Hostname: page-component-cd9895bd7-8ctnn Total loading time: 0 Render date: 2024-12-26T17:36:07.686Z Has data issue: false hasContentIssue false

Heat stress in dairy calves from birth to weaning

Published online by Cambridge University Press:  03 August 2020

Mikolt Bakony
Affiliation:
Department of Animal Hygiene, Herd Health and Mobile Clinic, University of Veterinary Medicine, Budapest, Hungary
Viktor Jurkovich*
Affiliation:
Department of Animal Hygiene, Herd Health and Mobile Clinic, University of Veterinary Medicine, Budapest, Hungary
*
Author for correspondence: Viktor Jurkovich, Email: jurkovich.viktor@univet.hu
Rights & Permissions [Opens in a new window]

Abstract

This Research Reflection collects current knowledge on the effects of heat stress in dairy calves. Chapters cover the concept of foetal programming, animal-based and environmental indicators of heat stress in the postnatal period, and methods of heat stress abatement. Conclusions for further research about economic efficiency, research methodology and an integrated approach of pre- and postnatal heat stress are also proposed.

Type
Research Reflection
Creative Commons
Creative Common License - CCCreative Common License - BY
This is an Open Access article, distributed under the terms of the Creative Commons Attribution licence (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted re-use, distribution, and reproduction in any medium, provided the original work is properly cited.
Copyright
Copyright © The Author(s), 2020. Published by Cambridge University Press on behalf of Hannah Dairy Research Foundation

Heat stress is one of the main challenges facing the dairy production industry. Physiological and behavioural coping mechanisms of lactating dairy cows are well documented (Polsky and von Keyserlingk, Reference Polsky and von Keyserlingk2017). However, the thermal status of hutch reared calves receives less attention from a scientific (Roland et al., Reference Roland, Drillich, Klein-Jöbstl and Iwersen2016), and even less so from a management standpoint. This research reflection aims to gather current knowledge about the effects of heat stress and the methods of heat alleviation in preweaned Holstein friesian dairy calves. Biological and environmental indicators of heat stress and methods of heat abatement are discussed, and targets of future research are proposed.

Indicators of heat stress in the prenatal period

There is growing evidence that the uterine environment of dry cows can convey an indirect effect of environmental stress and evoke adaptive mechanisms in the calf foetus. Signs of adaptation are present also in the postnatal period, which lead to the concept often called ‘foetal programming’. Earlier studies have observed that sensitivity to thermal stress is higher in periods of reproduction and neonatal life, as compared to other phases of the life cycle (Collier et al., Reference Collier, Beede, Thatcher, Israel and Wilcox1982). Effects of maternal heat stress on the growing foetus have been extensively studied by researchers at the Calf Unit of the University of Florida (Gainesville, USA). In the past years, the adaptive responses of the foetus have been elucidated in more detail.

Lower birth weight and adult height

Foetal growth is compromised due to hyperthermia-induced placental insufficiency. Reduced placenta size and function limit the maternal-foetal exchange of oxygen and nutrients. Even a few days shortening of gestation length, that often occurs in times of heat stress (Dahl et al., Reference Dahl, Tao and Monteiro2016), shortens the period of rapid foetal growth resulting in reduced birth weight. Calves born from dams exposed to heat stress had lower weaning weight than calves from cooled dams. However, pre-weaning weight gain, and body weight in the prepubertal period were not different (Tao et al., Reference Tao, Monteiro, Thompson, Hayen and Dahl2012; Monteiro et al., Reference Monteiro, Tao, Thompson and Dahl2014). Despite the postpubertal rebound in weight gain, adult height of calves born from heat-stressed dams did not reach that of calves born from dams cooled in the dry period (Monteiro et al., Reference Monteiro, Tao, Thompson and Dahl2014).

Metabolic shift

Heat stress impairs not only the uterine supply of nutrients but also the heat exchange between the dam and the foetus. The foetus has double the metabolic rate as the mother, that is why a narrower temperature gradient can result in foetal hyperthermia. As seen in the sheep model, the foetus can develop adaptive mechanisms at the expense of growth. These include reduced protein accretion in favour of hepatic gluconeogenesis as well as an increased level of catabolic and reduced level of anabolic hormones. The same diet has induced higher insulin concentrations in calves born to heat-stressed dams than those born to cooled cows. The increased insulin response suggests a carryover effect of maternal heat stress (Tao and Dahl, Reference Tao and Dahl2013). Calves born from cows not cooled in the dry period showed similar pancreatic insulin sensitivity and systemic insulin clearance at weaning age to that of calves born from cows cooled in the dry period, but a more rapid glucose clearance during both a glucose tolerance test and an insulin challenge (Tao et al., Reference Tao, Monteiro, Hayen and Dahl2014). Dahl et al. (Reference Dahl, Tao and Monteiro2016) concluded that calves experiencing heat stress in utero are prone to develop a smaller mature body size and more fat reserves than counterparts in thermoneutrality.

Impaired immune function

In the first 28 d of life, serum IgG concentrations and apparent efficiency of IgG absorption were lower in calves born from heat-stressed dams relative to calves born from cooled dams. Heat stress in late gestation has no evident effect on IgG content of colostrum. It suggests that impaired IgG absorption is presumably due to the deficiency of passive transfer (Tao et al., Reference Tao, Monteiro, Thompson, Hayen and Dahl2012; Monteiro et al., Reference Monteiro, Tao, Thompson and Dahl2014). However, acute brief heat stress during late gestation did not alter passive antibody transfer capacity in calves (Strong et al., Reference Strong, Silva, Cheng and Eicher2015). The proliferation rate of mononuclear cells was lower in calves born from heat-stressed dams, as compared to the offspring of cooled dams. However, antibody production in an ovalbumin challenge at 28 d of age was similar in both groups. Both humoral immune response and cell-mediated immune function seem to be altered by heat stress.

