Science of the Total Environment 843 (2022) 156910

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Science of the Total Environment

journal homepage: www.elsevier.com/locate/scitotenv

Review

Blood biomarkers as diagnostic tools: An overview of climate-driven stress

responses in fish
⁎
Md Shahjahan a, , Md Jakiul Islam b, Md Tahmeed Hossain c, Moshiul Alam Mishu d,
Jabed Hasan a, Christopher Brown e
a
Laboratory of Fish Ecophysiology, Department of Fisheries Management, Bangladesh Agricultural University, Mymensingh 2202, Bangladesh
b
Department of Fisheries Technology and Quality Control, Faculty of Fisheries, Sylhet Agricultural University, Sylhet 3100, Bangladesh
c
Department of Biochemistry and Molecular Biology, Bangladesh Agricultural University, Mymensingh 2202, Bangladesh
d
Department of Biochemistry and Molecular Biology, Shahjalal University of Science and Technology, Sylhet 3114, Bangladesh
e
FAO-World Fisheries University Pilot Programme, Pukyong National University, 45 Yongso-ro, Nam-gu, Busan 48513, South Korea

H I G H L I G H T S G R A P H I C A L A B S T R A C T

• Global climate change due to anthropo-

genic activities affects the dynamics of
aquatic communities.
• Alterations in informative blood-based in-
dicators are used to monitor the physio-
logical fitness of an individual fish or an
entire population.
• Abnormal changes to temperature, light,
salinity, oxygen, acidification, and
inclusion of pesticides, heavy metals,
microplastics etc. in the habitat are ex-
pected reasons for the deviation of
hemato-biochemical parameters in fish.
• The review summarizes the blood analyses
those provide a comprehensive picture of
physiological status of fish under various
environmental and treatment conditions.

A R T I C L E I N F O A B S T R A C T

Editor: Julian Blasco Global climate change due to anthropogenic activities affects the dynamics of aquatic communities by altering the
adaptive capacities of their inhabitants. Analysis of blood provides valuable insights in the form of a comprehensive
Keywords: representation of the physiological and functional status of fish under various environmental and treatment conditions.
Aquaculture
This review synthesizes currently available information about blood biomarkers used in climate change induced stress
Antioxidants
responses in fish. Alterations in informative blood-based indicators are used to monitor the physiological fitness of in-
Oxygen
Physiology
dividual fishes or entire populations. Specific characteristics of fish blood, such as serum and plasma metabolites, cell
Salinity composition, cellular abnormalities, cellular and antioxidant enzymes necessitate adapted protocols, as well as careful
Stress attention to experimental designs and meticulous interpretation of patterns of data. Moreover, the sampling technique,
Temperature transportation, type of culture system, acclimation procedure, and water quality must all be considered for valid inter-
pretation of hemato-biochemical parameters. Besides, blood collection, handling, and storage time of blood samples
can all have significant impacts on the results of a hematological analysis, so it is optimal to perform hemato-
biochemical evaluations immediately after blood collection because long-term storage can alter the results of the anal-
yses, at least in part as a result of storage-related degenerative changes that may occur. However, the scarcity of high-

⁎ Corresponding author.
E-mail address: mdshahjahan@bau.edu.bd (M. Shahjahan).

http://dx.doi.org/10.1016/j.scitotenv.2022.156910
Received 24 March 2022; Received in revised form 12 June 2022; Accepted 19 June 2022
Available online 24 June 2022
0048-9697/© 2022 Elsevier B.V. All rights reserved.
M. Shahjahan et al. Science of the Total Environment 843 (2022) 156910

throughput sophisticated approaches makes fish blood examination studies promising for climate-driven stress re-
sponses in fish.

Contents

1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Methodology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3. Effects of environmental factors on hemato-biochemical parameters of fish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3.1. Temperature . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3.2. Photoperiod/light . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
3.3. Salinity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
3.4. Dissolved oxygen (DO)/hypoxia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
3.5. pH/acidification . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
4. Effects of environmental contaminants on hemato-biochemical parameters of fish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
4.1. Pesticides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
4.2. Heavy metals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
4.3. Microplastics (MPs) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
5. Conclusions and future prospects . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
CRediT authorship contribution statement . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
Declaration of competing interest . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16

1. Introduction
Climate change in response to anthropogenic activities (burning fuel,
deforestation, and discharge of industrial wastes, atmospheric accumula-
tion of greenhouse gases, drugs and chemicals use in agriculture, etc.) is a
global concern. Due to climate change, extreme weather events have been
occurring with increasing frequently and intensively throughout the
world (Cox et al., 2018; Imada et al., 2018). Temperature and precipitation
analyses indicate that the total area that experienced extreme cold, warmth,
and precipitation over the global land surface in 2016 was record-breaking,
exceeding the previous highest record in 2010 (Jorgenson et al., 2019; Xie,
2020). Climate change models predict further increases in anthropogenic
activities and extreme events which magnify the deterioration of water
quality parameters, which in turn generates hydrological stress (Albouy
et al., 2012; Nowicki et al., 2019). Although climate change-induced
water quality effects tend to be stronger over land than over the ocean,
these extreme events will also impact near-shore water salinity, where
most marine aquaculture occurs, by modifying precipitation patterns
(Cadiz et al., 2018; Servili et al., 2020). Consequently, abnormal changes
to temperature, light, salinity, oxygen, acidification, and inclusion of pesti-
cides, heavy metals, microplastics etc. in the habitat are among expected
reasons for the deviation of hemato-biochemical parameters in fish.
Fish are sensitive and vulnerable to environmental alterations, climate
change-induced events, and anthropogenic pollution (Rudneva et al., 2012;
Shahjahan et al., 2017). Farmed and wild fish encounter a multitude of envi-
ronmental stressors. Studies on fish physiological responses to the environ-
ment, aquaculture practices, feed, and contamination have been conducted
and have progressed in a largely independent manner (Ashaf-Ud-Doulah
et al., 2021; Burgos-Aceves et al., 2019; Wade et al., 2019). Using more com-
prehensive testing methods to understand the impacts of environmental con-
ditions has the potential to improve our ability to resolve the likely causes
and dynamics of disruptions and even harmful confounding factors, enabling
a more comprehensive grasp of relevant diagnostic biomarkers for fish health
(Rahman et al., 2019a, 2019b; Seibel et al., 2021).
Blood-based assays are routinely used as predictive and diagnostic
methods to assess fish health. The variations of blood-based hemato-
biochemical parameters serve as indicators of physiological changes that
correlate with environmental alterations in response to climate change
and/or human activities (Table 1). Some hematological parameters,
namely hemoglobin (Hb), hematocrit (Hct), red blood cell (RBC) and
white blood cell (WBC), platelet count (PLT), packed cell volume (PCV),
mean corpuscular volume (MCV), mean corpuscular hemoglobin (MCH),
and mean corpuscular hemoglobin concentration (MCHC) etc. contribute im-
portantly to the study of fish physiology. In addition, blood biochemical pa-
rameters relevant to metabolism, immunity, hormones, ions, and ionic
regulation such as glucose (Glu), total protein (TP), triglyceride (TG), alanine
transaminase (ALT), aspartate transaminase (AST), alkaline phosphatase
(ALP), sodium hydroxide dismutase (SOD), catalase (CAT), lactate dehydroge-
nase (LDH) etc. are viewed as important indicators of fish health and physiol-
ogy. These parameters have been used to monitor the health of fish in
response to variable diets, disease states, and environmental settings, in
order to monitor alterations in fish health (Ashaf-Ud-Doulah et al., 2019;
Kumar and Banerjee, 2016; Zarejabad et al., 2010). On the other hand, cellular
and nuclear damage of erythrocytes are employed for qualitative evaluation
(Fig. 1) caused by short or long term exposure to mutagenic agents
(Shahjahan et al., 2019).
The use and validation of standard non-lethal and inexpensive methods
are essential to monitor fish health and physiological fitness (Clauss et al.,
2008). One obstacle to the standardization of methods and the interpreta-
tion of results is the diversity of fishes and their evolutionary adaptation
to very different water quality conditions. A temperature or salinity that
is optimal for one fish may be lethal to another species, so diagnostic uses
of biomarkers must be applied in a well-informed, species-specific manner.
Blood analysis is a viable option for the assessment of physiological fitness
in fish quickly without killing them. Blood is a complex mixture of erythro-
cytes (red blood cells), leukocytes (white blood cells), and thrombocytes
(analogs to mammalian platelets), electrolytes, and nutrient contents
(Fazio, 2019; Rudneva et al., 2012; Zafalon-Silva et al., 2017). Fish blood
is present throughout somatic tissues and organs and behaves similarly to
established mammalian patterns. Since blood transports a range of critically
important substances (gases, water, minerals, nutrients, hormones, im-
mune effectors, toxins, and waste products), its examination can provide in-
sights into the status of fish physiology and health (Beyea et al., 2005;
Hedayati and Tarkhani, 2013; Zarejabad et al., 2010). Alterations in infor-
mative blood-based indicators like metabolic and enzymatic contents, hor-
monal profiles, and transcript abundances is reflective of systemic reactions
to changes or disturbances in homeostasis, and can therefore be used to
monitor the physiological fitness of an individual fish, a cohort, or an entire
population (Grant, 2015; Seibel et al., 2021). Fish blood tests have been
used in laboratory and field settings for decades for the appraisal of

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Table 1
Blood biomarkers used in studies of climate-driven stress responses in fish.
Parameters Units Indicators/diagnosis/abnormalities

Red blood cell indices Red blood cells (RBC) count ×106/mm3 Anemia, iron deficiency, respiration status
Hematocrit (Hct) % Anemia, iron deficiency, respiration status
Hemoglobin (Hb) g/100 mL Anemia, iron deficiency, respiration status
Packed cell volume (PCV) Ht Anemia, iron deficiency, respiration status
Mean corpuscular volume (MCV) Ht × 100/RBC Anemia, iron deficiency, respiration status
Mean corpuscular hemoglobin (MCH) Hb × 10/RBC Anemia, iron deficiency, respiration status
Mean corpuscular hemoglobin
Hb × 100/Hct Anemia, iron deficiency, respiration status
concentration (MCHC)
Anemia, abnormal morphology, stress, iron deficiency, inflammation, respiration
Erythrocytic cellular abnormalities (ECA) %
status, metabolic status
Anemia, abnormal morphology, stress, iron deficiency, inflammation, respiration
Erythrocytic nuclear abnormalities (ENA) %
status, metabolic status
White blood cell indices White blood cells (WBC) count ×103/mm3 Inflammation, phagocytosis, diseases
Lymphocytes % Inflammation, phagocytosis, diseases
Monocytes % Inflammation, phagocytosis, diseases
Neutrophils % Inflammation, phagocytosis, diseases
Eosinophils % Inflammation, phagocytosis, diseases
Basophils % Inflammation, phagocytosis, diseases
Blood metabolites Protein mg dL−1 Metabolic status, metabolic disorder, overall physiological status
Lipid mg dL−1 Metabolic status, metabolic disorder, overall physiological status
Triglycerides mg dL−1 Metabolic status, metabolic disorder, overall physiological status
Glucose mg dL−1 Metabolic status, metabolic disorder, overall physiological status
Lactate mmol dL−1 Anaerobic metabolic status
Cortisol ng mL−1 Neuroendocrine status, stress status
Urea nitrogen mg dL−1 Kidney, gill functions, osmoregulatory performance
Total cholesterol mg dL−1 Metabolism, energy, and nutritional status
Total bilirubin mg dL−1 Liver functions and physiological status
Creatinine mg dL−1 Kidney, gill functions, osmoregulatory performance
Blood immunological Serum lysozyme activities (LSZ) U mL−1 Immune status, disease status
parameters Phagocytic respiratory burst (RB) – Immune status, disease status
Enzymes GOT/AST U/L Liver functions, metabolic status, cellular functions
GPT/ALT U/L Liver functions, metabolic status, cellular functions
LDH U/L Liver functions, metabolic status, cellular functions
ɣGGT U/L Liver functions, metabolic status, cellular functions
Lipase activities Digestive activities performances
Amylase activities Digestive activities performances
mol.min−1. mL−1 of
Catalase (CAT) Antioxidant activities, physiological homeostasis
protein
mol.min−1. mL−1 of
Glutathione peroxidase (GPx) Antioxidant activities, physiological homeostasis
protein
mol.min−1. mL−1 of
Glutathione reductase (GR) Antioxidant activities, physiological homeostasis
protein
mol.min−1. mL−1 of
Superoxide dismutase assays (SOD) Antioxidant activities, physiological homeostasis
protein
mol.min−1. mL−1 of
Glutathione (GSH) Antioxidant activities, physiological homeostasis
protein
mol.min−1. mL−1 of
Total antioxidants values Antioxidant activities, physiological homeostasis
protein
Ions Na+ mmol dL−1 Osmoregulatory functions
Cl- mmol dL−1 Osmoregulatory functions
K+ mmol dL−1 Osmoregulatory functions
Mg++ mmol dL−1 Osmoregulatory functions
HCO3- mmol dL−1 Osmoregulatory functions

