Effects of dissolved oxygen concentration on freshwater fish: A review

Abstract

Dissolved Oxygen (DO) is one of the most important factors for aquatic animals as especially for those who derive dissolved oxygen from the water. DO levels indicate the quality of water. Many biotic and abiotic factors may influence DO concentration like mixing of different water bodies, upwelling, atmospheric exchange, respiration, photosynthesis, ice cover, pollution and some physical factors like salinity and temperature. The fluctuations in DO levels in water affect fish physiology. In this review, we will focus on the impact of DO on freshwater fish-physiology. A detailed literature survey is given based on DO and freshwater fish swimming, feeding, disease management, survival, respiration, metabolism, growth, reproduction, health parameters, immunity and stress of freshwater fishes.

Full text

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International Journal of Fisheries and Aquatic Studies 2022; 10(4): 113-127
E-ISSN: 2347-5129
P-ISSN: 2394-0506
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IJFAS 2022; 10(4): 113-127
© 2022 IJFAS
www.fisheriesjournal.com
Received: 21-03-2022
Accepted: 11-05-2022
Bulbul Ali
Department of Zoology,
Babasaheb Bhimrao Ambedkar
University, Lucknow, Uttar
Pradesh, India
Anuska
Department of Zoology,
Babasaheb Bhimrao Ambedkar
University, Lucknow, Uttar
Pradesh, India
Abha Mishra
Department of Zoology,
Babasaheb Bhimrao Ambedkar
University, Lucknow, Uttar
Pradesh, India
Corresponding Author:
Bulbul Ali
Department of Zoology,
Babasaheb Bhimrao Ambedkar
University, Lucknow, Uttar
Pradesh, India
Effects of dissolved oxygen concentration on freshwater
fish: A review
Bulbul Ali, Anuska and Abha Mishra
DOI: https://doi.org/10.22271/fish.2022.v10.i4b.2693
Abstract
Dissolved Oxygen (DO) is one of the most important factors for aquatic animals as especially for those
who derive dissolved oxygen from the water. DO levels indicate the quality of water. Many biotic and
abiotic factors may influence DO concentration like mixing of different water bodies, upwelling,
atmospheric exchange, respiration, photosynthesis, ice cover, pollution and some physical factors like
salinity and temperature. The fluctuations in DO levels in water affect fish physiology. In this review, we
will focus on the impact of DO on freshwater fish-physiology. A detailed literature survey is given based
on DO and freshwater fish swimming, feeding, disease management, survival, respiration, metabolism,
growth, reproduction, health parameters, immunity and stress of freshwater fishes.
Keywords: Dissolved oxygen, hypoxia, hyperoxia, fresh water fish, fish physiology
1. Introduction
The term "dissolved oxygen (DO)" refers to gaseous oxygen that has been dissolved in water
and is available for the aquatic organism for their oxygen dependency. Fish and other aerobic
aquatic creatures require oxygen to survive and reproduce (Caldwell and Hinshaw, 1994) [1].
Oxygen dissolves in water by the process of diffusion from the atmosphere, that has tumbled
over falls and rapids movement, or by photosynthesis through aquatic plants (Singh and
Kumar, 2014) [2] oxygen is the most vital factor in aquaculture for their maintenance of
metabolism and growth (Doudoroff and Shumway, 1970; Kutty, 1981; Davis, 1975) [3, 4, 5]. In
developing countries aquaculture plays an important role to feed the undernourished people. In
India, 1.2 million tonnes of freshwater fish were consumed annually (Singh and Kumar, 2014)
[2]. The requirement of DO differs from species to species in fish. Generally, the DO
concentration is measured in mg/L or percent saturation (Wilson, 2010) [6]. 8-8.5 mg/L of DO
supports healthy growth rates (Hicks, 2002) [7] lower than 8 mg/L concentration will affect the
mature eggs and the larval development (Davis, 1975; Bjornn and Reiser, 1991) [5, 8]. Thus,
dependent on habitat, fishes frequently experience fluctuating O2 availability, which can range
from hypoxia (low O2 availability) to hyperoxia (O2 supersaturation) (Diaz & Breitburg, 2009)
[9].
When the DO concentration gets below 5-6 mg/L in fresh water, then the required level of an
aquatic organism it gets in hypoxic condition (Dong et al., 2011) [10]. The Anoxia and hypoxia
are known to be a primary cause of stress, poor appetite, slow growth, illness susceptibility,
and mortality (Timmons et al., 2001) [11] can create a large reduction in abundance, diversity
and harvest of fishes within affected water (Breitburg, 2002) [12]. Mild hypoxia only resulted in
a decrease in blood oxygen saturation and pO2. At all-time points, both acute and chronic
moderate and intense hypoxia resulted in a drop in blood pH, pO2, total oxygen content,
plasma Na+ and Cl (Aboagye and Allen, 2018) [13]. Certain species are far more hypoxia
tolerant than others, resulting in differences in long-term survival (Poon et al., 2002) [14]. In
general fishes avoid the areas where oxygen concentration is below the specific level
(Agostinho et al., 2021) [15]. Hypoxia enhanced haemoglobin oxygen affinity, the lowering in
temperature will also improve oxygen uptake (Petersen and Steffensen, 2003) [16].
In aquaculture practice DO above the air saturation is considered as hyperoxia. When fishes
are subjected to hyperoxia (<200% saturation), no detrimental consequences or anomalous
behaviour are observed (Dejours et al., 1977) [17] albeit there are some alterations in the acid
balance of the fish blood (Ruyet et al., 2002; Edsall and Smith, 1990; Wilkes et al., 1981) [18,
19-20].

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At high ammonia levels (Foss et al., 2003) [21] and stocking
density (Wilkes et al., 1981) [18] there are no negative
consequences. Because of enhanced rate of photosynthesis,
hyperoxia occurred at about noon or early afternoon (Richards
et al., 2009) [22].
In some species such as trout and fast-swimming fish, the
bladder acts as a storehouse of oxygen for use in the absence
of oxygen. when the water is oversaturated (hyperoxia), The
bladder becomes overstretched causing buoyancy issues,
especially in small fish (Groot et al.,1995) [23]. Also, cause
chloremia and respiratory acidosis (Heisler, 1993) [24],
oxidative damages (Brauner et al., 2000) [25]. Hyperoxia can
arise in shallow-water fish due to the photosynthetic activity
of phytoplankton, algae, seaweeds, and macrophytes, or in
aquaculture due to O2 supplementation (McArley et al., 2021)
[26]. When compared to normoxia, hyperoxia had similar but
less effective influences, with weight gain and decreasing in
growth rate and increasing in feed conversion ratio, but all of
these differences were less than hypoxia (Aksakal and Ekinci,
2021) [27]. The long-term implications of hyperoxia are still
unknown (Polymeropoulos et al, 2019) [28].
