In dairy cattle, late gestation is a critical period for fetal growth and physiological transition into the next lactation. Environmental factors, such as temperature and light, exert dramatic ...effects on the production, health, and well-being of animals during this period and after parturition. The aim of this review was to introduce effects of heat stress during late gestation on dairy cattle, and discuss the biological mechanisms that underlie the observed production and health responses in the dam and her fetus. Relative to cooled cows, cows that are heat stressed during late gestation have impaired mammary growth before parturition and decreased milk production in the subsequent lactation. In response to higher milk yield, cows cooled prepartum undergo a series of homeorhetic adaptations in early lactation to meet higher demand for milk synthesis compared with heat-stressed cows, but no direct effect of environmental heat stress on metabolism exists during the dry period. Prepartum cooling improves immune status of transition cows and evidence suggests that altered prolactin signaling in immune cells mediates the effects of heat stress on immune function. Late-gestation heat stress compromises placental development, which results in fetal hypoxia, malnutrition, and eventually fetal growth retardation. Maternal heat stress may also have carryover effects on the postnatal growth of offspring, but direct evidence is still lacking. Emerging evidence suggests that offspring from prepartum heat-stressed cows have compromised passive immunity and impaired cell-mediated immune function compared with those from cooled cows.
Heat stress during the dry period affects the cow’s mammary gland development, metabolism, and immunity during the transition period. However, the effect of late-gestation heat stress on calf ...performance and immune status is unknown. Our objective was to evaluate the effect of heat stress during the final ∼45d of gestation on growth and immune function of calves. Calves (17/treatment) were born to cows that were exposed to cooling (CL) or heat stress (HT) during the dry period. Only heifer calves (CL, n=12; HT, n=9) were used in measurements of growth and immune status after birth. Heifer calves were managed under identical conditions. All were fed 3.78L of colostrum from their respective dams within 4h of birth and were weaned at 2mo of age (MOA). Body weight (BW) was obtained at weaning and then monthly until 7 MOA. Withers height (WH) was measured monthly from 3 to 7 MOA. Hematocrit and plasma total protein were assessed at birth, 1, 4, 7, 11, 14, 18, 21, 25, and 28d of age. Total serum IgG was evaluated at 1, 4, 7, 11, 14, 18, 21, 25, and 28d of age, and apparent efficiency of absorption was calculated. Peripheral blood mononuclear cells were isolated at 7, 28, 42, and 56d of age, and proliferation rate was measured by 3H-thymidine incorporation in vitro. Blood cortisol concentration was measured in the dams during the dry period and in calves in the preweaning period. Gestation length was 4d shorter for HT cows compared with CL cows. Calves from CL cows had greater BW than calves from HT cows at birth (42.5 vs. 36.5kg). Compared with CL heifers, HT heifers had decreased weaning BW (78.5 vs. 65.9kg) but similar BW (154.6 vs. 146.4kg) and WH (104.8 vs. 103.4cm) from 3 to 7 MOA. Compared with CL, heifers from HT cows had less total plasma protein (6.3 vs. 5.9g/dL), total serum IgG (1,577.3 vs. 1,057.8mg/dL), and apparent efficiency of absorption (33.6 vs. 19.2%), and tended to have decreased hematocrit (33 vs. 30%). Additionally, CL heifers had greater peripheral blood mononuclear cell proliferation relative to HT heifers (23.8 vs. 14.1 fold). Compared with CL, late-gestation HT did not affect the blood cortisol concentration of dams during the dry period or that of the calves in the preweaning period, but CL calves tended to have increased circulating cortisol at birth (7.6 vs. 5.7µg/dL). We conclude that heat stress of the dam during the dry period compromises the fetal growth and immune function of offspring from birth through weaning.