physiological status, reproductive performance, immune status, and pat-
terns of hormonal regulation (Duman et al., 2019). Moreover, blood is the
subject of more fully-established analytical methods than those available
for other tissue or fluid samples (Burgos-Aceves et al., 2019; Seibel et al.,
2021). Nonetheless, deciphering the obtained data in order to derive prac-
tically useful knowledge about the individual level can be challenging. A
number of recent reviews have updated our comprehension of the use of he-
matological techniques in fish production and welfare (Burgos-Aceves
et al., 2019; Fazio, 2019; Seibel et al., 2021). Surprisingly, other blood bio-
marker measurements, rather than hematological parameters, are rela-
tively underrepresented in fish physiology analyses using systems
biochemical parameters, biological approaches, PCR, and omics-based
techniques. The intent of this review is to provide an overview of currently
available information about blood biomarkers used in climate change-
induced stress responses in fish under controlled laboratory conditions,
aquaculture systems, and in the wild, and to discuss their potential
consequences for fitness and performance in the context of potential eco-
logical consequences. Emphasis is also given to the modulation of
hemato-biochemical factors in fishes as induced by a range of climatic pa-
rameters.
2. Methodology
In this review, we interpret results published with a focus on contribu-
tions since 2000 regarding hemato-biochemical disorders in fish due to
climate-induced factors. We conducted the literature survey using Web of
Science, NCBI-PUBMED, Research Gate and Google Scholar search engines,
and Mendeley as the reference manager. Google Scholar was chosen above
other academic search engines because it includes materials from both aca-
demic and grey literature. Mendeley was chosen as an open source, collab-
orative platform that allows for the collection, organization, annotation,
and labeling of search results for analysis.

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M. Shahjahan et al. Science of the Total Environment 843 (2022) 156910

a b c

d e f

Fig. 1. Cellular and nuclear abnormalities in Giemsa stained blood smears of Nile tilapia exposed to high temperature, pesticides and heavy metals; (a) tear-drop, (b) twin,
(c) spindle, (d) karyopyknosis, (e) nuclear bridge, and (f) notched.

3. Effects of environmental factors on hemato-biochemical parame-
ters of fish
3.1. Temperature
Temperature can be viewed as the master abiotic environmental factor
associated with adaptation of hemato-biochemical parameters among ecto-
therms. Thermal fluctuations can trigger compensatory metabolic changes
at the cellular and molecular levels, resulting in considerable alterations
in blood chemistry. In fish, metabolic rate generally increases approxi-
mately 2 to 3 fold in response to a 10 °C increase in the ambient tempera-
ture (Pörtner and Peck, 2010). Consequently, metabolic rate, enzymatic
reactions, cellular respiration, and oxygen consumption vary with temper-
ature (Kamunde et al., 2019; Liu et al., 2019; Volkoff and Rønnestad,
2020). Blood is systemically engaged in these physiological responses.
Any change in temperature-induced physiological signatures is reflected
by a range of hemato-biochemical parameters (Table 2), which can serve
analytical and diagnostic purposes. A wealth of data is available on
temperature variations in hemato-biochemical parameters, indicating that
temperature change is among the most important factors influencing fish
hemato-physiology. This is primarily due to seasonal changes in ambient
temperature and dissolved oxygen content, which cause variability in fish
hemato-biochemical parameters.
Adverse environmental temperatures affect the numbers, size and
shapes of circulating erythrocytes and can also change the composition of
circulating leukocytes. European seabass, Dicentrarchus labrax exposed to
extreme warm temperatures exhibited significantly decreased RBC, Hb,
and Hct, whereas WBC counts were found to increase during extreme
warm exposure (Islam et al., 2020a, 2020b, 2020c). In other studies, the
leucocyte population rose dramatically during the warm exposure period,
but the opposite tendency was observed during the cold exposure period,
as observed in common carp Cyprinus carpio (Engelsma et al., 2003) and
channel catfish Ictalurus punctatus (Martins et al., 2011). Fish become sus-
ceptible to physiological impairments and have higher WBC counts when
exposed to stressfully-high temperatures for an extended period. Fish in
cold water, on the other hand, have low WBC counts because changes in
water temperature might interfere with immune response owing to ecto-
thermic metabolism (Tavares-Dias and Moraes, 2007). The effects of adre-
nocorticotrophic hormone and the resultant cortisol synthesis and
secretion, as well as cell mitogenic activity, all influence the reduction in
lymphocyte number at low temperatures (Castillo et al., 2009). These hor-
mones tend to result in immunosuppression by suppressing immune cell
production, (Adeyemo et al., 2012; Arjona et al., 2010).
Annually cyclic fluctuations of blood parameters reveal a significant in-
crease in the warm spring and summer months and a drop in the autumn
and winter months (Ibrahim and Erdeml, 2002). De Pedro et al. (2005)

Table 2
Effects of temperature on hemato-biochemical parameters of fish.
Species Temperature (°C) Exposure time (days) Hemato-biochemical alterations References

Increase Decrease

Dicentrarchus labrax 8 °C 20 RB and LSZ RBC, Hct, Hb Islam et al., 2021b

Labeo rohita 36 °C – WBC, Glu Hb, RBC Ashaf-Ud-Doulah et al., 2020
Dicentrarchus labrax 32 °C 30 WBC, TG and Lactate Glu, TP and RBC Islam et al., 2020c
Oreochromis niloticus >34 °C 30 WBC, Glu Hb, RBC Islam et al., 2020b
Cyprinus carpio 35 °C 14 WBC, Glu Hb, RBC Shahjahan et al., 2020b
Betta rubra 32 °C 14 Glu Nur et al., 2020
Labeo rohita >33 °C 60 WBC, Glu Hb, RBC Ashaf-Ud-Doulah et al., 2019
Danio rerio 35 °C 4 Glu Hb Shahjahan et al., 2019
Epinephelus akaara 31, 34 °C 42 RBC and WBC, neutrophil lymphocytes Rahman et al., 2019a, 2019b
Pangasianodon hypophthalmus 36 °C 30 WBC, Glu Hb, RBC Islam et al., 2019a, 2019b
Pangasianodon hypophthalmus 36 °C 7 WBC, Glu Hb, RBC Shahjahan et al., 2018
Salmo salar 18 °C 14 Glu, Lactate, Hct TG Sambraus et al., 2018
Acipenser brevirostrum 15 °C 21 Glu, Lactate MCHC, Na+, Cl- Zhang and Kieffer, 2014
Prochilodus scrofa 20 °C 4 Hct Hb, RBC Carvalho and Begnis, 2006

Glu; glucose, Hb; hemoglobin, Hct; hematocrit, LSZ; serum lysozyme activities, MCHC; mean corpuscular hemoglobin concentration, RBC; red blood cell, TP; total protein,
TG; triglycerides, WBC; white blood cell.

4
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reported variations in hemato-biochemical characteristics in tench Tinca
tinca at cold temperatures, with decreased mean levels of RBC, Hb, and
Hct. Low temperatures alter hormonal and metabolic processes, which
tend to inhibit erythropoiesis, resulting in decreased values of RBC-
related parameters during cold seasons (Huusko et al., 2007). Fish of differ-
ent ages and genders also show a selective pattern of hemato-biochemical
responses to rearing temperatures. For example, different age groups of
pike Esox lucius exhibited the highest value of RBC, Hct, Hb during warm
summer than cold winter (Fallah and Barani, 2014). Sayed et al. (2016) re-
ported that Hb, Hct, RBC, WBC, MCV, MCH, and MCHC levels were greater
in the summer and lower in the winter in both male and female snow fish
Schizothorax plagiostomus. From the above literature, it can be concluded
that a majority of fish exhibit trends of up-regulation in hematological
values (RBC, WBC, Hct, and Hb) during warm temperatures/seasons, and
decreases during cold exposure periods. In contrast, some fishes exhibit in-
creases in WBC counts, MCV, and MCH during cold exposure and winter
(Rahman et al., 2019a, 2019b).
Temperature also affects blood biochemistry, including glucose and cor-
tisol levels. For European seabass, glucose and cortisol content were found
to increase during extreme warm exposure, while serum protein and tri-
glycerides exhibited decreasing trends (Islam et al., 2020a, 2020b,
2020c). Cold temperature stress was also found to increase glucose, triglyc-
erides, cortisol, and concurrently decreasing protein content in European
seabass, D. labrax (Islam et al., 2020a, 2020b, 2021c). The changes in me-
tabolites in fish in response to stressful temperature exposure can be
interpreted as indicators that the fish are adjusting metabolically in order
to address increased energy demands. Fish become susceptible to physio-
logical impairments and have higher WBC counts when exposed to stressful
high temperatures for an extended period. Fish in cold water, on the other
hand, have low WBC counts because changes in water temperature can po-
tentially interfere with immune responses in cases of ectothermic metabo-
lism (Tavares-Dias and Moraes, 2007). The action of adrenocorticotrophic
hormone and cortisol synthesis, as well as cell mitogenic activity, impacts
the decrease in lymphocyte number in low temperatures (Castillo et al.,
2009). These hormones tend to inhibit immune cell production, resulting
in immunosuppression (Adeyemo et al., 2012; Arjona et al., 2010). WBC
profile comprises a differential count in the bloodstream of an animal, in-
cluding fish stressed by high temperatures (Davis et al., 2008). Circulating
cortisol levels influence the leukocyte profile in fish and other animals.
When under stress, teleosts produce and secrete a large quantity of gluco-
corticoids into the circulation, resulting in a decrease in the number of cir-
culating lymphocytes and an increase in WBC (Davis et al., 2008; Tort,
2011).
Stressful temperature exposure provokes enormous antioxidant and en-
zymatic responses in fish blood. The increase in water temperature caused
cellular and physiological stress in fish (Fuat Gulhan and Selamoglu, 2016;
Zafalon-Silva et al., 2017), associated with oxidative stress (lipid oxida-
tion). Changes in membrane permeability, fluidity, and ion transport result
from such disturbances (Acar, 2018; Chen et al., 2004; Abdel-Aziz et al.,
2016). Furthermore, inhibition of metabolic processes and changes in
transmembrane potential (depolarization) result in the release of mitochon-
drial calcium, which causes uncoupling and activation of caspase-3, result-
ing in DNA fragmentation, which can lead to erythrocytic cellular and
nuclear abnormalities. Furthermore, higher water temperatures promote
DNA damage by releasing DNase enzymes from lysosomes and thermally
inactivating the enzymes responsible for DNA repair, resulting in changes
in cellular nuclear morphology. Warm temperature-induced intracellular
damage is directly connected to lipid peroxidation, which promotes cross-
linking and polymerization of membrane components, hence affecting
membrane activities. The unsaturated fatty acids in membranes (phospho-
lipids, glycolipids, and sterols) and trans-membrane proteins are particu-
larly vulnerable to free radical damage. These findings highlight the
importance of membrane integrity as a barrier to intracellular homeostasis.
Cellular and nuclear disorders originate from structural damage to mem-
brane lipids or protein components. In European seabass, erythrocytic ab-
normalities, enzymatic antioxidants (catalase, superoxide dismutase,
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Science of the Total Environment 843 (2022) 156910
glutathione peroxidase, glutathione reductase), cellular antioxidants (as-
partate transaminase, alanine transaminase, lactate dehydrogenase,
gamma glutamyl transferase) reportedly increase during and after extreme
warm exposure (Islam et al., 2020a, 2020b, 2020c). When the production
of free radicals exceeds the capacity of antioxidants, fish are confronted
with oxidative problems caused by thermal stress (Oksala et al., 2014).
To function under conditions of warm stress, fish activate antioxidants for
defensive purposes (Rossi et al., 2017). Antioxidants and radical-
scavenging enzymes (CAT, SOD, GPx, GR, GSH) work together to counter-
act the potentially harmful effects of ROS (Banh et al., 2016; Rossi et al.,
2017). Thermal exposure increases antioxidant enzymes in the gills, liver,
and muscle of European seabass, D. labrax (Islam et al., 2020a), largemouth
bass, Micropterus salmoides (Sun et al., 2020), and Rock goby, Gobius
paganellus (Vinagre et al., 2014). Cellular and nuclear damage of erythro-
cytes (Fig. 1) are considered to be indicative of an organism's exposure to
stressful thermal events (Cavas et al., 2005; Gomes et al., 2015;
Harabawy and Mosleh, 2014; Zafalon-Silva et al., 2017). Cold temperature
stress was also found to increase blood cell abnormalities, and the abun-
dance of antioxidant enzymes in European seabass (Islam et al., 2020b,
2021a, 2021c). Lysozyme and complement components are largely gener-
ated by leukocytes and the liver, respectively, and are released into the
blood circulation to destroy pathogens. Quantification of blood lysozyme
activities (LSZ) and phagocytic respiratory bursts (RB) are considered im-
portant as immune system indicators in fish. Significantly, higher LSZ and
RB have been observed in European seabass exposed to both high and
low temperatures (Islam et al., 2020c, 2021a, 2021c). Temperature, how-
ever, is not the sole inducer of changes in hemato-biochemical parameters
in fish; other parameters must also be considered. Increased water temper-
ature reduces dissolved O2 content and also alters pH in the water, resulting
in increased metabolic activity, to which fish adapt by increasing hemoglo-
bin content.
3.2. Photoperiod/light
Another environmental component that profoundly influences fish
physiological status is light (Solomon and Okomoda, 2012). Because multi-
ple organs in fish have photo-sensory capacity, photoperiod is understood
to play a key role in maintaining physiological equilibrium. In fish, both
the eye and the pineal gland contain photoreceptor cells (Davies and
Bromage, 2002; Taranger et al., 2003). Artificial photoperiod manipulation
is a commonly used technique in aquaculture to optimize fish production
and to regulate the timing of reproductive performance (Badruzzaman
et al., 2020; Bromage et al., 2001; Guerrero-Tortolero et al., 2010). For
fish, due to high blood sensitivity, a range of blood-based biomarker studies
have been conducted to understand the potential effects of light and
changes in seasonal photoperiod (Burgos-Aceves et al., 2019; Seibel et al.,
2021).
The effects of photoperiod manipulation-induced stress on fish physio-
logical responses have been investigated (Table 3). Prolonged exposure to
light adversely affects hemato-biochemical responses. Fish exposed to
24 h of light had the lowest Hb, RBC, WBC, Hct, and PCV values compared
to fish reared at a photoperiod of 24 h of darkness or 12 h of light. In the
case of sturgeon (Huso huso), changes in hematological parameters were
found as a result of stress from changes in photoperiod. An increase in
Hct was seen in fish subjected to 24 h light and dark regimes, which was
followed by a decrease in Hb and erythrocytes (Bani et al., 2009), indicating
the possible onset of anemia. Rainbow trout Oncorhynchus mykiss exposed
to 24 h of photoperiod exposure exhibited increased Hct and RBC during
a 30-day study period (Valenzuela et al., 2008). In another study,
(Valenzuela et al., 2007) reported that rainbow trout exposed to continuous
light exhibited considerably greater Hct levels and RBC counts, but reduced
leukocyte numbers. Contrasting results have also been reported. For exam-
ple, in Indian catfish subjected to photoperiod of 24 h of light and 24 h of
darkness, RBC, and WBC did not differ in either photoperiod regime. How-
ever, fish subjected to 24 h light had altered leukocyte counts, revealing
lymphopenia and neutrophilia (Srivastava, 2010). Low levels of PCV, Hct,
M. Shahjahan et al. Science of the Total Environment 843 (2022) 156910