Fig 1: Range of tolerance for dissolved oxygen in fish
2. Factors which affect the concentration of dissolved
oxygen in water
2.1 Biotic factors
A sufficient amount of DO is required for optimal water
quality to support aerobic living forms (Devis, 1975) [5]. But
abiotic certain factors like diffusion and aeration,
photosynthesis, respiration, and decomposition have an
impact on DO concentration (Hargreaves, et al., 2003) [29].
Photosynthesis by aquatic plants and algae provides dissolved
oxygen to water bodies during the day.
6CO2 + 12H2O +energy C6H12O6 + 6O2 + 6H2O
The net effect on DO concentrations is usually very low
because these same organisms consume comparable amounts
of DO through respiration during the night when
photosynthesis is not active (Wilson, 2010) [6]. Aerobic
respiration obtains energy from energy-rich carbon molecules
by oxidising carbon in to CO2 and reducing oxygen to H2O to
sustain life. As a result, due to respiration needs, dissolved
oxygen concentrations will be highest in the mid to late
afternoon, when photosynthesis rates are highest, and lowest
immediately before the sun rises the next morning when
photosynthesis rate at lowest. This pattern of fluctuation is
known as the "diurnal oxygen cycle" (Talling, 1957) [30]. In
addition to photosynthetic organisms' demands during
darkness, other organisms such as aquatic vertebrates and
invertebrates, as well as bacterial and fungal communities
involved in decomposing dead plants and animals, use oxygen
within the system through aerobic respiration (Boyd &
Hanson, 2010) [31]. Biological oxygen demand (BOD) is a
measure of the potential for DO inside a water body to
become low and might became anaerobic due to microbial
biodegradation of organic molecules (Ultsch et al., 1978) [32].
When management operations may enhance the available
carbon within a system, such as aquatic weed management
with aquatic herbicides, BOD considerations are especially
essential. As the plants are decomposed by microbial
organisms, they will become a source of BOD in the system.
When water body gets enriched with nutrients caused by the
nutrients runoff from nearby lands and drainage systems leads
increased in algal production, frequently known as algae
blooms. As microorganisms decompose these algae blooms,
BOD levels will rise dramatically (Francis-Floyd, 2006) [33].
DO level is frequently used to indicate freshwater quality,
stream and river health, and the severity of aquatic pollution
(Kannel et al., 2007) [34]. A stream with a saturation rating of
more than 80% oxygen is regarded to be healthy. Warmwater
fish should have a minimum DO concentration of 5 mg/L,
while cold water fish should have a minimum DO
concentration of 6 mg/L (Doudoroff and Shumway, 1970) [3].
Fish growth, reproduction, physiology, biochemistry, and
behaviour can all be affected by low DO levels (Davis, 1975)
[5]. Oxygen depletion can occur as a result of sewage plant
discharges, abattoir wastes, sawdust, feedlot manure, and food
processing plant and paper mill use. These contaminants serve
as a food source for bacteria, which use dissolved oxygen to
break down these organic molecules, lowering the DO level in
the environment. Other factors that contribute to oxygen
depletion include the release of anoxic (lack of oxygen)
bottom water from dams, the turnover of oxygen-deficient
hypolimnetic water, an abundance of aquatic vegetation, and
huge algal blooms. Fish swimming at or near the surface
gulping air are signs of possible oxygen depletion, as are
sudden changes in watercolour to brown, black, or grey, a
rotting odour from the water, extended periods of hot, gloomy
weather, algae die-offs, and thunderstorms. Poor DO
concentrations can lead to the extinction of more vulnerable
species in an environment, resulting in a loss of species
diversity (Kibria, 2004) [35].
Overpopulation of bacteria and over fertilization of water
plants could all contribute to decrease in DO levels in a body

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of water (Bansal and Jaspal, 2009) [36]. If DO level in a water
body falls down below than 4-5.0 mg/L then the aquatic life
gets under stress (Dey, 2017) [37]. Even the most tenacious
fish would perish if DO drops below 3 mg/L. the requirement
of DO for eggs and the immature stages is more than normal
DO concentration. However, the amount of DO require by an
aquatic organism is determined by its species, physical state,
water temperature, contaminants present, and other factors
(Mallya, 2007) [38]. Fish are cold-blooded in nature because of
that their metabolic rates increases at higher temperatures;
hence they consume more oxygen at higher temperature. High
DO concentrations can pose issues just as low DO
concentrations can. Gas bubble disease in fish and
invertebrates can be caused by supersaturated water, however,
this is a very unusual event. When DO maintains above 115%
- 120% air saturation for an extended period of time,
significant death rate occurs (Geist et al., 2013) [39]. At 120
per cent DO saturation, total mortality occurs in young
salmon and trout in less than three days. Gas bubble illness
affects invertebrates as well. While invertebrates are
susceptible to gas bubble sickness, they can typically
withstand higher amounts of supersaturation than fish. The
bubbles, also known as emboli, obstruct the passage of blood
via the blood arteries, resulting in death. External bubbles
(emphysema) can form and be visible on fins, skin, and other
tissue (Weitkamp and Katz, 1980) [40]. Eutrophication and
organic pollution are one of the main factors of water,
especially in the heavily populated region (Pollock et al.,
2007) [41].
2.2 Abiotic factors
The total amount of oxygen that can be broken up in the water
is depend upon several factors, including water temperature
(Rajwa et al., 2014) [42] salinity (Moon et al., 2003) [43] and
atmospheric pressure. Higher water temperature cause
increase in molecular vibrations, which leads decreasing in
the intermolecular spaces between the water molecules.
That’s why the amount of DO is higher in cold water and low
in warm water (Khani et al., 2017) [44]. Altitude makes also
difference in DO concentration in water (Paz et al., 2020) [45].
