International Journal of Livestock Research ISSN 2277-1964 ONLINE www.ijlr.

org

Vol 3(2) May13

Physiological, Biochemical and Molecular Responses to Thermal Stress in Goats

Mahesh Gupta1*, Sachin Kumar2, S.S. Dangi3, Babu Lal Jangir4
Indian Veterinary Research Institute, Izatnagar, UP, India
1 & 3- Ph. D Scholar, Physiology & Climatology Division;
2- Ph. D Scholar, Animal Nutrition Division;
4- Ph. D. Scholar Pathology Division
* Corresponding Author : mahesh_bwm@rediffmail.com
Rec.Date: Sep 09, 2012 11:05; Accept Date: May 02, 2013 08:11

Abstract
Livestock undergo various kinds of stress such as physical, nutritional, chemical, psychological and
thermal stress. Among all thermal stress is most concerning now a day in the ever changing climatic
scenario. Thermal stress redistributes the body resources including protein and energy at the cost of
decreased growth, reproduction, production and health. Goats (Capra hircus) are relatively resistant to
harsh environmental conditions. Thermal stress stimulates sort of complex responses which are
fundamentals in the preservation of cell survival. Physiological responses to thermal stress are rectal
temperature, respiration rate, heart rate, skin temperature; hormonal changes in terms of thyroxin, triiodothyronine, cortisol, electrolyte concentration; hematological responses like hemoglobin level and
packed cell volume. Biochemical responses include changes in antioxidants level, plasma or serum
enzymes and metabolites like blood glucose, total cholesterol level. At molecular level changes in gene
expression of heat shock proteins and many other proteins ensure protections. Altogether these
physiological, biochemical and molecular responses makes the goats to survive in harsh environment.
Key words: Goat, Thermal stress, thyroxin, serum enzymes, heat shock proteins.
Introduction
Globally goat plays an important role in the economy of thousands poor livestock owners who earn their
livelihood by rearing them in different terrain and climatic condition. In the developing countries, goats
make a very valuable contribution, especially to the poor in the rural areas. Goat rearing is a traditional
occupation of small, marginal farmers and landless laborers in semiarid, arid and hilly and mountain
regions of developing countries, inhospitable to conventional crop cultivation. They produce food and
fiber at relatively low cost from feed materials from land that often cannot be used for conventional crop
production. The various methods of utility of goat render the animal to be labeled as a poor mans cow.
Perhaps, it is the only livestock that fits well for effective utilization in the diverse socio-economic

1990). Domestic animals undergo various kinds of stress such as physical, nutritional, chemical,
psychological and thermal stress. Among all, thermal stress is most concerning now a day in the ever

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Stress is reaction of body to stimuli that disturb homeostasis often with detrimental effects (David et al.,

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situations of the developing countries.

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changing climatic scenario. In tropical and sub tropical regions high ambient temperature is the major
constraint on animal production (Marai et al., 2007; Nardone et al., 2010) whereas extreme low
temperature in temperate regions is also detrimental to livestock. Thermal stress is the perceived
discomfort and physiological strain associated with an exposure to an extreme hot or cold environment.
Thermal stress includes both heat stress, during extreme summer season as well as cold stress, during
extreme winter season. High environmental temperature is the major concern in tropical and arid areas
whereas at the same time very low environmental temperature in temperate areas is also lethal that
challenges the animals ability to maintain energy, thermal, water, hormonal and mineral balance
Silanikove, 1992) Temperature determines metabolic rates, heart rates and other important factors within
the bodies of animals, so an extreme temperature change can easily distress the animal body. The effect of
high temperature is further aggravated when heat stress is accompanied by high ambient humidity.
It has been consistently shown in that goats perform better than other domesticated ruminants in harsh
environments (Shkolnik et al., 1981; Devendra, 1990). According to Shkolnik et al. (1972) the
adaptability of goats to thermal stress is due to water conservation capability, higher sweating rate, lower
basal metabolism rate (BMR), higher respiration rate, higher skin temperature, constant heart rate and
cardiac output (Shkolnik et al., 1972) whereas small body size, low metabolic requirement, ability to
reduce metabolism, efficiency of utilization of high forage diet i.e. high digestive efficiency, ability to
economise nitrogen requirement, efficient use of water etc. are also important chacteristics which help
them to cope up with harsh environmental conditions (Silanikove, 2000).
Although goats are resistant to thermal stress at a greater extent but they suffer from heat and cold stress
beyond their comfort zone, which is environmental temperature 13-270C for Indian goats (Mishra, 2009).
Browsing of goats to open fields during most of the day hours makes them susceptible to environmental
stress.
The purpose of the present review is to provide an integrative explanation of thermoregulatory responses
of goat at cellular, physiological, biochemical and molecular level during thermal stress.
Adverse effects of thermal stress
Thermal stress affects almost all systems of body. Heat stress reduces libido by reducing level of
testosterone, sperm output, decreasing sperm motility and by increasing up proportion of morphologically