Table 3
Effects of light on hemato-biochemical parameters of fish.
Species Light:Dark Exposure time Hemato-biochemical alterations References
(days)
Increase Decrease

Labeo rohita 18:06 30 WBC, Glu Hb, RBC Shahjahan et al., 2020a
Salmo salar 16 L:08D 24 weeks Cortisol – van Rijn et al., 2020
Megalobrama amblycephala 16 L:8D 56 Cortisol – Tian et al., 2019
Oncorhynchus mykiss 24 L:0D 90 – Neutrophil and Monocyte Lone et al., 2017
Heterobranchus bidorsalis 24/18 L:0/6D 84 Lymphocytes – Osho et al., 2016
18 L:6D Lymphocytes Hct
Brycon orbygnianus 90 Machado et al., 2016
6 L:18D Neutrophils, Lymphocytes –
Lophiosilurus alexandri 24 L:0D 36 Cortisol – Kitagawa et al., 2015
Rutilus rutilus caspicus 24 L:0D 56 RBCs and Hb – Shahkar et al., 2015
Acipenser persicus 0 L:24D 56 Hct Cortisol, WBC Falahatkar et al., 2012
24 L:0D – WBC, Hb, RBC, PCV
Clarias gariepinus 42 Solomon and Okomoda, 2012
0 L: 24D WBC, Hb, RBC, PCV
Clarias batrachus 24:00 1 Neutrophils Lymphocyte Srivastava and Choudhary, 2010
Onchorhynchus mykiss 24:00 30 Hct, Hb, RBC, MCH, MCHC WBC, Lymphocyte Valenzuela et al., 2008
Onchorhynchus mykiss 24:00 150 – Lymphocytes, thrombocytes Valenzuela et al., 2007
Acipenser baerii 16:08 1 WBC Ruchin, 2007
Onchorhynchus mykiss 24:00 60 Immature RBC – Valenzuela et al., 2006
Cyprinus carpio 12:12 1 Lymphocyte Monocyte Ruchin, 2006
Pagrus major 16/24:8/0 56 Cortisol – Biswas et al., 2006
Oreochromis niloticus 6:6 (Light phase) and 6:6 (Dark phase) 90 Cortisol, Lymphocyte Biswas et al., 2004
Oncorhynchus mykiss 24 L:0D 60 Cortisol – Leonardi and Klempau, 2003

Glu; glucose, Hb; hemoglobin, Hct; hematocrit, MCH; mean corpuscular hemoglobin, MCHC; mean corpuscular hemoglobin concentration, RBC; red blood cell, WBC; white
blood cell,

and Hb appear to be associated with a drop in RBC, which also appears to
be associated with ATP depletion (Felix et al., 2009) and failure of cell
membranes to discharge excess sodium, leading to hemolysis (Guyton
and Hall, 2005).
Cortisol levels and immune cells in fish blood have been reported to be
influenced by photoperiod. This appears to be a stress-related fish charac-
teristic, either as a direct cytolytic action of cortisol on lymphocytes or as
immunological cell dispersion in lymphoid regions (Grzelak et al., 2017).
For instance, rainbow trout (Oncorhynchus mykiss) exposed to 24 h light
showed a significant decrease in B and T lymphocytes and increased plasma
cortisol. Significantly elevated plasma cortisol levels indicate stress and im-
munosuppression (Leonardi and Klempau, 2003). According to Leonardi
and Klempau (2003), continuous light boosts cortisol levels for up to two
months after the light is turned off, which may explain the low number of
leucocytes found in the other studies. However, contrasting results were
also reported in several studies. Biswas et al. (2004), for example, found
no signs of stress (acute or chronic) in Nile tilapia Oreochromis niloticus, ex-
posed to different artificial light regimes. The hypothalamic/hypophysial/
thyroid axis is also exceedingly sensitive to photoperiod cues, as involved
with the regulation of reproductive cyclicity (Norberg et al., 2004), and
metabolic status (Vera Cruz and Brown, 2009).
There is conflicting evidence on the effect of artificial photoperiod re-
gimes on many hemato-biochemical parameters in fish. As a result,
hemato-biochemical parameters in fish cannot always serve as a good indi-
cator of physiological status when subjected to prolonged photoperiod and
artificial light changes. Given that other factors, which are not necessarily
related to photoperiod changes, can also affect hemato-biochemical param-
eters, consideration of other biomarkers that are responsive to photoperiod
changes would be advisable.
3.3. Salinity
Salinity tolerance is among the most important environmental/physio-
logical variables in fish and it has been investigated intensively since it is
regarded as a decisive determinant of fish growth and survival (Fiúza
et al., 2015; Saillant et al., 2003). Effects of salinity on hemato-

Table 4
Effects of salinity on hemato-biochemical parameters of fish.
Species Salinity (ppt) Exposure time (days) Hemato-biochemical alterations References

Increase Decrease

Nibea albiflora 60 7 Na+, Cl−, Ca2+ MDA, ACP, AKP Tian et al., 2020
Pangasianodon hypopthalmus >10 60 WBC, Glu Hb, RBC Jahan et al., 2019
Heteropneustes fossilis 9 90 Glu Glycogen, ALP, ATPase Ahmmed et al., 2017
Oreochromis niloticus 12 14 Hct, Hb TP, RBC Elarabany et al., 2017
Oreochromis spp 20 42 SOD, MDA, Glu, TG RBC He et al., 2017
Periophthalmus waltoni 17 1 WBC RBC Soltanian et al., 2016
Notopterus notopterus 1.6 30 WBC, ALT, AST, – Kavya et al., 2016
Barbonymus gonionotus >9 3 WBC, Glu RBC Fahima et al., 2015
Colossoma macropomum 15 84 Glu, Hct, Hb – Fiúza et al., 2015
Monopterus albus 20 g/L 6 Plasma [Na+], [K+] – Pedersen et al., 2014
Mugil cephalus 25 20 Lactate, AST, ALT Glu, Hb, RBC, Hct Fazio et al., 2013
Fundulus heteroclitus 35 4 SOD, GST – Loro et al., 2012
Sarotherodon melanotheron 0.34 % 7 WBC, MCV Hb, RBC, PCV, MCHC Anyanwu et al., 2007
Rachycentron canadum 30 70 – Hct, plasma protein Denson et al., 2007
Rhombosolea tapirina 3, 7, 15 4h – Plasma Na, Mg Girling et al., 2003
Acipenser naccarii 35 20 Hb, Hct, RBC – Martínez-Alvarez et al., 2002

ALP; alkaline phosphatase, ALT; alanine transaminase, AST; aspartate transaminase, Glu; glucose, Hb; hemoglobin, Hct; hematocrit, RBC; red blood cell, TP; total protein,
WBC; white blood cell.