Since the density of atmospheric O2 for dissolution at higher
altitude is low than at sea level (Zaker, 2007) [46]. Oxygen
transmission across the air-water interface is facilitated by
increased surface area exposed to the atmosphere. The surface
area of a water body in contact with the atmosphere is
increased through wind-driven waves and ripples, as well as
forcing water into droplets by splashing over obstacles or
forcing via a fountain (Connell and Miller, 1984) [47]. The
surface area to volume ratio is crucial for determining a water
body's baseline oxygen status, since aeration is the most
prevalent means of adding oxygen to an aquatic system
(Singh and Kumar, 2014) [2]. Deepwater bodies with a smaller
surface area will have less possibility for O2 dissolution into
the water than shallow water bodies with a larger surface area
exposed to the atmosphere (Araoye, 2009) [48].
Fig 2: Biotic and abiotic factors affecting dissolved oxygen in water
3. Physiological Changes
3.1 Swimming
It is generally known that fish's aerobic swimming
performance is limited by aquatic hypoxia. Hypoxia affects
spontaneous swimming activity, with sluggish species
slowing down and active species speeding up. Fish can escape
hypoxia by actively searching for well-oxygenated areas.
Several investigations using swim tunnels and an incremental
technique “critical swimming speed" (Ucrit) have shown that
fish in hypoxia has a lower Ucrit than fish in normoxia. Most
of fresh water fish drastically diminished swimming skills
when exposed to hypoxia, owing to the high metabolic cost of
aerobically propelled swimming as well as the physiological
difficulty of hypoxia (Jones, 1971; Herbert and Steffensen,
2005; Smit, 1965; Fu et al., 2011; Hanke and Smith, 2011;
Fitzgibbon et al., 2007) [49, 50, 51, 52, 53, 54]. Chronic exposure to

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hypoxia reduces some of the hypoxia's swimming-limiting
effects. Therefore, chronic hypoxia acclimation provides
long-term benefits (Ackerly et al., 2018) [55]. Hypoxia
tolerance and swimming ability in fish subjected to chronic
hypoxia are improved by transient gill remodelling, increased
haematocrit, the haemoglobin with stronger O2 binding
affinity, increased anaerobic capacity, and increased cardiac
output (Chippari-Gomes et al., 2005; Petersen and Gamperl,
2010; Fu et al., 2011) [56, 57, 52]. Larger gill surface area in
swamp-dwelling fish, could represent a compensatory
technique for the physiological constraints of hypoxia
(Chapman and Hulen, 2001) [58]. An individual's anaerobic
metabolism can have a considerable impact on performance,
particularly at speeds near Ucrit, where aerobic contribution to
power performance is often higher (Wilson and Egginton,
1994; Svendsen et al., 2010) [59, 60]. The non-invasive measure
of excess post-exercise oxygen consumption (EPOC) can be
used to quantify anaerobic activity. EPOC measures an
individual's rate of oxygen consumption during the time
frame during which excess waste (e.g., lactate) originating
from anaerobic metabolism is eliminated from the body
(Peake and Farrell, 2004; Svendsen et al., 2010) [61, 60].
Recovery from anaerobic activity requires more oxygen, so
recovery from Ucrit can take much longer in hypoxic
conditions (Svendsen et al., 2011) [62].
Hypoxia also affects predator-prey interactions, lowering fast-
start performance in particular species (Domenici et al., 2013)
[63]. The effects of hypoxia on aerobic swim performance and
sensory information acquisition, as well as the ability of fish
to enhance aerobic performance through acclimation
processes, might affect performance months after first
exposure (Ackerly et al., 2018) [55]. Hypoxia tolerance was
assessed using the aquatic surface respiration (ASR50) and
loss of equilibrium (LOE50) values, swimming performance
was assessed using the critical swimming speed (Ucrit),
aerobic capacity or space was assessed using the maximum
metabolic rate. The hypoxia tolerance drops with decreasing
temperature (fish usually showed high hypoxia tolerance due
to decreased oxygen demand and environmental oxygen
tension at low). Low hypoxia tolerance and poor swimming
performance may lead to problem in adoptability (Zhou et al.,
2019) [64]. At normal DO concentrations, fish can continue to
swim at modest rates. Only at lower DO concentrations,
around or below 5 mg/1 and some warm water fishes show
limited locomotion. Fish swimming is more linked to DO
concentration as compared to dissolved CO2 concentration of
aquatic body. Even considerably greater CO2 concentrations,
which have a significant influence on the sustained swimming
speeds of coho salmon, Oncorhynchus kisutch, at high levels
of DO, are ineffective at very low levels of O2. The
acclimatisation of goldfish, Carassius auratus, to O2 shortage
has little effect on their maximal speeds at low levels of O2. In
nature, very quick swimming is probably more common than
continuous swimming at maximum sustainable speeds
(Doudoroff & Shumway, 1970) [3].
The fast-start movement occurs in seconds and is primarily
limited by ATP and phosphocreatine in muscle tissues, the
lack of a DO influence on maximum speed is simply
understood. However, constant acceleration test (Ucat) was
more susceptible to DO alteration than Ucrit, with the former
showing a substantial decrease at a modest DO level (5 mg /l),
but the latter did not. Ucrit is more likely to be aerobic
swimming, whereas Ucat is more likely to be anaerobic
swimming. The reason for this could be because the
mobilisation, transportation, and use of energy fuels, rather
than the availability of oxygen, caused the limiting of Ucrit in
some fish species. In species like black barbel catfish
Pelteobagrus vachelli (Fu et al., 2009) [65], common carp
Cyprinus carpio (Zhang et al., 2010) [66], and Crucian carp
(Zhang et al., 2012) [67], this conclusion has been thoroughly
demonstrated (a so-called additive metabolic mode compared
to a locomotion priority mode in species whose swimming
activity can occupy all of their cardio-respiratory capacity;
(Fu et al., 2011) [52]. A minor fall in DO may not influence
Ucrit in these types of organisms. Ucrit was shown to be present
in mulloway Argyros omus japonicus and dark barbel catfish
(Pang et al., 2012) [68]. There has been no modification in the
current investigation, MO2 active in crucian carp did not
decrease at a moderate DO level, indicating that Ucrit was not
limited by respiratory capacity in normoxia. Furthermore,
instead of a component at the top end of the performance
range, Ucat consists of two components: an aerobic component
of steady swimming supported by aerobic 'red muscle' fibres,
and a component at the bottom end of the performance range
(Peake, 2008) [69]. Although Ucrit focuses primarily on
anaerobic metabolism, Ucat is governed by both aerobic and
anaerobic swimming, with the aerobic components of Ucat
relying on oxygen availability rather than substrate transit and
use due to its shorter duration. As a result, even little changes
in DO will reduce Ucat. It is important to conduct more
research into the effects of DO alterations on Ucat and Ucrit in
fish with distinct metabolic modes like additive vs. priority
mode (Penghan et al., 2014) [70].