conception rate and embryonic survival in animals (Edwards and Hansen, 1997; Sakatani et al., 2004). It

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impair follicular and oocyte development and reduces steroid production (Zeron et al., 2001; Ozawa et al.,

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abnormal spermatozoa in the ejaculate (Perez-Crespo et al., 2008). In female, it lowers fertility,

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2005). Primary effect of environmental stress in neonates is increased incidence of disease associated
with decrease in immunity by reducing immunoglobulin content in plasma. It also reduces fetal growth
and alters endocrine status of the dam. Carryover effects of heat stress during late gestation on postpartum
lactation and reproduction are also detectable (Collier, 1982).
Various kinds of stress including heat and cold stress leads to the production of reactive oxygen species
(ROS) such as superoxide, peroxide, hydroxyl radical, singlet oxygen etc. ROS are also generated during
normal body functions (Boveris and Chance, 1973). Although, low levels of ROS are essential for many
biochemical processes but their accumulation due to over production or a decreased antioxidant defense
mechanism causes oxidative stress which leads to damage of biomolecules viz. DNA/RNA, proteins and
lipid peroxidation of membranes and disruption of normal cell metabolism (Spurlock and Savage, 1993).
Heat stress may lead to over production of transition metal ions (TMI), which can make electron
donations to oxygen forming superoxide or H2O2 which is further reduced to an extremely reactive
OH radical causing oxidative stress (Agarwal and Prabhakaran, 2005).
At the molecular and cellular levels, temperature beyond the comfort zone reduces the rates of enzymatic
reactions, diffusion, transport and induce the denaturation and misaggregation of proteins. It also slows
down progression through cell cycle, inhibit transcription, translation, disrupt cellular cytoskeleton
elements and change membrane permeability (Fujita, 1999; Sonna et al., 2002).
Body defense mechanism against stress
Physiological Mechanism
Respiration rate, pulsation rate and rectal temperature are the parameters which illustrate the mechanism
of physiological adaptation. Several researchers studied physiological adaptation mechanisms such as
rectal temperature, pulse rate and respiration rate in small ruminants (Sevi et al., 2001; Srikandakumar et
al., 2003; Maurya et al., 2004; Marai et al., 2007; Otoikhian et al., 2009; Phulia et al., 2010, Sharma et al.
2013). Body temperature is good measure of heat tolerance in animals. It represents the resultant of all
heat gain and heat loss processes of the body. Rectal temperature is considered as a good index of body
temperature even though there is a considerable variation in different parts of the body core at different
times of the day (Srikandakumar et al., 2003). Rectal temperature of goats was found to be elevated with

77.33 respectively when goats were kept for 6 hours in hot ambient temperature in summer. Pulsation
and respiration rate per minute was found to be increased by the effect of environmental temperature.

