Nutritional Strategies for Managing the Heat-Stressed Dairy Cowl Joe W. West Animal and Dairy Science Department, University of Georgia, Coastal Plain Station, Tifton 31793-0748 ABSTRACT: Heat stress results from the animal's nutrients are used less efficiently. An excess of degradable dietary protein is undesirable because of energy costs to metabolize and excrete excess N as urea. Optimizing ruminally undegraded protein improves milk yield in hot climates. Mineral losses via sweating (primarily K ) and changes in blood acidbase chemistry resulting from hyperventilation reduce blood bicarbonate and blood buffering capacity and increase urinary excretion of electrolytes. Theoretical heat production favors feed ingredients with a lower heat increment, such as concentrates and fats, whereas forages have a greater heat increment. Improved dietary energy density and the lower heat increment associated with the inclusion of dietary fat must be coupled with limitations t o fat feeding to avoid ruminal and metabolic disorders. Numerous nutritional modifications are used for hot weather feeding; however, many need further investigation to achieve specific recommendations. inability to dissipate sufficient heat to maintain homeothermy. Environmental factors, including ambient temperature, radiant energy, relative humidity, and metabolic heat associated with maintenance and productive processes, contribute to heat stress. The focus of this article is to identify environmental and metabolic factors that contribute to excessive heat load, describe how disruption of homeothermy alters physiologic systems of the cow, and discuss nutritional modifications that help to maintain homeostasis or prevent nutrient deficiencies that result from heat stress. Changes in diet are needed during hot weather to maintain nutrient intake, increase dietary nutrient density, or to reestablish homeostasis. Formulation for adequate nutrient intake is challenging because of the competition between nutrient density and other needs for the cow, including energy density and adequate dietary fiber. Lower DMI during hot weather reduces nutrients available for absorption, and absorbed Key Words: Dairy, Heat Stress, Intake, Metabolism, Milk 01999 American Society of Animal Science and American Dairy Science Association. All rights reserved. Introduction adaptations and may be favorable or unfavorable to the economic interests of humans, and that, nevertheless, they are essential for survival of the animal. The impact of heat stress on livestock is broad in geographic terms. Dairy cattle across the southern United States and in many subtropical and tropical regions are subject to high ambient temperatures and(or) high relative humidity for extended periods. In the southeastern United States, high ambient temperature and relative humidity exceeding the temperature-humidity index ( THI) associated with heat stress, persist for 4 to 6 mo of each year. In an excellent review, Beede and Collier (1986) identify three management strategies that minimize the effects of thermal stress: 1) physical modification of the environment, 2 ) genetic development of heattolerant breeds, and 3 ) improved nutritional manage- High ambient temperature, relative humidity, and radiant energy compromise the ability of the lactating dairy cow to dissipate heat, resulting in heat stress. These environmental factors, coupled with metabolic heat, create difficulties in maintaining thermal balance. This results in elevated body temperature, which in turn initiates compensatory and adaptive mechanisms to reestablish homeothermy and homeostasis. Stott ( 1981) stated that these readjustments to maintain homeostasis are referred t o as lThe author gratefully acknowledges the assistance of Brett Norman for assistance in literature review and June Womack for assistance in manuscript preparation. 27 22 WEST ment practices. Nutritional modifications to compensate for the effects of heat stress have been developed, but the response to dietary changes may be improved by environmental modifications. Feeding excessive quantities of nutrients, such as crude protein, can contribute to reduced efficiency of energy utilization, potentially adding to stress levels. Therefore, a thorough understanding of dietary modifications to minimize heat stress is necessary. The objectives for this paper are to summarize environmental factors that cause heat stress, describe the physiologc or metabolic effects of the stressors, and discuss the nutritional practices developed to compensate for the effects of the stress. Effects of Environmental Stress on Physiologic Responses of the Cow Reactions of cattle and other homeotherms to moderate climatic changes generally are compensatory and are directed at maintaining or restoring thermal balance (Kibler and Brody, 1953). The numerous physiologic mechanisms for coping with heat stress have been reviewed (Blackshaw and Blackshaw, 1994). These include