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Physiological, Biochemical and Molecular Responses to Thermal Stress in Goats

Mahesh Gupta 1 *, Sachin Kumar 2 , S.S. Dangi 3 , Babu Lal Jangir 4 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

Sep 09, 2012 11:05; Accept Date: May 02, 2013 08:11 Abstract Livestock undergo various kinds of

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, tri- iodothyronine, 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.

stress, thyroxin, serum enzymes, heat shock proteins. Introduction Globally goat plays an important role in the

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 man’s cow”.

Perhaps, it is the only livestock that fits well for effective utilization in the diverse socio-economic

situations of the developing countries.

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

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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International Journal of Livestock Research

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Vol 3(2) May’13

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 animal’s 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 13-27 0 C 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 abnormal spermatozoa in the ejaculate (Perez-Crespo et al., 2008). In female, it lowers fertility, conception rate and embryonic survival in animals (Edwards and Hansen, 1997; Sakatani et al., 2004). It impair follicular and oocyte development and reduces steroid production (Zeron et al., 2001; Ozawa et al.,

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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 H 2 O 2 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 high environmental temperature in several studies (Devendra, 1987; Marai et al., 2007). Phulia et al., (2010) reported increase in rectal temperature and respiration rate from 38.97 0 C and 43.66 to 39.35 0 C and 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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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 flow 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 the hypothalamo-pituitary-adrenal cortical axis (HPA) and sympatho-adrenal medullary axis (Minton, 1994). Sivakumar et al. (2010) reported that there was increase in plasma concentration of prolactin from 11.73 to 26.37 μg/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 thermal stress, 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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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 2 O 2 produced during different metabolic processes and also in stressful conditions by reducing it to H 2 O and O 2 (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 O 2 - 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).

Blood glucose and total serum cholesterol levels are physiological adaptation mechanisms that can be affected by high ambient temperatures. Blood glucose and total serum cholesterol levels show greater differences in hot conditions than in the comfort zone. Some researchers reported that hot climatic

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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.

Significant 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 nonspecific cellular response to stress, whereas others may eventually found to be specific to certain types of stress. In mammalian cells, nonlethal heat shock produces changes in gene expression and in the activity of expressed proteins, resulting in what is referred to as a cell stress response. This response

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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 denatured proteins. HSP70 is mostly found in the cytosol and nucleus. Its functions are nascent protein folding, cytoprotection and as molecular chaperones. HSP90 is mostly found in the cytosol, ER and 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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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- denatured) proteins (Figure 1) .

repair and refolding of damaged (e.g., stress- denatured) proteins (Figure 1) . Genes other than HSPs

Genes other than HSPs during heat stress

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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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