Remarks

The key findings of research studies on heat stress in utero bring deserved attention to dry cow management and urge active cooling throughout the nonlactating period. Given that environmental stressors can induce the compensatory hypervascularization of the uterine horn and the alteration of ovarian activity (Collier et al., Reference Collier, Beede, Thatcher, Israel and Wilcox1982), further studies are needed to distinguish the maternal and foetal components of hyperthermia-induced intrauterine growth retardation. The metabolic shift in calves heat-stressed in utero makes them prone to preserve energy and acquire less lean tissue. Such a phenomenon is also observed in heat-stressed lactating cows, where increased insulin action is linked to the ‘leaky gut’ syndrome. It is worth investigating in which ways are the two mechanisms different. A clear distinction between the effects of prenatal and perinatal stress, if possible, would also contribute to improved newborn calf management practices.

Indicators of heat stress in the postnatal period

Just as subtle differences in the uterine environment of cooled and noncooled pregnant cows can induce prolonged effects in the calf foetus, severe heat load experienced after birth may also affect performance in the rearing period. However, the term heat stress is used quite loosely. Accurately assessing the amount of strain environmental conditions impose on dairy calves is challenging. As opposed to dairy cows, no clearly defined thresholds of biological or environmental indicators are commonly accepted for dairy calves that would reliably pinpoint the onset of production losses and thus necessitate cooling interventions. The animal-based indices of assessing thermal status proposed in the literature are discussed below.

Acute stress response parameters

Heart rate variability analysis confirmed that calves exposed to solar radiation had a higher sympathetic tone than shaded calves (Kovács et al., Reference Kovács, Kézér, Ruff, Jurkovich and Szenci2018c). Endocrine changes also suggest an increased level of stress due to heat exposure. In a study on preweaned calves exposed to heat load, salivary and plasma cortisol concentrations were elevated indicating an increased level of stress (López et al., Reference López, Mellado, Martínez, Véliz, García, de Santiago and Carrillo2018; Kovács et al., Reference Kovács, Kézér, Ruff, Szenci, Bakony and Jurkovich2019). Plasma concentrations of thyroid hormones T3 and T4 were lower in heat stress (López et al., Reference López, Mellado, Martínez, Véliz, García, de Santiago and Carrillo2018).

Behavioural responses

Altered behaviour is the first sign of thermal discomfort. Calves seek shade, change posture, move less during the hottest hours of the day and bunch to provide shade for each other (Roland et al., Reference Roland, Drillich, Klein-Jöbstl and Iwersen2016). Frequency of changing posture is reduced in hot conditions as a sign of discomfort (Kovács et al., Reference Kovács, Kézér, Bakony, Jurkovich and Szenci2018a) similarly to cows (Allen et al., Reference Allen, Hall, Collier and Smith2015). Hutch vs. pen preference or lying vs. standing provides valuable information about the heat-absorbing nature of hutch material or thermal conductive properties of the bedding.

Increased respiratory rates

Increased respiratory frequency promotes evaporative heat loss. Textbooks, publications and online guides describe rates of 20–40 to even 50–70 breaths/min as physiological (Rosenberg, Reference Rosenberg1979; Piccione et al., Reference Piccione, Caola and Refinetti2003). Studies on adaptive responses of calves in neutral/shaded vs. hot/noncooled thermal environments have reported an approx. 50% increase in average respiratory rates as a sign of increased evaporative cooling efforts [from 47 to 53 (de Lima et al., Reference de Lima, de Souza, de Lima, de Sousa Oliveira, Domingos, Tholon and de Miranda2013), from 50–78 to 73–105 (Peña et al., Reference Peña, Risco, Kunihiro, Thatcher and Pinedo2016) or from 30–50 to 70–140 (Kovács et al., Reference Kovács, Kézér, Ruff, Jurkovich and Szenci2018b)]. Heavier breathing is induced by an increase in ambient and consequently, body surface temperature. The elevation of respiratory frequency thus precedes the rise in core body temperature, which must be taken into account when assessing heat stress status.

Elevated rectal temperature

In thermoneutrality, mammals can maintain their physiological body temperature without increased efforts of heat dissipation or heat production. Most sources consider 38.5–39.1(39.5)°C as the range of healthy body temperature in calves (Rosenberg, Reference Rosenberg1979; Piccione et al., Reference Piccione, Caola and Refinetti2003). Consistently, studies on calves exposed to high ambient temperatures report on maximal body temperatures of 39.7°C (de Lima et al., Reference de Lima, de Souza, de Lima, de Sousa Oliveira, Domingos, Tholon and de Miranda2013), 40.1°C (Peña et al., Reference Peña, Risco, Kunihiro, Thatcher and Pinedo2016), 40.4°C (Kovács et al., Reference Kovács, Kézér, Ruff, Jurkovich and Szenci2018b), and 39.8°C (Hill et al., Reference Hill, Bateman, Suarez-Mena, Dennis and Schlotterbeck2016).

Water consumption

Water requirement is elevated in hot weather (Broucek et al., Reference Broucek, Kisac and Uhrincat2009), as calves may lose water via increased respiration and sweating. As the ambient temperature rises from 0 to 35°C, water intake increases almost 4-fold, from 1.4 l/d to around 4 l/d, in addition to the amount of milk replacer (Quigley, Reference Quigley2001). Making sure each calf is aware that water is available is crucial in preventing dehydration. Moreover, Wiedmeier et al. (Reference Wiedmeier, Young and Hammon2005) showed that increased frequency of changing and rinsing water buckets resulted in a higher average daily gain in the preweaning period.

