October. 2023, Vol. 13, No. 5 https://doi.org/10.1093/af/vfad046 This is an Open Access article distributed under the terms of the Creative Commons Attribution License (https://creativecommons.org/licenses/by/4.0/), which permits unrestricted reuse, distribution, and reproduction in any me- dium, provided the original work is properly cited. © Chauhan, Zhang, Osei-Amponsah, Clarke, Sejian, Warner, Dunshea Feature Article Impact of heat stress on ruminant livestock production and meat quality, and strategies for amelioration Surinder S. Chauhan,† Minghao Zhang,‡ Richard Osei-Amponsah,†,|| Iain Clarke,† Veerasamy Sejian,$ Robyn Warner,† and Frank R. Dunshea†,¶ †School of Agriculture, Food and Ecosystem Sciences, Faculty of Science, The University of Melbourne, Parkville, Melbourne, VIC 3010, Australia ‡Department of Food and Nutrition, Provincial Hospital, Shandong First Medical University, China ||Department of Animal Science, School of Agriculture, University of Ghana, Accra, Ghana $Animal Physiology Division, ICAR-National Institute of Animal Nutrition and Physiology, Bangalore 560030, India ¶Faculty of Biological Sciences, The University of Leeds, Leeds LS2 9JT, UK Key words: adaptation, animal breeding, climate change, meat safety, thermotolerance Introduction The Intergovernmental Panel on Climate Change (IPCC) projects a 1.5  °C to 2  °C increase in global warming in the 21st century, meaning that heat stress (HS) will continue to affect the sustainability of livestock farms worldwide nega- tively (Ciliberti et al., 2022), including reduced productivity of rangeland, shortage of nutritional feed, compromised animal welfare, and high energy costs for cooling (Ortiz-Colón et al., 2018). Additionally, heatwaves are becoming more frequent and of higher magnitude and are persisting for longer periods, compromising animal welfare and production in the absence of suitable interventions to counteract HS. For instance, the Australian Bureau of Meteorology (BOM) has recorded an an- nual increase in heatwaves in recent years (Figure 1), a trend that is expected to continue. Over millennia, ruminant livestock have developed behaviors to cope with high temperatures, through natural or artificial se- lection. For example, ruminants tend to rest during the hottest parts of the day and become more active during cooler times. Dairy cattle graze less during the hottest period of the day, seeking shade and spending more time at the watering points to conserve energy and reduce heat production (Figure 2). Direct and Indirect Effects of HS on Ruminant Production Ruminant livestock are raised within diverse cultural and environmental production systems globally and contribute to food security. They have well-developed thermoregula- tory mechanisms but adaptation to increased environmental temperature compromises animal performance production (Gaughan et  al., 2019). Like other homeotherms, rumin- ants maintain their core body temperature within a narrow zone by maintaining a balance between heat gain and heat loss. Ruminants reduce their feed intake, production, and growth rate, to reduce metabolic heat production, a significant Implications • Climate change will continue to accentuate the negative impact of heat stress (HS) on ruminant livestock pro- duction, compromising animal welfare and meat quality. • Mitigation strategies, including providing shade on farms, modifications of animal housing (heat extract- ors, fans, water sprinklers, and cool drinking water), and nutritional interventions, are important short-term measures to reduce the negative effects of HS. • Climate-smart breeding for thermotolerance and matching of adapted ruminant breeds to appropriate production systems should be considered for more sustainable livestock production systems. D ow nloaded from https://academ ic.oup.com /af/article/13/5/60/7311134 by U niversity of G hana, Legon user on 04 D ecem ber 2023 https://creativecommons.org/licenses/by/4.0/ 61October. 2023, Vol. 13, No. 5 Key words: adaptation, animal breeding, climate change, meat safety, thermotolerance Introduction The Intergovernmental Panel on Climate Change (IPCC) projects a 1.5  °C to 2  °C increase in global warming in the 21st century, meaning that heat stress (HS) will continue to Figure 1. Frequency of extreme heat events in Australia. Source: Australian Bureau of Meteorology. contributor to the total heat load on animals (Yadav et  al., 2016; Sejian et al., 2017). To maximize radiant heat loss from the body, heat-stressed animals redistribute blood supply away from the gastrointestinal tract to the periphery resulting in hyp- oxia and cellular damage to intestinal membranes leading to leaky gut and inflammation, all of which can result in localized and general oxidative stress (OS) (Chauhan et al., 2014). The OS is an imbalance between oxidants and antioxidants at the cellular level or the entire organism, which damages macromol- ecules such as lipids and proteins and may result in reduced meat quality and product deterioration(Chauhan et al., 2014). HS in animals during the preslaughter period over summer reduces liveweight and carcass weight; causes pale, soft, and exudate (PSE), and dark, firm, and dry (DFD) meat in live- stock; and increases the foodborne disease outbreaks, when compared to winter conditions (Gonzalez-Rivas et al., 2020). Similarly, exposure of pregnant animals to HS leads to signifi- cant adverse effects on their innate and adaptive immune func- tions and their offspring, influencing morbidity, mortality, and growth rate in heifers (Dahl et al., 2020). HS also has indirect effects on ruminant production through a reduction in the quantity and quality of fodder while increasing the spread of new vector-borne diseases (Sejian et  al., 2018). Generally, exposure to high ambient temperat- ures requires at least a week to influence sheep body weight. It takes 7 d of HS (28  °C to 40  °C cyclic HS) to negatively affect the bodyweight of lambs when compared to animals in a thermoneutral (TN: 18 °C to 21 °C) environment (Chauhan et al., 2016). For older sheep, this threshold may be longer due to conditioning to temperature change in previous seasons. Breed is another important determinant of heat tolerance in livestock, with hair sheep having thinner coats, more efficient Figure 2. Dairy cattle at the University of Melbourne Dookie Robotic Dairy Farm resting under trees on a hot day during summer of 2018 (Osei-Amponsah et al., 