6
M. Shahjahan et al.
biochemical parameters of fish are summarized in Table 4. In hypo- and
hyper-saline conditions, fish spend more energy to maintain physiological
homeostasis, affecting physiology, growth, development, and survival.
Some species are genetically capable of adapting to more extreme salinities
(euryhaline fishes; for example see Piermarini and Evans, 1998) while
others are constrained to more narrowly defined osmotic environments
(stenohaline fishes; Nakamura et al., 2020). Fish adaptation to various en-
vironmental salinities causes changes in the blood osmoregulatory, various
ions and glucose levels, thereby having an impact on physiological homeo-
stasis (Laiz-Carrión et al., 2005; Vargas-Chacoff et al., 2019). Salinities to
some levels have been shown to improve growth and development, as re-
ported for some freshwater fish species (Imanpoor et al., 2012; Luz and
Dos Santos, 2010; Overton et al., 2008). In those studies, several hemato-
biochemical parameters have been used as stress indicators to assess salin-
ity impacts on fish physiology (Abdel-Rahim et al., 2020; Fazio et al., 2013;
He et al., 2017; Sardella et al., 2004).
There is also a link between salinity stressors and hemato-biochemical
changes, which can have a substantial influence on fish physiology and
the immune system (Evans and Kültz, 2020; Silveira et al., 2019;
Zarejabad et al., 2010). In Nile tilapia (O. niloticus), Hct and Hb were
found to decrease, accompanied by a decrease in RBC, most likely resulting
from changes in blood water content in response to the increasingly
hyperosmotic environment. Meanwhile, WBC does not show any substan-
tial change in response to increased salinities (Bosisio et al., 2017). How-
ever, this is not the case in settings with decreasing salinity, where
lymphopenia, neutrophilia, and monocytosis can be seen, leading to immu-
nological dysregulation. In contrast, the euryhaline flathead grey mullet
subjected to salinities of 25 and 45 PSU had the lowest levels of RBC, Hb,
and Hct compared to fish exposed to a salinity of 35 PSU, while WBC
level was greatest in fish at 25 PSU and lowest at 45 PSU (Fazio et al.,
2013). The drop in RBC, Hb, and Hct levels might be attributed to osmoreg-
ulatory dysfunction caused by changes in salinity (Girling et al., 2003), and
a decrease in WBC implies a possible immunosuppressive impact from hem-
orrhagic damage induced by salinity fluctuation (Anyanwu et al., 2007). In
bronze featherback Notopterus notopterus, hematological parameters (Hb,
Hct, RBC, and WBC) reportedly increased following exposure to increasing
salinity, possibly reflecting adaptations to the changing salinity (Kavya
et al., 2016). In several teleosts, Hct levels decreased following exposure
to hypersalinities (Denson et al., 2007; Fiúza et al., 2015; Islam et al.,
2020a; Islam et al., 2021a, 2021b, 2021c, 2021d, 2022; Jahan et al., 2019).
Salinity levels have a significant impact on antioxidants and cellular
defense systems, and can induce oxidative stress (Fazio et al., 2013; Islam
et al., 2020a, 2020c, 2021a). In fish, salinity stress-induced functional
tissues and organ damage lead to the release of specific cellular
enzymes into the bloodstream (Krajnovic-Ozretic and Ozretic, 1987;
Roychowdhury et al., 2020). The increased activity of these enzymes in
serum can induce cellular and tissue damage, which could be caused by dis-
ruptions in normal cellular antioxidants processes and subsequent enzyme
leakage into the bloodstream (Samanta et al., 2014). Plasma SOD, GST,
and GSH content in olive flounder (Paralichthys olivaceus) were significantly
increased at low (2 PSU) salinity (Antony et al., 2021). Total protein, SOD,
CAT, RBC, and triglycerides in juvenile Nile tilapia O. niloticus, decreased
significantly as salinity increased (Elarabany et al., 2017; He et al., 2017;
Semra et al., 2016). Increases in total oxidative scavenging capacity
(TOSC) in killifish (Fundulus heteroclitus) occurred at increasing salinities,
which were associated with increases in SOD, GST, and GSH activities
(Loro et al., 2012). In recent studies, European seabass exposed to cold tem-
perature while at 30 PSU salinities exhibited significantly higher concentra-
tions of serum transaminase activities (GPT, GOT, LDH, γGGT) and
antioxidant activities (SOD, GPx, GR, CAT) in fish exposed at 3 and 30
PSU compared to fish at 6 and 12 PSU (Islam et al., 2021a, 2021c). Changes
in salinity also induce changes in osmotic balance and blood-based bio-
chemical parameters in fish. In recent studies, European seabass exposed
to cold temperature held at 30 PSU salinities exhibited significantly higher
concentrations of serum Na+, Cl−, and K+ compared to fish at 3 PSU
(Islam et al., 2021a, 2021c). Cortisol is an ion pump stimulator and is
7
Science of the Total Environment 843 (2022) 156910
commonly referred to as a salinity-adapting stress hormone because bran-
chial Na + secretion in fish acclimatized to both freshwater and seawater
needs a higher degree of ion pumping activities (Bystriansky and Schulte,
2011; Cheng et al., 2009; De Boeck et al., 2000).
Salinity stress can cause changes in membrane permeability and alter-
ations of the rates of cellular metabolic activities. To keep pace with these
osmotic stress-induced metabolic demands fish spend stored lipid reserves,
ultimately hampering cellular homeostasis and resulting in cellular and nu-
clear damage of erythrocytes (Fig. 1). The net effects of embryonic or larval
exposure to even minute alterations in salinity include serious developmen-
tal abnormalities and often acute or lethal morphological aberrations
(Brown et al., 2010). These aberrant cellular and nuclear structures might
have been caused by changes in lipid layer viscosity, protein-lipid phase
distribution-interaction, and enhanced lipid peroxidation in erythrocytes
(Islam et al., 2021a; Meena et al., 2014). Cell metabolism is hampered by
salinity, and cell membranes can be damaged. Through homeo-viscous ad-
aptation, salinity alters the protein-lipid interaction phase and the fatty acid
content in the erythrocytic cellular-nuclear membrane (Gracey et al., 2004;
Islam et al., 2020a). Thus, cellular and nuclear damage of erythrocytes
might be caused by enhanced lipid peroxidation in erythrocytes of fish ex-
posed to stressful salinity (Jahan et al., 2019; Klein et al., 2017; Martínez-
Álvarez et al., 2005; Roche and Bogé, 2000). Salinity stress has also been re-
ported to change metabolites. Blood examination in cobia Rachycentron
canadum found that fish kept at a salinity of 5 ppt had considerably lower
plasma protein levels than fish acclimatized at higher salinities (Denson
et al., 2007).
3.4. Dissolved oxygen (DO)/hypoxia
Dissolved oxygen (DO) is required for all aerobic organisms, including
fish, to support metabolic activity and biological functions such as respira-
tion (Maulu et al., 2021). Stress-induced by low DO in water has negative
impacts on overall fish performances (Dagoudo et al., 2021). Acceptable
performance and production of fish are dependent on the presence of suffi-
cient DO concentrations in the water (Araújo-Luna et al., 2018; Christensen
et al., 2018). Hypoxia occurs in aquatic habitats when the DO concentration
goes below the levels necessary to support the organisms in the water
(Delaney and Klesius, 2004; Lefevre et al., 2014). Hypoxia has been defined
as DO concentration < 2.0–3.0 mg/L for fish (Chen et al., 2007; Lundgreen
et al., 2008; Nivelle et al., 2019), although some aquatic organisms are far
more sensitive to hypoxia than others. Hypoxia can occur naturally or as a
result of anthropogenic activities in both marine and freshwater habitats
(Friedrich et al., 2014). Hypoxia is an established cause of stress reactions
in fish (Bernier et al., 2012; Segner et al., 2012; Welker et al., 2007). Despite
the fact that most fish have some degree of hypoxia tolerance, mortality oc-
curs commonly after exposure to severe hypoxia (Cook and Herbert, 2012;
Fitzgibbon et al., 2007). Fish increase oxygen intake in response to hypoxia
through a complex combination of physiological, behavioral, biochemical,
and molecular pathways (Guderley, 2004; Sørensen et al., 2014). Both
acute and chronic hypoxic stress can have a detrimental impact on the
fish's physiological performance (Table 5). Interestingly, in response to hyp-
oxia, fish attempt to maintain oxygen supply by adjusting RBC and Hb
levels (Jensen, 2004; Lefevre et al., 2014; Lundgreen et al., 2008). As a re-
sult, increasing Hct restores RBC function to normal levels of viscosity and
enhances oxygen transfer via Hb (Wells et al., 2005). Hb is a red protein in
the blood that transports oxygen. The considerable increase in Hb levels
after 12 h of acute hypoxia may be regarded as energetically cost-
effective. As reported in several studies, in fish during hypoxia exposure,
hematological parameters rise briefly before dropping again. In conclusion,
the time necessary to replenish fresh Hb production is regulated by the de-
gree of hypoxia as well as the species (Tran-Duy et al., 2008). In addition,
WBCs play a role in disease diagnosis and defense in response to tissue dam-
age, changes in cellular immunity, and foreign invaders (Qiang et al.,
2013). The immune system of yellow catfish is damaged by hypoxia stress,
as demonstrated by substantial reductions in WBC at 96 h. A similar pattern
M. Shahjahan et al. Science of the Total Environment 843 (2022) 156910

Table 5
Effect of DO/hypoxia on hemato-biochemical parameters of fish.
Species Oxygen Exposure time (h) Hemato-biochemical alterations References

Increase Decrease

Pelteobagrus fulvidraco × P. vachelli 1.5 mg/L 96 Hct, Hb, Glu, TC, TG, ALT, AST RBC, WBC, TP and gly Dagoudo et al., 2021
Oreochromis niloticus 0.64 mg/L 72 RBC, Hb, Ht WBC, lysozyme Sheng et al., 2019
Micropterus salmoides 1.20 mg/L 24 Glu – Yang et al., 2019
Polyodon spathula 59 mmHg 36 mmHg 72 Hb, RBC, Hct, Glu, Lactate Na+, Cl− Aboagye and Allen, 2018
Megalobrama amblycephala 3.5 and 1.0 mg/L 6 Hb, Glu, Lactate – Chen et al., 2017
Acipenser schrenckii 3 mg/L 6 Glu, cortisol, RBC, WBC, Hb, NEUT – Ni et al., 2014
Pseudosciaena crocea 1.5 mg/L 96 RBC, Hb, Ht – Xiaolian and Zhaoli, 2011
Pelteobagrus fulvidraco 1.5 mg/L 24 WBC, RBC, Hb, Hct – Xiaolian and Zhaoli, 2011
Oncorhynchus mykiss 25 % 1.5 Lactate – Omlin and Weber, 2010
Chiloscyllium punctatum
anoxia 12 Hb, RBC, Hct MCHC Chapman and Renshaw, 2009
Hemiscyllium ocellatum
Cyprinus carpio
Sarotherodon mossambicus 40 % 21 days – ATP Pollock et al., 2007
Salmo gairdneri
Oncorhynchys mykiss 2.57 mg/L 28 Hct Landman et al., 2006
Acipenser oxyrinchus
30 mmHg 1 Glu, Cortisol, Lactate – Baker et al., 2005
Acipenser brevirostrum
Scorpaena porcus 3 Hct, Glu, Na+ K+
Autogenic hypoxia Silkin and Silkina, 2005
Diplodus annularis 3 Hct, Glu, K+
Poecillia latipinna 1.0 mg/L 42 days Hb and RBC – Timmerman and Chapman, 2004
3.52 mg/L 2 Glu, Lactate –
Labeo rohita 2.46 mg/L 2 Lactate Glycogen Padmavathy et al., 2003
0.48 mg/L 6 Glu, Lactate –
Gymnotus carapo 1.0 mg/L 3 Hb, RBC, Glu, Lactate – Moraes et al., 2002
Hemiscyllium ocellatum 1.0 mg/L 4–5 Lactate – Routley et al., 2002

ATP; Adenosine triphosphate, Glu; glucose, Hb; hemoglobin, Hct; hematocrit, RBC; red blood cell, WBC; white blood cell, GPT; Glutamic-Pyruvic Transaminase, and NEUT;
Neutrophiic granulocytes.

of response has been reported in Nile tilapia O. niloticus exposed to hypoxia
at 0.64 mg/L for 48 to 72 h (Sheng et al., 2019).
Stress raises catecholamine levels, leading to glycogenolysis and gluco-
neogenesis, producing a compensatory increase in blood glucose levels
(Qiang et al., 2016). Under hypoxic stress, fish initially utilize glucose and
triglycerides as energy sources rather than serum protein, which decreases
later. Investigations on sockeye salmon (Oncorhynchus nerka) and yellow
croaker (Larimichthys crocea) revealed that proteins become the main
source of metabolic energy during acute hypoxia once other energy sources
have become depleted. The decrease in TP suggests that protein is being
used as a fuel for energy (Ding et al., 2020; Mommsen et al., 2011). LDH,
a key biomarker of anaerobic metabolism, rose considerably during hyp-
oxia. Significant increases in LDH were reported in Nile tilapia and yellow
catfish subjected to hypoxia at 0.64 mg/L during the first hours before a
progressive decrease (Sheng et al., 2019). In another study, Xiaolian and
Zhaoli (2011) reported a marked increase in LDH in yellow croakers
while exposed to hypoxia for 96 h.
An increase in the hepatic activity of protein-metabolizing enzymes re-
ported in fish treated with hypoxic stress might be related to the usage of
excess hydrocarbons from amino acids to fulfill energy needs, as well as
liver deterioration (Abdel-Tawwab et al., 2019). Serum glucose, total cho-
lesterol, triglyceride, ALT, and AST levels were shown to rise in the hypoxia
group, whereas total protein levels remained stable (Abdel-Tawwab et al.,
2019). AST and ALT are commonly used as hepatocellular injury markers
or as indicators of vertebrate liver activity (Pan et al., 2010). The levels of
AST and ALT in yellow catfish Pelteobagrus fulvidraco increased signifi-
cantly, indicating damage or a weakening of normal hepatic function.
Circulating cortisol levels are also useful as indicators of hypoxia stress.
Cortisol levels, for example, rose over 48 h before settling at 96 h, showing
the persistence of stress on the fish. GIFT tilapia subjected to hypoxia at
0.64 mg/L exhibited comparable results (Sheng et al., 2019). Juvenile
Amur sturgeon Acipenser schrenckii demonstrated a constant rise in cortisol
when exposed to hypoxia (Ni et al., 2014). In an adverse environment,
glycogen metabolism is a vital mechanism for the organism's energy
acquisition (Bacca et al., 2005). In hypoxia, carbohydrate metabolism is
crucial for energy provision. According to existing studies, boosting cortisol
activity during hypoxia can increase hepatic glycogen breakdown and
provide more glucose for energy demands. Large yellow croaker reported
a total glycogen reduction of 2.0 mg/L after 96 h of hypoxia exposure
(Ding et al., 2020). At 1.20 mg/L, a similar outcome was obtained in
largemouth bass M. salmoides (Yang et al., 2019).
3.5. pH/acidification
Water pH regulates fish physiological responses, and hemato-
biochemical measures (Table 6); are key physiological markers for
assessing fish health and stress status subjected to pH changes (Kim et al.,
2020a, 2020b, 2020c). Low pH stress increases H+ inflow into the fish
blood via the gills, and suppression of metabolically produced H+ ions
and CO2 excretion induces acidemia, limiting hemoglobin's capacity to
carry oxygen (Fromm, 1980). Low pH promotes hematological problems
owing to gill ion-regulatory abnormalities produced by ion loss, according
to (Costa et al., 2015). Furthermore, an increase in H+ ion inflow might
promote plasma acidosis, spleen contraction, and erythrocyte enlargement.
In contrast, high water pH decreases the fish's ability to transport oxygen
from the blood to tissues (Aride et al., 2007). A pH over 9.5 produces ele-
vated blood pH and ammonia content in the plasma, which can result in
fish mortality (Ghanbari, 2012). pH changes affect physiological and bio-
chemical processes in fish, and a considerable shift in pH induces unique re-
active responses in aerobic metabolism, culminating in oxidative stress
(Eissa and Wang, 2013; Kim et al., 2020a, 2020b). Excessive ROS formation
produced by pH variations affects the DNA, proteins, and lipids of aquatic
species, leading to toxicity and the development of hereditary illnesses
(Eissa and Wang, 2013; Kim et al., 2020a, 2020b). ROS are hypothesized
to induce Ca2+ influx from the extracellular environment into the cyto-
plasm, and an increase in cytoplasmic Ca2+ levels results in chromatin
damage and cell death (Ermak and Davies, 2002). In fish, antioxidant re-
sponses are an important defensive mechanism for avoiding ROS-induced
DNA damage produced by oxidative stress (Hegde et al., 2008). Antioxidant
responses in fish are an important defensive mechanism for avoiding ROS-
induced DNA damage caused by oxidative stress (Sun et al., 2019). pH
stress lowers fish immunity via influences on cellular or humoral immune
responses (Li and Chen, 2008). Tilapia mossambicus (currently known as
Oreochromis mossambicus) treated to acidic pH 5.5 and alkaline pH 9.5