sweating, a more rapid respiratory rate, and greater vasodilation with increased blood flow to the skin surface. A reduced rate of metabolism, decreased DM and nutrient intake, and altered water metabolism all occur in response to heat stress. Unfortunately, responses to heat stress often have negative effects on the physiology of the cow and on milk yield. Environmental Effects on Nufrient Irttake and M i l k Yield Feed DMI starts to decline and maintenance expenditures increase when environmental temperatures exceed 25°C (NRC, 1981). However, THI may describe more precisely the effects of the environment on the cow's ability to dissipate heat. Milk yield and TDN intake declined slightly when the THI exceeded 72 and declined sharply when 76 was exceeded (Johnson et al., 1963). Milk yield declined when body temperature exceeded 38.9"C, and, for each 0.55"C increase in rectal temperature, milk yield and intake of TDN declined 1.8 and 1.4 kg, respectively (Johnson et al., 1963). Consequently, minimizing the increase in rectal temperature can improve nutrient intake and subsequent milk yield. Genetic differences exist for heat tolerance of cattle. Bos indicus breeds are more heat-tolerant than Bos taurus breeds because of greater sweating capacity and lower metabolic rate (Blackshaw and Blackshaw, 1994). Milk yield decline for dairy cows exposed to temperatures of 24 or 34°C and to low ( 3 8 t o 46%) or high ( 7 6 to 80%) relative humidities, was greatest for Holstein, intermediate for Jersey, and least for Brown Swiss cows (Johnson and Vanjonack, 1976). Holstein cows are larger and have less surface area relative to internal body mass and would be expected to dissipate heat less effectively than would Jerseys. Brown Swiss are generally acknowledged as being more heattolerant than Holsteins, but generally produce less milk than Holsteins. Milk temperatures for Holstein and Guernsey cows increased as production increased from low, medium, and high levels during hot weather (Igono et al., 19851, an indication that high yielding cows are more sensitive to hot conditions. Heat production was about 14% greater for cows subjected to hot ( 3 5 ° C ) vs cool (20°C) conditions and was reflected in elevated rectal temperatures (Robinson et al., 1986). Environmental conditions such as temperature and humidity are interrelated, and the combined effects must be considered when determining effects on intake and milk yield. The mean daily temperature had the greatest effect on milk yield and rectal temperature when compared with minimum and maximum temperatures (Kabuga and Sarpong, 1991). Holter et al. ( 1 9 9 6 ) reported that the minimum daily THI was more closely correlated with DMI than maximum THI in Jersey cows. Reductions in DMI commenced when minimum THI exceeded 56 to 57 and continued until THI reached 72. Corresponding maximum THI ranged from 71 to 85; DMI declined 4.4 kg/d, or 22%. Similarly, correlation of DMI with minimum and maximum THI for Holsteins was -.63 and -.62, and maximum THI had a .95 correlation with minimum THI (Holter et al., 1997). During heat stress, DMI was reduced 22% for multiparous cows and 6% for primiparous cows, probably reflecting smaller body size for the primiparous cows as well as lower DMI and less metabolic heat production. Effects of weather conditions on intake and milk yield are most likely mediated through changes in body temperature. Across early, middle, and late stages of lactation, 9% of the variation for milk yield, 13% for milk fat, 5% for feed intake, and 65% for rectal temperature were attributable to weather conditions (Maust et al., 1972). However, weather conditions during the 2 to 3 d preceding the day of temperature measurement were most closely associated with milk yield, and composition and comparisons using a single day of temperature measurement J. h i m . Sci. Vol. 77, Suppl. 2/J. Dairy Sci. Vol. 82, Suppl. 2/1999 MANAGING HEAT-STRESSED DAIRY COWS could be misleading. In addition, DMI and milk yield for cows in midlactation were reduced most by hot weather conditions (Maust et al., 19721, possibly because early-lactation cows rely on body stores for a portion of the nutrients for production whereas latelactation cows consume fewer total nutrients. Araki et al. (1984) reported that lactating cows were more sensitive to the effects of heat than were nonlactating cows, consistent with the greater metabolic heat production for the former. Metabolic Effects of Heat Stress That Affect Nutrient Intake Johnson et al. ( 1967) termed cows that experienced sharp increases in body temperature upon exposure to high ambient temperatures as heat-intolerant, and those with lesser increases were termed heat-tolerant. When ambient temperatures were increased from 18°C to 29"C, the heat-intolerant cows exhibited a 1.4"C increase in body temperature and a 4-kg decline in daily milk yield, compared with a .7"C increase and 2-kg decline for heat-tolerant cows. Such responses