2020). D ow nloaded from https://academ ic.oup.com /af/article/13/5/60/7311134 by U niversity of G hana, Legon user on 04 D ecem ber 2023 62 Animal Frontiers sweating and shedding capacity, leading to superior heat tol- erance, compared to sheep with wool (Sejian et  al., 2018). Furthermore, high-production temperate breeds have poorer heat tolerance due to inherently higher metabolic rate and heat production than tropical breeds. Thus, we have shown that 2 wk of cyclic HS impacted Merino × Border Leicester lambs’ growth performance and feed intake but had no impact on Dorper lambs (Figure 3), which are adapted to hot climates. Effect of HS on Meat Quality of Climate- Adapted Ruminants Climate-adapted animals have evolved physiological and be- havioral mechanisms that allow the maintenance of body tem- perature within a narrow range, even when exposed to high temperatures. Climate-adapted phenotypes may be procured through either natural or artificial selection. Physiological mech- anisms and behavioral changes conferring heat tolerance reduce the negative impact of HS on meat quality leading to reduced muscle fatigue which may improve meat tenderness. Additionally, changes in metabolism minimize the breakdown of fats, which can improve the flavor and aroma of the meat. However, se- vere, or prolonged HS can impact even climate-adapted animals. Under such conditions, the animal’s coping mechanisms may be- come overwhelmed, negatively impacting meat quality. In general, reduction in meat production has been demon- strated in both long-term and short-term HS studies (Gregory, 2010; Thornton et  al., 2022) (Table 1). Pragna et  al. (2018a) found that the average body weight gain was lower in goats sub- jected to 45 d of summer conditions (73.5 to 86.5 Temperature Humidity Index [THI]) than in animals that were housed at a lower THI (69.9 to 74.9), confirming findings by Chauhan et al. (2016). Our current research indicates that less than 5 d of HS (28 °C to 40 °C, 30 % to 40% RH, cyclic temperature from day to night) does not affect body weight and daily weight gain of Merino crossbred lambs (Zhang et  al., 2022). Therefore, the negative effects of HS on meat production (growth) do not manifest until at least 7 d of heat exposure with normal feeding, although feedstuffs with low antioxidant contents may aggravate this situation. Carcass quality is affected by HS, as reflected in the quantity of subcutaneous and intramuscular fat (IMF). Limited research (Park et al. (2018) suggests that mild HS may be beneficial to fat deposition, conferring less subcutaneous fat and more IMF, but studies in small ruminants (Archana et al., 2018; Zhang et al., 2021) did not observe any impact of HS on subcutaneous fat thickness or IMF content. Similarly, a recent study in goat breeds indigenous to India (Devapriya et al., 2021) showed that, despite a higher magnitude of HS exposure (THI = 94.76) for a prolonged duration of 45 d, both the major carcass traits and meat quality variables remained unaffected. Although mild HS of less than 2 mo is unlikely to change the carcass fat content of ruminants, the effect of severe HS to reduce the subcutaneous fat thickness and IMF due to the loss of body weight requires further research. Meat color, which depends on the concentration and chem- ical state of myoglobin, is influenced by a change in meat pH and oxidation state under HS conditions. Current data suggest that HS tends to increase the redness of meat (a*) and reduce lightness (L*) and color stability due to the higher ultimate pH and oxidation state, which causes meat to appear darker than normal (Zhang et al., 2020). It should be noted, however, that the increase of a* is not defined as “problem meat.” Thus, in a recent study, meat from lambs exposed to 2 wk of HS (2 wk; 28  °C to 40  °C, 40% to 60%) had better color stability (less browning; Figure 4) compared with meat from animals under TN conditions (Zhang et al., 2021). Therefore, unlike the sea- sonal impact of HS on meat quality, production and animal welfare outcomes are more likely to be compromised during the short-term high-ambient-temperature environment conditions. As indicated above, effects of HS are mitigated in climate- adapted animals. Our unpublished data show that wool- shedding breeds of sheep displayed higher meat color stability compared with wool breeds under cyclic HS conditions (Zhang et al., 2022). Compared to water holding capacity (WHC) and texture, meat color parameters are more sensitive to HS. For instance, in a recent study (Zhang et al., 2021), we showed that after 2-wk HS (28 °C to 38 °C, 40% to 60% RH; cyclic tempera- ture) an increase in lamb meat redness (a*) becomes apparent but in other studies, the influence on WHC and texture was observed only after 1 mo of exposure to hot conditions as in- dicated in Figure 5 (Kadim et al., 2008; Archana et al., 2018). One of the most consistent negative consequences of HS in ruminants is the increased frequency of DFD meat (Gregory, 2010) based on the higher meat ultimate pH. However, heat- stressed ruminants produce meat with lower WHC (Kadim et al., 2008; Archana et al., 2018), which is usually associated with lower ultimate pH and may be due to oxidative modification of proteins because of OS induced by HS. In addition to WHC, HS may also affect meat tenderness. For example, seasonal HS (35  °C, 47% RH) in sheep reduced the myofibrillar fragmen- tation index (MFI; lower MFI indicating less tender meat) of Figure 3. The effect of 2-wk HS or TNon feed intake of Poll Dorset × (Merino × Border Leicester; high production) and Dorper (high heat tolerance) lambs (n = 12 for each group). “a” and “b” indicate levels that are significantly different at 5% level of SED (Zhang et al., 2021). SED is shown as vertical bar. D ow nloaded from https://academ ic.oup.com /af/article/13/5/60/7311134 by U niversity of G hana, Legon user on 04 D ecem ber 2023 63October. 