the Aleutian Pressure Index
(ALPI) to estimate the
regime concept and natural trends in the production of Pacific
salmon. Beamish et al. (1997)
investigated the effects of the seven el Niño events from 1960
to 1993, concluding that the smolt
production during three events (1965-1966, 1969-1970, and
1982-1983) on average was
significantly higher than in two other events (1972-1973 and
1992-1993).
! 12!
Conclusions
Pacific salmon and Atlantic salmon have been studied for many
generations through
physiological, morphological, and behavioral experiments.
These findings have discovered
many different critical life stages, which conservation efforts
need to be focused. The spatial and
temporal distribution of anadromous and semelparous species

occurs in a variety of rivers,
estuaries, and oceans around the world. The development of
hydropower systems is a critical
bottleneck, which has depleted many populations of salmon,
thus, pacific salmon specifically
have been continuously degreasing for many generations. The
Endangered Species Act (ESA)
enforces conservation of distinct populations of pacific salmon
in iconic rivers and streams of the
Pacific Northwest (USNMFS, 1995). Through the distinction of
specific salmon populations by
the ESA, restrictions and regulations for recreation have been
assessed and enforced by many
state agencies. Climate change is also depleting many
populations of salmon, the warming of the
atmosphere, is melting snow and ice on mountains which flows
down the mountain into the
rivers creating increased flow and greater surface area
(flooding). The increased flow is in part a
reason for lack of survival of avelin and fry during the
emergence of spawning beds; however,
the decreased flow of rivers from the introduction of dams has
also created situations that affect
the survival of juveniles migrating downstream. The effects of

global warming in relation to
increased water temperatures around the world has also had
significant negative effects on
spawning and migration.
Salmon have incredible evolutionary significance for osmo-
conformity and regulation.
The resilience and evolution of salmon is fascinating, their life
cycle is complex in that there are
many different critical aspects of development and survival.
Thus, it has been challenging for
scientists to fully understand their complexity. Finally,
protection efforts help with conservation,
! 13!
however, even with all the protection, salmon populations will
continue to decrease to a point of
no return if humans don’t step up and promote awareness of
pollution, deforestation, and the
introduction of water control systems in respect to salmon and
more importantly all forms of life
that humans cherish.

! 14!
Literature Cited
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96

Final review paper--The paper should discuss a topic related to
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5) Literature cited (20 pts)
Review Paper Rubric Break Down:
Along with the following rubric,Writing should be clear,
articulate and typo-free to be considered a perfect paper.
Title Page (5 points)
· A good title should concisely describe the topic area of the
review. You may be "scientific" in your title accuracy (using
physiology-related jargon) or you may be creative. Many

review titles are written creatively, or in layman's terms, in
order to compel readers outside of the field to read the review. I
encourage you to look up reviews in physiology to compare and
contrast title types.
· Also include your name, class, and lab section and the date.
Introduction (20 points)
· What historical research inspired this work? Find some papers
that inspired the research of the focal paper.
· Why is this interesting?
· What is the goal of this review?
· Make sure you cite all introductory articles correctly (see
"Literature Cited" section, below).
Body (40 points)
· What is the current state of the topic area?
· What new research has been done that refines or refutes the
discoveries of your topic?
· Papers that you cite throughout may refine or refute
highlights of your review article.
· Make sure you cite all journal articles correctly (see
"Literature Cited" section, below).
· You may include up to 3 figures from other literature sources
(NO WEBSITES OR REVIEWS) but please make sure that they
are properly cited (under the figure and in the "Literature Cited"
section.
· You may receive extra points for making your own figure
synthesis! Many review articles do this (most with the help of a
graphic artist).
· Your review should be well organized with major points
flowing in an order that makes sense.
Conclusions(15 points)
· The conclusion should articulate your thoughts on the topic
areas in a clear manner.
· The conclusions section should also describe future directions
for this research topic. Based on your review, provide

suggestions for what research might be done next to further
develop the progression of ideas you have summarized.
· You may also need to cite papers in this section so make sure
the format is correct, throughout.
Literature Cited (20 points)
Grading criteria for citations:
· Cite at the very least 8 research articles. NO WEBSITES OR
REVIEWS. Books may be included AFTER the 8 research
article quota are met but make sure you cite them correctly. (5
points)
· Use APA format. The following example uses the APA format
for the journal citation:
Goldschneider, F. K., Waite, L. J., & Witsberger, C. (1986).
Nonfamily living and the erosion of traditional family
orientations among young adults. American Sociological
Review, 51(4), 541-554.
· Put literature cited in alphabetical order and be consistent
down to the punctuation used in each. Go through,
painstakingly, to make sure that all elements of each review
paper are cited correctly and consistently. (5 points)
· Make sure that the cited paper accurately describes the section
of the paper where it was cited. I'll be spot-checking. (5 points).
· Make sure that you use acceptable citation format throughout
the manuscript body (5 points). See the following website for
reference:
http://blog.apastyle.org/apastyle/2011/01/writing-in-text-
citations-in-apa-
style.html?_ga=2.201041902.1527685899.1520435603-
2053305827.1520435603
· See this website for further details:
http://www.bibme.org/citation-guide/apa/

**While I encourage you to look over actual reviews on your
topic, please do not use review articles as citation sources in
your paper or copy a published review papers format. Reviews
are a synthesis of the current state of the researched primary
literature that was written by a potentially biased or incomplete
perspective, so if you cite a review, you are just taking the
reviewers word for a particular finding and many reviewers
misinterpret the data. This task is to make you draw your own
conclusions on the state of the research, even if you are wrong.
*Tips on avoiding plagiarism:
Paraphrase- Read a paper or a section of the paper that has the
concept that you want to communicate. Look away from the
paper and then write it in your own words. Refer back to the
original paper and if your writing is too similar, change it. If
you use more than two words in a row from the original paper,
it must be put into quotations. Use a thesaurus and use your own
writing style. This is a great exercise and should always be
done when writing. Never copy and past from your own
writings. This is self plagiarism and it is not acceptable. Once
you’ve written it, it's an original piece of work and should not
be copied. As a rule, the more you force yourself to re-write
(even the same exact methods), the better the description gets
each time and the better you get a writing.
Cite - After paraphrasing, the work must be cited at the end of
the sentence or section referring to the original article. This is
one of the most effective ways to avoid plagiarism. Use the
APA format for citations within the text and in your reference
section. Citing is really that simple. Not citing properly can
constitute plagiarism.
As a note - strive to cite original papers. Your Literature Cited
section should contain predominantly original research articles,
from historic articles to the most recent articles in the topic. A
scholar should interpret original data for himself. However,
citing a review article may be necessary in the case of well
established topics and concepts.

Quote - If a particular concept or hypothesis needs to be
included verbatim, use quotes followed by a proper reference.
When quoting a source use the quote exactly as it appears.
Block quotes (of 40 words or more) are unacceptable under
most circumstances. A scholar should be able to effectively
paraphrase most material. This process takes time but the effort
pays off.
For other resources on the topic of plagiarism see:
http://isites.harvard.edu/icb/icb.do?keyword=k70847&pageid=ic
b.page342057

! 1!A Scientific Review of the Physiology of Pacific Salmo

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