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(2010) reported increase in rectal temperature and respiration rate from 38.970C and 43.66 to 39.350C and

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high environmental temperature in several studies (Devendra, 1987; Marai et al., 2007). Phulia et al.,

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Increased respiration is an attempt to increase heat loss by evaporative cooling. Devendra (1987) reported
that changes of metabolism and muscle activity of goats also changes pulsation and respiration rates. Heat
loss via high respiration rate was reported as higher than that via other ways (Devendra, 1987). Increase
in heart rate and pulse rate is attributed to two causes. One is the increase in muscular activity controlling
the rate of respiration, concurrent with elevated respiration rate. The second is the reduction in resistance
of peripheral vascular beds and arteriovenous anastomoses. Increase in pulsation rate increases blood ow
from the core to the surface as a result of it more heat is lost by sensible (loss by conduction, convention
and radiation) and insensible (loss by diffusion water from the skin) means (Marai et al., 2007). The
increase in cardiac output and cutaneous blood flow by heat stress, due to blood redistribution from deep
splanchnic to more peripheral body regions, have been implicated in goat [Silanikove, 1987 & 2000)
Blight (1985) reported that a daily change in respiration rate per minute from the effect of environmental
temperature may not be parallel with change in body temperature and pulsation number. Higher values of
means of these parameters (RR, RT, HR) have been reported than that of values in thermoneutral zone
(Mc Dowell and Woodward, 1982; Al-Tamimi, 2007).
Hormonal changes
Hormones involved in thermal adaptations include viz prolactin, growth hormone, thyroxine,
glucocorticoids, mineralocorticoids, catecholamines and antidiuretic hormone. These are either involved
with nutrient partitioning and homeorhesis or for homeostatic regulation, augmented by thermal stressors.
Hormonal changes are mediated through
sympatho-adrenal

medullary

the hypothalamo-pituitary-adrenal cortical axis (HPA) and

axis (Minton, 1994). Sivakumar et al. (2010) reported that there

was

increase in plasma concentration of prolactin from 11.73 to 26.37 g/L and cortisol from 25.27 to 40.57
nmol/L in heat stressed goats whereas T3 and T4 level decreased from 4.55 and 21.27 to 3.21 and 16.70
pmol/L respectively. There is decline in T3 and T4 levels during short and long long term exposure to
solar radiation in goats (Helal et al., 2010). Thyroid function declines as an acclimation response to
alleviate heat stress. This reduction may be due to effect of heat on HPA to decrease in thyrotropin
releasing hormone which enable animal to reduce basal metabolism (Johnson, 1987). Similarly T3 and T4
level increases during cold stress resulting in increase in oxygen consumption and heat production by
cells to increase BMR (Todini, 2007; Ocak et al., 2009; Bernabucci et al., (2010). Relationship among
plasma aldosterone level and urine electrolyte concentration have been documented.

Prolonged heat exposure decreased plasma aldosterone level was as well as significant fall in serum
and urinary K+ (El-Nouty, et al., 1980; Kumar et al., 2011).
Haematological parameters

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stress,

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thermal

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Thermal stress is known to alter the homeostatic mechanism of animals resulting in impaired
erythropoiesis. High environmental temperature increases oxygen consumption of animals by increasing
respiration rate. The higher oxygen intake increases the partial pressure of oxygen in blood, decreases
erythropoiesis which in turn reduces the number of circulating erythrocytes and thus PCV and Hb values
(Sivakumar et al., 2010; Maurya et al., 2007;Temizel et al., 2009; Kumar et al., 2011a). Other explanation
of decrease in hemoglobin and PCV levels during thermal stress could be increased attack of free radicals
on the erythrocyte membrane, which is rich in lipid content, and ultimate lysis of RBC or inadequate
nutrient availability for hemoglobin synthesis as the animal consumes less feed or decreases voluntary
intake under heat stress. During summer stress a significant depression in PCV may also be due to
haemodilution effect where more water is transported into the circulatory system for evaporative cooling
(EL-Nouty et al., 1990).
Biochemical Changes
Antioxidants: Body has antioxidants in the form of enzymatic (viz. superoxide dismutase, glutathione
peroxidase & catalase) and nonenzymatic (vitamins C, E and A, glutathione, pyruvate etc.) They provide
protection against ROS generated due to thermal stress. Catalase detoxifies H 2O2 produced
during different metabolic processes and also in stressful conditions by reducing it to H2O and O2
(Fridovich, 1978). SOD along with catalase and glutathione peroxidase (GPx) scavenges both
intracellular and extracellular superoxide radicals and prevents lipid peroxidation (Agarwal and
Prabhakaran, 2005). Kumar et al. (2011b) reported increase in SOD level in goats during heat stress. GPx
reacts with peroxides and requires glutathione (GSH) as the reductive

