Pertemuan 2. Gupta2019

Biological Rhythm Research
ISSN: 0929-1016 (Print) 1744-4179 (Online) Journal homepage: https://www.tandfonline.com/loi/nbrr20
Heat stress and thermoregulatory responses of

goats: a review
Mokshata Gupta & Tanmay Mondal
To cite this article: Mokshata Gupta & Tanmay Mondal (2019): Heat stress and thermoregulatory
responses of goats: a review, Biological Rhythm Research
To link to this article: https://doi.org/10.1080/09291016.2019.1603692
Published online: 12 Apr 2019.
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BIOLOGICAL RHYTHM RESEARCH
https://doi.org/10.1080/09291016.2019.1603692
ARTICLE
Heat stress and thermoregulatory responses of goats: a

review
Mokshata Guptaa and Tanmay Mondalb
a
Division of Animal Nutrition, ICAR- Indian Veterinary Research Institute, Izatnagar, India; bDivision of
Physiology & Climatology, ICAR- Indian Veterinary Research Institute, Izatnagar, India
ABSTRACT ARTICLE HISTORY

Good health is central to animal welfare as it indicates their Received 18 March 2019
physiological functioning. Sustainability of livestock is largely Accepted 3 April 2019
affected by prevailing climatic conditions. Animals undergo KEYWORDS
a variety of stress, among which heat stress is the most concern- Alleviation; growth; heat
ing. The vulnerability of animals to heat stress varies according to stress; physiology;
species, genetic potential, management system and nutritional production; reproduction
status. The ability of goats to cope with heat stress without harm-
ing their health and production performance has been often
glorified. Although goats are resistant to heat stress at a greater
extent but they suffer from heat stress beyond their comfort zone.
An imbalance between metabolic heat production inside their
body and its dissipation to the surroundings leads to heat stress.
The foremost reaction of goats includes behavioral changes,
increases in respiration rate, rectal temperature and heart rate. It
directly affects feed intake thereby, lowers growth rate, produc-
tion, reproduction, and even death in severe cases. The purpose of
the present review is to provide an integrative explanation of
thermoregulatory responses of goats at behavioral, physiological,
biochemical, productive and reproductive level during heat stress
and also the strategies for its alleviation.
Abbrevations: ACTH: Adrenocorticotropic hormone; ADH:
Antidiuretic hormone; ALP: Alkaline phosphatase; ALT: Alanine
transaminase; AST: Aspartate transaminase; BHBA: Beta-
hydroxybutyric acid; CRH: Corticotropin-releasing hormone;
DMI: Dry matter intake; FSH: Follicle stimulating hormone; Hb:
Hemoglobin; HPA axis: Hypothalamo-pituitary-adrenal axis; LH:
Luteinizing hormone; NE: Norepinephrine; NEFA: Non-esterified
fatty acids; PCV: Packed cell volume; T3: Triiodothyronine; T4:
Thyroxine; TSH: Thyroid stimulating hormone
Introduction
Developing countries like India constitute substantial small ruminant population and
they are considered the major source of the rural economy (Agrawal et al. 2014). Among
small ruminants, goat plays a predominant role in the sustenance of the livelihoods of
poor families (Okoruwa 2014). Goats are among the species with a wide geographical
distribution and have an important role in the agricultural revolution and advancement
CONTACT Mokshata Gupta mokshtagupta1407@gmail.com

© 2019 Informa UK Limited, trading as Taylor & Francis Group
2 M. GUPTA AND T. MONDAL
of human civilization. They produce food and fiber at relatively low cost from unconven-
tional feed materials. That is why referred as “poor man’s cow” taking a broader picture
of small and marginal farmers. The economic importance of goat production has been
increased predominantly in countries that are routinely exposed to harsh climates
(Banerjee et al. 2014), as they perform better than other domesticated ruminants
(Silanikove and Koluman 2015; Shilja et al. 2016; Darcan and Silanikove 2018). There
are more than 600 goat breeds in the world (Dwyer 2009) and these breeds differ in
their capacity to overcome harsh climatic conditions. Indigenous breeds are well
adapted to the agro-climatic conditions where they have evolved and are hardy enough
to thrive well on poor forage and stressful conditions where high yielding exotic and
crossbred animals succumb (Akinyi 2008). The smaller size of indigenous goat breeds is
a genetic adaptive strategy synchronized with the available nutrient resources as
compared to exotic and crossbred goats (Nyamushamba et al. 2017). Goats with loose
skin and floppy ears are more heat tolerant (Naandam and Assan 2014), indicating
proper breed selection is a very valuable tool for sustaining goat production under an
increasingly challenging environment that is heat stress.
The goats are the species of preference kept by animal producers. This interest arises
due to several encouraging characteristics of goats over other livestock as highlighted in
Figure 1. Goats are more resilience than other ruminants (Silanikove 2000b). Resilience is
defined as “the ability of a species to survive and recover from a perturbation” (Williams
et al. 2008). The resilience is influenced by species ecology, physiology and genetic
diversity. The effective urea recycling in the rumen of goats allows them to effectively
Figure 1. Unique characteristics of goats.