3.2 Feeding
The concentration of dissolved oxygen in the water exerts a
significant impact on the metabolic rate of fish. Feeding
activities and other bodily functions decrease as the
concentration of dissolved oxygen drops. As a result, the
growth rate slows down and the fish become unable to absorb
the nutrients (Tom, 1998; Buentello et al., 2000; Andrews et
al., 1973) [71, 72, 73]. High DO condition leads to increment in
movement and digestion (Dey, 2017) [37]. Under hypoxia, both
feed intake and growth rate are significantly lower than under
normoxia (Chabot, 2003) [74]. When water oxygen saturation
goes below 70%, salmonids lower their food intake and stop
developing (Jobling, 1993) [75].
For non-aerated freshwater catfish ponds, a link between
feeding rate and dissolved oxygen was observed (Tucker et
al., 1979) [76]. Minimum dissolved oxygen concentrations
were about 1-0, 1-7, and 3-7 mg/litre at maximum feeding
rates of 78, 56, and 34 kg/ha day, respectively. Another study
used overnight DO prediction models to provide emergency
aeration to catfish ponds stocked at six densities and fed from
0 to 224 kg/ha day (Cole and Boyd, 1986) [77]. At larger
feeding rates, the ponds' aeration requirements increased.
There were significant relationship between low dissolved
oxygen and feeding rate (Boyd et al., 1979) [78].
3.3 Disease management
The availability of dissolved oxygen is the most important, as
it is influenced by temperature, water source, and biological
demand (i.e., high concentrations of bacteria and decaying
matter). The water's pH should be consistent and just below 7.
The level of waste products should be kept low, with special
attention paid to the presence of excessive carbon dioxide,
which is poisonous to most fish, and a build-up of ammonia,
which can cause the pH to rise above 7.5 (Cawley, 1983) [79].

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When DO levels are low and ammonia concentrations are
high that may cause mortality from disease breakout
(Chitmanat, 2013) [80]. Reduced respiration rates, diminished
reproductive activity, forced habitat shifts, and ultimately
reduced fish population sizes are all detrimental effects of a
lack of oxygen on fish and other lake biota (Garside, 1959;
Pollock et al., 2007) [81, 41].
Environmental stress, such as hypoxia, can damage fish
immune systems, making them more prone to disease
(Wedemeyer et al., 1976) [82]. Hypoxia makes certain endemic
diseases more harmful (Mellergaard and Nielsen, 1995) [83].
Fish sensitivity to low dissolved oxygen concentrations varies
by species, life stage (eggs, larvae, adults), and life processes
(feeding, growth, and reproduction) as well as different types
of activity (resting, swimming, digesting, etc.). Young fish are
most sensitive at the time of hatching, resulting in significant
losses when dissolved oxygen levels are decreased to 2–3
mg/l for many days. In the lack of oxygen, newly hatched
larvae can survive for one hour, while free-swimming larvae
can only survive for a few minutes (Alabaster and Lloyd,
1980) [84].
While sub-optimal dissolved oxygen levels are not instantly
fatal, they can stress fish and cause delayed mortality. A high
level of dissolved oxygen is essential for the fish to quickly
recuperate from the stress of catching and handling (Rottmann
et al., 1992) [85]. Handling stress, poor water quality, fast
temperature fluctuations, high stocking density, and
inadequate nutrition are all risk factors that can contribute to
the spread of viral infections. In intensive aquaculture
systems, which are typically highly stocked with fish, these
conditions may exist (Akoll & Mwanja, 2012) [86]. Due to
overcrowding or chemical pollutant exposure, fish face stress
that aggregated due to decreased DO. The situation supports
indigenous diseases spread in the aquatic body (Hershberger
et al, 1999; Carls et al, 1998) [87, 88]. The DO level in pond
water is important because it is directly associated with
disease outbreaks (Null et al., 2017; Domenici et al., 2017;
Gallage et al., 2016, 2017) [89, 90, 91, 92]. Fish development and
productivity will be reduced as a result of disease outbreaks
due to hypoxia, (Lovell, 1998; Shoemaker et al., 2000) [93, 94].
Adverse water quality in terms of anthropogenic activities or
adverse environmental conditions including hypoxia may
damage the immune system, resulting in diminished
resistance to pathogen infections (Di Marco et al., 2008) [95].
The majority of bacteria are opportunistic pathogens. As a
result, environmental stresses such as high temperature, low
dissolved oxygen, high ammonia content, and others are
primarily responsible for the initiation and severity of
bacterial infections (Plumb et al.,1976; Walters and Plumb,
1980) [96, 97]. Low levels of dissolved oxygen inhibit nitrifying
bacteria's capacity to convert ammonia and nitrite, it's critical
to keep an eye on dissolved oxygen levels. A strong water
quality management programme will help to prevent disease,
boost growth, and eliminate the need for chemical treatments
(Francis-Floyd et al., 2009) [98].
3.4 Survival
The most essential water quality variable in fish culture is
dissolved oxygen content (DO). If dissolved oxygen
concentration is continuously declining, then in aquatic
animals’ growth will be hampered, they will become prone to
infectious disease, finally fish will perish out. The smaller fish
consumes dissolved oxygen at a higher rate than the larger
fish, which explains why the larger fish perished faster
(Nimesh et al., 2012) [99]. Size-dependent mortality, like
oxygen deprivation, happens naturally but can be altered by
human actions (Lorenzen, 1996; Sogard, 1997; Gislason et
al., 2010) [100, 101, 102]. Total dissolved gas (TDG) caused by
the rapid overflow of water from the dam may threaten the
survival of fish. The increasing TDG level can decrease the
tolerance of juvenile fish. Large juvenile fish has weaker
tolerance to TDG supersaturated water than small juvenile
fish. TDG supersaturation can cause abnormal behaviours in
fish like loss of balance, loss of ability to swim and faster
breathing (Fan et al., 2020) [103]. The temperature has a
twofold effect: it decreases the solubility of oxygen while also
raising the metabolic requirement for oxygen in ectotherms
(Portner & Knust, 2007; Holt and Jorgensen, 2015) [104, 105].