In dairy cattle, the final weeks before parturition are physiologically challenging and an important determinant of subsequent production performance. External stressors should be carefully managed ...during this period to avoid adding strain on the animals. Late-gestation heat stress impairs productivity in the dam and exerts transgenerational effects on progeny. Physiological responses are complex and detriments to performance are multifaceted. Late-gestation heat stress blunts mammary gland involution in the first half of the dry period and impairs cell proliferation as calving approaches. Moreover, cows that were exposed to prepartum heat-stress exhibit reduced adipose tissue mobilization and a lower degree of insulin resistance during early lactation. Prepartum heat exposure also depresses immune function and evidence links this decrease to altered prolactin signaling under heat stress. Placental functions are also impaired as reflected in a higher cotyledon mass but lower maternal circulating estrone sulfate concentrations, potentially resulting in lower nutrient supply and reduced calf birth weight. In addition, calves born to heat-stressed dams show impaired immune function and therefore higher disease susceptibly. Novel evidence reported that intrauterine heat stress alters the methylation profile of liver and mammary DNA, which may also contribute to the poorer performance during adulthood of calves exposed to heat stress in utero. Understanding the contribution of all altered biological systems during late-gestation heat stress can be used as a basis for improving cow management during the dry period. This article provides a review of the impacts of late-gestation heat stress and of the emerging understanding of the biological mechanisms that underlie the observed impairments of performance.
•Heat stress in late gestation negatively impacts the dam’s productivity and health.•In utero heat stress alters mammary development of the dam and the fetus.•In utero heat stress programs a lower productivity phenotype in offspring.
Heat stress reduces cow milk yield and results in a significant economic loss for the dairy industry. During lactation, heat stress lowers milk production by 25 to 40% with half of the decrease in ...milk synthesis resulting from the reduced feed intake. In vitro studies indicate that primary bovine mammary epithelial cells display greater rates of programmed cell death when exposed to high ambient temperatures, which may lead to a decrease in the total number of mammary epithelial cells in the mammary gland, partially explaining the lower milk production of lactating cows under heat stress. The function of mammary cells is also altered by heat stress. In response to heat stress, mammary cells display higher gene expression of heat shock proteins, indicating a need for cytoprotection from protein aggregation and degradation. Further, heat stress results in increased gene expression without altering protein expression of mammary epithelial cell junction proteins, and does not substantially influence the integrity of mammary epithelium. These data suggest that the mammary gland strives to maintain cell-to-cell junction integrity by synthesizing more proteins to compensate for protein losses induced by heat stress. During the dry period, heat stress negatively affects mammary gland development by reducing mammary cell proliferation before parturition, resulting in a dramatic decrease in milk production in the subsequent lactation. In addition to mammary growth, the mammary gland of the heat-stressed dry cow has reduced protein expression of autophagic proteins in the early dry period, suggesting heat stress influences mammary involution. Emerging evidence also indicates that heifers born to cows that experience late-gestation heat stress have lower milk yield during their first lactation, implying that the maternal environment may alter mammary gland development of the offspring. It is not clear if this is due to a direct epigenetic modification of prenatal mammary gland development by maternal heat stress. More research is needed to elucidate the effect of heat stress on mammary gland development and function.
Environmental effects on pathogen abundance and access are precursors to mastitis. Indeed, high heat and humidity, and unsanitary housing and equipment, are associated with greater pathogen load and ...exposure. Although less is known about effects of environment on a cow's ability to resist infection, several indicators suggest that it can affect pathogen responses. Mastitis incidence and bulk tank somatic cell count vary with season, typically peaking in summer. Recent controlled studies have revealed that heat stress exposure results in changes in the microbiome of the cow and her environment, which may relate to negative effects on milk quality and cow health. Alternatively, specific pathogen loads may vary based on housing dynamics rather than associations with physical environment. Indeed, housing-related stressors, such as overcrowding and social group challenge, influence secretion of glucocorticoids, thus affecting pathogen resistance in the cow. Two key seasonal variables are photoperiod and temperature, specifically the heat stress consequent to elevated temperature and humidity. Shifts in light duration regulate immune function in other species, but apparently have limited effect on udder health of lactating cows. In contrast, in dry cows, short days increase peripheral blood mononuclear cell number and are associated with lower somatic cell count in the next lactation, compared with long days. With heat stress, elevated body temperature directly affects expression of immune-related genes in mammary tissue. Responses depend on duration of exposure and feature acute upregulation of immune-signaling pathways, followed by enrichment of other immune-related pathways after prolonged exposure. Most responses are transient and recover within 1 wk. Functionally, heat stress impairs some aspects of acquired immunity in dry cows, including antigen responses and lymphocyte proliferation, but apparently not innate immune function. However, heat stress in late gestation reduces neutrophil phagocytosis and killing in vitro, and neutrophils in circulation are reduced in vivo as are responses to pathogen challenge in the subsequent lactation. A holistic understanding of the complex interplay of environment, pathogens, and host is needed to inform advances in this area.