BIOLOGICAL RHYTHM RESEARCH 3
digest low-protein feeds. The spacious rumen volume of goats plays an important role
by serving as a huge fermentation vat and water reservoir. The water stored in the
rumen is utilized during dehydration, and thus it serves as a container, which accom-
modates the ingested water upon rehydration. Goats usually browse tree-leaves and
shrubs, which ensure a steady supply of food all year around. One of the most
noteworthy features of goats is characterized by seasonal changes in the volume and
anatomy of the digestive tract, which provide rapid acclimatization to changes in forage
quality (Maitra et al. 2014). The corresponding morpho-physiological adaptations include
larger salivary glands, higher respiration rate, higher skin temperature, constant heart
rate and cardiac output.
Stress is described as the reaction of body to external forces that disturb homeostasis,
usually associated with detrimental effects on health and performance (Rosales 1994).
Sejian et al. (2018) established the superiority of goats over sheep in terms of a tolerance
to heat stress. Goats undergo different types of stress such as physical, nutritional,
chemical, psychological and heat stress. Among these environmental variables, heat stress
is the major constraint that shows deleterious effects due to ever-changing climatic
scenario (Silanikove and Koluman 2015; El-Tarabany et al. 2017; Shaji et al. 2017; Mondal
et al. 2019). Climate change is the most serious long-term challenge faced by small
ruminants’ owners in much of the world, as it impacts animal production, health and
welfare (El-Tarabany and El-Bayoumi 2015; Silanikove and Koluman 2015). The developing
countries are more prone to extreme climatic events as they largely depend on climate-
sensitive sectors like agriculture and forestry (IPCC 2007). Recently, Silanikove and
Koluman (2015) also stated that the severity of heat stress issue is an increasing problem
in near future because of global warming progression. Marai and Habeeb (2010) defined
heat stress as the state at which the animal body physiological mechanisms activate to
maintain the body’s thermal balance, when exposed to elevated temperature. The vulner-
ability of livestock to heat stress varies according to species, genetic potential, life stage,
management or production system and nutritional status (Das et al. 2016). Heat stress
influences the hematological and physiological, and biochemical parameters of goat
(Ocak and Guney 2010; Phulia et al. 2010; Alam et al. 2011; Sharma and Kataria 2011). In
addition, it negatively affects their feed intake, growth, production, reproduction as well
as natural immunity and thus, making them more vulnerable to diseases, and even death
(Rojas-Downing et al. 2017). The negative effect of heat stress in goats has been high-
lighted in Figure 2. Consequently, heat stress leads to economical impacts on the goat
industry, emphasizing the necessity to objectively assess their welfare. Although goats are
resistant to heat stress at a greater extent but they get affected when exposed beyond
their comfort zone. Browsing of goats to open fields during most of the day hours makes
them susceptible to heat stress. The purpose of the present review is to provide an
integrative explanation of thermoregulatory responses of goat at behavioral, physiologi-
cal, biochemical, productive and reproductive level during heat stress and also the
strategies for its alleviation.
Thermoregulatory mechanism
Goats are homeotherms and can maintain near-constant body temperature under
a wide range of environmental conditions (Morrison 1983). All animals have a thermal
Figure 2. Negative impacts of heat stress on goats (Abbreviations: HPA axis: Hypothalamo-pituitary
-adrenal axis, CRH: Corticotropin-releasing hormone, ACTH: Adrenocorticotropic hormone, Hb:
Haemoglobin, PCV: Packed Cell Volume, DMI: Dry Matter Intake, T3: triiodothyronine, T4: thyroxine)
(Modified from Pragna et al. 2017).
comfort zone, which is a range of ambient environmental temperatures that are bene-
ficial to physiological functions (FAO 1986). The thermoneutral zone for goats is about
12–24°C. Changes in ambient temperature cause imbalance between the level of heat
production and heat loss and thus, alter metabolism. The severity of heat stress on goats
depends on temperature, humidity, breed, genetic potential, life stage, outer body
surface and nutritional status. The exposure of goats to high environmental temperature
induces an increase in the dissipation of excess body heat so as to counteract the
excessive heat load. The first defense layer protecting the goats from direct sunlight is
provided by the coat, and this protection differs according to coat color, depth and
length (Naandam and Assan 2014). Goats with a dark coat absorb more thermal radia-
tion and are more prone to heat than those with a light colored coat. Helal et al. (2010)
mentioned that the thermal insulation increases by increasing the coat depth and
attributed to the air space between the hair fibers. The long hair serves as an insulator
from the heat, providing a hair buffer zone between the outer environment and the
animal’s body (Naandam and Assan 2014).
Animal has a series of mechanisms to counter heat stress by dissipating heat to the
environment and reducing the production of metabolic heat (Silanikove 2000a). Animal
dissipates body heat via different mechanisms: 1) Conduction, where they conduct heat
to a cooler surface. 2) Convection, where thermal currents leave their body. 3) Radiation,
where the animal radiates heat to a cooler environment. 4) Evaporation, where moisture
is evaporated from the body surface (sweating) and lungs (panting). Physiological
thermolytic pathways under heat stress incorporate an increase in respiratory and
cutaneous evaporative cooling (Katamoto et al. 1998), an increase in peripheral to
splanchnic blood flow ratio (Silanikove 2000a) and the suppression of thermogenic
hormone production (Baccari et al. 1983). In addition, multiple behavioral adaptive
mechanisms play a major role, including seeking of shade and cooler surfaces, changing
the pattern of feed and water intake and change in posture and reduced muscular
activity (Mitlohner et al. 2002).
Exposure of goats to high ambient temperature augments the thermoregulation
by increasing respiration, sweating and rectal temperature that consequently causes
disturbances in the water, protein, energy and minerals metabolism. Generally, the
heat stressed animals reduces their feed intake and slows down their basal metabo-
lism causing hypo-function of thyroid gland in order to prevent the additional
metabolic heat production (McManus et al. 2009). Further, the animals reduce the
production of triiodothyronine (T3) and thyroxine (T4) during exposure to high
ambient temperatures (Rhoads et al. 2009). The final result of these changes is
impairment of appetite, feed intake, feed efficiency and utilization, growth, milk
yield and reproduction.
There are several phenotypic and genotypic characters which impart the adaptive
potential to goats, thereby allowing them to cope with harsh conditions. These may arise
over generations through slow modifications as goats adapt to environmental challenges.
The adaptive process can be expanded to include (i) Behavioural, (ii) Physiological (iii)
Hematological and Biochemical (iv) Neuro-endocrine (v) Productive, and (vi) Reproductive
responses, which combine to promote survival in a specific environment. They have been
highlighted in Figure 3 and further discussed in detail in following section.
Behavioral adaptations
Among the several responses of the animal, behavioral adaptation is the first and
foremost response adopted to reduce heat load (Shilja et al. 2016). There is little
literature published about the behavioral changes in goats during heat stress.
However, goats behave in various ways during heat stress as indicated in Figure 3.
One of the most quick and profound behavioral adaptation in heat stressed goats is
seeking shade (Curtis et al. 2017). If shade is not available, they will change their posture
to the vertical position or urination and defecation frequency.
Posture
In severe heat stress, goats reduce their grazing time, tend to lie down to reduce their
locomotion and spend more time in the shade. Standing and lying are behavioral
adaptive mechanisms associated with higher ambient temperatures (Darcan et al.
2008; Provolo and Riva 2009). Generally, heat-stressed goats spend more time standing
so that they can reorient themselves in different directions to avoid direct solar radiation
and ground radiation (Alam et al. 2011). In addition, the standing position also obstructs
the conductive heat transfer into the animal body due to the presence of a layer of air
adjacent to the skin, and also facilitates the dissipation of body heat load to the
Figure 3. Thermo-regulatory responses of goats to heat stress (Modified from Sejian et al. 2018).
surroundings by increasing the amount of skin exposed to airflow or wind. However,

divergent results have also been reported, indicating the adaptation to heat stress
conditions (Shilja et al. 2016).
Urination and defecation