There is limited work has been done hypoxia influence on
reproduction, but it can be hampered by excessive degrees of
hypoxia (Wu et al., 2003; Landry et al., 2007; Chabot and
Claireaux, 2008) [106, 107, 108]. Demersal fish populations are
declining due to a lack of oxygen and overfishing (Diaz and
Rosenberg, 2008) [109]. Because the minimal oxygen available
is dedicated to maintenance rather than somatic growth, low
oxygen saturation in water is a proximate factor causing
reduced asymptotic maximal size (Pauly, 1981; 2010; Van
Dam and Pauly, 1995; Chabot and Claireaux, 2008) [110, 111, 112,
108].
3.5 Respiration
For optimal fish production, proper oxygen management is
critical. The maintenance of healthy fish and bacteria that
break down the waste produced by the fish, and the fulfilment
of the biological oxygen demand (BOD) in the culture system
all necessitate the presence of oxygen. Fish respiration can be
hampered by low dissolved oxygen levels, as well as
ammonia and nitrite toxicity (Mallya, 2007) [113]. In hypoxic
environments, species specialised for aerial and surface film
respiration dominated the fish assemblage. Other animals
found in hypoxic environments have developed hypoxia-
specific behavioural and morphological adaptations. The
banded pygmy sunfish (Elassoma zonatum Jordan) and pirate
perch are both solitary species with low activity levels
(Robison and Buchanan, 1988) [114]. Which results in lower
respiration rates (Killgore and Hoover, 2001) [115].
Many fishes are exposed to seasonal temperature fluctuations
in their natural environment, rise in environmental
temperature causes an increase in metabolism in teleosts, as
seen by oxygen intake (Fry and Hart, 1948; Beamish, 1964;
Heath and Hughes, 1973) [116, 117, 118]. This increased oxygen
demand creates additional demands on the respiratory and
cardiovascular systems, which are partially addressed by
increased ventilation volume and heart output (Hughes and
Roberts, 1970; Stevens et al., 1972; Watters and Smith, 1973)
[119, 120, 121]. Any variation in O2 concentration is likely to cause
respiratory or cardiovascular compensations in fish. These
difficulties, which include variations in respiratory rhythm
(opercular rate), are adaptive and do not indicate any
impairment of ecologically important functions. Incipient
respiratory compensation may give idea of the dissolved O2
requirements of fish (Doudoroff and Shumway, 1970) [3].
During aquatic hypoxia, fish can maintain oxygen intake by
increasing gill ventilation. Increases in breathing rate and
stroke volume (Smith & Jones, 1982) [122]. Because marked
bradycardia is countered by an increase in stroke volume,
cardiac output is maintained during hypoxia. Systemic
resistance increases, increasing both dorsal and ventral aortic

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blood pressure (Short et al., 19979) [123]. Pulse pressure rises
dramatically as a result of the increased heart-stroke volume
and lower heart rate. The levels of dopamine, adrenaline, and
noradrenaline in the blood of sandy dogfish Scyliorhinus
canicula also increases in response to hypoxia (Butler et al.,
1978) [124]. Both arterial and venous blood oxygen contents
decrease (Holeton & Randall, 1967) [125]. (Soivio et al., 1980)
[126] found that erythrocytic ATP levels are lowered, resulting
in a significant increase in haemoglobin oxygen affinity.
Hypoxia increases the number of circulating erythrocytes,
which expand in some fish, resulting in a significant increase
in haematocrit. In sandy dogfish (S. canicula), however, no
rise in haematocrit has been seen in response to hypoxia
(Butler et al., 1979) [124]. Hypoxia cause increased activity in
vagal cholinergic fibres innervating the heart leads decrease in
heart rate (Randall, 1982) [127]. The afferent arm of this reflex
originates from receptors that respond to hypoxia and are
found on the first gill arch in the region of the various vessels
(Daxboeck & Holeton, 1978) [128]. When water flow through
the gills increases in proportion to oxygen absorption, and in
hypoxia, it is raised to sustain oxygen supply (Smith & Jones,
1981) [122].
3.6 Metabolism
The concentration of oxygen in the rearing environment has a
significant impact on the metabolic rate of fish (Tom, 1998)
[71]. Metabolism transforms meals or stored energy into the
energy required to carry out daily tasks. This process
eventually necessitates the use of oxygen, which must be
obtained from the environment (Nelson, 2016) [129]. Fish have
evolved respiratory and circulatory systems to perform this
role over a wide range of ambient oxygen levels, from above
air saturation to levels below which oxygen-demanding
activities cannot be sustained and death occurs (Chabot et al.,
2016) [130]. Hypoxia has been shown to have an impact on a
variety of physiological systems in fish, including metabolism
as a result, a reduction in feeding and growth (Chabot &
Dutil, 1999) [131].
Only a stable essential level of O2 can be ecologically
important, below which metabolic rates sustained by fish in
natural situations is still unknown or limited studies has been
done (Doudoroff and Shumway, 1970) [3]. In limited DO fish
groups had lower protein and lipid content in their bodies
because of metabolism requires more energy to cope with
hypoxic stress. Lipid and protein may be employed as an
energy metabolic substrate to adapt to low DO conditions and
sustain overall metabolism, resulting in a decrease in lipid and
protein content. Normal DO groups had more protein and fat
content, which could be linked to lower energy requirements
for feed absorption and other physiological processes. Fish
with normal DO levels appear to be able to lower the
proportions of metabolic energy and energy loss, saving more
energy for growth and lipogenesis (Duan et al., 2011) [132].
When a fish is under hypoxia stress its aerobic metabolism is
suppressed, whereas anaerobic metabolism is boosted and
metabolism of steroid hormones is slowed. Cell cycle arrest
occurs in liver cells, genes involved in cell development are
down-regulated in aerobic metabolism, while genes involved
in anaerobic metabolism are up-regulated. Uncoupling
proteins 2 and 3 are upregulated and may help to reduce
mitochondrial activity (Randall et al., 2006) [133]. Hypoxia-
tolerant animals have long been thought to be able to extend
their length of survival under extreme hypoxic conditions by
lowering their basal metabolic rate, which restricts the level of
activation of O2-independent ATP generation pathways
(Richards, 2009) [134].