2023, Vol. 13, No. 5 meat, compared with that of animals in the cool season (21 °C, 59%RH) (Kadim et al., 2008). Similarly, 45 d of HS exposure in goats increased shear force of meat as compared to meat pro- duced from goats housed at lower THI (Archana et al., 2018). Macías-Cruz et al. (2020) confirmed these results, showing that Warner–Bratzler shear force of Dorper × Katahdin lambs’ Longissimus thoracis (LT) muscle from animals subjected to a month of summer conditions (28.4 ± 4.0 °C, 55.2 ± 18.2% RH) was higher than that of meat from animals subjected to winter conditions (19.2 ± 2.6 °C, 41.7 ± 11.0% RH). At least 2 wk of HS in ruminants is required before any impact on WHC and shear force of their meat occurs (Zhang et al., 2021). A sum- mary of the literature on effects of HS on meat quality is given in Figure 5. How Does HS Change Meat Quality in Ruminants? Until now, studies of the impact of HS on meat quality have focused mainly on the effect on postmortem muscle glycolysis. HS alters the physiology and metabolism of animals, including effects on redox balance and apoptosis. Thus, to develop suit- able strategies to alleviate HS, it is important to consider how HS changes biological processes that affect meat quality. Lower glycogen concentrations due to reduced feed intake and hyper- thermia under HS can predispose muscle to a higher ultimate pH, which is close to the definition of DFD meat (pH ≥ 5.8 or 6.0). If glycogen concentrations are above the threshold for postmortem glycolysis utilization, exposure to acute HS before Table 1. The effect of heat stress on growth in small ruminants Author Breed HS condition Body weight change Observation Pragna et al. (2018b) Osmanabadi goats Malabari goats Salem Black goats Summer exposure: 73.5 to 86.5 THI Shed feeding: 69.9 to 74.9 THI 45-d study ADG1: HS significantly reduced the body weight gain among all heat exposed groups, but the reduction of feed intake of HS group was not significant (except for Malabari goats). Osmanabadi: Exposure −39.63a g Shed 48.02b g Malabari: Exposure−25.00a g Shed 39.29b g Salem Black: Exposure −21.03a g Shed 34.53b g Nicolás- López et al. (2021) Dorper × Katahdin male lambs (34.6 ± 1.4 kg; 4.5 mo) Summer: 28.3 ± 4.0 °C, 77.2 ± 5.4 THI Winter: 19.2 ± 2.6 °C, 64.0 ± 3.0 THI 30-d study ADG: The average body weight gain and feed efficiency of summer group were significantly lower than winter group.Dorper × Katahdin: Summer 226a g Winter 302b g Zhang et al. (2021) Poll Dorset × (Border Leicester × Merino) lambs Dorper lambs HS: 28 °C to 38 °C, 40% to 60% RH cyclic TN: 18 °C to 21 °C, 45% to 55% RH 2-wk study ADG: Two weeks of cyclic HS had a significant negative influence on feed intake and body weight gain of wool breed lambs (2nd cross), but the influence of HS was not significant for hair breed (Dorper lambs). Dorper: HS −50.6a g TN 5.95a g 2nd cross: HS −92.3a g TN 101.0b g Chauhan et al. (2016) White Suffolk × Merino × Border Leicester lambs (42 ± 2.0 kg; 7 mo) HS: 28 °C to 40 °C, 30% to 40% RH TN: 18 °C to 21 °C, 40 % to 50% RH Feed intake: HS 959a g/d TN 1266b g/d One week of HS had a significant impact on 2nd cross lambs’ feed intake compared with the TN group. 1ADG, average body weight gain. Figure 4. Effect of 2-wk HS or TN on meat retail display of Poll Dorset × (Merino × Border Leicester; high production) lambs’ longissimus thoracis et lumborum muscle in 10-d high-oxygen-modified atmosphere packaging (80% O2: 20% CO2; n = 12 for each group; Zhang et al., 2021). D ow nloaded from https://academ ic.oup.com /af/article/13/5/60/7311134 by U niversity of G hana, Legon user on 04 D ecem ber 2023 64 Animal Frontiers slaughter may lead to a more rapid decline in muscle pH re- sulting in PSE meat (Gonzalez-Rivas et al., 2020). Generally, more than 2 wk of HS is required to elevate the ultimate pH of meat from sheep and beef cattle (Kadim et al., 2004; Rana et al., 2014; Zhang et al., 2021). HS is also known to cause OS. In healthy biological sys- tems, animals maintain a balance between reactive oxygen species (ROS) and antioxidants at the cellular level (Chauhan et  al., 2014). However, HS leads to excessive ROS produc- tion and reduced antioxidants due to lower feed intake and increased requirements to scavenge ROS. For example, Shi et al. (2020) reported that 28 d of HS (25.25 °C, 74.49% RH) significantly reduced glutathione peroxidase activity and raised the malondialdehyde concentration of crossbred lambs (Dorper × Mongolian) compared with TN. Similarly, in a study of Merino crossbred lambs, we (Chauhan et al. (2016) observed that oxidation products in plasma were higher in lambs subjected to HS (28 °C to 40 °C, 30% to 40% RH) for 7 d than in lambs maintained under TN (18 °C to 21 °C, 40% to 50% RH) conditions, influencing the color stability of meat in the former. Moreover, high levels of ROS production during HS are likely to cause protein oxidation (such as myosin, sarco- plasmic proteins, calpain, myoglobin, and glycolytic enzymes) in meat leading to a change in muscle structure by cross-linking or denaturation, resulting in a reduction in the water space in muscle and decreased tenderness (Mitra et al., 2017). Therefore, further studies on antioxidant supplementation or the effects of adding natural herbs to the ruminant diet to reduce OS and improve meat quality are warranted. Effect of HS on Meat Safety It is generally accepted that meat with high ultimate pH and intestinal tissue damage in live animals poses safety risks for meat and by-products. Some psychrotrophic bacteria (which grow at refrigeration temperatures and lead to the spoilage of product) are inhibited at normal pH (5.6) of meat but can grow on meat with a high ultimate pH, including Acinetobacter and Altermonas putrefaciens (Newton and Gill, 1981). On the other hand, tissue damage and inflammation of the intestines of ru- minants induced by HS increases the intestinal permeability and luminal attachment of bacteria (Bailey et  al., 2004). An increase in the levels of stress hormones such as catecholamine and glucocorticoids may also change intestinal barrier func- tion and the microbial environment in ruminants (Verbrugghe et al., 2012). For example, HS impairs intestinal integrity and increases intestinal permeability to endotoxins and transloca- tion of pathogenic bacteria into the bloodstream (Salmonella, Escherichia coli, and Campylobacter) into the bloodstream (Lian et  al., 2020). Such intestinal bacteria may be taken up by muscle and liver (dependent on the health of the animal), which can lead to foodborne illness in humans. Thus, HS can compromise meat safety, especially in low- to medium-input production systems in the developing world where ruminant production significantly contributes to food security. Strategies for the Amelioration of HS Climate change induced HS, and its challenge to livestock production requires adaptation and mitigation strategies. There are some adaptive practices that have been developed in