8
M. Shahjahan et al. Science of the Total Environment 843 (2022) 156910

Table 6
Effects of pH on hemato-biochemical parameters of fish.
Species pH Exposure time (days) Hemato-biochemical alterations References

Increase Decrease

Seriola lalandi 6.58 27 Glu Lactate Abbink et al., 2012

Oreochromis niloticus 5.30 1.5 Antioxidant enzyme, ROS, intracellular Ca+ Blood cells Mai et al., 2010
Litopenaeus vannamei 5.60 & 9.30 1 DNA damage in hemocytes, Ca+ in hemocytes – Wang et al., 2009
Prochilodus scrofa 4.50 & 8.00 30 Hct, RBC, Hb MCHC Carvalho and Begnis, 2006
Tribolodon hakonensis 3.50 2–7 – Plasma Na+ Hirata et al., 2003
Salmo salar 5.70 25 Cortisol, Hct Plasma Cl−, ATPase activity Fivelstad et al., 2003
Colossoma macropomum 3.00 3 Cortisol, Glu, TP Plasma Na+ Wood et al., 1998
Oncorhynchys mykiss 4.50 7 Cortisol Plasma Na+ Kakizawa et al., 1996
Enneacanthus obesus 3.50 1 Hct – Gonzalez and Dunson, 1987
Perca fluviatilis 3.80–4.50 3 Hct Plasma Na+ Rask and Virtanen, 1986
Coregonus clupeaformis 4.10 14 Cortisol, Glu Plasma Na+ Scherer et al., 1986
Salmo trutta 6.00 42 – Hct, plasma Na+, Cl− McWilliams, 1980

ATP; Adenosine triphosphate, Glu; glucose, Hb; hemoglobin, Hct; hematocrit, RBC; red blood cell, ROS; reactive oxygen species, WBC; white blood cell, TP; total protein.

revealed a large drop in RBC count, Hb, and Hct, as well as a significant rise
in WBC, monocytes, and neutrophil count (Arain and Rauf, 2013).
4. Effects of environmental contaminants on hemato-biochemical pa-
rameters of fish
4.1. Pesticides
Pesticides (insecticides, herbicides and fungicides) are used worldwide
to control and eradicate pests in intensive agricultural production, includ-
ing aquaculture farms (Oruç, 2010; Rahman et al., 2020; Saravanan et al.,
2011; Shahjahan et al., 2021). Effects of various insecticides, herbicides
and fungicides on hemato-biochemical parameters of fish are summarized
in Table 7. Production of pesticides is dependent in part on the use of
phenol-derived chemicals, and these chemicals ultimately accumulate in
the aquatic environment (Michałowicz and Duda, 2007; Bojarski and
Witeska, 2020). Phenol exposure resulted in a significant increase in Hct,
Hb, and RBC in pacu Piaractus mesopotamicus (Clauss et al., 2008). This
rise might be ascribed to spleen constriction or erythropoiesis (Moraes
et al., 2015). Octylphenol is a widely-used surfactant phenolic compound
that can contaminate the aquatic environments (Ying et al., 2002).
Octylphenol has been reported to cause hematological stress in acara
Cichlasoma dimerus, resulting in increased RBC, Hb, and MCV levels, possi-
bly leading to anemia (Rey Vázquez and Lo Nostro, 2014). Common carp
juveniles' continuous exposure to phenoxy acid causes a decrease in RBC al-
though the WBC count trend was opposite (Lutnicka et al., 2018a, b), indi-
cating that this herbicide may be causing an inflammatory response and
immunosuppression. Similarly, in juvenile common and white carp,
cypermethrin induces a considerable drop in RBC, Hb, and PCV, with a
varying propensity for WBC (Neelima et al., 2015, 2016). Quinalphos, a
highly toxic organophosphate classified as a yellow label pesticide that
has caused concern, is another synthetic pesticide widely used in agricul-
ture (Das and Mukherjee, 2000).
Chlorpyrifos, an organophosphate insecticide commonly used in agri-
cultural, residential, and commercial settings, is very hazardous to fish
(Biswas, 2018; Deb and Das, 2013; Sunanda et al., 2016). Chlorpyrifos ex-
posure has been found to significantly reduce RBC, Hb, Hct, and leucocyte
counts in dose- and time-dependent manners, causing physiological issues
such as leukemia and immune dysfunction in the spotted snakehead
Channa punctatus (Ali and Kumar, 2012; Malla et al., 2009). Clomazone,
an agricultural herbicide, has also been shown to alter hematological pa-
rameters in juvenile streaked prochilod Prochilodus lineatus in a dose-
dependent manner. Calmozone appears to induce a considerable decrease
in Hct, Hb, HCM, and MCHC, implying an anemic condition. However, an
increase in RBC was found at low doses, which might be attributable to
the release of new red blood cells into the circulation as a result of splenic
constriction in response to the stressor agent (Pereira et al., 2013). Common
carp treated to sub-acute concentrations of either fipronil or buprofezin, or
both exhibited substantial decreases in RBC, TC, Hct, and Hb but an
increase in WBC (Qureshi et al., 2016). The curzate® 60DF (Cymoxanil
60 %) fungicide appears to be capable of generating a gradual and consid-
erable decrease in RBC, Hb, RVC, and MCHC in Mozambique tilapia while
increasing WBC, MCV, and MCH (Desai and Parikh, 2012). Similarly, in the
African catfish (Clarius gariepinus), the two pesticides, chlorpyrifos and
DDFORCE cause a significant decrease in RBC, PVC, and Hb while increas-
ing WBC and MCHC (Adewumi et al., 2018). Juvenile African catfish ex-
posed to glyphosate herbicides exhibited significantly higher RBC, MCV,
MCH, PLT, and WBC levels when compared to controls (Alohan et al.,
2014). This might be related to the activation of defensive systems
(Cazenave et al., 2005). Saravanan et al. (2011) found that lindane had
comparable effects to those of furadon on hematological parameters in
the same species, as did (Islam et al., 2022). Hb, Hct, RBC, MCV, MCH,
and MCHC levels were lower in the exposed fish, whereas WBC levels
were higher. When exposed to lindane, the morari Aspidoparia morar
showed a considerable drop in RBC, Hb, Hct, and WBC, with varying values
for MCV, MCH, and MCHC (Sachar and S.R, 2014).
Bisphenol-A, a phenolic compound classified as a potent endocrine
disruptor, has been shown to have toxic effects on fish physiology when de-
tected in water environments by sewage discharges (Faheem and Lone,
2013; Liu et al., 2011, 2014). Bisphenol-A significantly decreased the he-
matological parameters Hb, Hct, MCV, and MCH while increasing the
WBC value in rohu Labeo rohita (Kaliappan et al., 2017). The observed
drop in hematological values might be due to a reduction in the rate of
erythrocyte production, destruction, and/or anemia in the fish caused by
Bisphenol-A toxicity (Jenkins et al., 2003; Seth and Saxena, 2003). A de-
crease in RBC content may cause the development of a hypoxic situation,
which may accelerate RBC destruction, or a decrease in RBC production
rate owing to a lack of Hb content (Chen et al., 2004). Such a decrease in
hematological parameters indicated anemia, which was most likely caused
by hemolysis and erythrocyte destruction, resulting in a decrease in O2 car-
rying capacity by blood and impaired gas exchange (Jenkins et al., 2003;
Seth and Saxena, 2003). The increase in WBC levels may indicate that the
fish can develop a defensive mechanism to combat toxic stress during the
exposure period (Desai and Parikh, 2012). The increase in WBC count sug-
gests immune stimulation against Bisphenol-A toxicity, as seen in yellow
perch Perca flavescens (Rogers and Mirza, 2013).
4.2. Heavy metals
Heavy metals contamination in aquatic ecosystems has become a rising
ecological and worldwide problem due to the danger of toxicity and bioac-
cumulation in fish (Shahjahan et al., 2022; Taslima et al., 2022). The gen-
eral process of heavy metals toxicity in fish is described in Fig. 2. Effects
of various heavy metals on hemato-biochemical parameters of fish are sum-
marized in Table 8. African catfish Clarias gariepinus had higher RBC, Hb,
Hct, and WBC values with elevated concentrations of copper (Cu), iron
(Fe), lead (Pb), cadmium (Cd), manganese (Mn), and zinc (Zn) originated
from high sewage discharge compared to individuals of the same species

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M. Shahjahan et al. Science of the Total Environment 843 (2022) 156910

Table 7
Effect of pesticides (insecticides, herbicides and fungicides) on hemato-biochemical parameters of fish.
Species Pesticides Concentration Exposure Hemato-biochemical alterations References
time
Increase Decrease
(days)