3.7 Growth
Dissolved oxygen had a substantial effect on fish growth, and
low oxygen levels combined with reduced feed intake led to
decreased growth and changes in to stress response (Lakani et
al., 2013) [135]. Low DO has a negative impact on fish growth
and feed intake (Abdel-Tawwab et al., 2015) [136]. Growth
rates reduced and became progressively less as the dissolved-
oxygen concentrations decreased (Carter, 2005) [137]. Fish
metabolism and growth are dependent on the availability of
ambient oxygen. The growth rate of fish is fastest at high
dissolved oxygen and the growth rate of fish is slowest at low
dissolved oxygen. It is clearly shown that the different level
affects fish growth (Tsadik and Kutty, 1987) [138].
Some fish species are particularly sensitive to oxygen
saturation, and increasing DO to 100% enhances their growth
rate. In tropical freshwater fish, 5 mg per litre (80%
saturation) is indicated as the minimum and most effective
dose (Mallya, 2007) [38]. Fish development and feed utilisation
were harmed by low DO levels. A lack of oxygen available
for fish growth may reduce the growth observed under low
DO conditions. Fish development and feed efficiency were
affected by DO availability, according to (Bergheim et al.,
2006; Duan et al., 2011) [139, 132], When fed at a high enough
DO in water, fishes were always showed good feed
efficiency. according to (Abdel-Tawwab et al., 2014) [140]
Low DO levels significantly reduced the development of Nile
tilapia, Fish appetite and digestibility were both reduced in
low DO circumstances, resulting in low feed intake and
growth (Tran-Duy et al., 2012; Gan et al., 2013) [141, 142]. As a
result, under typical DO conditions, significant growth was
mostly owing to increased feed consumption and nutrient
digestibility. Spotted wolfish, Anarhichas minor (Foss et al.,
2002) [143], Nile tilapia, O. niloticus (Tran-Duy et al., 2008,
2012; Abdel-Tawwab et al., 2014) [144, 141, 140], striped bass,
Morone saxatilis (Brandt et al, 2009) [145] Atlantic halibut,
Hippoglossus (Thorarensen et al., 2010) [146]; Japanese
flounder, Paralichthys olivaceus (Duan et al., 2011) [132] and
grass carp, Ctenopharyngodon idella (Gan et al., 2013) [142]
Under hypoxic conditions, all of these fish had lower feed
intake and growth.
Smaller fish ate a low-calorie diet and developed faster than
larger fish at normal DO (Tran-Duy et al., 2008; Abdel-
Tawwab et al., 2010) [144, 147]. Larger fish were found to
tolerate low DO better than smaller fish. Small ones are
substantially less hypoxia-tolerant than larger ones (Almeida-
Val et al., 2000; Sloman et al., 2006) [148, 149]. As big body has
lower metabolic rate and body size affect a fish's ability to
take up oxygen under hypoxic situations (Nilsson and
Ostlund-Nilsson, 2008) [150].
3.8 Reproduction
The most significant abiotic elements impacting aquatic
species breeding efficiency are temperature (T) and dissolved
oxygen (DO). Maintaining the best combination of
temperature and DO will aid in improving breeding efficiency
and ensuring the largest quantity and quality of fingerlings are
produced (Qiang et al., 2019) [151]. Dissolved oxygen
concentrations in freshwater streams must be enough for fish
viability. To oxygenate the blood and meet their metabolic
demands, fish have evolved highly efficient physiological
processes for acquiring and utilising oxygen in the water.

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Reduced amounts of dissolved oxygen, can affect the growth
and development of eggs, and fry, as well as the swimming,
eating, and reproductive abilities of juveniles and adults
(Tang et al., 2020) [152]. By modifying embryo incubation
durations, decreasing fry size, increasing the chance of
predation, and decreasing feeding activity, such changes can
have an impact on fitness and survival. Low dissolved oxygen
concentrations can be fatal for fish under extreme
circumstances (Carter, 2005) [137].
Hypoxia can affect courtship behaviours, mate choice, and
reproductive efforts in fish. It can cause major reproductive
impairments by inhibiting testicular and ovarian development,
affecting sperm and egg production and quality, reducing
fertilisation and hatching success, and affecting larva survival
as well as the quality of fitness of juveniles (Spence et al.,
1996) [153]. In many fish species, hypoxia has been
demonstrated to delay embryonic growth and hatching. Under
hypoxia, fish embryos lose their usual synchronisation, and
defects in spinal and vascular development are prevalent (Wu,
2019) [154]. Hypoxia has been shown to affect fish endocrine
systems, slowing gonadal development and lowering
spawning, fertilisation, and larval growth success (Zhou et al.,
2001; Wu et al., 2003) [155, 106]. Zebrafish exposed to hypoxia
(0.5-0.8 mg/L) produced just 9 eggs per fish after the first
day, compared to 52 eggs per fish in the control group (Zhou
et al., 2001) [155]. Fertilized zebrafish eggs hatch 48 to 60
hours after fertilisation in normoxia, with 93.8 per cent of the
eggs hatching. Fertilized eggs took 96 to 260 hours to hatch in
hypoxia, with only 4.9 per cent hatching and the rest dying.
Fertilized eggs growing in hypoxia were pale, indicating that
they lacked pigment. Growth was slowed, and there were
numerous anomalies (Randall and Yang, 2003) [156].
Fish embryonic and larval stages are especially vulnerable to
low dissolved oxygen levels (Chapman, 1986) [157]. When
dissolved oxygen levels are below saturation (but over a
critical level), embryos can survive, although development is
often disrupted. When dissolved oxygen levels were below
saturation throughout development, embryos were found to be
smaller than normal, and hatching was either delayed or
preterm (Bjornn and Reiser, 1991; Wu et al., 2002; Zhou et
al., 2001) [158, 159, 155]. (Jones and Reynolds, 1999) [160]
observed that low DO did not affect hatching success or the
size of the young; nevertheless, hatching began one day later
on average in hypoxic environments. Similarly, (Lissaker et
al., 2003) [161] observed no link between decreased DO and
increased filial cannibalism.
Fish won't spawn at hypoxic condition. There were no eggs
laid at 1.0 mg/L DO, and there were fewer laid at 2.0 mg/L
DO than at control levels of 5.9–9.9 mg/L DO (Brungs, 1971)
[162]. The time it took for the eggs to hatch varied with
temperature and DO concentrations, with the latter increasing
as the former decreased. All of the fry died within 6-13 days
at 2.0 mg/L, while only 6% of the fry survived 30 days at 3.0
mg/L. Approximately 18% of the fry that survived the 4.0
mg/L treatment had spine curvatures. When it comes to fry
survival, fish exposed to 5.0 mg/L DO were comparable to
controls. At 2.0 mg/L, fry length was drastically reduced.