Early mortality

The biological cost of adaptation to prolonged heat exposure can impact calf welfare and the profitability of rearing. High ambient temperature, especially in calves housed outdoors proved to be a risk factor for early calf mortality in veal calves (Renaud et al., Reference Renaud, Duffield, LeBlanc, Ferguson, Haley and Kelton2018). Extreme heatwaves can cause excess death of different cattle subpopulations, including dairy calves (Morignat et al., Reference Morignat, Perrin, Gay, Vinard, Calavas and Hénaux2014). Research data are, however, inconsistent, as others showed mortality of 1–21 d old Holstein calves to be higher in moderate conditions than in the hot season (Mellado et al., Reference Mellado, Lopez, Veliz, De Santiago, Macias-Cruz, Avendaño-Reyes and Garcia2014). The term ‘early mortality’ is rather unspecific, though. Death occurring in the preweaning period has multiple causes. Further data on the prevalence of different causes of death could highlight areas that need more attention in periods of hot weather.

Reduced feed intake and weight gain

The small number of studies seasonal effects of growth in dairy calves all agree on a lower average daily weight gain in seasons with higher ambient temperature (Donovan et al., Reference Donovan, Dohoo, Montgomery and Bennett1998; Broucek et al., Reference Broucek, Kisac and Uhrincat2009; López et al., Reference López, Mellado, Martínez, Véliz, García, de Santiago and Carrillo2018). The reduced growth rate is attributed mainly to reduced starter intake in the hottest periods of the year (Bateman et al., Reference Bateman, Hill, Aldrich, Schlotterbeck and Firkins2012; Holt, Reference Holt2014), rather than consequences of maternal heat stress. Given that dry cow heat abatement is an overlooked area in dairy farming, it is tempting to speculate that prenatal heat stress effects possibly mask postnatal heat stress responses. However, it was shown that postnatal thermal conditions (cooling vs. no cooling) dominate in calf welfare and performance in the preweaning period, irrespective of prenatal thermal status (cooled vs. not cooled dams) (Dado-Senn et al., Reference Dado-Senn, Vega Acosta, Torres Rivera, Field, Marrero, Davidson, Tao, Fabris, Ortiz-Colón, Dahl and Laporta2020).

Remarks

Heat stress affects many physiological and production indicators in calves. It is essential to determine which of these is considered valid for defining heat stress. Animal-based indices are the primary measures of animal welfare. Having the primary indicators identified, the physiological thresholds (if not already available) for defining heat stress should be determined. Production indicators are also relevant; however, the authors are of the opinion that importance is only secondary to animal-based indicators. We believe that the principles of animal welfare should prevail and that this will be economical in the long term.

Thermoneutral zone and measurements of environmental heat load

Heat stress abatement of hutch-reared dairy calves is largely ignored in dairy management. However, maintaining constant body temperature in conditions of high ambient temperature, intense solar radiation and high relative humidity is not possible without expending extra energy. Knowing the factors that affect thermoregulation of the calf promotes housing and environmental modifications that could save energy for growth and health.

Thermoneutral zone not clearly defined

There is far more information on the effects of cold on welfare than there is on the upper critical temperature of dairy calves. Several different ambient temperatures were reported as set points of increased evaporative heat dissipation. Gebremedhin et al. (Reference Gebremedhin, Cramer and Porter1981) observed increased respiration as temperature exceeded 20°C. Other researchers agreed on 26°C as the upper critical temperature of preweaned calves (Spain and Spiers, Reference Spain and Spiers1996; Holt, Reference Holt2014; Collier et al., Reference Collier, Baumgard, Zimbelman and Xiao2019). Neuwirth et al. (Reference Neuwirth, Norton, Rawlings, Thompson and Ware1979) observed the first signs of heat stress at 32°C, 60% relative humidity.

Environmental indicators of heat stress

Ambient temperature is accepted to be the sole reliable indicator of the thermal environment of calves in most heat stress studies. In dairy cows, the effect of relative humidity on heat dissipation capacity is well documented, and that knowledge has been incorporated in the temperature-humidity index (THI), the weighted estimator of environmental heat load. It shows a strong correlation to biomarkers of heat stress (Bouraoui et al., Reference Bouraoui, Lahmar, Majdoub, Djemali and Belyea2002; Dikmen and Hansen, Reference Dikmen and Hansen2009; Bernabucci et al., Reference Bernabucci, Lacetera, Baumgard, Rhoads, Ronchi and Nardone2010). Attempts have been made to adopt the THI in calf studies (Peña et al., Reference Peña, Risco, Kunihiro, Thatcher and Pinedo2016; Manriquez et al., Reference Manriquez, Valenzuela, Paudyal, Velasquez and Pinedo2018); however, its reliability is limited. Little is known about how relative humidity affects the heat dissipation of dairy calves. THI formulas and thresholds – originally adapted for lactating cattle – do not carry more information than ambient temperature alone.

Quantifying radiant heat

In outdoor conditions, radiant heat and wind speed are determining factors in the operative temperature, that is the temperature perceived by the animal. The THI incorrectly estimates the environmental heat load in hutch reared calves, as it does not incorporate the radiant temperature and airspeed. The use of complex environmental indices was proposed for outdoor measurements by several reports (Gaughan et al., Reference Gaughan, Mader, Holt and Lisle2008; Mader et al., Reference Mader, Johnson and Gaughan2010; Hammami et al., Reference Hammami, Bormann, M'hamdi, Montaldo and Gengler2013); however, it has not yet been adopted in studies on dairy calves.

Remarks

The environmental thresholds to decide between thermoneutrality or heat stress in dairy calves have not been sufficiently defined. Adequate definitions and limits are needed to judge the real effects of heat stress and to measure the efficiency of heat stress control.