It has long been known that season of the year has major impacts on dairy animal performance measures including growth, reproduction, and lactation. Additionally, as average production per cow has ...doubled, the metabolic heat output per animal has increased substantially rendering animals more susceptible to heat stress. This, in turn, has altered cooling and housing requirements for cattle. Substantial progress has been made in the last quarter-century in delineating the mechanisms by which thermal stress and photoperiod influence performance of dairy animals. Acclimation to thermal stress is now identified as a homeorhetic process under endocrine control. The process of acclimation occurs in 2 phases (acute and chronic) and involves changes in secretion rate of hormones as well as receptor populations in target tissues. The time required to complete both phases is weeks rather than days. The opportunity may exist to modify endocrine status of animals and improve their resistance to heat and cold stress. New estimates of genotype×environment interactions support use of recently available molecular and genomics tools to identify the genetic basis of heat-stress sensitivity and tolerance. Improved understanding of environmental effects on nutrient requirements has resulted in diets for dairy animals during different weather conditions. Demonstration that estrus behavior is adversely affected by heat stress has led to increased use of timed insemination schemes during the warm summer months to improve conception rates by discarding the need to detect estrus. Studies evaluating the effects of heat stress on embryonic survival support use of cooling during the immediate postbreeding period and use of embryo transfer to improve pregnancy rates. Successful cooling strategies for lactating dairy cows are based on maximizing available routes of heat exchange, convection, conduction, radiation, and evaporation. Areas in dairy operations in which cooling systems have been used to enhance cow comfort, improve milk production, reproductive efficiency, and profit include both housing and milking facilities. Currently, air movement (fans), wetting (soaking) the cow's body surface, high pressure mist (evaporation) to cool the air in the cows’ environment, and facilities designed to minimize the transfer of solar radiation are used for heat abatement. Finally, improved understanding of photoperiod effects on cattle has allowed producers to maximize beneficial effects of photoperiod length while minimizing negative effects.
The impacts of late gestation heat stress on the dam and her subsequent lactation are well-recognized. However, more recent research has demonstrated the long-lasting and severe negative consequences ...on the in-utero heat-stressed progeny. Dairy calves born to late gestation heat-stressed dams weigh less at birth and up to one year of age and have compromised metabolism and immune function. In-utero programming of these offspring may coordinate alterations in thermoregulation, mammary development, and milk synthetic capacity at different developmental windows. Thus, prenatally heat-stressed dairy heifers will produce less milk across multiple lactations and have a lower herd survival rate, potentially negatively impacting the U.S. dairy economy. Dry period heat stress abatement strategies should be considered not only for the productivity and welfare of the pregnant dam but also for the developing calf.
•Heat stress in late gestation negatively impacts calf health.•In utero heat stress alters the trajectory of mammary development.•In utero heat stress decreases productivity of the offspring.
The cessation of lactation (i.e., dry-off) in dairy cattle is an area of research that has received much focus in recent years. The dry period is necessary to optimize tissue remodeling of the ...mammary gland, but represents a stressful event, incorporating several changes in daily routine, diet, and metabolism. Moreover, the high milk yields achieved by modern cows in late gestation exacerbate the need for relevant manipulations in the days around dry-off, as excessive accumulation of milk might jeopardize the success of the dry period, with potential negative effects on future lactation. Production levels over 15 kg/d are an additional risk factor for udder health, delay mammary involution, and worsen metabolic stress and inflammatory responses. Furthermore, the pressure to reduce antibiotic usage in farm animals has resulted in increased attention on the dry period, given that historically most dairy cattle were provided prophylactic intramammary antibiotic treatment at dry-off as a means to reduce the risk of intramammary infections in the subsequent lactation. Several strategies have been proposed over the years to cope with these challenges, aiming to gradually reduce milk yield before dry-off, promoting at the same time the start of mammary involution. Among them, the most common are based on feed or nutrient restriction, a decrease in milking frequency, or administration of prolactin inhibitors. These practices have different capacities to reduce milk yield through different mechanisms and entail several implications for udder health, animal welfare, behavior, endocrine status, metabolism, and inflammatory conditions. The present review aims to provide a comprehensive overview of the dry-off phase in high-yielding cows and of the impact of high milk production at dry-off, and to describe possible strategies that might be implemented by farmers and veterinarians to optimize this critical phase in an integrated way.