substance

donating

an

electron. Glutathione reduces oxygen toxicity by preventing O2- formation. Vitamin C and vitamin E are
water-soluble antioxidants in mammalian tissues and biological fluids which are needed in high
concentrations to protect the cells from the oxidants produced by respiratory burst (Weiss et al., 2004).
Vitamin E is a free radical scavenger on the cell membrane. It protects biological membranes from
oxidative degeneration and makes up an integral part of the enzyme GPx of the antioxidant system.
Vitamin C is a water soluble, extra-cellular, natural antioxidant and is involved in a number of oxidation
and reduction reactions in the body. Concentration of vitamin C and vitamin E lowered during heat stress
in goats (Sivakumar et al., 2010; Kumar et al., 2007; Kumar et al., 2011a).

differences in hot conditions than in the comfort zone. Some researchers reported that hot climatic

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affected by high ambient temperatures. Blood glucose and total serum cholesterol levels show greater

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Blood glucose and total serum cholesterol levels are physiological adaptation mechanisms that can be

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conditions decrease blood glucose and total serum cholesterol levels (Marai et al., 1995; Ocak et al.,
2009) and some reports are contrary to these (Webster, 1976). Bahga et al. (2009) and Ocak et al. (2010)
reported that blood glucose and total cholesterol level decreases during summer season and increases
during winter season in goats. Determination of blood parameters may be important in establishing the
effect of heat stress. The marked decrease in total serum cholesterol levels may have a relation with the
increase in total body water or the decrease in acetate concentration which is the primary precursor for the
synthesis of cholesterol.
Signicant decrease in total protein concentration and increase in HSP concentrations in goats have been
reported during heat stress (Dangi et al., 2012). The total plasma protein, albumin, globulin decreased
from 6.56, 2.53, 4.04 to 5.88, 2.08 and 3.80 g/dl respectively in balady goats subjected to short term heat
stress for two days (Helal et al., 2010).This may be due to increase in plasma volume as a result of heat
shock which causes results in decreases plasma protein concentration. Prolong exposure of solar
radiations increased plasma total protein, albumin, and globulin. This might be due to vasoconstriction
and decreased plasma volume during heat stress (Helal et al., 2010).
Enzymes Activity: Metabolic regulators are important in elucidating a picture of modulation in
physiological mechanisms during stressed conditions and are best assessed by determining the enzymes
governing various metabolic reactions in plasma or serum. The level of these enzymes reflects the
metabolic activities during stress.

Heat stress lowers the alkaline phosphatase (ALP) and lactic

dehydrogease (LDH) activity (Sevi et al., 2001; Helal et al., 2010). Decrease in these enzymes during heat
stress is due to decrease in thyroid activity during heat stress (Helal et al., 2010). Serum level of aspartate
transaminase (AST) and alanine tranaminase (ALT) is helpful in diagnosis of welfare of animals. Serum
ALT value found to be increased during heat stress in goats (Sharma. and Kataria, 2011). No significant
changes were observed in AST level in goats during heat stress (Ocak et al., 2009; Sharma and Kataria,
2011).
Molecular mechanism: It is widely accepted that changes in gene expression are an integral part of the
cellular response to thermal stress. Although the HSPs are perhaps the best-studied examples of genes
whose expression is affected by heat shock, it has become apparent in recent years that thermal stress also
leads to induction of a substantial number of genes not traditionally considered to be HSPs. Some of these
genes are affected by a wide variety of different stressors and probably represent a nonspecic cellular

mammalian cells, nonlethal heat shock produces changes in gene expression and in the activity of

32

expressed proteins, resulting in what is referred to as a cell stress response. This response

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response to stress, whereas others may eventually found to be specic to certain types of stress. In

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Vol 3(2) May13

characteristically includes an increase in thermotolerance (i.e., the ability to survive subsequent, more
severe heat stresses) that is temporally associated with increased expression of HSPs. Thermal-induced
changes in gene expression occur both during hyperthermia as well as hypothermia. About 50 genes not
traditionally considered HSPs have been found to undergo changes in expression during or after heat
stress. Many of these genes will likely to be proved as important mediators and effectors of the cell stress
response.