breeding and pasture management programs including selection for more heat-resistant genotypes (Gaughan et al., 2019), silvopasturing (the practice of integrating trees, forage, and the grazing of do- mesticated animals in a mutually beneficial way), and diversifica- tion of forage production and optimization production practices to reduce HS (Ortiz-Colón et al., 2018). Another strategy is to in- crease the energy and nutrient densities (reduced fiber, increased concentrates, and supplemental fat) of the diet since feed intake is reduced under HS. Increasing ration concentrates should be Figure 5. Summary of literature on effects of HS for different periods on meat quality in ruminants. considered with care; however, as heat-stressed cows, for in- stance, are highly prone to rumen acidosis (Baumgard et  al., 2014). The risk of increased concentrate feeding can be miti- gated by feeding slowly fermentable grains such as maize or by treatment of rapidly fermentable grains to reduce rumen starch fermentation (Gonzalez-Rivas et al., 2016, 2017, 2021; Prathap et al., 2022). Another strategy to mitigate the effects of HS in ruminant livestock is the physical modification of animal’s envir- onment to reduce direct exposure to heat ranging from the pro- vision of shade, and cooling animals down using sprinklers, fans, mists, and perforated air ducting systems (Gaughan et al., 2019). Nutritional interventions to protect against oxidative damage re- sulting from HS-induced ROS production and antioxidant sup- plementation with betaine, lipoic acid, selenium, Vitamins C and E can also be utilized in combination with physical modifica- tion of the environment (Osei-Amponsah et al., 2019). However, these strategies often depend on the availability of electrical energy, adequate water supply, availability and affordability of nutritional supplements, and more capital investment often not available in all production systems. A key component of adaptation is the innate genetic ability of an organism to survive under stressful conditions (Osei- Amponsah et  al., 2019). Climate-adapted animals (Pereira et al., 2014; Cardoso et al., 2015) have evolved specific physio- logical and behavioral mechanisms that help regulate their body temperature to maintain homeostasis. For example, sweating allows the animals to release heat from their bodies through evaporation, while panting increases airflow over the respiratory system, allowing heat dissipation. Animals reduce their metabolic activity and heat production to cope with HS (Gonzalez-Rivas et  al., 2020). In cattle, thermo-tolerance is directly associated with sweat gland density and sweating rate. Cattle adapted to arid regions possess smooth, short, and thin hair (slick hair gene) coat, which enhances heat dissipation from the body (Dikmen et  al., 2018). Sheep with light coat colors, which are sleek and shiny, reflect greater solar radiation than hair coats that are dark and dense or woolly (Sejian et al., 2018). Furthermore, sheep with carpet wool are better protected from solar radiation by the facilitation of cutaneous heat dissipa- tion (Sejian et al., 2018). Sheep with longer, thicker, and darker coats are more substantially stressed and exhibit higher rectal temperatures and sweating rates in tropical regions than white- haired sheep (Gaughan et al., 2019). Goats are proficient desert- dwelling animals as their physiological characteristics provide them with an advantage over other ruminants in harsh envir- onmental conditions. Their small body size, short pointed erect ears, fleece structure, light coat, and high digestive efficiency help them survive extreme climatic conditions (Silanikove, 1997). Genetic selection for heat tolerance could provide a sus- tainable means of augmenting feeding and housing modifica- tions. For instance, selection signatures for thermotolerance can be identified through functional genomics, and productive breeds improved through crossbreeding with resilient genotypes and the incorporation of stress-tolerant genes. An example is the SLICK haplotype (https://omia.org/OMIA001372/9913) initially identified in Senepol cattle and other South America D ow nloaded from https://academ ic.oup.com /af/article/13/5/60/7311134 by U niversity of G hana, Legon user on 04 D ecem ber 2023 https://omia.org/OMIA001372/9913 65October. 2023, Vol. 13, No. 5 Figure 5. Summary of literature on effects of HS for different periods on meat quality in ruminants. considered with care; however, as heat-stressed cows, for in- stance, are highly prone to rumen acidosis (Baumgard et  al., 2014). The risk of increased concentrate feeding can be miti- gated by feeding slowly fermentable grains such as maize or by treatment of rapidly fermentable grains to reduce rumen starch fermentation (Gonzalez-Rivas et al., 2016, 2017, 2021; Prathap et al., 2022). Another strategy to mitigate the effects of HS in ruminant livestock is the physical modification of animal’s envir- onment to reduce direct exposure to heat ranging from the pro- vision of shade, and cooling animals down using sprinklers, fans, mists, and perforated air ducting systems (Gaughan et al., 2019). Nutritional interventions to protect against oxidative damage re- sulting from HS-induced ROS production and antioxidant sup- plementation with betaine, lipoic acid, selenium, Vitamins C and E can also be utilized in combination with physical modifica- tion of the environment (Osei-Amponsah et al., 2019). However, these strategies often depend on the availability of electrical energy, adequate water supply, availability and affordability of nutritional supplements, and more capital investment often not available in all production systems. A key component of adaptation is the innate genetic ability of an organism to survive under stressful conditions (Osei- Amponsah et  al., 2019). Climate-adapted animals (Pereira et al., 2014; Cardoso et al., 2015) have evolved specific physio- logical and behavioral mechanisms that help regulate their body temperature to maintain homeostasis. For example, sweating allows the animals to release heat from their bodies through evaporation, while panting increases airflow over the respiratory system, allowing heat dissipation. Animals reduce their metabolic activity and heat production to cope with HS (Gonzalez-Rivas et  al., 2020). In cattle, thermo-tolerance is directly associated with sweat gland density and sweating rate. Cattle adapted to arid regions