suggest metabolic or physiologic differences among cattle that could be exploited if the mechanisms were identified. Hormonal changes that occur in response t o heat stress may play an integral role in the decline in productivity. Plasma growth hormone concentration and growth hormone secretion rate declined with hot temperatures (35°C) (Mitra et al., 1972). Growth hormone content in milk of low, medium, and high production groups declined when THI exceeded 70, possibly reflecting suppressed production of growth hormone so that metabolic heat production is reduced (Igono et al., 1988). Plasma growth hormone reductions that occurred with heat-stressed cows did not occur in thermoneutral conditions for cows fed restricted intakes that were similar to those consumed during heat stress (McGuire et al., 1991). The change in plasma growth hormone was probably in response to effects of high temperature rather than to reduced nutrient intake. The thyroid hormones triiodothyronine and thyroxine declined in response t o heat stress (Johnson et al., 1988; Magdub et al., 19821, which is probably an attempt to reduce metabolic heat production in the cow. However, reduced metabolic rate compromised productive capability as well. Greater plasma content of epinephrine and norepinephrine with high ambient temperatures is an indicator of stress response (Alvarez and Johnson, 1973) and reduces rate of passage in the digestive tract. Slower passage rate can lead to greater gut fill and limited 23 intake but improved digestibility because of greater residence time in the gut. Mean total-tract retention time for cattle was 36.6 and 43.2 h when the ambient temperatures were 18 and 32°C (Warren et al., 1974). Digestibility of DM, ADF, and NDF was 6.7, 11, and 8.1% greater for cattle in the hot environment, partially offsetting the reduced nutrient intake that occurred with reduced DM consumption. Cortisol response to heat stress challenge is often variable, but workers evaluating the effects of increasing fiber content of the diet on the response to hot, humid weather reported a linear increase in plasma cortisol with increasing fiber content of the diet (West et al., 19981, suggesting that greater stress associated with greater dietary fiber content occurred. Reduced blood flow during heat stress may reduce nutrient uptake and mammary blood flow. Cows given ad libitum access to feed or with intake restricted to 75% of the ad libitum amount in a thermoneutral environment or with ad libitum access in heat stress had intakes of 15.1, 11.5, and 11.1kg/d, respectively. Portal vein plasma flow was reduced by 14% in the thermoneutral environment for cows fed the restricted diet compared with ad libitum consumption (McGuire et al., 1989). In addition, net flux of alpha amino-N declined by 20 and 35% for cows fed restricted diets and for cows in the heat stress environment-ad libitum feeding treatment. The authors indicated that a portion of the negative effects of heat stress could be explained by reduced nutrient intake and reduced nutrient uptake by the portal drained viscera of the cow. Similarly, cows with restricted intake in thermoneutral conditions or ad libitum intake in heat stress conditions had reduced blood flow to the mammary gland when compared with cows on ad libitum consumption in thermoneutral conditions (Lough et al., 1990). Changes in blood flow that alter the supply of nutrients for milk production are at least partially related to reduced nutrient intake during hot weather; however, a direct effect of heat stress on blood flow to the viscera may be involved. Interestingly, exogenous compounds can alter circulatory flow and contribute to greater stress. Beef heifers fed endophyte-infected tall fescue (Al-Haidary et al., 1994) or administered ergovaline (Al-Haidary et al., 1995) and exposed to hot environmental temperatures had greater core temperatures and respiratory rates. Heifers consuming ergovaline had lower temperatures for the back and hips; the authors suggested that decreased peripheral blood flow reduced heat loss via the skin. Endophyte-infected tall fescue has been associated with greater heat stress, which is consistent with these findings. Each avenue of heat dissipation is necessary to the maintenance of homeothermy. J. h i m . Sci. Vol. 77, Suppl. 2/J. Dairy Sci. Vol. 82, Suppl. 