Insecticides
Chlorfenapyr,
Cirrhinus mrigala dimethoate, 3 ppb 30 WBC, MCV, MCH Hb, RBC, PCV Ghayyur et al., 2021
acetamiprid
Oreochromis
Sumithion 2.579 mg/L 4 WBC, Glu Hb, RBC Sharmin et al., 2021
niloticus
Heteropneustes
Envoy 0.015, 0.0075 mg/L 4 – RBC Akter et al., 2020
fossilis
Clarias gariepinus Diazinon 0.73–1.46 mg/L 30 RBC, Hb, Hct, Glu, AST, ALT WBC Al-Otaibi et al., 2019
Rhamdia quelen Fipronil 300–400 μg/L 4 AST, ALT, ALP, GGT Hct, PLT Fredianelli et al., 2019
TP, Glob, Cholesterol, TG, ALP,
Cyprinus carpio Chlorpyrifos 25–50 μg/L 21 Glu, AST, ALD, LDH Hatami et al., 2019
CK, GGT
Pangasianodon
Sumithion 5.886 mg/L 4 WBC, Glu Hb, RBC Islam et al., 2019b
hypophthalmus
Danio rerio Sumithion 3.58 mg/L 4 Glu Hb Shahjahan et al., 2019
Hypophthalmichthys MCV, MCH, WBC, K, Cholesterol, RBC, Hb, Hct, MCHC, Na, Cl,
Deltamethrin 2 μg/L 4 Ullah et al., 2019
molitrix Cortisol PO34, TP, Alb
Brycon amazonicus Cypermethrin 1.8 μg/L 4 RBC, Hb, Ht, Glu, Na, Cl de Moraes et al., 2018
Labeo rohita Chlorpyrifos 73.8–221.4 4 WBC RBC, Hb, Hct Ismail et al., 2018
RBC, Hb, Hct, MCV, MCH,
Labeo rohita Cypermethrin 0.5–1.5 4 WBC, PLT, Glu Khan et al., 2018
MCHC
Ctenopharyngodon Swarna Kumari et al.,
Dichlorvos 9.8 mg/L 4 WBC RBC, Hb, Hct
idella 2018
Kalita and Khatun,
Clarias magur Dimethoate 6.5–13 mg/L 7 WBC RBC, Hb
2018
Carassius gibelio Trichlorfon 0.5–2.0 μg/L 4 CAT, SOD – Lu et al., 2018
Oreochromis
Chlorpyrifos 15–75 μg/L 14 Hct, MCV, WBC, Neu, Lym RBC Zahran et al., 2018
niloticus
Kalaimani and
Labeo rohita Phosalone 2.5 mg/L 4 Glu Hb
Kandeepan, 2017
Glu, TP, Alb, Glob, RBC, Hb,
Clarias batrachus Chlorpyrifos 1.65 mg/L 15 – Narra et al., 2017
Hct
MCHC, WBC, Neu, Eos, Bas, Glu, RBC, Hb, Hct, MCV, Lym, TP,
Clarias batrachus Dimethoate 1.245 mg/L 56 Narra, 2017
Creat, AST, GTP, Lys, Perox Alb, Glob, NBT, IgM
Clarias batrachus Sumithion 5.25 mg/L 1 – Hb, RBC and Hct Naqvi, 2017
Labeo rohita Profenofos 10 μg/L 16 WBC RBC, Hb, Hct, MCV, MCH Nagaraju et al., 2017
Malathion
Cirrhinus mrigala Dimethoate 0.1–1.0 μg/L 60 Na, K Ca, Cl Rani et al., 2017
Chlorpyrifos
RBC, Hb, Hct, WBC, MCH, MCHC,
Clarias gariepinus DDT 155–310 μg/L 30 TP, Ca Ahmad et al., 2016
Glu, AST, ALT
Danio rerio Sumithion 0.5, 1.0 & 2.0 mg/L 7 Glu Hb Ahmed et al., 2016
Barbus sharpeyi Diazinon 1.0 mg/L 21 – RBC, Hb, Hct, WBC Alishahi et al., 2016
Ashaduzzaman et al.,
Channa punctatus Carbofuran 1–10 mg/L 4 Neu Hb, WBC, Mono, PLT
2016
Ctenopharyngodon
Azadirachtin 73–219 μg/L 30 – TP, Alb, AST, ALT, ALP, LDH Gholami et al., 2016
idella
Chanos chanos Endosulfan 18.5–22.5 μg/L 4 Glu, Cortisol NBT Kumar et al., 2016
RBC, Hb, Hct, PLT, Glu, TP,
Cyprinus carpio Fipronil 400 μg/L 4 WBC
Glob Qureshi et al., 2016
Cyprinus carpio Buprofezin 100 mg/L 4 WBC, Glu RBC, Hct, PLT
Cyprinus carpio Malathion 1.5, 3.0 mg/L 4 WBC, Glu RBC, Hb Sharmin et al., 2016
Bhanu and Deepak,
Cyprinus carpio Cypermethrin 200 μg/L 28 Glu TP, Alb, Glob
2015
RBC, Hb, Hct, MCV, MCH, Binukumari et al.,
Cirrhinus mrigala Endosulfan 400 μg/L 3 WBC, Lym, Eos
MCHC, Neu, PLT 2015
Cirrhinus mrigala Deltamethrin 0.8 mg/L 15 MCH, WBC RBC, Hb, Ht David et al., 2015
Catla catla Dimethoate 20.5–24 mg/L 2 Glu Gly, TP, Trigl Hussain et al., 2015
Cyprinus carpio Sumithion 0.85–1.70 mg/L 21 WBC, Glu Hb, RBC Hossain et al., 2015
Cyprinus carpio Sumithion 0.8, 1.6, 3.2 mg/L 4 Glu, MCV and MCH Hb, RBC and Hct Salam et al., 2015
Cyprinus carpio Azadirachtin 20–60 μg/L 4 – Hb, Hct, MCH, MCHC Murussi et al., 2016
Tor putitora Cypermethrin 63 μg/L 4 WBC RBC Ullah et al., 2015
Cyprinus carpio Monocrotophos 4.4–13.5 mg/L 30 WBC RBC, Hb, Hct Vaiyanan et al., 2015
Barbonymus
Quinalphos 0.47–0.97 mg/L 28 MCV, MCHMCHC, WBC, Glu RBC, Hb, Hct, Glu Mostakim et al., 2015
gonionotus
Capoeta damascina Malathion 0.76–1.52 mg/L 10 MCV, MCH RBC, Hb, Hct, WBC Shahbazi et al., 2015
Oreochromis Abu Zeid and Khalil,
Sumithion 4.7 mg/L 4 DNA, TM, SOD, MDA, ALT, Cortisol GSH and IgM
niloticus 2014
Heteropneustes
Sumithion 0.24 mg/L 240 Glu RBC, WBC and Hb Ali et al., 2014
fossilis
RBC, Hb, Hct, MCV, MCH,
Cirrhinus mrigala Diazinon 0.815, 1.63 mg/L 30 Glu, AST, ALT Haider and Rauf, 2014
WBC, TP, Alb, Glob, LDH
Labeo rohita Imidacloprid 120 mg/L 8 Glu RBC, Hb, Hct, MCV, PLT, TP, Qadir et al., 2014

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Table 7 (continued)

Species Pesticides Concentration Exposure Hemato-biochemical alterations References

time
Increase Decrease
(days)

Alb, TG, AST, ALT

Cyprinus carpio Diazinon 5–80 mg/L 4 MCV, MCH, Neu RBC, Hb, Hct, WBC, Neu Al-Ghanim, 2013
Cirrhinus mrigala Diazinon 8.15 mg/L 4 MCV, MCH RBC, Hb, Hct, WBC Rauf and Arain, 2013
Rhamdia quelen Deltamethrin 0.1–1.5 mg/L 4 WBC AST, ALT, FA Galeb et al., 2013
RBC, Hb, Hct, MCV, MCH,
Cyprinus carpio Chlorpyrifos 40–80 μg/L 10 WBC Yonar et al., 2012
MCHC
Catla catla Cypermethrin 0.443 μg/L 60 – RBC, Hb, Hct, WBC, TP, Alb Vani et al., 2012
Amin and Hashem,
Clarias gariepinus Deltamethrin 0.75 μg/L 2 AST, ALT TP, Alb
2012
RBC, Hb, WBC, TP, Alb, Glob,
Channa punctatus Endosulfan 1.2 μg/L 90 – Sarma et al., 2012
NBT
Nandan and Nimila,
Etroplus maculatus Lindane 1–5 μg/L 30 WBC RBC, Hb, Hct, MCV, MCH
2012
RBC, Hb, Hct, MCV, MCH. Chowdhury and Haq,
Clarias batrachus Malathion 8.8 mg/L 21 –
MCHC, WBC 2012
Channa punctatus Malathion 0.8 mg/L 4 WBC RBC, Hb Dube, 2012
Cyprinus carpio Propoxur 5 mg/L 4 WBC, TP, Na, Cl, AST RBC, PLT Gül et al., 2012
WBC, Hb, Hct, MCV, MCH, MCHC, RBC, Hb, Hct, MCV, MCH.
Cyprinus carpio Lindane 38 μg/L 1 Saravanan et al., 2011
Glu MCHC, TP
Prochilodus lineatus Endosulfan 1.2–2.4 μg/L 4 WBC, PLT, Mono, Glu Hb, MCH, Lym, Neu, TP Bacchetta et al., 2011
Cichlasoma dimerus Endosulfan 2–4 μg/L 4 – MCV, MCH, MCHC Da Cuña et al., 2011
Catla catla Deltamethrin 1.61 μg/L 45 – RBC, WBC, Hb, TP, Alb, Glob Vani et al., 2011
Oncorhynchus
Diazinon 0.1, 0.2 mg/L 28 Glu, CK, AST, ALT TP, Alb, Glob Banaee et al., 2011
mykiss
Oncorhynchus RBC, Hb, Ht, MCV, MCH, WBC,
Dimethoate 73.5–735 μg/L 30 Glu Dogan and Can, 2011
mykiss TP
Sepici-Dinçel et al.,
Cyprinus carpio Cyfluthrin 10 μg/L 7 MCV, Cortisol RBC, WBC, PLT
2009
Oncorhynchus
Bifenthrin 1.47 μg/L 4 Glu, LDH, CK, ALP MCV, MCH, Neu, NH3 Velisek et al., 2009
mykiss
Clarias gariepinus Diazinon 15 μg/L 4 Neu RBC, Hb, Ht, WBC Adedeji et al., 2009
Maduenho and
Prochilodus lineatus Diflubenzuron 25 mg/L 4 Glu RBC, Hb
Martinez, 2008
Hb, MCHC, Na, K, P, Creat, Glu,
Rhamdia quelen Cypermethrin 80–120 μg/L 8 TP, TG Borges et al., 2007
Chol, AST, ALP, ALT
Oreochromis RBC, WBC, Hb, Hct, Lym, Chol,
Deltamethrin 15 μg/L 4 Neu, TP, Alb, Glu El-Sayed et al., 2007
niloticus AST, ALT, ALP
Ancistrus multispinis Deltamethrin 0.1–0.31 μg/L 4 RBC, WBC, Hb Pimpão et al., 2007
Monopterus albus Endosulfan 0.05–0.7 μg/L 4 – RBC, Hb, Ht, WBC Hii et al., 2007
Sepici-Dinçel et al.,
Cyprinus carpio Sumithion 10 mg/L 2 Cortisol, Glu Hct, RBC, plasma Cl, Na
2007
Salmo salar Endosulfan 500 μg/L 35 Hb, Hct, MCH Petri et al., 2006
Oncorhynchus
Cypermethrin 3.14 μg/L 4 AST, LDH, CK Neu Velisek et al., 2006
mykiss
RBC, Hb, Ht, MCV, MCH,
Channa punctatus Monocrotophos 1.86 mg/L 60 Neu, Mono, Eos Agrahari et al., 2006
MCHC, WBC, Lym, Bas, PLT
Oreochromis
Monocrotophos 1.15 μg/L 30 ALT, AST, ACP, ALP Rao, 2006
mossambicus
Sebastes schlegeli Cypermethrin 0.08 μg/L 56 Glu, Bil, AST, ALT Hb, TP, Alb, Chol, ALP, Lys Jee et al., 2005
Clarias RBC, Hb, Hct, MCV, MCH,
Actellic 0.3–1.0 μg/L 18 WBC Mgbenka et al., 2005
albopunctatus MCHC
Salvelinus alpinus Toxaphene 10 μg/L 104 – RBC Blanar et al., 2005
Chandrasekara and
Cyprinus carpio Trichlorfon 0.5 mg/L 1 Neu, Mono WBC, Lym
Pathiratne, 2005
Venkataramana et al.,
Glossogobius giuris Malathion 0.5 mg/L 4 Urea Glu
2005
Labeo rohita Cypermethrin 0.16–0.8 μg/L 28 MCV, MCH, WBC RBC, Hb Adhikari et al., 2004
Tilapia guineensis Chlorpyrifos 0.6–5 μg/L 56 – RBC, WBC Chindah et al., 2005
Das and Mukherjee,
Labeo rohita Cypermethrin 3–14 μg/L 45 WBC, Glu TP
2003
Cyprinus carpio Deltamethrin 3.25 μg/L 4 – RBC, Hb, Ht, TP Svobodová et al., 2003
Channa punctatus Fenvalerate 0.15–0.4 mg/L 30 WBC RBC, Hb, Ht Seth and Saxena, 2003
Oncorhynchus Atamanalp et al.,
Cypermethrin 1.0–4.1 μg/L 15 Na PO3−
4
mykiss 2002a
Oncorhynchus Atamanalp et al.,
Cypermethrin 1.0–4.1 μg/L 21 RBC, Hb, MCH, MCHC, PLT MCV, WBC
mykiss 2002b
Channa punctatus Cypermethrin 0.1–0.35 mg/L 30 – RBC, Hct, Hb, TP Saxena and Seth, 2002
Cyprinus carpio Diazinon 19.5 mg/L 4 Neu RBC, Hb, Hct, WBC, Lym Svoboda et al., 2001

Herbicides
Cyprinus carpio Linuron 0.0315 mg/L 14 RBC, Hct, MCV, WBC, A, NA, Cort Hct, Lym Lutnicka et al., 2019
Oreochromis Hb, RBC, Hct, MCV, MCHC,
Penoxsulam 0.90, 1.79 mg/L 30 – Galal et al., 2018
niloticus WBC, Neu, Lym, Lys, IgM
Cyprinus carpio Glyphosate 0.02 mg/L 7 Hct, Chol, TG WBC, NBT Kondera et al., 2018

(continued on next page)

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M. Shahjahan et al. Science of the Total Environment 843 (2022) 156910

Table 7 (continued)

Species Pesticides Concentration Exposure Hemato-biochemical alterations References

time
Increase Decrease
(days)