HSP: Expression of many HSPs including HSP32, HSP40, HSP60, HSP70, HSP90, HSP110 and many
others are found to be increased during hyperthermic stress (Patir and Upadhyay, 2010; Gade et al, 2010;
Dangi et al., 2012, Sharma et al., 2013 )
Functions of HSPs
HSP acts as molecular chaperones by participating in the assembly of proteins without being part of the
final protein structure (Ellis, 1987). HSPs contribute to cell survival by reducing the accumulation of
damaged or abnormal polypeptides within cells (Parsell and Lindquist, 1993). They possess crucial role in
intracellular transport, the maintenance of proteins in an inactive form and the prevention of protein
degradation (Neuer et al., 2000). They possess important protective role in vivo as they can protect heart
and brain against ischaemia and lungs and liver against sepsis. HSPs play crucial roles in cell-cycle
control, signaling and protection of cells against apoptosis (Zihai and Pramod, 2003). During the period
of hyperthermia and shortly thereafter, HSPs become the predominant proteins synthesized by cells
(Lindquist, 1986). Interestingly, most HSP genes lack introns (Lindquist, 1986), which may facilitate
their rapid expression and which may also help explain how they can be expressed in the presence of
stressors (such as heat) that can interfere with RNA splicing.
HSPs possess three principal biochemical activities:1) Chaperonin activity -HSPs with this function help prevent misaggregation of denatured proteins and
assist the refolding of denatured proteins back into native conformations.
2) Regulation of cellular redox state eg. HSP32
3) Regulation of protein turnover (Parsell and Lindquist, 1993). An example is ubiquitin, which is
expressed in unstressed cells, upregulated by heat shock and serves as a molecular tag to mark proteins
for degradation by proteasomes.
HSP60 is mostly found in the mitochondria. It helps in refolding of proteins and prevents aggregation of

nucleus. Its main functions are protein translocation and regulation of steroid hormone receptors.
Ubiquitin is a small HSP of 8.5 kDa and found in cytosol and perform selective degredation of proteins.

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folding, cytoprotection and as molecular chaperones. HSP90 is mostly found in the cytosol, ER and

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denatured proteins. HSP70 is mostly found in the cytosol and nucleus. Its functions are nascent protein

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Mechanism of HSP expression during stress

Heat shock factors (HSFs), present in the cytosol, are bound by heat shock proteins (HSPs) and
maintained in an inactive state. A broad array of physiological stimuli (stressors) are thought to activate
HSFs, causing them to separate from HSPs. HSFs are phosphorylated by protein kinases and translocate
into nucleus. In the nucleus trimerization occurs. These HSF trimer complexes bind to heat shock
elements (HSE) in the promoter region of the HSP gene. HSP mRNA is then transcribed and leaves the
nucleus for the cytosol, where new HSP is synthesized. Proposed mechanisms of cellular protection for
HSPs include their functioning as molecular chaperones to assist in the assembly and translocation of
newly synthesized proteins within the cell and the repair and refolding of damaged (e.g., stress-

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Genes other than HSPs during heat stress

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denatured) proteins (Figure 1) .

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Several transcriptional systems other than HSF-1 are affected by heat shock. At least three mechanisms
have been identified through which heat shock specifically affects these systems: 1) changes in the level
of expression of transcription factors themselves (through a variety of mechanisms, 2) changes in activity
of transcription factors that are already expressed (for example, by phosphorylation), and 3) changes in
cellular location of transcription factors (such as translocation to the nucleus or sequestration in the
cytoplasm) (Sonna et al., 2002).
Some of these genes are C-reactive protein (CRP), p53, p21, interleukin-6, interleukin-8, VEGF(Vascular
endothelial growth factor), cNOS, iNOS and many others (Sonna et al., 2002).
Conclusion
Thermal stress is a cause of great concern among livestock owners in tropical countries. It causes change
in the antioxidant level and various hormones and electrolyte concentration but increases lipid
peroxidation in vivo. Animal shows various responses to thermal stress at physiological, cellular,
biochemical and molecular level. Changes in various genes expression helps in amelioration from stress
at cellular level. At molecular level increased expression of stress related genes and hence their protein
products will help to combat the stress.

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