possess smooth, short, and thin hair (slick hair gene) coat, which enhances heat dissipation from the body (Dikmen et  al., 2018). Sheep with light coat colors, which are sleek and shiny, reflect greater solar radiation than hair coats that are dark and dense or woolly (Sejian et al., 2018). Furthermore, sheep with carpet wool are better protected from solar radiation by the facilitation of cutaneous heat dissipa- tion (Sejian et al., 2018). Sheep with longer, thicker, and darker coats are more substantially stressed and exhibit higher rectal temperatures and sweating rates in tropical regions than white- haired sheep (Gaughan et al., 2019). Goats are proficient desert- dwelling animals as their physiological characteristics provide them with an advantage over other ruminants in harsh envir- onmental conditions. Their small body size, short pointed erect ears, fleece structure, light coat, and high digestive efficiency help them survive extreme climatic conditions (Silanikove, 1997). Genetic selection for heat tolerance could provide a sus- tainable means of augmenting feeding and housing modifica- tions. For instance, selection signatures for thermotolerance can be identified through functional genomics, and productive breeds improved through crossbreeding with resilient genotypes and the incorporation of stress-tolerant genes. An example is the SLICK haplotype (https://omia.org/OMIA001372/9913) initially identified in Senepol cattle and other South America breeds, which has been introgressed to Holsteins to improve their thermotolerance (Dikmen et  al., 2008). Inheritance of the SLICK1 allele of the prolactin receptor gene improves the thermotolerance of lactating Holstein cows under hot-humid environmental conditions (Carmickle et al., 2022). Furthermore, local breeds, particularly those from the Near East and Africa, are known for their better adaptation to high temperatures and harsh conditions and generally have a greater adaptive capacity to stressful environments than exotic breeds (Osei-Amponsah et  al., 2019). Therefore, breeding for thermotolerance and matching adapted ruminant breeds to suitable environments and production systems are useful strategies. A combination of strategies (Baumgard et al., 2014; Sejian et al., 2018; Collier et al., 2019; Gaughan et al., 2019) will be needed to mitigate HS in ruminants: • Provision of shade—Access to shaded areas can reduce dir- ect exposure to sunlight and the animal’s body temperature. In the extensive dairy systems, shade from trees can form part of a long-term strategy for managing climate variability on farms. The shade and shelter provided by trees in pad- docks and laneways can reduce the radiant heat load by 50% or more. • Provision of adequate water to maintain hydration during hot weather. • Adjusting feeding patterns—Feeding animals during cooler times, such as early morning or late evening, to reduce HS by lowering the heat produced during digestion. • Adjusting feed composition—Feeding diets with a lower proportion of fiber and a higher proportion of more readily digestible carbohydrates can reduce heat production during digestion. • Sprinklers or misting fans—Sprinkling or misting animals with water can help to cool them down and reduce HS. • Improving ventilation—Adequate ventilation in housing can help to reduce heat build-up and improve air quality. • Convective cooling—Fans can circulate air and reduce heat load on animals, which can help lower the core body tem- perature of animals. • Provision of cool, dry bedding. • Nutritional supplementation—In the short term, antioxi- dant supplements can reduce OS and betaine can reduce en- ergy requirements. • Genetic Interventions—In the longer term, selection for heat tolerance is an effective long-term strategy, as is intro- gression of specific (SLICK) genes. The application of OMICS technologies offers considerable scope for identifi- cation of more reliable biomarkers for HS resilience which could be incorporated in marker-assisted selection (MAS) to redefine the existing breeding goals for thermotolerance. Conclusions Ruminant livestock are important contributors to global food production and an important income source for many farmers in developing countries. HS remains a major threat D ow nloaded from https://academ ic.oup.com /af/article/13/5/60/7311134 by U niversity of G hana, Legon user on 04 D ecem ber 2023 https://omia.org/OMIA001372/9913 66 Animal Frontiers to ruminant livestock production, reducing feed intake, LWG, limiting meat and milk production, damaging the intestinal wall, compromising nutrient digestion and absorption, and affecting meat quality. Unfortunately, these harmful conse- quences of HS in terms of animal health and production are likely to be an ongoing issue across the globe, and a multidis- ciplinary approach integrating animal breeding, nutrition, and housing is required to counteract the impact of climate change on livestock. Amelioration strategies such as provision of shade, sprinklers, fans, energy-efficient cooling systems, water provision, feeding changes, and nutritional interventions can be utilized in the short term. However, it is important to note that some of these strategies may not be applicable to extensive sys- tems and most of these require the use of electricity and water, which might not be a sustainable option in certain regions. In extensive systems, crossbreeding high-producing exotic breeds with local climate-adapted breeds, planting more trees in pad- docks and laneways, and encouraging early morning and late afternoon grazing could be more useful. In the short term, modification of the animal’s environment and nutritional interventions can be considered mitigation strategies to sustain livestock production in a changing cli- mate. In the long term, genetic selection for thermotolerance and climate adaptation must be prioritized in future ruminant livestock breeding programs to improve animal comfort and performance in a changing climate. The integration of new technologies and capacity building in less-developed countries offers many opportunities for further development of strategies to reduce the negative impact of climate change on livestock. Conflict of interest statement. None declared. References Archana,  P., V.  Sejian, W.  Ruban, M.  Bagath, G.  Krishnan, J.  Aleena, G. Manjunathareddy, V. Beena, and R. Bhatta. 2018. 