Cyprinus carpio Pendimethalin 0.0025 mg/L 14 RBC, WBC, Mono, A, NA, Cort Hct, MCV, Lym, Neu Lutnicka et al., 2018a
Carassius auratus Prometryn 2.5 4 – Hct, LDH Mosiichuk et al., 2018
Anabas testudinius Glyphosate 12.21 30 MCV, MCH, WBC, PLT RBC, Hb, Hct, Lym Samanta et al., 2019
Heteropneustes
Glyphosate 12.21 mg/L 30 Ht, MCV, MCH, MCHC, WBC, PLT RBC, Hb Samanta et al., 2019
fossilis
Oreochromis
Glyphosate 12.21 30 MCV, MCH, MCHC, WBC, PLT RBC, Hb, Hct Samanta et al., 2019
niloticus
Oreochromis Abubakar et al., 2018;
Propanil 0.44. 0.87 mg/L 56 WBC, Lym, Neu, TP, GTP, TG RBC, Hb, Hct, Chol
niloticus Yaji et al., 2018
0.00188–0.00191 + Golombieski et al.,
Rhamdia quelen Imazapyr + Imazapic 4 Mono, AST MCV, Lym
0.00072–0.00074 mg/L 2016
Cyprinus carpio Ethofumesate 1.1 mg/L 7 Hb, MCHC, Lym Neu Bojarski et al., 2015
Felix and Saradhamani,
Catla catla Glyphosate 0.19 mg/L 4 MCHC, WBC MCV, MCH
2015
Labeo rohita Butachlor 0.75, 1.0 mg/L 4 MCV, WBC, Lym, Mono RBC, Hb, Hct, MCHC, Neu Ghaffar et al., 2015
Oncorhynchus Ahmadivand et al.,
Butachlor 0.39 mg/L 10 Neu Hb, RBC, WBC, Lym
mykiss 2014
Cyprinus carpio Atrazine 0.02–0.03 mg/L 4 Glu, ALT, AST, ALP, LDH Hb, Hct, WBC, Lym, Lact, Blahova et al., 2014
Oreochromis
Pendimethalin 2.5 mg/L 4 TP, Glob, Glu Hct, MCV, MCH, WBC El-Sayed et al., 2015
niloticus
Carassius auratus 2,4-D 10, 100 mg/L 4 TP, LDH Lym Kubrak et al., 2013
Hb, MCV, WBC, LCT, Lym, TP,
Cyprinus carpio Metribuzin 175.1 mg/L 4 Mono, Neu, Bas, Glu, NH3, Ca2+ Velisek et al., 2013
ALB, Glob, TG, Lact
Cyprinus carpio Prometryn 0.008–0.08 60 Hb, MCH, MCHC, Glu, Lact, ALT AST, ALT, Creat Velisek et al., 2013
Cyprinus carpio Simazine 0.00006–0.004 mg/L 90 – WBC, ALT, ALP Velisek et al., 2012
S-metolachlor +
Cyprinus carpio 3.9 + 2.26 4 Glu, NH3, AST, LDH Ht, WBC, P, Cl-, TG Dobšíková et al., 2012
terbuthylazine
Ctenopharyngodon Abdali et al., 2011;
Atrazine 0.008–8 mg/L 30 – TP, Alb, Glu, Chol, TG
idella Soorena et al., 2011
Cyprinus carpio Terbutryn 0.0002–0.002 mg/L 90 RBC, MCHC, Glu, AST, LDH Creat, Mg Velisek et al., 2011
Leporinus obtusidens Glyphosate 0.48, 2.4 90 – RBC, Hb, Hct, TP Salbego et al., 2010
Cyprinus carpio Terbutryn 0.002–0.04 mg/L 28 RBC, NH3, AST, LDH, CK, Lact MCV, MCH, Creat Velisek et al., 2010
Anguilla anguilla Molinate 11.15 4 – RBC, Hb, Hct, WBC, TP, ChE Sancho et al., 2000

Fungicides
Danio rerio Boscalid 0.1–1.0 mg/L 28 – Glu Qian et al., 2019
Oreochromis RBC, Hb, Hct, MCV, MCH,
Paclobutrazol 1.18, 2.36 mg/L 28 – Ghane et al., 2017
mossambicus MCHC, WBC
Ctenopharyngodon Hb, RBC, Hct, MCHC, WBC, Mohammadalikhani
Captan 0.26–0.68 mg/L 4 MCV, MCH, Neu, Mono
idella Lym et al., 2017
Labeo rohita Propiconazole 0.89 μL/L 28 MCV, MCHC, Glu, RBC, Hb, Hct, MCHC, WBC, TP Hemalatha et al., 2016
Cyprinus carpio Mancozeb Prochloraz 1.0 mg/L 14 WBC, Neu RBC, Hb, Hct, MCHC, Lym Lutnicka et al., 2016
Cyprinus carpio Isoprothiolane 0.0027–0.027 mg/L 4 WBC, Alb RBC, Hb, Hct, Glu, TG, Chol Saravanan et al., 2015
Oreochromis
Copper-oxychloride 32.3 mg/L 90 ALT, AST, UA, Creat Hb, Hct Hassaan et al., 2014
niloticus
Oncorhynchus
Propiconazole 0.5 mg/L 30 MCH, Glu, TP, CK, LDH, ALT, AST RBC, Hb, Hct, WBC Li et al., 2011
mykiss

A; adrenaline, ACP; acidic phosphatase, ALP; alkaline phosphatase, ALT; alanine transaminase, AST; aspartate transaminase, Bas; basophils, ChE; cholinesterase, Chol; cho-
lesterol, CK; creatine kinase, Cort; cortisol, Creat; creatinine, Eos; eosinophils, Glob; globulin, Glu; glucose, GTP; phosphoglycerate kinase, Hb; hemoglobin, Hct; hematocrit,
IgM; immunoglobulin M, LDH; lactate dehydrogenase, Lym; lymphocytes, Lys; lysozyme, MCV; mean cell volume, MCH; mean corpuscular hemoglobin, MCHC; mean cor-
puscular hemoglobin concentration, Mg; magnesium, Mono; monocytes, NA; noradrenaline, NBT; oxidative metabolic activity of phagocytes, NO; nitric oxide, P; phosphorus,
Neu; neutrophils, PLT; thrombocyte count, RBC; red blood cell, TG; triglyceride, TP; total protein, UA; uric acid, WBC; white blood cell.

Zn Pb
As
Ni
Heavy
Cd metals Hg
Cr Fe
Cu
Use reserved energy (protein,
Feed supplement lipid, carbohydrate) to cope
up with the stress

Bioaccumulation in different
Food chain tissues/organs
Fish under
Plankton stress

Fig. 2. Routes of heavy metal accumulation in different tissues / organs in fish and created stress.

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Table 8
Effects of heavy metals on hemato-biochemical parameters of fish.
Species Heavy metals Concentration Exposure Hemato-biochemical alterations References
(mg/L) time
Increase Decrease
(days)

Pangasianodon
Cr 0.8, 1.6, & 3.2 30 WBCs Hb, RBC, and Glu Suchana et al., 2021
hypophthalmus
Pichhode and Gaherwal,
Clarias batrachus As 84 4 – TP
2020
Pangasianodon 10, 20, 30, 40, 50
Cr 4 WBC, MCV, MCH, Glu, ECA, ENA RBC, Hb, PCV Islam et al., 2020a
hypophthalmus & 60
Cd 0.316
Ni 0.302
Clarias gariepinus – AST, ALP, ALT, Cort, Glu, MCH CK, TLC, MCV Banday et al., 2019
Cr 0.203
Cu 0.073
Clarias gariepinus Cu 100 4 TP, Albumin, globulin Javed and Usmani, 2019
Cd 0.001
Channa striata Cr 0.001 – HDL, LDL, TP, AST and ALT Glu Phoonaploy et al., 2019
Pb 0.005
Fe 9.00
Mn 2.50
Zn 0.54
LDL, TP, lipid and chol., SOD, CAT, Albumin, triglyceride, HDL and
Channa punctatus Co 0.26 – Javed et al., 2017
GST and LPO VLDL
Ni 0.10
Cu 0.08
Cr 0.08
Fe 8.71
Cu 0.86
Zn 0.30
Channa punctatus Mn 0.21 – WBC RBC, Hb, Javed and Usmani, 2014
Ni 0.12
Co 0.11
Cr 0.10
Tilapia mossambica As 21 WBC, MCH, MCHC Hb, RBC, PCV, Soundararajan et al., 2014
Hct, Hb, RBC, WBC, MCH, and
Cyprinus carpio, Cr 2 28 Glu Abedi et al., 2013
MCHC
Hb, MCV, MCH, MCHC, Hct,
Tilapia mossambicus Zn 1.0, 2.5, 5.0 14 RBC, lymphocyte, WBC Çelik et al., 2013
lysozyme & MPO
Fe 8.71
Cu 0.86
Zn 0.30
Mastacembelus armatus Mn 0.21 – Glu – Javed and Usmani, 2013
Ni 0.12
Co 0.11
Cr 0.10
7, 14, 21, Parthipan and Muniyan,
Cirrhinus mrigala Ni MCHC RBC, WBC, Hb, Ht, MCV
28 2013
Cd 0.826
Zn 1.46
Labeo rohita – WBC RBC, Hct, Chandanshive et al., 2012
Pb 0.756
Hg 0.0987
RBC, Hb, PCV, MCH, MCHC, Glu,
Cyprinus carpio Cr(vi) 25–150 180 WBC, MCV, AST, ALT, ALP, ESR, AP Shaheen and Akhtar, 2012
protein, chol.
4
Colisa fasciatus Cd 1.86 – Protein, gly, nucleic acid Tripathi et al., 2012
30
RBC, WBC, Hct and Hb, MCV, MCH,
Cyprinus carpio Ni 6,9,12,15,18 4 – Al-Ghanim, 2011
MCHC
Zn 5.0
Oreochromis niloticus 7 & 14 ALT, Cort., Glu and TP cholesterol levels Firat and Kargin, 2010
Cd 1.0
RBC, WBC, Hct, Hb, MCV, MCH and
Clarias gariepinus Ni 4, 6, 8, 10, 12 4 – Ololade and Oginni, 2010
MCHC
0.01, 0.02, 0.04,
Acanthopagrus lates Hg 21 Hb, Hct, monocyte WBC, lympho, eosinophil Safahieh et al., 2010
0.08
Cd + Pb + Cr Rajamanickam and
Cyprinus carpio 5.0 32 Glu, TC, serum iron and copper Hb, Hct, RBC
+ Ni Narayanan, 2009
Clarias batrachus Hg 0.6 35 WBC RBC, Hb Maheswaran et al., 2008
Channa punctatus Cu 0.36 15, 30, 45 WBC, MCV, MCHC, MCH Hb, RBC, PCV Singh et al., 2008
Catla catla Cd 1 to 8 4–7 Glu Glycogen, TP Sobha et al., 2010
Cu 0.87
Zn 0.7
Pb 3.6 Vosylienė and Jankaitė,
Oncorhynchus mykiss 4 – RBC
Ni 0.47 2006
Cr 0.367
Mn 14.0
Cyprinus carpio Cd 0.05–1.0 10 Glu Glycogen Cicik and Engin, 2005
Labeo rohita Cr + 6 39.40 4 – RBC, Hb, MCH, lipids, TP Vutukuru, 2005
Cyprinus carpio Pb 10 4 Hb, Hct, neutrophil WBC, leukocyte Witeska, 2005

(continued on next page)