Techniques to decrease heat load in calves

From birth until weaning, most calves are kept outdoors, in individual hutches with small fenced exercise pens. In summer, the microclimate of polyethylene hutches, even if placed under shade, is worse than that of plywood hutches (Lammers et al., Reference Lammers, VanKoot, Heinrichs and Graves1996; Peña et al., Reference Peña, Risco, Kunihiro, Thatcher and Pinedo2016). Rectal temperature and respiration rates were higher in calves housed in plastic hutches as compared to plywood, however, no differences in weight gain or general health status were observed (Lammers et al., Reference Lammers, VanKoot, Heinrichs and Graves1996; Peña et al., Reference Peña, Risco, Kunihiro, Thatcher and Pinedo2016). The practicality of durable and more hygienic plastic and fibreglass hutches make them the most popular type of housing for outdoor reared calves worldwide. Thermal properties of the plastic used during the manufacturing of hutches are improving, however, it is still necessary to reduce heat load and heat absorption of plastic and fibreglass hutches in summer, by means of additional shade (Andrews and Davison, Reference Andrews and Davison2002).

Increasing airflow

Increasing airspeed could help heat dissipation of the calves. Elevation of the rear side of the hutches is showed to increase airspeed and decrease CO2 concentration inside the hutch, making it apparently more comfortable for the indwelling calf (44 vs. 58 breaths/min, compared to the control) (Moore et al., Reference Moore, Duprau and Wenz2012). The specific design of hutches to maximize ventilation – including ridge-top vents and adjustable vent doors – are practical alternatives to labour-intensive manipulation of hutches. The use of fans provide a favourable microclimate (Hill et al., Reference Hill, Bateman, Aldrich and Schlotterbeck2011; Dado-Senn et al., Reference Dado-Senn, Vega Acosta, Torres Rivera, Field, Marrero, Davidson, Tao, Fabris, Ortiz-Colón, Dahl and Laporta2020), but this is limited to indoor conditions and thus not widespread.

Orientation of hutches

The orientation of calf hutches affected inner microclimate and consequently, the heat load of calves on sunny days (Bakony et al., Reference Bakony, Kiss and Jurkovich2019). Respiration rate was elevated in all four groups, being highest in south-facing hutches (103/min vs. 90/min). Probabilities of a calf lying, being inside the hutch or seeking shade at the time of observation, respectively, were highest in the group of hutches facing south. The level of heat stress in south-facing hutches is attributed to exposure to the most intense solar radiation and the least amount of shade during the day. Individual calf hutches should be positioned to face north or east in the summer period. Oriented alignment of calf hutches could serve as a no-cost measure of improving calf welfare.

Reflective covers

Friend et al. (Reference Friend, Haberman and Binion2014) tested different radiant barriers. The silver painting was practically ineffective, while laminates and aluminized plastic covers decreased black globe temperature by 2–4°C in empty hutches. Carter et al. (Reference Carter, Friend, Garey, Sawyer, Alexander and Tomazewski2014) found that reflective covers provided a more favourable inner climate at both low and high ambient THI. Increase in respiration rate and ear canal temperature of the calves, relative to THI, were moderate in insulated hutches. Average daily gain did not differ between calves housed in covered or uncovered hutches. Other studies doubt the advantages of reflective covers. Manriquez et al. (Reference Manriquez, Valenzuela, Paudyal, Velasquez and Pinedo2018) found that average THI and ambient temperature were higher (68.6 vs. 67.6, and 23.2 vs. 22.8°C, respectively) in the hutches covered with aluminized plastic material. However, rectal temperature and respiratory rate were not different in control and experimental calves. The authors supposed that reflective covers impede the cooling of the hutch material in the evening hours.

Shading structures

Shading is evidentially more effective in decreasing exposure to solar radiation than reflective covers. Shading reduces the temperature both inside and outside the hutch (Coleman et al., Reference Coleman, Moss and McCaskey1996; Spain and Spiers, Reference Spain and Spiers1996; Gu et al., Reference Gu, Yang, Leng, Xu, Tang, Liu, Gao and Mao2016; Kovács et al., Reference Kovács, Kézér, Ruff, Szenci, Bakony and Jurkovich2019). Respiratory rate of calves is usually lower under shade (Spain and Spiers, Reference Spain and Spiers1996; Gu et al., Reference Gu, Yang, Leng, Xu, Tang, Liu, Gao and Mao2016) and calves spend more time lying in shaded areas (Gu et al., Reference Gu, Yang, Leng, Xu, Tang, Liu, Gao and Mao2016). Shading also provides more comfortable conditions for caretakers in all seasons (Coleman et al., Reference Coleman, Moss and McCaskey1996). In practice, greenhouse shade nets (80–85% shade rate) installed at the height of 2 m are one way of providing shade (Coleman et al., Reference Coleman, Moss and McCaskey1996; Spain and Spiers, Reference Spain and Spiers1996; Kovács et al., Reference Kovács, Kézér, Ruff, Szenci, Bakony and Jurkovich2019). Thatch shading or well-grown trees can also be effective (Kamal et al., Reference Kamal, Dutt, Patel, Dey, Chandran, Barari, Chakrabarti and Bhusan2014). A built roof is a bigger investment for dairy operations; however, it offers protection from precipitation and build-up of radiant heat, while maintaining adequate airflow. Calf mortality rates were observed to drop after installation of a roof (personal observation).

Remarks

Of the techniques used to reduce the heat load, shading is the most effective, however, not widespread. The low-cost measures are flexible but rarely hard-wearing. A permanent solution is, still, a more considerable financial investment that requires proven results on the favourable effects. In the authors' opinion, roof construction is certainly a desirable goal in protecting the calves against solar radiation. Longitudinal studies on the impacts of shading would support the economic feasibility of roof installations.

Nutritional management in heat stress

We have shown that increased energy demands of heat dissipation coupled with reduced starter intake often result in the reduced growth rate of calves in summer months. In support of weight gain, increasing the plane of nutrition or the use of different feed additives are the main strategies for nutritional interventions.