Goats under severe heat stress moisten their body surface with water, saliva or nose
secretions. The loss of body heat to environment by evaporation of respiratory secre-
tions, sweat or saliva might lead to severe dehydration. This leads to decrease in the
number of urination in heat stressed goats (Alam et al. 2011; Shilja et al. 2016). In
addition, heat stress also reduces defecation frequency, considered to be an adaptive
mechanism to conserve body water (Shilja et al. 2016).
Inference
The range of behavioral responses affects heat exchange between the goats and its
environment by reducing heat gain from radiation and increasing heat loss via convec-
tion and conduction. It includes shade seeking, change in posture, urination and
defecation frequency.
Physiological adaptations
Heat stress has significant changes on some physiological parameters of goat that result
in a incredible economic loss for the goat industry (Panda et al. 2016). Physiological
parameters like respiration rate, heart rate and rectal temperature demonstrate the
mechanism of physiological adaptation. Increased temperature and pulse rate are the
important indicators of heat stress of goats (Sanusi et al. 2011). Changes in respiration
rate, pulse rate and rectal temperature have been frequently used as indicators of
physiological adaptability of goats to heat stress (Al-Dawood 2017).
Rectal temperature
Body temperature is good measure of heat tolerance in animals. It represents the
consequential of all heat gain and heat loss processes of the body. Maintenance of
body temperature is under neuronal control in a negative feedback system (Fecteau and
White 2014). Rectal temperature is an important index of body temperature that can be
used to assess the severity of heat stress. An increase in rectal temperature by 1°C or less
is enough to reduce animal performance. Rectal temperature of goats was found to be
higher during heat stress conditions (Marai et al. 2007; Phulia et al. 2010; Okoruwa 2014;
Wang et al. 2016). This has been considered a natural mechanism by which they prevent
body temperature from raising during heat stress help prevent death from heat stroke.
In contrast, some studies reported no changes in rectal temperature of heat-stressed
goats (Alam et al. 2011; Panda et al. 2016). This difference might be due to variation in
experimental conditions and/or breed and age of goats. In addition to rectal tempera-
ture, scrotal temperature is also a better indicator in assessing the impact of heat stress
in goats (Shilja et al. 2016).
Pulse rate
The pulse rate is the regular beat rate of the arteries as the blood is pumped through
them to the heart. It primarily reflects the homeostasis of circulation along with the
general metabolic status. It can be quickly altered due to external factors like high
ambient temperature (Phulia et al. 2010). Normal pulse rates for goats range from 90
to 95 beats/min. But, it increases during heat stress in an attempt to dissipate more
heat to its surroundings by increasing the blood flow to the body surfaces (Shilja
et al. 2016). Increase in pulse rate is attributed to additional two causes: a) increase in
muscular activity controlling the rate of respiration, concurrent with elevated respira-
tion rate and b) reduction in resistance of peripheral vascular beds and arteriovenous
anastomoses.
Respiratory rate
Respiration rate is a practical and reliable indicator of heat stress (Okoruwa 2014).
Normal respiration rates for goats range from 15 to 30 breaths/min. Heat stress is
characterized by an increase in respiration rate (Phulia et al. 2010; Hamzaoui et al.
2013; Wang et al. 2016). The severity of heat stress can be qualified according to panting
rate (breaths/min) (low: 40–60, medium: 60–80, high: 80–120, and severe: >200), as it
appears to be the most accessible and easiest way for evaluating the impact of heat
stress (Silanikove 2000a, 2000b). The increased respiration rate is probably indicating an
effort of animals to maintain their normal body temperature by increasing their heat
dissipation through evaporative cooling (Hamzaoui et al. 2013). However, respiration
rate can be indirectly influenced by the changes of metabolism and muscle activity as
well as environmental conditions (Silanikove 2000a).
Inference
The general homeostatic responses of goats to heat stress include increase in rectal
temperature, pulse rate and respiration rate. Excessive heat stress may have several
physiological side effects and economical impacts on the goat industry. But, the com-
plexity of these physiological changes varies from species to species, individual to
individual, age of goats and the environmental conditions.
Effects on intake and growth