211999 24 WEST Effects of Cooling on Intake and Milk Yield Environmental modification may be necessary if productivity is to be maintained by the dairy cow with high genetic potential. Maintenance of DMI is critical to production, and minimizing the increase in body temperature during hot conditions translates directly into greater feed consumption. The most obvious environmental modification is the use of shade. Shading in a hot, humid climate reduced rectal temperature by 2 to 4.1%, respiratory rate by 29 to 60%, improved DMI by 6.8 to 23.2%, and improved milk yield by 9.4 to 22.7% compared with unshaded cows (Mallonee et al., 1985; Schneider et al., 1984, 1986). Cows shaded during the dry period yielded 4.5 and 13.6% more milk at 100 and 305 d postpartum and delivered calves weighing 3.1 kg more compared with cows receiving no shade while nonlactating, even though cows were handled similarly postpartum (Collier et al., 1982b). This suggests that cows may have had greater energy reserves entering the lactation. Energy use by the cow is affected by the degree of heat stress. Grazing cows that were unshaded had higher respiratory rates and rectal temperatures than unshaded cows, and shaded cows with no concentrate supplementation had milk yield similar to that of unshaded cows fed 3.5 kg/d of concentrate (Valtorta et al., 1996). Shading was found to be profitable, whereas concentrate supplementation without shade was not. Additional benefits from cooling cattle with fans and sprinklers can be derived in hot or hot and humid climates. Reports as early as the 1940s (Seath and Miller, 1948) demonstrated that, used independently, fans and sprinklers reduced rectal temperature and respiratory rate but when used together the effects were synergistic. Sprinklers used in combination with fans also reduced diurnal variation of body temperature. Much of the response t o cooling is a direct result of increased DMI. Bucklin et al. (1991) summarized studies from Florida, Kentucky, and Missouri and reported improvements in DMI of 7.1, 9.2, and 7.1% and milk yields of 11.6, 15.8, and 8.6% for the three states, respectively. Rectal temperatures for the Kentucky and Missouri cows were reduced by 0.5 and 0.4"C. High producing cows in early lactation were most sensitive to heat stress, and milk yield declined significantly when rectal temperatures exceeded 39°C for more than 16 h (Igono and Johnson, 1990). Because genetic potential for milk yield has improved and the predominant dairy breed has become Holstein, the impact of heat stress on production has increased. Physiologic improvements with cooling included greater circulating growth hormone concentrations and lower prolactin concentrations (Igono et al., 1987). Accompanying these physiologic changes were a 2.3 and 2.0 kg/d increase in DMI and milk yield. Similar to shade work with nonlactating cows (Collier et al., 1982b), cooling with fans and sprinklers during the dry period increased mean 150-d milk yield at 150 d by 3.5 kg/d, and the effect was greater with increasing age (Wolfenson et al., 1988). Milk yield was 0.9, 2.5, and 7.3 kg per day greater in lactation 2, 3, and 4+ for cows that were cooled during the dry period. It is apparent that with increasing production, body size, and maturity, cows become more sensitive to the effects of environmental stress. Effective nutritional programs need to be implemented with some environmental modification if high production is to be maintained. Heat Production and Heat Increment The processes associated with maintenance, digestion, activity, metabolism, and production create a large amount of heat. Coppock (1985) reported that heat production for a 600-kg cow yielding 40 kg of 4% fat milk accounted for 31.1% of consumed energy, second only t o fecal energy losses of 35.3%. Heat energy is further partitioned into heat associated with waste formation and excretion ( 3 % 1, product formation (52.9%), fermentation (8.3%), digestion (12.2%), and maintenance (23.5%) (Coppock, 1985). Heat production, though advantageous during cool temperatures, is a liability during hot weather. Obvious advantages to reducing metabolic heat production in the cow during hot weather are improved metabolic efficiency and reduced heat to be dissipated. High milk yield requires the intake of large quantities of nutrients and greater heat production. Cows at high (31.6 kg/d) and medium (18.5 kg/ d ) milk yield had 48.5 and 27.3% greater heat production when compared with dry cows (Purwanto et al., 1990). However, cows in mid and late lactation were more adversely affected by hot weather compared with early lactation cows, despite greater yield for the early lactation cows (Maust et al., 1972). The higher producing early-lactation cows were probably less sensitive to the effects of high ambient temperatures because of less total DMI and the more eficient use of mobilized body tissue reserves. The heat production associated with the use of body tissue stores is approximately one-half of that produced by direct conversion of dietary ME to milk (Coppock, 1985). Identifying dietary modifications to reduce heat production and improve efficiency of energy use has J. h i m . Sci. Vol. 7 7 , Suppl. 215. Dairy Sci. Vol. 82, Suppl. 