Preference for liquid feed and water

Considering that calves prefer liquid feed over solids in hot weather, a promising approach is to increase the energy content of milk replacer. Increased feeding rate of milk replacer (0.66 kg vs. 0.44 kg dry matter/d, 21% crude protein, 21% fat) increased average daily gain and hip width in calves raised in summer (Hill et al., Reference Hill, Bateman, Aldrich and Schlotterbeck2012). An accelerated milk replacer feeding programme (0.66 or 0.77 kg dry matter of milk replacer daily, 26% crude protein, 17% fat) improved energy intake and weight gain in 3–56 d-old calves during summer (Orellana Rivas et al., Reference Orellana Rivas, Komori, Beihling, Marins, Bernard and Tao2020). Increasing dietary fat content of the milk replacer from 10 to 20% (besides 20% crude protein) yielded higher body weight body size of weaned calves (Blair, Reference Blair2015). Adding water to dry calf starter was also shown to increase palatability. Starter intake and average daily gain of calves increased when feeding 75 or 50% dry matter vs. 90% dry matter diets (Beiranvand et al., Reference Beiranvand, Khani, Omidian, Ariana, Rezvani and Ghaffari2016). As mentioned in an earlier chapter, merely the regular provision of fresh, clean water yielded higher weight gain in hutch-reared calves (Wiedmeier et al., Reference Wiedmeier, Young and Hammon2005)

Vitamins, minerals, yeasts

Trials on the use of different feed additives yielded varying results. Supplementation with fat-soluble and B-vitamins, omega fatty acids and minerals did not provide additional benefit to summer reared calves (Blair, Reference Blair2015). A combination of vitamins A and E and microelements increased growth performance post-weaning, enhanced immune functions and antioxidant capacity (Bordignon et al., Reference Bordignon, Volpato, Glombowsky, Souza, Baldissera, Secco, Pereira, Leal, Vedovatto and Da Silva2019). An exciting approach was adding Saccharomyces boulardii to milk replacer to calves between 1–28 d of age (Lee et al., Reference Lee, Kacem, Kim, Peng, Kim, Joung, Lee and Lee2019). It resulted in higher dry matter intake and improved gut health during thermoneutrality and higher dry matter intake, but lower rectal temperature and cortisol levels in experimental heat stress than that of untreated controls (Lee et al., Reference Lee, Kacem, Kim, Peng, Kim, Joung, Lee and Lee2019). The authors explained the results with the balancing effect of yeast supplementation on intestinal flora that lowered lipopolysaccharide absorption due to the leaky gut syndrome in times of heat stress.

Similarly to dairy cows, the effects of chromium supplementation was also studied in dairy calves. It is known to potentiate insulin action and enhance glucose metabolism. In the study of Kargar et al. (Reference Kargar, Mousavi, Karimi-Dehkordi and Ghaffari2018), oral supplementation with chromium in a dose of 0.05 mg/kg body weight was associated with greater meal sizes and longer meal durations. The respiratory rates of supplemented calves were lower than that of unsupplemented calves in high ambient temperatures. Overall average daily gain and body weight at weaning was higher in Cr supplemented calves; however, the difference gradually diminished after weaning.

Remarks

Strategies for nutritional alleviation of heat stress are, at present, not numerous in calves, contrary to that in dairy cows. However, results are encouraging, since certain cost-effective adjustments in the feeding regime can promote appetite or provide the extra energy in times of increased maintenance requirements. While, in dairy cows, nutritional interventions can only be secondary to adequate cooling technologies, seasonal adjustments in the feeding protocol are advisable in calf rearing. The physiological chromium requirements of ruminants are not precisely known. Thus the growth-promoting effects of chromium supplementation can partially be due to satisfying by chromium-deficiency of control animals; however, this hypothesis needs further testing. Despite the promising results, trivalent chromium is not authorized as a feed additive in the European Union, based on the scientific opinion of the European Food Safety Authority (EFSA, 2009).

Conclusions for future research

Economic efficiency

Despite the growing body of evidence of adverse effects of heat stress on dairy calves from as early as the prenatal period, most dairy operations carry on without any cooling interventions for dry cows or preweaned calves. Translating the biological cost to financials could convince farm owners to invest in some kind of heat abatement. It could also speed up the much-needed change of thinking in dairy (calf) management, that non-lactating animals require as much attention as lactating animals do.

Understanding the basics

Scarce literature on the upper end of the thermoneutral zone warns that there is room for improvement in understanding thermal requirements and heat dissipation capacities of dairy calves. The indices initially developed for indoor conditions, like dry bulb temperature or the temperature-humidity index, can be misleading when assessing the thermal environment of outdoor reared calves. Upper critical thresholds should be formulated in a manner that suits the housing environment of calves. It necessitates a better understanding of how radiant heat, relative humidity and wind speed contribute to the thermal load of dairy calves.

Integrating the concept of in utero heat stress

Besides dry cow management, heat stress abatement in calf rearing is another overlooked area in dairy management. A longitudinal study involving a larger number of calves could shed light on whether adverse effects of pre- and postnatal heat stress are comparable and whether these effects add up when occurring. It would be interesting to study whether postnatal heat exposure without maternal heat stress results in the same adaptive metabolic and immune responses as that of the calf foetus. It is also worth investigating whether improved calf management and nutrition strategies could prove useful in mitigating the effects of heat stress on growth and passive transfer of immunoglobulins.

Methodology

Real-time recording of environmental indices (radiant heat, humidity, wind speed) is feasible and could be easily integrated into precision livestock farming technologies (Fournel et al., Reference Fournel, Rousseau and Laberge2017; Koltes et al., Reference Koltes, Koltes, Mote, Tucker and Hubbell2018). Automated monitoring of physiological parameters in outdoor kept calves is currently not widely available, due to high costs or limited time of recording (10–14 d for indwelling thermometers). Respiratory rate can only be measured by labour-intensive visual observation. Adaptation of automated methods of measuring breathing rate – designed initially for cattle – would improve reliability and facilitate the determination of upper critical temperatures (Eigenberg et al., Reference Eigenberg, Hahn, Nienaber, Brown-Brandl and Spiers2000; Strutzke et al., Reference Strutzke, Fiske, Hoffmann, Ammon, Heuwieser and Amon2019).