With lowering DO, the number of spawning attempts per
female dropped (Brungs 1971) [162].
3.9 Immunity
Fish can adjust to low dissolved oxygen levels in water by
boosting blood flow and red blood cell concentration. By this
way fish can increase the oxygen-carrying capacity of the
blood per unit volume and in the long run, by releasing excess
blood cells from the spleen (Svobodova and colleagues, 1993)
[163]. Hypoxia significantly affects the physiological and
immune responses of fish, making them more vulnerable to
disease. It leads to a functioning acute inflammatory response
to bacterial stimulation, and mildly down regulated gene
expression (FTH1, HIF1A, and NR3C1) (Abdel-Tawwab et
al., 2019; Schafer et al., 2021) [164, 165].
Individual heterogeneity in disease susceptibility in fish has
been connected to fish species (Yuasa et al., 1999; Evans et
al., 2000) [166, 167], genetic diversity, and immune response
(Suanyuk et al., 2008; Mian et al., 2009; Zamri-Saad et al.,
2010; Sarder et al., 2001) [168, 169, 170, 171]. Fish immunity was
found to be stronger in larger fish than in smaller fish,
showing that bacterial infection and innate immunity in
farmed fish may be influenced by fish weight or age.
Similarly, individual fish coping techniques may have an
impact on the fish susceptibility to infection (MacKenzie et
al., 2009; Huntingford et al., 2010) [172, 173].
Hypoxia, or lack of oxygen saturation, is one of the most
serious stressors in intensive aquaculture. The impact of
hypoxic conditions (3.2 mg/L DO) grown in RAS on their
health and immunological system. In fish, low dissolved
oxygen (DO) causes primary, secondary, and tertiary stress
reactions. Furthermore, the duration and intensity of hypoxic
conditions, as well as the animal's susceptibility to low
oxygen saturation, determine the result of the triggered
reaction (Schafer et al., 2021) [165].
3.10 Stress
In fish, hypoxia causes primary, secondary, and tertiary stress
responses (Bernier and Craig, 2005; Welker et al., 2007;
Bernier et al., 2012; Segner et al., 2012) [174, 175 176, 177]. Fish in
captivity are constantly exposed to recurrent and chronic
stressors (e.g., confinement, crowding, handling, and
changing water quality, including hypoxia) from which they
have no method of escaping. As a result, fish must adapt to
any of these husbandry stressors. As a result, the DO level
should be kept near saturation to improve fish development
and feed intake, as well as growth, development, disease
resistance, behaviour, and reproduction (Mallya, 2007;
Thorarensen et al., 2010) [38, 146].
Two hormonal systems, that produce corticosteroids
(primarily cortisol) and catecholamines (adrenaline and
noradrenaline and their precursor dopamine) which controls
the stress response (Reid et al., 1998) [178]. These components
work together to regulate secondary stress response, factors
that affect the delivery of vital resources such as energy and
oxygen to important parts of the body, as well as the
hydromineral balance and the immune system (Seibel et al.,
2021) [179]. If a fish survives a stressor, it returns to normal
state by homeostatic balance. The long-term effects of
repeated or protracted stress exposures are maladaptive,
compromising other vital life functions (growth, development,
disease resistance, behaviour, and reproduction), because of
high energy cost generates stress response (Schreck and Tort,
2016) [180].
Acute stress from capture, handling, transport, forced
exercise, hypoxia, osmotic and temperature shocks, or social
stressors, as well as exposure to water pollutants like acid
water containing aluminium, has been shown to cause a rapid
rise in muscle and plasma lactate, as well as a drop in blood
pH and oxygen content. These alterations were typically
accompanied by a significant increase in ventilation, branchial

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blood flow, gas exchange, and blood glucose levels, as well as
enormous catecholamines (CA) release from the chromaffin
cells (Barton and Iwama, 1991; Brown, 1993; Mazeaud and
Mazeaud, 1981; Pickering et al., 1987; Randall and Perry,
1992; Witters et al., 1991) [181, 182, 183, 184, 185, 186].
Acute hypoxia generated a release of epinephrine over
norepinephrine (Perry and Reid, 1994) [187], with the reduction
of arterial oxygen content/ saturation (rather than PO2) being
a key trigger inducing catecholamines to release. Gill injury
can also reduce arterial oxygen content, which may contribute
to catecholamines release in response to stress (Duthie and
Hughes, 1987) [188]. The fact that when hyperoxic and
normoxic rainbow trout were made acidotic by hypercapnia
concentration, blood pH decreased more in the hyperoxic than
in the normoxic fish, whereas CA levels increased only in the
normoxic animals, who had a lower arterial oxygen content,
suggests that acidosis is insufficient to induce CA to release
(Perry et al., 1989) [189]. However, it is unclear whether rapid
CA release during acute stress is only mediated by the
stressor's effects on arterial oxygen levels, or if it might also
be induced by the perception of external sensory cues.
The immunological response of fish is to boost innate
function when the stressor is acute and a short-term response
is stimulated (Tort, 2011) [190]. Acute stress causes an increase
in the number of circulating leukocytes (Barcellos et al.,
2004) [191]. This is due to the sympathetic nervous system's
activation and the production of catecholamines, which
mobilise both erythrocytes and leukocytes (Nardocci et al.,
2014) [192]. When a stressor is long-term, the immune system
is inhibited, which raises the risk of infection (Niklasson et
al., 2011) [193]. The negative effects of stress on the immune
response are thought to be mediated mainly by the
suppressive effects of glucocorticoids (i.e., cortisol) and are a
result of a failure to adjust to chronic stresses (Nardocci et al.,
2014) [192]. This could happen because coping with the
stressor has an allostatic cost that interferes with the
immunological response that is required.
3.11 Behaviour
Fish constantly expend energy on perfusion, typically on
ventilation, and frequently on movement during the process of
oxygen acquisition. These expenditures, as well as the risk of
predation, will change depending on the amount of oxygen
available and the sort of behavioural response demonstrated.
The four primary behavioural reactions to diminishing
external availability of dissolved oxygen are (1) changes in
activity, (2) increased use of air-breathing, (3) increased use
of aquatic surface respiration, and (4) vertical or horizontal
habitat alterations. Fish should select the response
combination that reduces the expense of supplying their
oxygen demands (Kramer, 1987) [194].