2/1999 MANAGING HEAT-STRESSED DAIRY COWS merit during hot weather. Digestion and metabolism of nutrients creates heat, and heat increment is defined as energy expenditures associated with the digestion and assimilation of food (Baldwin et al., 1980). The conversion efficiencies of intermediate products, such as acetate and glucose, t o end products, such as fatty acids, is 68 to 72 and 82 to 85%, respectively, and partial efficiencies for the conversion of acetate and dietary fat to milk fat are in the ranges of 70 to 75 and 94 to 97%, respectively (Baldwin et al., 1985). Infused W A resulted in varying heat increment, depending on the specific W A infused (Armstrong and Blaxter, 1957; Moe, 1981). Differing efficiencies for nutrient utilization offers the potential t o formulate diets with a lower heat increment. Theoretical diets were formulated for beef cattle with high, low, and intermediate heat increments. Rations with high heat increment contained more forage fiber; those with low heat increment contained more grains. Using the NE,, NE,, and ME energy equations, the author determined that expected performance improvements with low heat increments more than offset greater ration costs during high temperatures (Brokken, 1971). Can these differences in energy efficiency be exploited in practical diets for animals in hot weather? Low efficiency for use of acetate may account for the low net energy of feeds high in fiber (Moe, 1981) and provides the rationale for feeding low fiber diets during hot weather. The high efficiency of fat utilization and its low heat increment led Coppock (1985) to comment that fats are undervalued by current feed evaluation systems when they are fed within or above the zone of thermoneutrality. Nutritional Modifications for Feeding During Hot Weather Fiber Feeding Because there is greater heat production associated with metabolism of acetate compared with propionate, there is a logical rationale for the practice of feeding low fiber rations during hot weather. Feeding more concentrate at the expense of fibrous ingredients increases ration energy density and should reduce heat increment. Kurihara (1996) fed diets of Italian ryegrass or corn silage plus soybean meal to cows exposed t o 18, 26, or 32°C. Ryegrass resulted in a greater heat increment at higher temperatures without changes in ME intake (because of force feeding via ruminal cannulae), probably resulting 25 from greater heat production associated with acetate production. The author reported that DMI decreased in the order of ryegrass hay, corn silage, beet pulp, and concentrates with high environmental temperature. The decline in ME intake over the range of 18 to 32°C was 22% for cows fed the ryegrass hay diet but was only 12.9% for the corn silage plus soybean meal diet, and energy retention was far less for the ryegrass hay diet. Cows fed diets containing 100, 75, or 50% alfalfa, with the remainder being a mix of corn and soybean meal, had a mean efficiency of conversion of ME to milk of 54, 61, and 65% (Coppock et al., 1964). Heat production for the three diets was 688, 647, and 620 kcal per megacalorie ME. The greater metabolic heat production for high fiber diets was confirmed by Reynolds et al. (19911, who fed pelleted diets of 75% alfalfa or 25% alfalfa, with the remainder as concentrate. Diets containing 75% alfalfa resulted in greater heat production and reduced retained energy, and the greater 0 2 intake by portal drained viscera and liver accounted for 44 and 72% of heat increment for the concentrate and alfalfa diets, respectively. Altered proportions of ruminal VFA may explain a part of the differences in heat increment with fiber feeding during heat stress. Volatile fatty acids constitute a large proportion of the energy available to the cow (Van Soest, 19821, and declining intake during heat stress reduces the quantity of VFA in the rumen because fermentable carbohydrate is reduced. However, heat-stressed cattle force-fed via ruminal cannulae to achieve constant intake had reduced total VFA production with altered molar proportions of VFA compared with cattle in thermoneutral conditions (Kelly et al., 1967). The mechanism for this decline is not understood but could relate to combined effects of heat stress on ruminal buffering and efficiency of microbial VFA production. Cows subjected to high environmental temperatures selectively decrease the quantity of forage consumed relative to concentrates (McDowell, 1972). Increased feeding of concentrates is a common practice during conditions conducive to heat stress, but maximal benefit from concentrates appears to be approximately 60 to 65% of the diet (Coppock, 1985). Excessive concentrate feeding leads to acidosis and the associated production, health, and metabolic difficulties. Does the greater heat production associated with high fiber diets actually translate into measurable performance effects? In early work, cows fed low fiber diets during hot weather had greater daily milk yield, lower body temperature, and slower respiratory rates compared with those fed high fiber diets (Leighton and Rupel, 1956; Stott and Moody, 1960; Tsai et al., J. h i m . Sci. Vol. 77, Suppl. 2/J. Dairy Sci. Vol. 82, Suppl. 