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Small Ruminant Res. 137:17–23. doi:10.1016/j. smallrumres.2016.02.011 Ciliberti,  M., M.  Caroprese, and M.  Albenzio. 2022. Climate resilience in small ruminant and immune system: an old alliance in the new sustain- ability context. Small Ruminant Res. 210(106662):10. doi:10.1016/j. smallrumres.2022.106662. Collier,  R.J., L.H.  Baumgard, R.B.  Zimbelman, and Y.  Xiao. 2019. Heat stress: physiology of acclimation and adaptation. Anim. Front. 9(1):12–19. doi:10.1093/af/vfy031 Dahl,  G.E., S.  Tao, and J.  Laporta. 2020. Heat stress impacts immune status in cows across the life cycle. Front. Vet. Sci. 7:116. doi:10.3389/ fvets.2020.00116 Devapriya,  A., V.  Sejian, W.  Ruban, C.  Devaraj, P.V.  Spandan, M.V.  Silpa, M.R. Reshma Nair, P.O. Nameer, and R. Bhatta. 2021. Analysis of car- cass traits and quantitative expression patterns of different meat quality governing genes during heat stress exposure in indigenous goats. 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Meat Sci. 162(108025):13. doi:10.1016/j.smallrumres.2021.106523 Gonzalez-Rivas, P.A., K. DiGiacomo, P.A. Giraldo, B.J. Leury, J.J. Cottrell, and F.R.  Dunshea. 2017. Reducing rumen starch fermentation of wheat with three percent sodium hydroxide has the potential to ameliorate the effect of heat stress in grain-fed wethers. J. Anim. Sci. 95(12):5547–5562. doi:10.2527/jas2017.1843 Gonzalez-Rivas, P.A., K. DiGiacomo, V.M. Russo, B.J. leury, J.J. Cottrell, and F.R. Dunshea. 2016. Feeding slowly fermentable grains has the potential to ameliorate heat stress in grain-fed wethers. J. Anim. Sci. 94:2981–2991. doi:10.2527/jas.2016-0295 Gonzalez-Rivas,  P.A., P.  Prathap, K.  DiGiacomo, J.J.  Cottrell, B.J.  Leury, S.S. Chauhan, and F.R. Dunshea. 2021. Reducing rumen starch fermen- tation of wheat with 3% NaOH does not reduce whole tract starch digest- ibility and increases energy utilization in wethers during heat stress. Small Ruminant Res. 204:8. doi:10.1016/j.smallrumres.2021.106523 Gregory, N.G. 2010. How climatic changes could affect meat quality. Int. Food Res. J. 43(7):1866–1873. doi: 10.1016/j.foodres.2009.05.018 Kadim,  I., O.  Mahgoub, D.  Al-Ajmi, R.  Al-Maqbaly, S.  Al-Mugheiry, and D.  Bartolome. 2004. The influence of season on quality characteris- tics of hot-boned beef m.  longissimus thoracis. Meat Sci. 66(4):831–836. doi:10.1016/j.meatsci.2003.08.001 Kadim,  I., O. Mahgoub, W. Al-Marzooqi, D. Al-Ajmi, R. Al-Maqbali, and S. Al-Lawati. 2008. The influence of seasonal temperatures on meat quality characteristics of hot-boned, m. psoas major and minor, from goats and sheep. Meat Sci. 80(2):210–215. doi:10.1016/j.meatsci.2007.11.022 Lian, P., S. Braber, J. Garssen, H.J. Wichers, G. Folkerts, J. Fink-Gremmels, and S.  Varasteh. 2020. Beyond heat stress: intestinal integrity disrup- tion and mechanism-based intervention strategies. Nutrients. 12(3):734. doi:10.3390/nu12030734 Macías-Cruz,  U., O.R.  Saavedra, A.  Correa-Calderón, M.  Mellado, N.G.  Torrentera, A.  Chay-Canul, M.A.  López-Baca, and L.  Avendaño- Reyes. 2020. Feedlot growth, carcass characteristics and meat quality of hair breed male lambs exposed to seasonal heat stress (winter vs. summer) in an arid climate. Meat Sci. 169:108202. doi:10.1016/j.meatsci.2020.108202 Mitra, B., A. Rinnan, and J. Ruiz-Carrascal. 2017. Tracking hydrophobicity state, aggregation behaviour and structural modifications of pork proteins under the influence of assorted heat treatments. Food Res. Int. 101:266– 273. doi:10.1016/j.foodres.2017.09.027 Newton, K.G., and C.O. Gill. 1981. The microbiology of DFD fresh meats: a review. Meat Sci. 5(3):223–232. doi:10.1016/0309-1740(81)90005-X Nicolás-López,  P., U.  Macías-Cruz, M.  Mellado, A.  Correa-Calderón, C.A. Meza-Herrera, and L. Avendaño-Reyes. 2021. Growth performance and changes in physiological, metabolic and hematological parameters due to outdoor heat stress in hair breed male lambs finished in feedlot. Int. J. Biometeorol. 65(8):1451–1459. doi:10.1007/s00484-021-02116-x Ortiz-Colón,  G., S.J.  Fain, I.K.  Parés, J.  Curbelo-Rodríguez, E.  Jiménez- Cabán, M. Pagán-Morales, and W.A. Gould. 2018. Assessing climate vul- nerabilities and adaptive strategies for resilient beef and dairy operations in the tropics. Clim. Change 146(1–2):47–58. doi:10.1007/s10584-017-2110-1 Osei-Amponsah, R., S.S. Chauhan, B.J. Leury, L. Cheng, B. Cullen, I.J. Clarke, and F.R. Dunshea. 2019. Genetic selection for thermotolerance in rumin- ants. Animals (Basel). 9(11):18. doi:10.3390/ani9110948 Osei-Amponsah, R., F.R. Dunshea, B.J. Leury, L. Cheng, B. Cullen, A. Joy, A. Abhijith, M.H. Zhang, and S.S. Chauhan. 2020. Heat stress impacts on lactating cows grazing Australian summer pastures on an automatic ro- botic dairy. 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Food Chem. 3:100052. doi:10.1016/j.fochms.2021.100052 Dikmen,  S., E.  Alava, E.  Pontes, J.M.  Fear, B.Y.  Dikmen, T.A.  Olson, and P.J. Hansen. 2008. Differences in thermoregulatory ability between slick- haired and wild-type lactating Holstein cows in response to acute heat stress. J. Dairy Sci. 91(9):3395–3402. doi:10.3168/jds.2008-1072 Dikmen, S., R.G. Mateescu, M.A. Elzo, and P.J. Hansen. 2018. Determination of the optimum contribution of Brahman genetics in an Angus-Brahman multibreed herd for regulation of body temperature during hot weather. J. Anim. Sci. 96(6):2175–2183. doi:10.1093/jas/sky133 Gaughan,  J.B., V.  Sejian, T.L.  Mader, and F.R.  Dunshea. 2019. Adaptation strategies: ruminants. Anim. Front. 9(1):47–53. doi:10.1093/af/vfy029 Gonzalez-Rivas, P., S. Chauhan, M. Ha, N. Fegan, and F.R. Dunshea. 2020. Effects of heat stress on animal physiology, metabolism, and meat quality: a review. Meat Sci. 162(108025):13. doi:10.1016/j.smallrumres.2021.106523 Gonzalez-Rivas, P.A., K. DiGiacomo, P.A. Giraldo, B.J. Leury, J.J. Cottrell, and F.R.  Dunshea. 2017. Reducing rumen starch fermentation of wheat with three percent sodium hydroxide has the potential to ameliorate the effect of heat stress in grain-fed wethers. J. Anim. Sci. 95(12):5547–5562. doi:10.2527/jas2017.1843 Gonzalez-Rivas, P.A., K. DiGiacomo, V.M. Russo, B.J. leury, J.J. Cottrell, and F.R. Dunshea. 2016. Feeding slowly fermentable grains has the potential to ameliorate heat stress in grain-fed wethers. J. Anim. Sci. 94:2981–2991. doi:10.2527/jas.2016-0295 Gonzalez-Rivas,  P.A., P.  Prathap, K.  DiGiacomo, J.J.  Cottrell, B.J.  Leury, S.S. Chauhan, and F.R. Dunshea. 