Goats have several nutrient requirements including water, energy, protein, minerals, and
vitamins (Thornton et al. 2009). Interaction between heat stress and nutrition is asso-
ciated with marked reduction in feed intake (Alam et al. 2011), higher drinking frequency
and increased water intake (Valente et al. 2015; Shilja et al. 2016). Increased water intake
is an attempt to boost up heat loss by sweating and panting (Hamzaoui et al. 2013).
Numerous studies reported lower feed intake during heat stress condition (Baumgard
and Rhoads 2012; Hamzaoui et al. 2013; Salama et al. 2014; Shilja et al. 2016; Bagath
et al. 2017; Curtis et al. 2017; Darcan and Silanikove 2018). This reduction in feed intake
is an adaptive response of the animal to reduce metabolic heat because the heat
increment of feeding is an important source of heat production (Sejian et al. 2018;
Pragna et al. 2018a).
Heat exposure increases the level of circulating arginine vasopressin, which may
contribute to the increased sympathetic tone in the visceral microvasculature
(Streefkerk et al. 2002). Consequently, it reduces visceral blood flow and may depress
feed intake (Renquist 2019) as depicted in Figure 4. Additionally, heat exposure stimu-
lates the peripheral thermal receptors to transmit suppressive nerve impulses to the
appetite center in the hypothalamus and thus, causing a decrease in feed intake so as to
minimize thermal load. Thus, less substrate becomes available for heat production.
Growth is defined as increase in the live body mass or cell multiplication, is controlled
genetically and environmentally by well-balanced available nutrients, hormones and
enzymes (Marai et al. 2007). Heat stress negatively affects the growth performance, i.e.
growth rate, daily weight gain, and live body weight (Gad 2013). Among the several
growth variables, body weight is an important parameter for explaining the adaptive
capability of goats. Several researches reported alteration in the growth parameters
during heat stress (Caulfield et al. 2014; Okoruwa 2014; Popoola et al. 2014; Habibu et al.
2016). The major reason for the growth retardation could be decreased anabolic activity
due to the reduced voluntary feed intake of essential nutrients, particularly metaboliz-
able energy for both maintenance and weight gain and enhanced tissue catabolism. The
Figure 4. Impact of heat stress on milk production.
growth reduction varies between different breeds of goats. This was supported by the
study of Pragna et al. (2018a), body weight reduction was higher in Osmanabadi breed
compared Malabari and Salem back goats breeds. This was consistent with the findings
of Helal et al. (2010) and Indu et al. (2014). Also, heat stress has negative effect on body
condition score of the animals (Pragna et al. 2018a), due to higher rate of mobilization of
fat storage for energy synthesis to support life-sustaining activities.
Inference
One of the most noticeable consequences of HS is decreased feed intake. It also leads to
a negative energy balance and reduced weight gain. Numerous nutritional changes are
needed (i.e. use high-quality fiber, add fats, increase mineral content and adjust vitamins) to
maintain feed intake, increase nutrient density and minimize the negative effect of heat stress.
Neuro-endocrine response and metabolic adaptations

The heat-stressed goats enter a bioenergetic state similar to the negative energy
balance (Moore et al. 2005), associated with a variety of metabolic and hormonal
changes. Metabolic adaptation is considered to be one of the important means to
overcome heat stress challenges by reducing the metabolic heat production (Pragna
et al. 2018a). The hypothalamo–pituitary–adrenal axis (HPA axis) is one of the principal
endocrine regulators of the stress responses. The products of HPA axis which control
stress in animals are corticotropin-releasing hormone (CRH), Adrenocorticotropic hor-
mone (ACTH) and cortisol as indicated in Figure 2.
Hormones
Several hormones (i.e. thyroxine, triiodothyroxine, prolactin, leptin, adiponectin, growth
hormone, glucocorticoids, mineralocorticoids, catecholamines and antidiuretic) are
involved in adaptation to heat stress and could be important indicators for assessment
of heat stress (Sivakumar et al. 2010). These hormonal changes are mediated through
HPA axis and sympatho-adrenal medullary axis (Minton 1994). The exposure of goats to
severe heat stress conditions suppresses the production of hormone releasing factors
from the hypothalamic centers causing a decrease in pituitary prolactin, somatotropin,
thyrotropin and leutinizing hormone, insulin and possibly parathormone and conse-
quently lowers the secretion of anabolic hormones (Habeeb et al. 1992). These are either
involved with nutrient partitioning and homeorhesis or homeostatic regulation augmen-
ted by heat stress.
Thyroid hormones play a vital role in metabolic adaptation and act as an indicator for
assessing the thermotolerance of goats (Haque et al. 2012). Thyroid hormones, specifi-
cally triiodothyronine (T3) and thyroxine (T4), play a vital role in metabolic adaptation
and growth performance of animals (Aleena et al. 2016). The TSH is an important
pituitary hormone that enhances the production of thyroid hormones and its blood
level is synchronized by the negative feedback mechanisms of thyroid hormones
(Lalsangpuii et al. 2015). Thyroid hormones increase oxygen consumption and cellular
ability for heat production, which elevate basal metabolic rate (Ocak et al. 2009).
Therefore, decreased thyroid hormone level during heat stress is an adaptive response
and occurs due to the direct effect on the hypothalamo pituitary and thyroid axis to
decrease the production of thyrotropin-releasing hormone, which enables goats to
reduce metabolic rate (Aleena et al. 2016; Pragna et al. 2018a). Additionally, energy
balance also plays a major role in decreasing thyroid hormone level (Kong et al. 2004).
Recently, it was stated that T3 could serve as an indicator of metabolic activity in goats
during heat stress (Pragna et al. 2018a). But, increased TSH concentration during heat
stress was also reported in goats (Pehlivan and Dellal 2017), indicating their superior
adaptive capability to the heat stress.
Cortisol is the primary stress hormone that acts as an important biological marker for
stress in goats (Binsiya et al. 2017; Shaji et al. 2017). The activation of HPA axis during
heat stress leads to enhanced production and release of cortisol into the circulation. The
association between heat stress and increased secretion of cortisol in small ruminants is
well documented (Shilja et al. 2016; Tajik et al. 2016; Chergui et al. 2017; Sejian et al.
2018). The significantly higher cortisol concentration indicates the sensitivity to heat
stress (Archana et al. 2018). This increase is an adaptive response to initiate hepatic
gluconeogenesis (Ghassemi Nejad et al. 2017). Further, cortisol is the principal stress
reliever in ruminant species and hence its higher level is beneficial to cope up with heat
stress condition (Shaji et al. 2017).
Aldosterone is a steroid hormone released from the cortex of the adrenal glands and
helps to maintain the sodium/potassium balance within the body. There is
a simultaneous relationship among heat stress, plasma aldosterone level and urine
electrolyte concentration (Phillips and Santurtun 2013). Severe dehydration in goats
during heat stress may lead to increased secretion of antidiuretic hormone (ADH)
through activation of rennin-angiotensin-aldosterone system (Kaliber et al. 2016). The
ADH hormone regulates the blood osmolarity by increasing the water absorption in the
kidneys, which also assists the excretion of concentrated urine in heat stressed goats
(Kaliber et al. 2016). Further, the plasma aldosterone level was reported to be higher
when goats have an ad libitum access to water (Shilja et al. 2016).
The growth hormone is a major peptide hormone that promotes the growth, cell
multiplication and cell regeneration. The plasma growth hormone may act as ideal
biomarkers for assessing the impact of heat stress on growth performance in goats
(Pragna et al. 2018a). The balance between the rate of energy production and utilization
determines the concentration of this hormone (Hirayama and Katoh 2004). During heat
stress, the level of this hormone increases (Pulina et al. 2012; Bagath et al. 2016), possibly
due to reduction in feed intake.
NEFA and BHBA