211999 26 WEST 1967). The addition of 30 or 10% molasses to diets containing 35 or 45% alfalfa, respectively, had little effect on milk energy produced per unit of feed energy (20.7 and 20.0%) for cows in an 18.3"C environment. However, in a 31.1"C ambient environment, the efficiencies were 19.2 and 17.3% for the 30 and 10% molasses diets (Wayman et al., 1962). In addition, total feed intake declined by 18.8 and 31.8% for the 30 and 10% molasses rations. Extra carbohydrate furnished by the molasses probably contributed to greater efficiency of energy use. It has been proposed that diets that contain sufficient concentrates for high propionate production supply adequate NADPH to enable acetate to be converted to fat, whereas, in high roughage diets, the larger amount of acetate and smaller quantity of propionate produced results in a metabolic excess of acetate, which must be utilized as heat as a result of a futile cycle (MacRae and Lobley, 1982; Parker, 1984). They suggested that extra N as crude protein provides for more efficient use of available VFA by providing amino acids for glucose synthesis; thus, ruminants might be able to efficiently utilize larger quantities of acetate from roughage feeds if an additional source of reduced coenzyme were available. This may be particularly true for subtropical and tropical regions where adapted forages often are very high in fiber and low in readily fermentable carbohydrate. Rarely are total forage diets fed to lactating dairy cows in the United States and the carbohydrate furnished in mixed diets, such as those in practical total mixed rations, may improve the efficiency of energy use. Nonlactating cows fed diets containing 100% alfalfa hay or 30% alfalfa hay plus concentrates used ME from infused acetic acid with 27% efficiency for 100% hay diets, which improved t o 69% efficiency for the 30% hay diets (Tyrrell et al., 1979). The large amount of highly fermentable carbohydrate fed in typical high-concentrate diets should minimize the heat production observed in the very high fiber diets, which were used in research settings. Although high fiber diets contribute to heat stress, the level of intake is far more critical to the total amount of metabolic heat produced. Growing heifers fed pelleted rations containing 75% alfalfa or 25% alfalfa produced 48.8 and 45.5 MJId of heat (Reynolds et al., 1991). However, when the low and high intake (4.2 and 7.1 kgld DMI) heifers were compared, heat production was 38.2 and 56.1 MJId. Therefore, intake effects have a substantial effect on heat production and must be considered in designing an effective nutritional and environmental management program. In sheep, heat production in the gut was not greatly affected by diet composition but increased exponentially with increasing ME intake (Webster et al., 1975). The authors concluded that 25 to 30% of total heat increment resulted from intake and digestion. Work in Alabama supports the concept of the interaction of high intake with greater heat stress. Cows were fed diets containing 20, 17, and 14% ADF across varying environmental conditions. Dry matter intake was greatest with the lower fiber diets in cool weather, yet there was a more rapid decline in intake with the low fiber, higher intake diets as the environmental temperature increased (Cummins, 1992). Cows fed the diet containing 14% ADF were more sensitive to changes in minimum ambient temperature than for those fed higher ADF diets, but total DMI was also greater for these cows. Intake declined for cows fed four diets with a range for NDF from 27 to 35% with increasing NDF content during cool weather, but the decline was less severe during hot weather, indicating that the level of intake was more critical than fiber content during hot weather (West et al., 1998). Research suggests that lower fiber, high grain diets may indeed reduce metabolic heat production and contribute to lower heat load in the animal. Further, the low fiber, high grain diets provide more efficiently used end products, which contribute to lower dietary heat increment. However, low fiber, high grain diets must be balanced with the need for adequate fiber to promote chewing and rumination to maintain ruminal pH and cow health. Research to compare high and low quality forages for effects on energy and heat increment should address the question of adequate fiber vs high concentrate diets for hot weather. Feeding Fats and Concentrates The reduction in DMI and related efficiency of nutrient utilization that occurs with heat stress requires greater dietary nutrient density if milk yield is to be maintained at a level comparable to that at thermoneutral conditions. The addition of fat to the diet of lactating dairy cows is a common practice, and the greater energy density and the potential to reduce heat increment of high-fat diets may be particularly beneficial during hot weather. The addition of 3 to 5% fat to the ration can be achieved without toxic effects t o ruminal microflora (Palmquist and Jenkins, 19801, and, in thermoneutral conditions, cows fed diets with 25% of ME from a protected tallow had 8 to 13.6% higher efficiency for use of ME for lactation than those not receiving supplemental tallow (Kronfeld et al., 1980). The conversion of dietary fat to body fat is highly efficient when compared with the conversion of J. h i m . Sci. Vol. 7 7 , Suppl. WJ. Dairy Sci. Vol. 82, Suppl. 