2021. Reducing rumen starch fermen- tation of wheat with 3% NaOH does not reduce whole tract starch digest- ibility and increases energy utilization in wethers during heat stress. Small Ruminant Res. 204:8. doi:10.1016/j.smallrumres.2021.106523 Gregory, N.G. 2010. How climatic changes could affect meat quality. Int. Food Res. J. 43(7):1866–1873. doi: 10.1016/j.foodres.2009.05.018 Kadim,  I., O.  Mahgoub, D.  Al-Ajmi, R.  Al-Maqbaly, S.  Al-Mugheiry, and D.  Bartolome. 2004. The influence of season on quality characteris- tics of hot-boned beef m.  longissimus thoracis. Meat Sci. 66(4):831–836. doi:10.1016/j.meatsci.2003.08.001 Kadim,  I., O. Mahgoub, W. Al-Marzooqi, D. Al-Ajmi, R. Al-Maqbali, and S. Al-Lawati. 2008. The influence of seasonal temperatures on meat quality characteristics of hot-boned, m. psoas major and minor, from goats and sheep. Meat Sci. 80(2):210–215. doi:10.1016/j.meatsci.2007.11.022 Lian, P., S. Braber, J. Garssen, H.J. Wichers, G. Folkerts, J. Fink-Gremmels, and S.  Varasteh. 2020. Beyond heat stress: intestinal integrity disrup- tion and mechanism-based intervention strategies. Nutrients. 12(3):734. doi:10.3390/nu12030734 Macías-Cruz,  U., O.R.  Saavedra, A.  Correa-Calderón, M.  Mellado, N.G.  Torrentera, A.  Chay-Canul, M.A.  López-Baca, and L.  Avendaño- Reyes. 2020. Feedlot growth, carcass characteristics and meat quality of hair breed male lambs exposed to seasonal heat stress (winter vs. summer) in an arid climate. Meat Sci. 169:108202. doi:10.1016/j.meatsci.2020.108202 Mitra, B., A. Rinnan, and J. Ruiz-Carrascal. 2017. Tracking hydrophobicity state, aggregation behaviour and structural modifications of pork proteins under the influence of assorted heat treatments. Food Res. Int. 101:266– 273. doi:10.1016/j.foodres.2017.09.027 Newton, K.G., and C.O. Gill. 1981. The microbiology of DFD fresh meats: a review. Meat Sci. 5(3):223–232. doi:10.1016/0309-1740(81)90005-X Nicolás-López,  P., U.  Macías-Cruz, M.  Mellado, A.  Correa-Calderón, C.A. Meza-Herrera, and L. Avendaño-Reyes. 2021. Growth performance and changes in physiological, metabolic and hematological parameters due to outdoor heat stress in hair breed male lambs finished in feedlot. Int. J. Biometeorol. 65(8):1451–1459. doi:10.1007/s00484-021-02116-x Ortiz-Colón,  G., S.J.  Fain, I.K.  Parés, J.  Curbelo-Rodríguez, E.  Jiménez- Cabán, M. Pagán-Morales, and W.A. Gould. 2018. Assessing climate vul- nerabilities and adaptive strategies for resilient beef and dairy operations in the tropics. Clim. Change 146(1–2):47–58. doi:10.1007/s10584-017-2110-1 Osei-Amponsah, R., S.S. Chauhan, B.J. Leury, L. Cheng, B. Cullen, I.J. Clarke, and F.R. Dunshea. 2019. Genetic selection for thermotolerance in rumin- ants. Animals (Basel). 9(11):18. doi:10.3390/ani9110948 Osei-Amponsah, R., F.R. Dunshea, B.J. Leury, L. Cheng, B. Cullen, A. Joy, A. Abhijith, M.H. Zhang, and S.S. Chauhan. 2020. Heat stress impacts on lactating cows grazing Australian summer pastures on an automatic ro- botic dairy. 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J. Biometeorol. 64(9):1613–1628. doi: 10.1007/s00484-020-01929-6 Zhang, M., R.D. Warner, F.R. Dunshea, K. DiGiacomo, A. Joy, A. Abhijith, R.  Osei-Amponsah, D.L.  Hopkins, M.  Ha, and S.S.  Chauhan. 2021. Impact of heat stress on the growth performance and retail meat quality of 2nd cross (Poll Dorset×(Border Leicester× Merino)) and Dorper lambs. Meat Sci. 181:108581. doi:10.1016/j.meatsci.2021.108581 Zhang, M., R.D. Warner, F.R. Dunshea, K. DiGiacomo, A. Joy, A. Abhijith, P.  Prathap, T.  Ma, and S.S.  Chauhan. 2022. Impact of Heatwaves on the Physiology and Retail Meat Quality of Lambs. Foods 11(3):414. doi:10.3390/foods11030414 About the Author(s) Surinder Singh Chauhan is a Senior Lecturer in Animal Science, School of Agriculture, Food and Ecosystem Sciences (SAFES), The University of Melbourne, Australia. Dr. Chauhan’s research program strives to deliver solutions to pro- mote sustainable and efficient livestock production systems cap- able of producing quality animal protein for consumers. Dr. Chauhan obtained his Bachelor and Master of Veterinary Science (Physiology) from HP Agriculture University Palampur, India, followed, PhD in Animal Sciences at The University of Melbourne and Postdoctoral in Muscle Biology and Meat Science at The Ohio State University, Columbus, USA. Dr. Chauhan is leading Australia–India col- laboration for sustainable livestock production. His current research is fo- cused on understanding the impact of climate change on farm animals and developing economic strategies to reduce the negative effects of heat stress on farm animal production and product quality. Surinder has many publi- cations, has presented papers, and has been keynote speaker at international conferences. Corresponding author: ss.chauhan@unimelb.edu.au Minghao Zhang is a Research Fellow in the Department of Food and Nutrition at the Provincial Hospital Affiliated with Shandong First Medical University. His main research focus is on the influence of cli- mate change on meat quality and human health. Michael obtained his PhD at School of Agriculture, Food and Ecosystem Sciences (SAFES), Faculty of Science, University of Melbourne, Australia, on comprehensive assessments of the effects of different ambient temperatures and heat exposure durations on sheep and chicken meat quality. He was also responsible for a research project on the relationship between climate change and body temperature management in patients with diabetes. D ow nloaded from https://academ ic.oup.com /af/article/13/5/60/7311134 by U niversity of G hana, Legon user on 04 D ecem ber 2023 https://doi.org/10.1016/j.smallrumres.2016.02.011 https://doi.org/10.1016/j.smallrumres.2016.02.011 https://doi.org/10.1016/j.smallrumres.2022.106662 https://doi.org/10.1016/j.smallrumres.2022.106662 https://doi.org/10.1093/af/vfy031 https://doi.org/10.3389/fvets.2020.00116 https://doi.org/10.3389/fvets.2020.00116 https://doi.org/10.1016/j.fochms.2021.100052 https://doi.org/10.3168/jds.2008-1072 https://doi.org/10.1093/jas/sky133 https://doi.org/10.1093/af/vfy029 https://doi.org/10.1016/j.smallrumres.2021.106523 https://doi.org/10.2527/jas2017.1843 https://doi.org/10.2527/jas.2016-0295 https://doi.org/10.1016/j.smallrumres.2021.106523 https://doi.org/10.1016/j.foodres.2009.05.018 https://doi.org/10.1016/j.meatsci.2003.08.001 https://doi.org/10.1016/j.meatsci.2007.11.022 https://doi.org/10.3390/nu12030734 https://doi.org/10.1016/j.meatsci.2020.108202 