They are the important metabolic regulator in plasma and serum during heat stress (Aleena
et al. 2016). In addition, they are the most indicative substances of the animal energy
status. Their concentration in blood reflects the degree of adipose tissue mobilization
(Pullen et al. 1989). As an adaptive mechanism to heat stress, goats decrease metabolic
rate through reduction in feed intake, which enhances the mobilization of body fat
reserves. Thus, the enhanced production of NEFA is an attempt to maintain the energy
balance (Al-Dawood 2017). But, Hamzaoui et al. (2013) stated that reduction in feed intake
and body weight during heat stress was not accompanied by body fat mobilization as
NEFA concentrations did not vary during heat stress. This might be an attempt to increase
glucose utilization which will result in lower metabolic heat production.
Ketone bodies come from volatile fatty acids (mainly butyrate that converted to BHB
across the rumen epithelium) and fat mobilization (Salama et al. 2014). Neither weekly
nor mean total concentrations of BHB were affected by heat stress, indicating no
mobilization of body fat (Al-Dawood 2017). It might be considered as metabolic adap-
tive responses during heat stress. Although some contradictory findings were observed
in heat-stressed dairy goats (Salama et al. 2014).
Enzymes
Metabolic regulators are important determinants of physiological mechanisms during
stressed conditions and are best assessed by determining the level of these enzymes
in plasma or serum. Acid phosphatase and alkaline phosphatase (ALP) are key
enzymes associated with the metabolic activities in animals. The levels of these
enzymes are generally low in heat stressed animals, which could be attributed to
a metabolic shift in the animals (Chaidanya 2015). But, significant variation in plasma
ACP (acid phosphatase) enzyme concentration but non-significant change in the
plasma ALP concentration were also observed during the heat stress (Pragna et al.
2018b). These differences could be correlated to the heat stress magnitude as well as
the breed differences.
Serum level of aspartate transaminase (AST) and alanine transaminase (ALT) is helpful
in assessing the animal welfare. The serum level of these enzymes increases in heat
stressed goats due to their higher adaptive capability (Banerjee et al. 2015). On the
contrary, no significant changes were observed in AST level of heat-stressed goats (Ocak
et al. 2009; Sharma and Kataria 2011).
Inference
Heat-stressed goats undergo various metabolic and hormonal changes. Hormones
produced from the adrenal and thyroid glands play a significant role in thermoregula-
tion and metabolic adaptation during heat stress. The NEFA, BHBA, T3 and T4 are
considered to be reliable indicators of metabolic adaptation during heat stress.
Hematological and blood biochemical adaptations

Hematological adaptations
Heat stress is known to alter the hematological parameters (Rojas-Downing et al. 2017).
There is a great variation in these parameters between different goat breeds. Heat stress
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, packed cell volume (PCV) and
hemoglobin (Hb) values (Temizel et al. 2009; Sivakumar et al. 2010; Alam et al. 2011; Kumar
et al. 2011). In addition, decrease in Hb and PCV levels could be due to 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 decreases
voluntary intake under heat stress. However, an elevated value of Hb was also reported in
heat stressed goats and the observed change may be attributed to higher Hb requirement
to meet the increased oxygen circulation during panting (Okoruwa 2014). Decrease in PCV
might be due to haemodilution effect where more water is transported into the circulatory
system for evaporative cooling (EL-Nouty et al. 1990). But, several experiments reported
higher levels of PCV during heat stress (McManus et al. 2009; Rana et al. 2014). This higher
PCV value could be an adaptive mechanism to provide water necessary for evaporative
cooling process (Al-Haidary 2004).
Neutrophil is the main defender of the body against infection and antigens. Its high levels
may indicate an active infection; a low count may indicate a compromised immune system.
Lymphocytes are involved in protection of the body from viral infections. Elevated levels
may indicate an exhausted immune system. Monocytes are helpful in fighting severe
infections and are considered the bodies’ second line of defense against infection and the
largest cells in the bloodstream. The total leucocytes reduce during severe heat stress
(Wojtas et al. 2014; El-Tarabany et al. 2017). This has been attributed to the increment in
cortisol level, which probably is responsible for the impairment of the cellular immune
response (Caroprese et al. 2012). This was inconsistent with the findings of Alam et al. (2011).
Inference
The regular assessment of blood constituents is useful in assessing the health status of
goats during heat stress conditions. These hematological parameters show varying trend
with an increase in environmental temperature.
Blood biochemical adaptations

Blood biochemical parameters reflect the health and the metabolic status of an animal.
The heat stress alters the serum level of several biochemical parameters like total
protein, albumin, globulin, glucose, cholesterol, etc.
Glucose and cholesterol

Physiologically, blood glucose and total serum cholesterol levels are an adaptation
mechanism that can be affected by high ambient temperatures. Their level shows
greater variations during heat stress condition than in the comfort zone. Some research-
ers reported that heat stress decrease blood glucose and total serum cholesterol levels
in goats (Ocak et al. 2009; Ocak and Güney 2010; Pandey et al. 2012; Hooda and
Upadhyay 2014; El-Tarabany et al. 2017) but, some studies are contrary to these findings
(Webster 1976; Hamzaoui et al. 2013). This decrease in glucose level could be related to
the decrease in availability of nutrients and lower rate of propionate production
(Mohamed 2012), or greater blood insulin activity (Rhoads et al. 2009; Baumgard and
Rhoads 2013). Additionally, it may be due to the increase in plasma glucose utilization to
provide energy for muscular expenditure required for high muscular activity associated
with increased respiration rate (Sejian and Srivastava 2010). The marked decrease in total
serum cholesterol levels has been attributed to increase in total body water or the
decrease in acetate concentration which is the primary precursor for the synthesis of
cholesterol. Additionally, it may be due to lower thyroid activity, decrease feed intake
and consequently, reduction in intake of dietary cholesterol (Pandey et al. 2012).
Protein
The level of total plasma protein, albumin and globulin decreases in heat stressed goats
(Helal et al. 2010; El-Tarabany et al. 2017; Shaji et al. 2017). This has been attributed to
increase in plasma volume as a result of heat stress. On the contrary, prolong exposure
of goats to heat stress increases plasma total protein, albumin, and globulin (Okoruwa
2014). This might be due to vasoconstriction and dehydration that occurs due to
increased respiration rate during heat stress (Helal et al. 2010).
Inference
Blood biochemical parameters reflect the health and the metabolic status of goats. The
serum level of total protein, albumin, globulin, glucose, and cholesterol shows greater
differences during heat stress. In addition, there have been conflicting results on their
serum level with different environmental conditions.
Heat stress and production performance