Hypoxia effects on fish behaviour like schooling which is
beneficial for their survival could have cause substantial
ecological consequences (Domenici et al., 2017) [195]. It also
impacts on anti-predator behaviour and fish escape responses,
as well as its modulation by ASR and schooling behaviour
(Domenici et al., 2017) [195]. When oxygen levels drop, fish
prey's metabolism and growth slow down, as well as their
overall health. This may have an adverse effect on their
locomotor and sensory abilities (Ackerly et al., 2018) [55].
Hypoxia can alter the circumstances of fish predators (growth
and metabolism). It can affect the preys escape performance
and predators attack performance (Lefrancois et al., 2005)
[196]. This could lead to a reduction in the number of predator-
prey encounters. Terrestrial predators, on the other hand,
could be able to take advantage of fish prey's reduced
performance in hypoxia, improving their chances of catching
a meal (Domenici et al., 2017) [195].
Fish appears to be able to safely avoid fatal levels of DO in
natural (Doudoroff and Shumway, 1970) [3]. Fish might be
seen swimming fast in a circular manner with a wide mouth
gape in other circumstances when the DO level was at its
lowest. As DO is restored to a normoxic level, this tendency
faded to normal swimming activity (Bowyer et al., 2014) [197].
This behaviour could be related to gill adaptation to hypoxia,
which includes decreased gas diffusion distance and increased
total respiratory surface (Saroglia et al., 2000) [198].
4. Consequences of Fluctuations in Dissolved Oxygen
In aquatic habitats, changes in oxygen levels are common; as
a result, organisms, including fish, have evolved a diverse
range of adaptations to both anoxia/hypoxia and hyperoxia.
Reactive oxygen species cause oxidative damage to cellular
components, affect glutathione status, and cause antioxidant
ant enzymes to respond to fluctuating oxygen supply in fish.
Antioxidant enzymes are increased in anticipation of
oxidative stress in anoxia- and hypoxia-tolerant species
during low-oxygen states, enhancing their antioxidant
capability for dealing with possible oxidative damage upon
restoration to normoxia. The glutathione system appears to
play an important adaptive role in hyperoxic environments.
Most stressful situations result in a rapid increase in lipid
peroxidation products, which are then quickly detoxified by
low- and high-molecular-weight antioxidants (Lushchak &
Bagnyukova, 2006) [199]. It should be noted that both an
excess and a deficiency of O2 cause oxidative stress. HIF-1 is
made up of a constitutively expressed HIF-1ß subunit and an
O2-regulated HIF-1 subunit. Prolyl hydroxylase domain
(PHD) proteins use oxygen as a substrate to hydroxylate HIF-
1 on proline 402 and/or 564. Proline hydroxylation allows
binding to the "von Hippel-Lindau" protein (VHL), which
recruits a ubiquitin ligase and promotes HIF-1 proteasomal
degradation (Kaelin et al., 2008) [200]. HIF-1α has been shown
to be activated faster by intermittent hypoxia than by
continuous hypoxia, though through different mechanisms
(Peers et al., 2007) [201]. Intermittent hypoxia induces ROS
production by NADPH oxidase, activating phospholipase
(PLC)γ, which generates inositol 1,4,5-triphosphate (IP3) and
diacylglycerol, thereby mobilizing intracellular Ca2+. Calcium
activates calcium-calmodulin kinase (CamK), protein kinase
C and finally mTOR, which facilitates HIF-1α synthesis and
inhibits PHD2-dependent degradation.
When a cell has given the treatment of hyperoxic conditions
results in an exponential release of ROS. Studies revealed that
the inhibition of mitochondrial complexes I & II by hyperoxia
leads releasing of ROS through ETC predominantly in early
phase and in late phase with more ROS being released by
NAD(P)H oxidase. Mitochondrial ROS initiates a calcium
(Ca2+) signal, that translocates Rac1 (a small GTPase) to the
plasma membrane, where it activates NAD(P)H oxidase
(Brueckl et al., 2006) [202]. Hyperoxia increases in ROS
affects a large number of intracellular signal transduction
proteins, including protein kinases, channels, transcription
factors, receptors and members of the apoptosis pathway
(Gore et al., 2010) [203]. Molecular responses to hyperoxia are
the redox-activated transcription factors, nuclear factor,
erythroid 2 related factor 2 (Nrf2), nuclear factor kappa B
(NF-KB) and activator protein-1 (AP-1) (Wright and
Dennery, 2009) [204].

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Fig 3: Cellular response to different oxygen condition, Abbreviations: activator protein 1 (AP1), calcium (Ca2+), Ca2+/calmodulin dependent protein kinase
(CamK), diacyl glycerol (DAG), electron transport chain (ETC), hypoxia-inducible factor (HIF), inositol 1,4,5-triphosphate (IP3), mammalian target of rapamycin
(mTOR), nicotine amide adenine dinucleotide phosphate (NADPH), nuclear factor ’kappa-light-chain-enhancer’ of activated B-cells (NFKB), nuclear factor
erythroid 2-related factor 2 (Nrf2), oxygen (O2), prolyl hydroxylase domain protein (PHD), phosphatidyl inositol (4,5)-bisphosphate (PIP2), phospholipase C
(PLC), protein kinase C (PKC), Rac, small GTPase;, reactive oxygen species (ROS), von Hippel-Lindau protein (VHL).
5. Conclusion
In this review we provide a concise overview on the influence
of higher and the lower concentration of dissolved oxygen on
freshwater fish. Dissolved oxygen is a critical premise for the
healthy growth of aquatic organisms, particularly in
aquaculture. Therefore, an accurate concentration of dissolved
oxygen is very important for the survival of fishes. The
fluctuations in DO are mainly caused by biotic and abiotic
factors like respiration, photosynthesis, organic waste
decomposition, aeration, temperature, salinity and
atmospheric pressure etc. Increasing and decreasing in these
factors make hypoxic and hyperoxic conditions in aquatic
ecosystem. These conditions cause drastic consequences in
fish physiology. Somewhere the anthropogenic activities for
the development of mankind is the root cause of variation in
DO concentration. To prevent these aquatic animals, get
under hypoxic and hyperoxic condition some preventive
measures should have taken like enhancement in the surface
area of an aquatic body for aeration, minimize in biological
oxygen demand, controlling weeds, lowering in nutrient
inputs. So, these practices can make an aquatic body healthier
for the survival of fish.
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