13
M. Shahjahan et al.
Table 8 (continued)
Species Heavy metals Concentration Exposure Hemato-biochemical alterations
(mg/L) time
Increase
(days)
Cu 5
Cd 10
or Zn 20
Prochilodus lineatus Pb 95 4 Glu
Cu 0.0075
Zn 0.064
Ni 0.0021
Oncorhynchus mykiss Cr 0.0028 28 – RBC, Hb, Hct, WBC
Pb 0.0142
Cd 0.00018
Mn 0.0099
Hct, RBC, Hb, WBC, neutrophils,
Prochilodus scrofa Cu 0.029 4 Lymphocytes, plasma Na+ and Cl−
plasma K+
ALP; alkaline phosphatase, ALT; alanine transaminase, AST; aspartate transaminase, GST; glutathione S transferase, Glu; glucose, Hb; hemoglobin, Hct; hematocrit, HDL; high
density lipoprotein, LPO; lipid peroxidation, RBC; red blood cell, TP; total protein, WBC; white blood cell, VLDL; very low density lipoprotein.
exposed to less sewage discharge (Gaber et al., 2013). Common carp ex-
posed to Pb, Cu, Cd, and Zn exhibited a rise in Hct without a substantial
change in RBC, as well as an initial increase in WBC that later fell to stay
low (Witeska, 2005). Aluminum (Al) is one of the most prevalent metals
on the globe, emitting both natural and manmade contaminants into the en-
vironment while serving no proven biological uses (Sjögren et al., 2007).
The hematological indices RBC, Hct, Hb, MCHC, MCH, and MCV in adult
tropical freshwater fish redbelly tilapia Coptodon zillii rose significantly
after Al exposure. In fish, these parameters progressively rose with increas-
ing Al concentration and time exposure, indicating that they may constitute
a protective mechanism against aluminum toxicity via erythropoiesis acti-
vation (Alwan et al., 2009). Mn, an important trace metal, is frequently
used in industry, and its waste is discharged into water bodies, where it de-
grades into an unrecognizable hazardous metal that disrupts organisms'
physiological equilibrium. After an acute Mn exposure, changes in blood
cells were found in goldfish, demonstrated a drop in leucocytes, weakening
the immune system, while an erythron profile revealed an increase in cellu-
lar and nuclear RBC alteration, leading to eryptosis, limiting the blood
oxygen-carrying capacity and therefore the fish health condition (Aliko
et al., 2018). A declining trend in RBC, Hb, and Hct was seen in the sucker
head G. gotyla, followed by Mn exposure. Because of Mn toxicity, hemopoi-
etic organs may be impacted and become unable to discharge normal RBC
into general circulation, resulting in a significant decrease in erythrocyte
life span and delayed erythropoiesis (Sharma and Langer, 2017). Mercury
(Hg) is a prominent and pervasive aquatic contaminant that bacteria can
convert into a more hazardous form (Schrope, 2001). As demonstrated in
the tench Tinca tinca, Hg may permeate the erythrocyte membrane, produc-
ing cell destruction and hemorrhages, as well as raised Hct, Hb, and RBC
levels in both acute and chronic exposure. These increases might be the re-
sult of splenic constriction (a frequent stress reaction) and subsequent re-
lease of blood cell reserves, or they could be the result of
contemporaneous erythropoiesis in response to an O2-CO2 transport re-
quirement (Shah and Altindağ, 2004). In the African catfish C. gariepinus,
Hg exposure produces a dose-dependent reduction in RBC and Hb
(Maheswaran et al., 2008). Cd and Pb are two other elements that can
have an effect on hematological variables, reducing the amounts of RBC,
WBC, and TC in the blood of flathead grey mullet (Mugil cephalus; Fazio
et al., 2015). Internal bleeding from kidney injury induced by Cd and Pb,
as well as decreased osmoregulation caused by Cd, may all contribute to
the drop in RBC content (Faggio et al., 2014; Kori-Siakpere et al., 2006).
While a reduction in WBC in striped mullet blood appears to be associated
with Cd and Pb bioaccumulation in the kidney and liver (Kori-Siakpere
et al., 2006), weakening the fish's immune system and making them
disease-prone. Common carp exposed to Pb for a short period displayed var-
iations in RBC, Hb, Hct, and MVC. However, this was followed by a contin-
uous rise in the proportion of erythrocyte abnormalities (Shah and
Altindağ, 2004).
14
Science of the Total Environment 843 (2022) 156910
References
Decrease
Lipid, plasma Na+, TP, Cholesterol Martinez et al., 2004
Vosyliene et al., 2003
Mazon et al., 2002
Transient leukopenia was also detected, weakening the fish's immune
system (Witeska, 2013). Furthermore, fish exposed to Cd exhibited reduced
RBC, Hb, and Hct levels (Al-Asgah et al., 2015). The drop in these values
might be due to the loss of mature RBC or an acute hemolytic crisis, either
of which would result in severe anemia (Faggio et al., 2014). Arsenic is
ranked among the most toxic elements, especially in its salt form. The
African catfish C. batrachus demonstrated a gradual decline in Hb, RBC,
and PVC when exposed to arsenic salt, leading to anemia (Kumar and
Banerjee, 2016). Ni exposure cause hematological abnormalities in goldfish
C. auratus, with substantial reductions in WBC, Hct, and Hb followed by in-
creases in MCH and RBC (Moosavi and Shamushaki, 2014). The drop in
WBC might be attributed to the production of adrenaline during stress,
which can promote splenic constriction. A reduction in Hct and Hb suggests
severe anemia induced by Ni in the water (Vinodhini and Narayanan,
2009). Several mechanisms may be associated with the advancement of
anemia, either through a decrease in RBC generation or an increase in
RBC loss (Shah and Altindağ, 2004).
4.3. Microplastics (MPs)
Microplastics (MPs) are plastic molecules derived from large plastic
components used in a range of mechanical and photo oxidative procedures
which are <5 mm in size (Arthur et al., 2009). Large amounts of non-
degradable and hazardous solid waste have accumulated in the oceans
and in other aquatic ecosystems due to extensive uses of plastics, which
are being discarded and are accumulating at a severely increasing rate
over the recent decades (Selvam et al., 2021). Several studies reported
the harmful effects of MPs, including physiological injuries, cellular toxicity
etc. in different fish species in the field and laboratory (Hossain et al., 2019;
Savoca et al., 2019; Strungaru et al., 2019). Effects of MPs on hemato-
biochemical parameters of fish are summarized in Table 9.
Decline of Hb, RBC and Hct may be attributed to hem-dilution of blood
due to hemolysis caused by MPs toxicity (Sharma and Langer, 2017). Sim-
ilarly, negative effect of MPs on lymphoid tissues may be the cause of de-
creases of WBC in the exposed fish (Alkaladi et al., 2015). Significant
decrease of alanine-amino-transferase (ALT) - an important serum marker
for hepatic damage - might be due to hepatic system disruption by MPs
(Solter et al., 2000; Oliveira et al., 2013). On the other hand, changes in cre-
atinine and uric acid indicated the dysfunction of kidney as these are used
as indices of the glomerular filtration rate, thereby serving as biomarkers of
kidney dysfunction (Abdel-Khalek et al., 2016). Various enzymes (ALT,
AST, ALP, LDH etc.) are present in cells of different organs and regarded
as indicators of cell membrane damage (Kumar et al., 2016). Therefore,
changes in different enzymes due MPs exposure have been interpreted as
indications that MPs potentially damaged plasma membranes in different
organs (Nematdoost Haghi and Banaee, 2017).
M. Shahjahan et al. Science of the Total Environment 843 (2022) 156910

Table 9
Effects of microplastics (MPs) on hemato-biochemical parameters of fish.
Species MPs Concentration Exposure Hemato-biochemical alterations References
time
Increase Decrease
(days)

Ctenopharyngodon
PS 0, 100, 1000 μg/L 21 – SOD, CAT, Liu et al., 2022
idella
Oncorhynchus 0.5, 2.0, 5.0 % in
PE 42 Hct Leukocytes, ALT, Lactate Hodkovicova et al., 2021
mykiss feed
Raw AST, ALT, Eos, Mono, Neu, RBC, Hb, Lymphocytes, Alb, Glob, SOD, TAC,
Clarias gariepinus 500 mg/kg diet 15 Sayed et al., 2021
powder TP, Glu GST, CAT
Danio rerio PS 0, 20, 100 μg/L 21 Hepatic pyruvate TG, Chol, Glu, ICD Zhao et al., 2020
0, 0.5 %, 1.5 %, 3.0
Clarias gariepinus PVC 45 MCH, MCHC RBC, WBC, Hb, Neu, MCV Iheanacho and Odo, 2020
%
Oreochromis Raw
0, 1, 10, 100 mg/L 15 ROS, AST, ALT, ALP, Glu, TP RBC, Hb, Hct, MCHC, WBC Hamed et al., 2019, 2020
niloticus powder
Cyprinus carpio PE 0, 250, 500 μg/L 30 Glu, Chol, TP, ALT, ALP GGT, AChE Banaee et al., 2019
0, 100, 500 mg/kg
Sparus aurata PVC 30 AST, CK, Alb, Glob Glu, Espinosa et al., 2017
diet
Nematdoost Haghi and Banaee,
Cyprinus carpio PE 1–2 mg/L 01 Glu, AST, ALP, CPK TP, Glob, Chol, TG
2017
Cyprinus carpio PE 0, 1, 2 mg/L 21 ALP, CPK, Creat Alb Haghi and Banaee, 2017
Clarias gariepinus LDPE 50, 500 mg/L 04 Alb:Glob ratio Chol, HDL, Glob Karami et al., 2016
Pomatoschistus
PE 0, 18.4, 184 μg/L 04 – AChE, IDH Oliveira et al., 2013
microps

AChE; Acetylcholinesterase, AST; aspartate aminotransferase, ALT; alanine aminotransferase, ALP; alkaline phosphatase, CPK; creatine phosphokinase, Creat; creatinine,
ChE; cholinesterase, Chol; cholesterol, CK; creatine kinase, Cort; cortisol, Eos; eosinophils, Glu; glucose, GGT; Gamma glutamyl-transferase, Glob; globulin, Hb; hemoglobin,
Hct; hematocrit, ICD; isocitrate dehydrogenase, LDPE; low density polyethylene, Neu; neutrophils, PE; polyethylene, PVC; poly vinyl chloride, PS; polystyrene, RBC; red blood
cell, ROS; reactive oxygen species, TG; Triglyceride, TP; total protein, WBC; white blood cell.

5. Conclusions and future prospects
Blood-based biomarkers play important roles in our understanding of fish
physiology and stress response status. Blood analyses provide a comprehen-
sive representation of physiological and functional status of fish under
various environmental and experimental treatment conditions (Fig. 3). Ex-
amining the landscape scale and inter-annual effects of environmental varia-
tion on fish health and physiological fitness at both the individual and
population levels is a complex and difficult task due to the limited ability
to track and monitor physiological states and responses over space and
time. A large number of variable parameters must be considered when inter-
preting fish blood biomarkers, including the reproductive cycle, age, sex,
feeding activity, stress, nutritional condition, and water quality. This is espe-
cially important for migratory and highly mobile fish. In addition, the lack of
reference ranges for the many aquatic species, the difficulties of minimizing
stress in the subjects, and the analytical instrumental difficulties must be
taken into consideration. Understanding how fish respond and adjust to
changing environments requires tracking fish and their responses to ecolog-
ically appropriate conditions. To comprehend the efficacy of blood based
Temperature
Cortisol Light Hypoxia
Increases
biomarkers in fish, it is necessary to conduct more species-specific studies
under a range of environmentally realistic conditions. In addition, variations
in fish sampling methods, transportation, culture systems, acclimation condi-
tions, and water quality should all be considered. Other variables such as fish
handling, blood sample collection, and storage time may all have a signifi-
cant impact on the blood biomarker-based results, so measurements should
be performed as soon as possible after blood collection. However, along
with hemato-biochemical parameters, employing other cellular and molecu-
lar parameters as an integrative approach could increase the degree of cer-
tainty in the interpretation physiological responses of fish.
It is also critical to select a broader range of climate stressors and some
model fish to predict unexpected outcomes under various climate scenarios.
More research is needed on non-lethal endpoints for tracing stress events in
fish. There have been no studies that investigate fish blood biomarker re-
sponses at different stages in the life cycle, variable culture conditions, and
feeding regimes using a comparative-collective approach. Therefore, it is
also essential to examine the effects of single environmental and anthropo-
genic factors species-by-species covering responses among physiological, im-
munological and molecular mechanisms. More studies are needed to clarify
the specific molecular mechanisms aiding a single factor or a combination
of factors to interact with organisms and causing DNA damage in different
fish, with genomic and transcriptomics technologies. However, this review
creates a basis for the development of hemato-biochemical sensors for instant
Protein checking of the occurrence of pesticides, heavy metals and microplastics in
the fish body. Moreover, it is needed to explore the bioremediation of
Decreases

Salinity pH
pesticides, heavy metals and microplastics toxicity and accumulation in the
Affected fish fish body. Finally this review illustrates the necessity for formulation or
Glucose Hb legislation of the laws for launching an industry with proper discharging
Microplastics protocols, considering their disturbing effects on fish and other aquatic
biota, and could also help to determine safe levels of pesticides, heavy metals
Pesticides Heavy metals and microplastics in the aquatic environment.
Increases

WBC RBC

CRediT authorship contribution statement

Erythrocytic cellular and nuclear abnormalities Md Shahjahan: conceptualization and edited the manuscript. Md Jakiul
Islam: preparation of the first draft of the manuscript. Md Tahmeed
Fig. 3. Schematic illustrations of the effects of different environmental parameters Hossain, Moshiul Alam Mishu: data collection and preparation of the Ta-
on hemato-biochemical parameters of fish. bles. Jabed Hasan formatted references in the manuscript. Christopher

15
M. Shahjahan et al.
Brown edited the manuscript. All authors have read the final version and
approved the manuscript.
Declaration of competing interest
The authors declare that they have no known competing financial inter-
ests or personal relationships that could have appeared to influence the
work reported in this paper.
Acknowledgments
This research was funded by the Aqua-drugs initiative of the Bangladesh
Fisheries Research Institute and the University Grants Commission of
Bangladesh (2018/666/UGC), Grants for Advanced Research in Education,
Ministry of Education, Government of the People's Republic of Bangladesh
(2017/503/MoE) and Bangladesh Agricultural University Research System
(2017/282/BAU & 2019/812/BAU) to the corresponding author (Md
Shahjahan) which are gratefully acknowledged.
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