211999 MANAGING HEAT-STRESSED DAIRY COWS acetate to fatty acids (Baldwin et al., 1980). Improved efficiency and lower heat increment should make fat especially beneficial during hot weather. However, results from the addition of dietary fat during hot weather have been mixed. Both Brahman cross and British crossbred steers fed diets containing 9.2% fat during hot weather had lower body temperatures (.3 to .4"C lower) than those fed diets containing 2.5% fat (O'Kelly, 19871, suggesting less heat production in those steers. Cattle fed diets containing 10% of concentrate as soybean oil or hydrogenated vegetable oil during hot weather had similar body temperatures and efficiency of energy utilization for lactation compared with controls (McDowell et al. 1969; Moody et al. 1967). Feeding 15% whole cottonseed or 3% of a ruminally protected fat to cows during hot weather resulted in a small increase in fat-corrected milk yield with protected fat, and no improvements in respiratory rate or rectal temperature (Saunders et al., 1990). Ruminally-protected fats allow the inclusion of a substantial quantity of fat in the diet, which could lower heat increment significantly. Holter et al. (1992) added 15% whole cottonseed, or 15% whole cottonseed plus 5 4 kg of a calcium salts of fatty acids and reported that heat production in excess of maintenance declined by 6.7 and 9.7%, and total heat loss declined by 4.9 and 7.0%, respectively. The high fat levels produced a measurable decline in heat production in thermoneutral conditions. Knapp and Grummer ( 199 1) offered diets containing 5% added fat (60% prilled fatty acids, 40% tallow) to cows held at thermoneutral (2O.S0C, 38% relative humidity) or hot environmental conditions ( 3 1.8"C, 56% relative humidity) and reported a nonsignificant increase in milk yield of 1.1 and .3 kg in the cool and hot environments, although fat-corrected milk increased by 2.7 and 1.8 kgld. No diet x environment interaction occurred. Feeding diets containing 4.6% or 7.4% fat (from 3% added prilled fatty acids) and housing in a shade or shade plus evaporatively cooled environment did not improve milk or fat-corrected milk yields (Chan et al., 1997). Cooling improved milk yield ( P < .08), but rectal temperatures were unchanged by either cooling or dietary fat content, and no interactions occurred, again indicating no apparent benefits from lower heat increment associated with fat feeding. The authors speculated that high rectal temperatures across treatments could have resulted from cow movement associated with the collection of rectal temperatures, but performance variables did not indicate added benefits from fats during hot weather. High levels of prilled fatty acids (5% of diet DM) fed to cows that were blocked according to calving 27 season (cool or warm seasons) improved milk yield only for the warm season (Skaar et al., 1989). Maximum ambient temperatures during the warm season ranged from 29.4 to 35.0°C, and fat levels were high relative t o other studies. Conversely, in Georgia cows that calved in fall or summer and were fed diets with or without added tallow to supply 1 kg/d had improved NE intake with added fat. Milk yield was unchanged and efficiency of NE1 utilization was reduced by added fat (Niango et al., 1991). Fatsupplemented cows lost more BW than did controls and had lower fiber digestion, which suggests that fat content impaired digestion. The literature for fat supplementation is inconclusive relative to the benefits of added fat during hot weather. Additional research should include a comparison of several levels of added fat imposed over an extended period of time, implemented before the exposure t o temperatures above the upper critical level for the cow. Protein Nutrition in Hof Climates As DMI declines, the quantity of consumed nutrients, including CP, also declines and a negative protein balance may occur (Hassan and Roussel, 1975; Ibrahim et al., 1970; Kamal and Johnson, 1970). During hot weather, dietary protein density is often increased to compensate for lower intake, and cows offered diets containing 20.8% CP during hot conditions had greater DMI and milk yield than those offered diets containing 14.3% CP (Hassan and Roussel, 1975). The low CP diet was calculated to be adequate for the level of production achieved, and improved milk yield was associated with greater DMI for the high protein diet. It should be noted that there is an energetic cost associated with feeding excess protein, and dietary N above requirement reduces ME by 7.2 kcaWg of N (Tyrrell et al., 1970). Milk yield declined from 24.5 