Acknowledgements

This article is based upon work from COST Action FA1308 DairyCare, supported by COST (European Cooperation in Science and Technology, www.cost.eu). COST is a funding agency for research and innovation networks. COST Actions help connect research initiatives across Europe and enable scientists to grow their ideas by sharing them with their peers. This boosts their research, career and innovation. The Project was supported by the European Union and co-financed by the European Social Fund (grant agreement no. EFOP-3.6.2-16-2017-00012, project title: Development of a product chain model for functional, healthy and safe foods from farm to fork based on a thematic research network). Mikolt Bakony was supported by the UNKP-18-3 New National Excellence Program of the Ministry of Human Capacities (MHC). Viktor Jurkovich was supported by the János Bolyai Research fellowship by the Hungarian Academy of Science (BO/29/16/4) and the UNKP-18-4 New National Excellence Program of the MHC.

References

Allen, JD, Hall, LW, Collier, RJ and Smith, JF (2015) Effect of core body temperature, time of day, and climate conditions on behavioural patterns of lactating dairy cows experiencing mild to moderate heat stress. Journal of Dairy Science 98, 118127.CrossRefGoogle ScholarPubMed
Andrews, J and Davison, T (2002) Dairy farm layout and design: building and yard design, warm climates. In Fuquay JW (ed.) Encyclopedia of Dairy Sciences, 2nd Edn., Cambridge, MA, USA: Academic Press, pp. 1328.Google Scholar
Bakony, M, Kiss, G and Jurkovich, V (2019) The effect of hutch orientation on primary heat stress responses of dairy calves. In Wickens S, Hubrecht R and Golledge H (Eds) Advancing Animal Welfare Science: How Do We Get There? – Who Is It Good For? Proceedings of UFAW International Animal Welfare Science Symposium, 3–4 July 2019, Bruges, Belgium, UFAW, Wheathampsterad, UK p. 52.Google Scholar
Bateman, HG, Hill, TM, Aldrich, JM, Schlotterbeck, RL and Firkins, JL (2012) Meta-analysis of the effect of initial serum protein concentration and empirical prediction model for growth of neonatal Holstein calves through 8 weeks of age. Journal of Dairy Science 95, 363369.CrossRefGoogle ScholarPubMed
Beiranvand, H, Khani, M, Omidian, S, Ariana, M, Rezvani, R and Ghaffari, MH (2016) Does adding water to dry calf starter improve performance during summer? Journal of Dairy Science 99, 19031911.CrossRefGoogle ScholarPubMed
Bernabucci, U, Lacetera, N, Baumgard, LH, Rhoads, RP, Ronchi, B and Nardone, A (2010) Metabolic and hormonal acclimation to heat stress in domesticated ruminants. Animal: An International Journal of Animal Bioscience 4, 11671183.Google ScholarPubMed
Blair, SJ (2015) Effects of milk replacer and multivitamin-mineral supplementation on performance of heat stressed dairy calves. Louisiana State University.Google Scholar
Bordignon, R, Volpato, A, Glombowsky, P, Souza, CF, Baldissera, MD, Secco, R, Pereira, WAB, Leal, MLR, Vedovatto, M and Da Silva, AS (2019) Nutraceutical effect of vitamins and minerals on performance and immune and antioxidant systems in dairy calves during the nutritional transition period in summer. Journal of Thermal Biology 84, 451459.CrossRefGoogle Scholar
Bouraoui, R, Lahmar, M, Majdoub, A, Djemali, M and Belyea, R (2002) The relationship of temperature-humidity index with milk production of dairy cows in a Mediterranean climate. Animal Research 51, 479491.CrossRefGoogle Scholar
Broucek, J, Kisac, P and Uhrincat, M (2009) Effect of hot temperatures on the hematological parameters, health and performance of calves. International Journal of Biometeorology 53, 201208.Google ScholarPubMed
Carter, BH, Friend, TH, Garey, SM, Sawyer, JA, Alexander, MB and Tomazewski, MA (2014) Efficacy of reflective insulation in reducing heat stress on dairy calves housed in polyethylene calf hutches. International Journal of Biometeorology 58, 5159.Google ScholarPubMed
Coleman, DA, Moss, BR and McCaskey, TA (1996) Supplemental shade for dairy calves reared in commercial calf hutches in a southern climate. Journal of Dairy Science 79, 20382043.CrossRefGoogle Scholar
Collier, RJ, Beede, DK, Thatcher, WW, Israel, LA and Wilcox, CJ (1982) Influences of environment and its modification on dairy animal health and production. Journal of Dairy Science 65, 22132227.CrossRefGoogle Scholar
Collier, RJ, Baumgard, LH, Zimbelman, RB and Xiao, Y (2019) Heat stress: physiology of acclimation and adaptation. Animal Frontiers 9, 1219.CrossRefGoogle ScholarPubMed
Dado-Senn, B, Vega Acosta, L, Torres Rivera, M, Field, SL, Marrero, MG, Davidson, BD, Tao, S, Fabris, TF, Ortiz-Colón, G, Dahl, GE and Laporta, J (2020) Pre- and postnatal heat stress abatement affects dairy calf thermoregulation and performance. Journal of Dairy Science 103, 48224837.Google ScholarPubMed
Dahl, GE, Tao, S and Monteiro, APA (2016) Effects of late-gestation heat stress on immunity and performance of calves. Journal of Dairy Science 99, 31933198.Google ScholarPubMed