Among dairy animals, goats are the most adapted species to imposed heat stress in
terms of production, reproduction and disease resistance (Silanikove and Koluman
2015). The contribution of the dairy goat sector to total milk production has been
popularized during the last 20 years (Gerosa and Skoet 2012). Globally, the majority of
goat milk production is obtained in developing countries where goat’s milk plays
a significant role in the nourishment of millions of humans (FAO 2013). The maintenance
requirements during heat stress increases by 30% (NRC 2007) and the energy intake
would not be sufficient to cover the daily requirements (Hamzaoui et al. 2013). As
results, animals enter into a stage of negative energy balance, consequently affects
growth, lactation, mammary development, in utero development (Renquist 2019), and
large economic losses (Polsky and von Keyserlingk 2017).
The decrease in milk production due to heat stress is multifaceted as summarized in
Figure 4. Briefly, heat stress reduces voluntary feed intake, which leads to utilization of
body reserve of fat and nitrogen to furnish energy through gluconeogenesis process at
the expense of the mammary tissues (Amaral-Phillips et al. 1993). In another systemic
mechanism, stress activates the HPA axis and stimulates the secretion process ACTH by
the anterior pituitary. The ACTH enhances the synthesis and emission of glucocorticoids
from the cortex of the adrenal gland (depicted in Figure 2). Ultimately, the cortisol
causes decrease in milk synthesis by blocking the uptake of glucose by the mammary
gland (Dwyer and Bornett 2004).
Heat stress can lead to “transient hypogalactia” or “chronic hypogalactia” (Renquist
2019). “Transient hypogalactia”, occurs during lactation and results in an acute decrease
in milk production in goats (Salama et al. 2014), although response is breed specific.
Lactating dairy goats are more sensitive to heat stress compared with non-lactating one,
due to milk production elevating metabolism. Moreover, because of the positive asso-
ciation between milk yield and heat production, higher yielding animals are more prone
to heat stress than lower yielding. Heat stress throughout gestation and lactation or only
during lactation decreases kit growth, whereas kits from does that were exposed to heat
stress only during gestation grew normally during lactation (Marco-Jiménez et al. 2017).
Consecutively, the seasonal effects on doe milk production and kit growth are likely
a result of transient hypogalactia. On the contrary, Hamzaoui et al. (2013) reported no
significant decrease in milk yield in heat stressed lactating goats, probably because
glucose was sufficient for the milk yield level at late lactation. Moreover, this may be
attributed to some degree of body muscle degradation under heat stress and the
resultant amino acids could be utilized for gluconeogenesis (Wheelock et al. 2010).
“Chronic hypogalactia”, results from depressed mammary gland development, indi-
cating heat stress affects lactation independent of depressed feed intake (Renquist
2019). Heat stress during the critical stages of mammary gland development, before
and immediately following parturition, can affect milk production throughout the entire
lactation. Milk yield depends on mammary cell numbers and secretory capacity of each
cell (Tao et al. 2011). Heat exposure in the periparturient period limits mammary gland
development by suppressing proliferation and has no effect on apoptosis. Mammary
blood flow positively regulates milk production by increasing the delivery of nutrients to
the mammary gland to produce milk (Davis and Collier 1985). But, during heat stress,
mammary blood flow decreases which in turn limits milk production (Delamaire and
Guinard- Flament 2006). The basics have been summarized in Figure 4.
Besides having an effect on milk yield, heat stress also alters the milk quality and
cause a decrease in the lactation period in dairy goats. Lactose is one of the most critical
components in milk that displays disturbances in secretory tissues. Its level reduces
dramatically as the severity of heat stress increases in goats (El-Tarabany et al. 2017). This
was in agreement with the study of Hamzaoui et al. (2012) and Salama et al. (2014),
heat-stressed goats produced milk with lesser lactose (−1 to −5%) contents. The heat
stressed goats produces milk with lesser protein percentage (−6.3%) (Hamzaoui et al.
2012), due to lower mammary synthesis of milk protein during heat stress (Bernabucci
et al. 2002). Moreover, this might be due to decreased protein intake and increased
sweat secretion that contains protein and urea might have limited the availability of
amino acids for milk protein synthesis (Joshi et al. 1968). Exposure to heat stress also
reduces milk fat, fat plus protein yield and solids-not-fat (SNF) content (Bouraoui et al.
2002; Kadzere et al. 2002; Menéndez- Buxadera et al. 2012).
Inference
Increasing the environmental temperature is associated with production losses in goats.
The response of goats to heat stress in terms of production varies according to breed.
Both heat stress and the advancement of lactation concur to reduce mobilization of
body reserve for milk synthesis, thus inducing a worsening of milk quantity and quality.
It can be overcome by providing shade during late gestation, cooling strategies, improv-
ing dry matter intake, etc.
Heat stress and reproduction efficiency

The animal reproductive system is more sensitive to temperature than other body parts.
High ambient temperature impairs various tissues or organs of the reproductive system
in both the sexes of the animal. In males, it 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 follicular growth, oocyte maturation (Cui et al. 2009; Mustafi et al. 2009),
impair of embryo development and pregnancy rate (Sirotkin 2010; Rojas-Downing et al.
2017). The following section discusses in detail the effects on heat stress on estrous
period and follicular growth, fertility, and embryonic development.
Estrous period and follicular growth