Although dairy calves are more thermotolerant relative to mature cows, they are still susceptible to heat stress, as demonstrated by elevated physiological responses and reduced feed intake under ...high ambient temperature and relative humidity. However, indicators of heat stress have not been well-characterized in calves. Herein, we evaluated associations between environmental and thermoregulatory and productive animal-based indicators of heat stress in dairy calves exposed to chronic heat stress or continuous cooling in a subtropical climate. Holstein calves were exposed to heat stress (HT; shade of barn, n = 24) or continuous cooling (CL; shade of barn plus 2 fans, n = 24) from 2 to 42 d of age. Environmental indices, including ambient temperature, relative humidity, temperature-humidity index (THI), and wind speed, and animal-based indices, including respiration (RR), heart rate (HR), rectal (RT), and skin temperature (ST) were recorded thrice daily (0900, 1300, and 1900 h). Milk replacer (MI) and grain intakes were recorded daily from 15 to 42 d of age. Using segmented regression models, we then estimated THI thresholds for significant changes in physiological responses. We found a strong, positive correlation between animal-based indicators (except for HR, MI, and grain intakes) and ambient temperature and THI, with the highest correlation obtained with ST and THI (r ≥ 0.72). Ambient temperature and ST and ambient temperature or THI and MI were the only correlations that differed between treatments. The coefficient of determination (R2) obtained from regression analyses to model animal-based indicators was substantially improved by the inclusion of environmental indicators, with the greatest improvement achieved with THI. Overall, continuous cooling by fans promoted calf heat loss as CL calves had lower RR, RT, ST, and higher feed intake compared with HT calves. Temperature-humidity index breakpoints could be determined for RT (THI = 67), RR (THI = 65), and MI (THI = 82) in HT calves, and only for RR (THI = 69) in CL calves. Skin temperature variables had no detectable breakpoints in either treatment due to the strong linear relationship to THI. Collectively, our results suggest that ST is appropriate to estimate chronic heat stress and that THI is the best environmental indicator of heat stress in dairy calves raised in a shaded, subtropical environment. At a practical level, calves should be closely monitored when THI reaches 65 to 69 to minimize the risk of heat stress–related impairments.
Heat stress during the dry period negatively affects hepatic metabolism and cellular immune function during the transition period, and milk production in the subsequent lactation. However, the ...cellular mechanisms involved in the depressed mammary gland function remain unknown. The objective of the present study was to determine the effect of heat stress during the dry period on various indices of mammary gland development of multiparous cows. Cows were dried off approximately 46 d before expected calving and randomly assigned to 2 treatments, heat stress (HT, n=15) or cooling (CL, n=14), based on mature equivalent milk production. Cows in the CL treatment were provided with sprinklers and fans that came on when ambient temperatures reached 21.1°C, whereas HT cows were housed in the same barn without fans and sprinklers. After parturition, all cows were housed in a freestall barn with cooling. Rectal temperatures were measured twice daily (0730 and 1430h) and respiration rates recorded at 1500h on a Monday-Wednesday-Friday schedule from dry off to calving. Milk yield and composition were recorded daily up to 280 d in milk. Daily dry matter intake was measured from dry off to 42 d relative to calving. Mammary biopsies were collected at dry off, −20, 2, and 20 d relative to calving from a subset of cows (HT, n=7; CL, n=7). Labeling with Ki67 antigen and terminal deoxynucleotidyl transferase deoxyuridine triphosphate nick-end labeling were used to evaluate mammary cell proliferation and apoptosis, respectively. The average temperature-humidity index during the dry period was 76.6 and not different between treatments. Heat-stressed cows had higher rectal temperatures in the morning (38.8 vs. 38.6°C) and afternoon (39.4 vs. 39.0°C), greater respiration rates (78.4 vs. 45.6 breath/min), and decreased dry matter intake (8.9 vs. 10.6kg/d) when dry compared with CL cows. Relative to HT cows, CL cows had greater milk production (28.9 vs. 33.9kg/d), lower milk protein concentration (3.01 vs. 2.87%), and tended to have lower somatic cell score (3.35 vs. 2.94) through 280 d in milk. Heat stress during the dry period decreased mammary cell proliferation rate (1.0 vs. 3.3%) at −20 d relative to calving compared with CL cows. Mammary cell apoptosis was not affected by prepartum heat stress. We conclude that heat stress during the dry period compromises mammary gland development before parturition, which decreases milk yield in the next lactation.
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