de Lima, PO, de Souza, JBFJ, de Lima, RN, de Sousa Oliveira, FC, Domingos, HGT, Tholon, P and de Miranda, MVFG (2013) Effect of time of day and type of shading on the physiological responses of crossbred calves in tropical environment. Journal of Animal Behavior and Biometeorology 1, 712.CrossRefGoogle Scholar
Dikmen, S and Hansen, PJ (2009) Is the temperature-humidity index the best indicator of heat stress in lactating dairy cows in a subtropical environment? Journal of Dairy Science 92, 109116.CrossRefGoogle Scholar
Donovan, GA, Dohoo, IR, Montgomery, DM and Bennett, FL (1998) Calf and disease factors affecting growth in female Holstein calves in Florida, USA. Preventive Veterinary Medicine 33, 110.Google ScholarPubMed
EFSA (2009) Scientific opinion of the panel on additives and products or substances used in animal feed (FEEDAP) on a request from the European Commission on the safety and efficacy of chromium methionine (Availa® Cr) as feed additive for all species. EFSA Journal 1043, 169.Google Scholar
Eigenberg, RA, Hahn, GL, Nienaber, JA, Brown-Brandl, TM and Spiers, DE (2000) Development of a new respiration rate monitor for cattle. Transactions of the ASAE 43, 723728.Google Scholar
Fournel, S, Rousseau, AN and Laberge, B (2017) Rethinking environment control strategy of confined animal housing systems through precision livestock farming. Biosystems Engineering 155, 96123.Google Scholar
Friend, TH, Haberman, JA and Binion, WR (2014) Effect of four different reflective barriers on black-globe temperatures in calf hutches. International Journal of Biometeorology 58, 21652168.CrossRefGoogle ScholarPubMed
Gaughan, JB, Mader, TL, Holt, SM and Lisle, A (2008) A new heat load index for feedlot cattle. Journal of Animal Science 86, 226234.CrossRefGoogle ScholarPubMed
Gebremedhin, KG, Cramer, CO and Porter, WP (1981) Predictions and measurements of heat production and food and water requirements of Holstein calves in different environments. Transactions of the ASAE 24, 715720.CrossRefGoogle Scholar
Gu, Z, Yang, S, Leng, J, Xu, S, Tang, S, Liu, C, Gao, Y and Mao, H (2016) Impacts of shade on physiological and behavioural pattern of Dehong buffalo calves under high temperature. Applied Animal Behaviour Science 177, 15.CrossRefGoogle Scholar
Hammami, H, Bormann, J, M'hamdi, N, Montaldo, HH and Gengler, N (2013) Evaluation of heat stress effects on production traits and somatic cell score of Holsteins in a temperate environment. Journal of Dairy Science 96, 18441855.CrossRefGoogle Scholar
Hill, TM, Bateman, HG, Aldrich, JM and Schlotterbeck, RL (2011) Comparisons of housing, bedding, and cooling options for dairy calves. Journal of Dairy Science 94, 21382146.CrossRefGoogle ScholarPubMed
Hill, TM, Bateman, HG, Aldrich, JM and Schlotterbeck, RL (2012) Case study: effect of feeding rate and weaning age of dairy calves fed a conventional milk replacer during warm summer months. Professional Animal Scientist 28, 125130.CrossRefGoogle Scholar
Hill, TM, Bateman, HG, Suarez-Mena, FX, Dennis, TS and Schlotterbeck, RL (2016) Short communication: changes in body temperature of calves up to 2 months of age as affected by time of day, age, and ambient temperature. Journal of Dairy Science 99, 88678870.Google ScholarPubMed
Holt, S (2014) Ambient temperature, calf intakes, and weight gains on preweaned dairy calves. Utah State University.Google Scholar
Kamal, R, Dutt, T, Patel, BHM, Dey, A, Chandran, PC, Barari, SK, Chakrabarti, A and Bhusan, B (2014) Effect of shade materials on microclimate of crossbred calves during summer. Veterinary World 7, 776783.Google Scholar
Kargar, S, Mousavi, F, Karimi-Dehkordi, S and Ghaffari, MH (2018) Growth performance, feeding behavior, health status, and blood metabolites of environmentally heat-loaded Holstein dairy calves fed diets supplemented with chromium. Journal of Dairy Science 101, 98769887.CrossRefGoogle ScholarPubMed
Koltes, JE, Koltes, DA, Mote, BE, Tucker, J and Hubbell, DS (2018) Automated collection of heat stress data in livestock: new technologies and opportunities. Translational Animal Science 2, 319323.Google ScholarPubMed
Kovács, L, Kézér, FL, Bakony, M, Jurkovich, V and Szenci, O (2018a) Lying down frequency as a discomfort index in heat stressed Holstein bull calves. Scientific Reports 8, 15065.CrossRefGoogle Scholar
Kovács, L, Kézér, FL, Ruff, F, Jurkovich, V and Szenci, O (2018b) Assessment of heat stress in 7-week old dairy calves with non-invasive physiological parameters in different thermal environments. PLoS ONE 13, e0200622.CrossRefGoogle Scholar
Kovács, L, Kézér, FL, Ruff, F, Jurkovich, V and Szenci, O (2018c) Heart rate, cardiac vagal tone, respiratory rate, and rectal temperature in dairy calves exposed to heat stress in a continental region. International Journal of Biometeorology 62, 17911797.CrossRefGoogle Scholar
Kovács, L, Kézér, FL, Ruff, F, Szenci, O, Bakony, M and Jurkovich, V (2019) Effect of artificial shade on saliva cortisol concentrations of heat-stressed dairy calves. Domestic Animal Endocrinology 66, 4347.CrossRefGoogle ScholarPubMed
Lammers, BP, VanKoot, JW, Heinrichs, AJ and Graves, RE (1996) The effect of plywood and polyethylene calf hutches on heat stress. Applied Engineering in Agriculture 12, 741745.CrossRefGoogle Scholar
