L.M.Whitehouse Science Reviews - Biology, 2024, 3(4), 24-30
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prematurely may survive longer without the pres-
ence of food, which is likely due to the increased
size of their yolk sacs when compared with fish that
hatched later (Laurel et al., 2008).
Physiological and cellular responses to thermal
stress
Elevated temperatures can also disrupt gene
expression and essential biochemical processes in
embryonic development (Podrabsky and Somero,
2004.) Research has shown that embryonic fish can
activate stress responses that could in theory be di-
recting energy away from other important pro-
cesses, resulting in the development of heightened
stress responses and increased chances of develop-
mental deformities that could impact later survival
(Van de pol et al., 2021).
One of the most universal responses to ther-
mal stress is the heat shock response (HSR; Hu et al.,
2022). This cellular stress response involves in-
creased production of heat shock proteins (hsps),
which under normal conditions help maintain cel-
lular proteostasis. Under stressful conditions, in-
cluding thermal stress, these proteins protect cellu-
lar function by repairing proteins that have been de-
graded and preventing the build-up of non-native
proteins. The HSR is present throughout the life his-
tory of fish, from embryogenesis to adulthood and
may play an important role in the ability of fish to
respond to increasing temperatures (Iwama et al.,
1999). Several studies have shown that hsp levels in-
crease in response to increased temperatures in em-
bryonic fish (Sales et al., 2019; Stefanovic et al., 2016;
Takle et al., 2005; Werner et al., 2001, Werner et al.,
2007; Whitehouse et al., 2017), and the amount of
HSPs present plays an important role in protecting
embryos against damage induced by exposure to
increased temperatures (Mirkes et al., 1999).
In addition to stress responses, increasing
temperatures can also result in physiological adjust-
ments in metabolism, respiration, and immune
function (Little et al., 2020). Research has shown
that the thermal conditions experienced during em-
bryogenesis can influence energy metabolism, im-
pacting oxygen consumption and leading to adjust-
ments in metabolic pathways such as glycolysis and
lipid metabolism. Additionally, temperature has
been shown to have long-term impacts on the ther-
mal optima of enzymes involved in these processes.
For example, zebrafish (Danio rerio) embryos raised
at higher temperatures had increased mitochon-
drial and glycolytic enzyme activity at hatch com-
pared to enzymes raised at optimal temperatures.
This difference persisted into adulthood demon-
strating that developmental environments can have
long-term impacts on metabolic mechanisms
(Schnurr et al., 2014). This notion is further sup-
ported by research that revealed that developmen-
tal temperature affects resting oxygen consumption
in larval zebrafish (Barrionuevo and Burggren,1999)
and resting and maximal oxygen consumption in
larval cyprinids (Wieser and Forstner, 1986; Kauf-
mann and Wieser, 1992).
Constant vs fluctuating temperatures
While most studies have focused on the ef-
fects of rearing fish at constant temperatures, there
is a growing body of research that is examining the
impact of heat shock events or diel cycles that ex-
pose developing fish to short bursts of increased
temperatures. Fluctuating or cycling temperatures
more closely mimic the natural environment as fish
developing in the wild experience natural fluctua-
tions because of seasonal variations. These fluctua-
tions are expected to increase because of climate
change. Moreover, anthropogenic activity such as
the release of thermal plumes from nuclear power
plants exposes embryos in their vicinity to frequent
temperature spikes (Reutter and Herdendorf, 1976.).
Research has shown that regular exposure to
temperatures above a fish’s optimal range can aid
in increasing thermotolerance and survival (Bilyk et
al., 2012; Grinder et al., 2020; Morgan et al., 2018).
For example, exposing juvenile Atlantic salmon
(Salmo salar) to one diel cycle increased their Criti-
cal Thermal Maxima (CTMax; Corey et al., 2017), a
measure used to assess thermotolerance in fish
(Becker and Genoway, 1979). Moreover, Colorado
Squawfish (Ptychocheilus Lucius) exposed to fluctu-
ating temperatures during development had a 10 –
30% increased chance of survival post-hatch com-
pared to those reared at control temperatures (Best-
gen and Williams, 2011).
Lim et al (2019) demonstrated that exposure
to temperature spikes had no impact on the devel-
opment of lake whitefish (Coregonus clupeaformis).
Exposing embryos to a + 3°C spike in temperature
for 1 hour, three times a week had little impact on
hatch timing, mortality, or yolk sac absorption.
Moreover, exposure to fluctuations in temperature
during development appeared to have no impact on
developmental processes. In contrast, Artic charr