High ambient temperature negatively affects the ability to display natural mating
behavior, as it reduces both the duration and intensity of estrous besides increases
the incidence of anestrous and silent heat in farm animals (Kadokawa et al. 2012; Singh
et al. 2013; Polsky and von Keyserlingk 2017). This reduction has been attributed to
reduce DMI and the subsequent effects on hormone production as it increases ACTH
and cortisol secretion (Singh et al. 2013). Luteinizing hormone (LH) regulates the
progression of medium follicles to large follicles as well as follicular dominance
(Bodensteiner et al. 1996). In addition, pulsatile LH secretions are known to drive

follicular steroidogenesis. But, when female goats are exposed to high ambient tem-
perature, follicular growth to ovulation suppresses, accompanied by decreased LH
receptor level and follicular estradiol synthesis activity (Ozawa et al. 2005). Thus, it
leads to regression of the follicles before ovulation.
Estradiol is important in the maintenance of ovarian function (Bayne et al. 2011). It
regulates follicle development and ovarian atresia, inhibits granulosa cell apoptosis and
promotes the division and growth of granulosa cells (Chun et al. 1996). The exposure to
heat stress decreases estradiol concentrations in follicles (Li et al. 2016) as well as in primary
cultures of granulosa cells (Luo et al. 2016). Low estradiol secretion suppresses signs of
estrus, gonadotropin surge, ovulation, transport of gametes and ultimately reduced ferti-
lization (Wolfenson et al. 2000). Granulosa cells are critical for normal ovarian function and
synthesize many factors essential for follicle development (Matsuda et al. 2012; Wang et al.
2012). Furthermore, the growth, differentiation, and maturation of oocytes are dependent
upon proliferation and differentiation of granulosa cells (Jančar et al. 2007). Aromatase
activity in granulosa cells is highly regulated by Follicle stimulating hormone (FSH) (Erickson
and Hsueh 1978). The heat-stressed follicles lack sufficient FSH stimulation to enhance the
aromatase activity owing to decrease FSH receptor expression. Thus, reduced granulosa
cells aromatase activity and viability also contributes to poor estradiol secretion (Ozawa
et al. 2005). Thus, a shift in the balance of the signaling pathway between granulosa cell
survival and death induces a decrease in the production of estradiol (Chen et al. 2004).
The nutrition has significant role in maintaining reproductive functions (Evans and
Anderson 2012). Acute nutrient restriction reduces the size of the dominant follicle and
causes lack of pre-ovulatory increase in estradiol and the ovulatory surge of LH, resulting in
ovulatory failure (Lents et al. 2013). On the other hand, a short period of improved nutrition
can stimulate ovulation rate, indicating that nutrient imbalance will be reason behind it.
Fertility: Studies on the reproduction performance indicated that the fertility
decreases during heat stress. Several mechanisms are involved in reducing the fertility
of goats, depending on the severity of heat stress. It includes changes in follicular
dynamics and steroidogenic activity (Ozawa et al. 2005; Shimizu et al. 2005), lower
oocyte competence (Cui et al. 2009; Mustafi et al. 2009; Singh et al. 2013; Lacerda and
Loureiro 2015), a reduction in the expression of estrus, increase in the circulating
prolactin level (Alamer 2011; Singh et al. 2013) and early embryonic death (Ozawa
et al. 2002). About 80% of estrus may be unnoticeable during summer which further
reduces fertility. During severe heat stress, secretion of endometrial PGF-2α increases,
thereby threatening maintenance of pregnancy and leads to infertility (Bilby et al. 2008).
Also, plasma FSH surge increases and inhibin concentrations decrease leading to varia-
tion in follicular dynamics and depression of follicular dominance that could also be
associated with low fertility (Roth et al. 2000). This increase in FSH secretion is probably
due to reduced inhibition of negative feedback from smaller follicles which ultimately
affect the reproductive efficiency of dairy animals (Khodaei-Motlagh et al. 2011).
Embryonic growth and development

Heat stress negatively affects embryonic growth and development. High ambient tem-
perature decreases the blood flow to the uterus together with the increase in body
temperature. Consequently, decrease in fertilization rate limits the embryonic develop-

ment and aggravate early embryonic fatalities (De Rensis et al. 2002). Low progesterone
secretion limits endometrial function and also embryo development (Wolfenson et al.
2000; Khodaei-Motlagh et al. 2011). Additionally, it causes embryonic death by interfer-
ing with protein synthesis, oxidative cell damage (Wolfenson et al. 2000), reducing
interferon-tau production for signaling pregnancy recognition (Bilby et al. 2008) and
expression of stress-related genes associated with apoptosis (Fear and Hansen 2011).
Further, during early organogenesis and fetal development, heat stress leads to various
teratologies that are most evident during early stages of development (Wolfenson et al.
2000; Demetrio et al. 2007). Fetal malnutrition and eventually fetal growth retardation
under heat stress were also reported (Tao and Dahl 2013).
Inference
Heat stress influences follicular growth, ovarian function and embryonic development
which results in decreased fertility. These effects are associated with reduced viability of
granulosa cells, impaired aromatase activity, reduced oestradiol concentration and LH
receptor levels. Proper nutrition may have an important role in improving reproductive
function.
Alleviation strategies
The strategies to alleviate heat stress on goats require a multidisciplinary approach that
has been summarized in Figure 5. It is important to understand the responses of goats
to high ambient temperature in order to design a suitable alleviation strategy. The
amelioration strategies can be broadly grouped into three categories: Physical modifica-
tions of environment, nutritional management and genetic development of thermoto-
lerance breeds.
Physical modifications of environment