https://doi.org/10.1016/j.foodres.2017.09.027 https://doi.org/10.1016/0309-1740(81)90005-X https://doi.org/10.1007/s00484-021-02116-x https://doi.org/10.1007/s10584-017-2110-1 https://doi.org/10.3390/ani9110948 https://doi.org/10.3390/ani10050869 https://doi.org/10.5713/ajas.18.0310 https://doi.org/10.5713/ajas.18.0310 https://doi.org/10.1016/j.jtherbio.2014.08.004 https://doi.org/10.1111/jpn.12892 https://doi.org/10.1111/jpn.12892 https://doi.org/10.1080/09291016.2017.1386891 https://doi.org/10.3390/ani12111396 https://doi.org/10.3329/bjas.v43i2.20717 https://doi.org/10.3329/bjas.v43i2.20717 https://doi.org/10.1017/S1751731118001945 https://doi.org/10.1016/j.jveb.2016.10.009 https://doi.org/10.1007/s00484-020-02000-0 https://agris.fao.org/agris-search/search.do?recordID=QC1997066135 https://doi.org/10.1016/S2542-5196(22)00002-X https://doi.org/10.1016/S2542-5196(22)00002-X https://doi.org/10.1016/j.vetmic.2011.09.012 https://doi.org/10.5713/ajas.15.0693 https://doi.org/10.1007/s00484-020-01929-6 https://doi.org/10.1016/j.meatsci.2021.108581 https://doi.org/10.3390/foods11030414 mailto:ss.chauhan@unimelb.edu.au?subject= 68 Animal Frontiers Richard Osei-Amponsah is a William Gamble Visiting Fellow at The University of Melbourne Dookie Campus, School of Agriculture, Food and Ecosystem Sciences (SAFES), Faculty of Science, The University of Melbourne, Australia and an Associate Professor of Animal Science, University of Ghana. He holds a Doctor of Philosophy (PhD) degree in Animal Science (Animal Breeding and Genetics) from the University of Ghana, Legon. His research focuses on sustainable improvement and management of animal genetic resources and their adaptation to their production and changing environments. He has contributed to the development of sustainable animal breeding programs for low input-livestock systems, served as resource person at various stakeholder training workshops, supervised many students, and participated in sev- eral international and local conferences. Richard is an animal genetics consultant, has many publications to his credit, is a member of the All-Africa Society of Animal Production (AASAP), and is the current President of the Ghana Society of Animal Production (GSAP). Iain Clarke has been a Research Professor in the School of Agriculture, Food and Ecosystem Sciences, The University of Melbourne since 2018. Prior to this, he held a Fellowship from National Health and Medical Research (Australia) as an in- dependent scientist at Prince Henry’s Institute of Medical Research and then moved to Monash University, rising to the position of Head of Department of Physiology (2017–2017). He has graduated 18 PhD students. His original research was in the field of the neuroendocrinology of reproduction and other areas of neuroendocrinology (stress, metab- olism). Later, he developed a strong interest in the neuroendocrinology of obesity. He has published several peer-reviewed papers/chapters/re- views and has received several national and international awards. His current research interest is in the field of animal science, with a special emphasis on the identification of thermotolerant animals. He is strongly committed to producing scientific data that may be translated into on-farm benefits. Veerasamy Sejian, MVSc, PhD, is the Dean, Rajiv Gandhi Institute of Veterinary Education and Research (RIVER), Puducherry, India. Dr. Sejian is the team leader who established the concept of “Multiple stresses impacting small ruminant production in the changing climate scenario.” His current research is focused on identifying molecular markers for different environmental stresses in small ruminants with an objective of developing agro-ecological zone specific thermo-tolerant breeds. He has been the leader for an Indo-French collaborative project “Emerging Crop-Livestock Production Systems adapted to a Changing Environment (ECLIPSE Project).” Dr. Sejian has pub- lished three international Springer books on collaborative mode “Environmental stress and amelioration in livestock production”; “Climate change impact on livestock: adaptation and mitigation” and “Sheep production adapting to climate change.” For his outstanding contribution in climate change and livestock production, Indian Council of Agricultural Research (ICAR) has bestowed him with the prestigious Lal Bahadur Shastri Outstanding Young Scientist Award. Robyn Warner is a Professor in Meat Science, Faculty of Science, The University of Melbourne, Australia. Her research uses fundamental muscle biochem- istry and structure to investi- gate issues of importance to the meat industry. Her focus is on integrating novel techniques in biometrics and sensory to deeply understand consumer response to meat; developing capability and equipment to quantitate the flavor fingerprint of red meat in order to leverage consumer demand for unique experiences; understanding the effects of heat stress and climate change on meat quality; and developing fundamental under- standing of how muscle proteins change during animal growth, nutrition, preslaughter stress, postmortem biochemistry, packaging, processing, and cooking. She is actively driving engagement with industry and serves on Meat Standards Australia assuring quality to Australian consumers of red meat and has won national and international prizes for this role. Robyn has many publications and has undertaken research with impact. Frank Dunshea is a Redmond Barry Distinguished Professor and Chair of Agriculture at the University of Melbourne and Professor of Animal Growth and Development at the University of Leeds. He has a BAg Sci (Hons) and PhD from La Trobe University and postdoctoral training at Cornell University. He has more than 35 yr of re- search in growth physiology and nutrition and the use of domestic animals in nutritional and bio- medical research. His research has had a high scientific impact, with many of the results being rapidly adopted by industry. Professor Dunshea is a respected research leader in global livestock industries and is committed to ensuring that livestock industries operate in a responsible and sustainable manner. In addition to many awards, he is a Fellow of the Australian Nutrition Society, the Australasian Pig Science Association, the Australian Society of Animal Sciences, and the International Union of Nutritional Sciences. D ow nloaded from https://academ ic.oup.com /af/article/13/5/60/7311134 by U niversity of G hana, Legon user on 04 D ecem ber 2023