Lee, JS, Kacem, N, Kim, WS, Peng, DQ, Kim, YJ, Joung, YG, Lee, C and Lee, HG (2019) Effect of Saccharomyces boulardii supplementation on performance and physiological traits of Holstein calves under heat stress conditions. Animals 9, 510.Google ScholarPubMed
López, E, Mellado, M, Martínez, AM, Véliz, FG, García, JE, de Santiago, A and Carrillo, E (2018) Stress-related hormonal alterations, growth and pelleted starter intake in pre-weaning Holstein calves in response to thermal stress. International Journal of Biometeorology 62, 493500.CrossRefGoogle ScholarPubMed
Mader, TL, Johnson, LJ and Gaughan, JB (2010) A comprehensive index for assessing environmental stress in animals. Journal of Animal Science 88, 21532165.CrossRefGoogle ScholarPubMed
Manriquez, D, Valenzuela, H, Paudyal, S, Velasquez, A and Pinedo, PJ (2018) Effect of aluminized reflective hutch covers on calf health and performance. Journal of Dairy Science 101, 14641477.CrossRefGoogle ScholarPubMed
Mellado, M, Lopez, E, Veliz, FG, De Santiago, MA, Macias-Cruz, U, Avendaño-Reyes, L and Garcia, JE (2014) Factors associated with neonatal dairy calf mortality in a hot-arid environment. Livestock Science 159, 149155.CrossRefGoogle Scholar
Monteiro, APA, Tao, S, Thompson, IM and Dahl, GE (2014) Effect of heat stress during late gestation on immune function and growth performance of calves: isolation of altered colostral and calf factors. Journal of Dairy Science 97, 64266439.CrossRefGoogle ScholarPubMed
Moore, DA, Duprau, JL and Wenz, JR (2012) Short communication: effects of dairy calf hutch elevation on heat reduction, carbon dioxide concentration, air circulation, and respiratory rates. Journal of Dairy Science 95, 40504054.Google ScholarPubMed
Morignat, E, Perrin, JB, Gay, E, Vinard, JL, Calavas, D and Hénaux, V (2014) Assessment of the impact of the 2003 and 2006 heat waves on cattle mortality in France. PLoS ONE 9, e93176.CrossRefGoogle ScholarPubMed
Neuwirth, JG, Norton, JK, Rawlings, CA, Thompson, FN and Ware, GO (1979) Physiologic responses of dairy calves to environmental heat stress. International Journal of Biometeorology 23, 243254.CrossRefGoogle ScholarPubMed
Orellana Rivas, RM, Komori, GH, Beihling, VV, Marins, TN, Bernard, JK and Tao, S (2020) Effects of milk replacer feeding levels on performance and metabolism of preweaned dairy calves during summer. Journal of Dairy Science 103, 313324.CrossRefGoogle ScholarPubMed
Peña, G, Risco, C, Kunihiro, E, Thatcher, M-J and Pinedo, PJ (2016) Effect of housing type on health and performance of preweaned dairy calves during summer in Florida. Journal of Dairy Science 99, 16551662.CrossRefGoogle ScholarPubMed
Piccione, G, Caola, G and Refinetti, R (2003) Daily and estrous rhythmicity of body temperature in domestic cattle. BMC Physiology 3, 7.Google ScholarPubMed
Polsky, L and von Keyserlingk, MAG (2017) Invited review: effects of heat stress on dairy cattle welfare. Journal of Dairy Science 100, 86458657.CrossRefGoogle ScholarPubMed
Quigley, J (2001) Calf Note # 68 – Predicting water intake in young calves. Calf Note.com, pp. 14.Google Scholar
Renaud, DL, Duffield, TF, LeBlanc, SJ, Ferguson, S, Haley, DB and Kelton, DF (2018) Risk factors associated with mortality at a milk-fed veal calf facility: a prospective cohort study. Journal of Dairy Science 101, 26592668.CrossRefGoogle Scholar
Roland, L, Drillich, M, Klein-Jöbstl, D and Iwersen, M (2016) Invited review: influence of climatic conditions on the development, performance, and health of calves. Journal of Dairy Science 99, 24382452.Google ScholarPubMed
Rosenberg, G (1979) Clinical Examination of Cattle. Berlin: Verlag Paul Parey.Google Scholar
Spain, JN and Spiers, DE (1996) Effects of supplemental shade on thermoregulatory response of calves to heat challenge in a hutch environment. Journal of Dairy Science 79, 639646.CrossRefGoogle Scholar
Strong, RA, Silva, EB, Cheng, HW and Eicher, SD (2015) Acute brief heat stress in late gestation alters neonatal calf innate immune functions. Journal of Dairy Science 98, 77717783.CrossRefGoogle ScholarPubMed
Strutzke, S, Fiske, D, Hoffmann, G, Ammon, C, Heuwieser, W and Amon, T (2019) Technical note: development of a noninvasive respiration rate sensor for cattle. Journal of Dairy Science 102, 690695.CrossRefGoogle ScholarPubMed
Tao, S and Dahl, GE (2013) Invited review: heat stress effects during late gestation on dry cows and their calves. Journal of Dairy Science 96, 40794093.CrossRefGoogle ScholarPubMed
Tao, S, Monteiro, APA, Thompson, IM, Hayen, MJ and Dahl, GE (2012). Effect of late-gestation maternal heat stress on growth and immune function of dairy calves. Journal of Dairy Science 95, 71287136.CrossRefGoogle ScholarPubMed
Tao, S, Monteiro, APA, Hayen, MJ and Dahl, GE (2014) Short communication: maternal heat stress during the dry period alters postnatal whole-body insulin response of calves. Journal of Dairy Science 97, 897901.Google ScholarPubMed
Wiedmeier, RD, Young, AJ and Hammon, DS (2005) Frequent changing and rinsing of drinking water buckets improved performance and health of hutch-reared Holstein beef calves. Bovine Practicioner 40, 18.Google Scholar