Modification of environment to enhance heat dissipation mechanism is one of the most
common approaches to alleviate heat stress. It includes provision of shade, evaporative
cooling system with water in the form of fog, mist or sprinkling with natural or forced air
movement, and possibly cooling ponds (Atrian and Shahryar 2012; Caulfield et al. 2014).
Among these, shading is the simplest and cost-effective methods and it is applicable
under extensive conditions. However, the use of shades, fans or evaporative cooling is
not possible in semi-intensive systems as goats are grazed in the open during most of
the day, and this necessitates other strategies to counteract the adverse effects of heat
stress (Silanikove et al. 2010). Trees and shrubs are very effective and natural shading
materials along with beneficial cooling as moisture evaporates from the leaves (Ayo
et al. 2006). If natural shelter is not available, goat producers can use Quonset huts,
plastic calf hutches, polydomes and/or carports to provide shelters for grazing goats. In
addition, hay or straw shades, solid shade provided by sheet metal painted with white
on top, and aluminum sheets are the most effective and cheap materials (Ghanem et al.
2008). A well-designed shade structure reduces heat load by 30–50% (Atrian and
Figure 5. Relevant amelioration strategies to counteract heat stress in goats (Modified from Sejian
et al. 2018).
Shahryar 2012). Providing shade to goats reduces rectal temperature and respiration
rate (Darcan and Guney 2008; Aggarwal and Upadhyay 2013). Goats handling should
also be kept at minimum and that too in the early morning or late evening time. Proper
site selection of animals housing is fundamental to decrease long-term exposure to solar
radiations (Sarangi 2018). Goat response positively to cooling measures as indicated by
improvement of milk yield in treated goats in comparison to non-cooled goats (Darcan
et al. 2008). Thus, improvement in thermoregulatory indices leads to increase in feed
efficiency and profit.
Nutritional management
Careful ration modifications can greatly help in reducing the negative impacts of heat
stress. Lower DMI during heat stress reduces availability of nutrients for absorption, and
absorbed nutrients are used less efficiently (West 1999). During summer, the feeding
should be done during the cooler periods of the day as it encourages them to maintain
their normal feed intake and prevents the climatic heat load (Onyewotu et al. 2003).
Also, feeding at more frequent intervals is beneficial to minimize the diurnal fluctuation
in ruminal metabolites and thus, increase feed utilization efficiency in the rumen (Sejian
et al. 2015). Decreasing the forage to concentrate ratio can result in more digestible
rations that may be consumed in greater amounts (Darcan and Güney 2008). Feed
containing low fiber content is preferred during heat stress since heat production is
highly associated with metabolism of acetate compared with propionate (Atrian and
Shahryar 2012; Samara et al. 2016). More nutrient-dense diets are usually preferred
during the heat stress period. Dairy goats supplemented with 4% fat during summer
had lower rectal temperature, enhanced milk production efficiency and yield (Linn et al.
2004). Feed additives have been proposed to offset the consequence of heat stress. For
example, supplementation of live yeast plays an important role in nutrient digestibility
by altering the volatile fatty acids production in the rumen, decrease the production of
ruminal ammonia and increase in ruminal microorganism population (Stella et al. 2007).
The antioxidants such as vitamin C and E protect the body defense system against
excessive production of free radicals, decreases rectal temperature and respiration rate
during heat stress (Sivakumar et al. 2010) and thus, stabilize the health status of the
animal (Silanikove et al. 2010). Another best way to reduce heat stress is to provide
adequate fresh and cool drinking water (Samara et al. 2016).
Genetic selection
Apart from nutritional and environmental modifications, genetic selection is an impor-
tant aspect to alleviate the impact of heat stress. However, long-term strategies have to
be evolved for adaptation to climate change. Differences in heat tolerance between
different goat breeds provide clues to select thermotolerant breeds using genetic tools.
Improvement of adaptability by simultaneous heterosis and crossbreeding is a better
option in the hand of goat breeders for producing better offspring capable of with-
standing heat stress. Genetic variability for the response to climatic change can be used
to select the most tolerant and robust animal to cope up with future climatic changes
(Sarangi et al. 2018). Identification of genes associated with thermotolerance and using
these genes as markers in the breeding program or marker-assisted selection should be
applied to identify animals adapted to thermal stress considering genotype–environ-
ment interactions in addition to higher productivity.
Inference
A variety of adjustments should be adopted by goat owners to overcome the negative
effects of heat stress. These adjustments include use of shades, changes in feeding
schedules (during cool hours, feeding intervals), grazing time, handling time, and ration
composition such as adjustment in dietary fiber, use of high-quality fiber forage,
increased energy density, use of feed additives and antioxidants. Thus, improvement
in thermoregulatory indices is beneficial in terms of health and economic benefits.
Conclusion
Among the environmental variables affecting goats, heat stress has emerged as the
major threat to the goat production. Although new facts about the responses of goats to
heat stress continue to be developed, but their management to reduce its impact
remains a challenge. Extended exposure to high ambient temperature compromises
their ability to dissipate excess body heat which ultimately affects their behavior,
physiological functions, feed intake, growth, milk production, and reproductive effi-
ciency, resulting in severe economic constraints for goat producers. These can be over-
come by adopting suitable alleviation strategies like modifications of environment,
nutritional management and genetic selection. Thus, a cavernous understanding of
various adaptive responses of goats may provide future directions for coping them to
the devastating effects of heat stress.
Disclosure statement
No potential conflict of interest was reported by the authors.
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Biological Rhythm Research

ISSN: 0929-1016 (Print) 1744-4179 (Online) Journal homepage: https://www.tandfonline.com/loi/nbrr20

Heat stress and thermoregulatory responses of

Mokshata Gupta & Tanmay Mondal

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Heat stress and thermoregulatory responses of goats: a

ABSTRACT ARTICLE HISTORY

CONTACT Mokshata Gupta mokshtagupta1407@gmail.com

Figure 1. Unique characteristics of goats.

surroundings by increasing the amount of skin exposed to airﬂow or wind. However,

Urination and defecation

Eﬀects on intake and growth

Figure 4. Impact of heat stress on milk production.

Neuro-endocrine response and metabolic adaptations

NEFA and BHBA

Hematological and blood biochemical adaptations

Blood biochemical adaptations

Glucose and cholesterol

Heat stress and production performance

Heat stress and reproduction eﬃciency

Estrous period and follicular growth

(Bodensteiner et al. 1996). In addition, pulsatile LH secretions are known to drive

Embryonic growth and development

temperature. Consequently, decrease in fertilization rate limits the embryonic develop-

Physical modiﬁcations of environment

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