to 23.1 kg when dietary CP increased from 19 to 23% (Danfaer et al., 1980); the energy cost of synthesizing and excreting urea accounted for the reduced milk output (Oldham, 1984). Cows offered diets of two protein solubilities (40 and 20.9%) during thermoneutral and heat-stress conditions consumed more feed and yielded more milk in both environments when fed the less soluble protein diet (Zook, 1982). Although there were no overt measures of increased heat stress for the highly soluble protein diets during heat stress, N balance was improved for the low solubility diet. Hassan and Roussel ( 1975) reported that blood nonprotein-nitrogen content was positively correlated with rectal temperature, which is consistent with reduced efficiency of energy use for excessive N J. Anim. Sci. Vol. 77, Suppl. 215. Dairy Sci. Vol. 82, Suppl. 2/1999 28 WEST (Tyrrell et al., 1970). Thus, the energy necessary to form urea from excess protein appears as heat production and decreases the proportion of NE1 in ME, and the energy lost in excreted urinary N from excess protein decreases the proportion of ME in DE (NRC, 1989). Until recently, research into the effects of excessive dietary protein content, solubility, or degradability has been lacking, especially under heat stress conditions; recent studies investigating dietary protein content as well as composition suggest an interaction between protein availability and environment. Diets with low (31.2% of CP) and high (39.2% of CP) ruminally undegradable protein fed during hot weather had no effect on DMI; however, milk yield increased by 2.4 kgld and blood urea protein declined from 17.5 to 13.3 mgl100 mL for the diet containing higher undegradable protein (Belibasakis et al., 1995). Arizona workers compared diets with high (18.4%) and low (16.1%) CP and high ( 5 7 to 60%) and medium (40%) protein degradability during hot weather (Higginbotham et al., 1989b). Milk yield and persistency was lowest for cows offered the high protein, high solubility diet, whereas blood urea N was highest. However, similar diets fed under moderate temperature conditions had little effect on performance (Higginbotham et al., 1989a1, suggesting that excessively high dietary protein content may have deleterious effects only in a heat stress environment. In two trials, cows were fed diets with high ( 6 1 to 64%) and low (47 to 5 5 % ) protein degradabilities to cows under shade or shade plus evaporative cooling environments (Taylor et al., 1991). They reported reduced milk yield for low degradability diets in the first trial but significant improvements in both DMI and milk yield for low degradability diets during the second trial. In the first trial, temperatures were cooler and a lower quality of escape protein source (corn gluten meal) was used, whereas, in the second trial, temperatures were hotter and a higher quality of ruminal escape protein (corn gluten meal and blood meal) was used, suggesting that the degree of heat stress as well as protein quality are factors during hot weather. Cows fed diets with a similar ruminally undegradable protein content but from high quality (blood, fish, and soybean meals) or lower quality (corn gluten meal) sources were kept in shade or shade plus evaporatively cooled environments (C hen et al., 1993). Lysine was -98 and 3 9 % of DM for high and low quality protein diets. Although protein quality x environment interaction was not significant, cows fed high quality ruminal escape protein gave 3.8 and 2.4 kg more milk in the evaporatively cooled and shaded environments, respectively, than those fed low quality proteins. The authors theorized that the greater response to high quality protein for cows in the cooled environment was because the amount of protein metabolized for energy was reduced and less energy was used in converting NH3 to urea. In addition, cows in the cooled environment had higher milk yield and thus greater protein demand. Huber et al. (1994) summarized protein research by concluding that, when cows are subject to hot environmental conditions, ruminally degradable protein should not exceed 61% of dietary CP and should not exceed NRC recommendations (NRC, 1989) by greater than 100 g N/d. One hundred grams of N is equivalent to about 3.1% CP in the diet, assuming 20 kg DMI/d. In addition, the authors reported that cows fed diets containing 241 g/d lysine (1%of DM) yielded 3 kg more milk than for diets containing 137 g/d lysine (.6% of DM). Electrolytes and Acid-Base Chemistry During hot weather, declining DMI and high lactation demand requires increased dietary mineral concentration. However, alterations in mineral metabolism also affect the electrolyte status of the cow during hot weather. The primary cation in bovine sweat is K (Jenkinson and Mabon, 1973), and sharp increases in the secretion of K through sweat occur during hot climatic conditions (Jenkinson and Mabon, 1973; Johnson, 1967; Mallonee et al., 1985). The absorption of macrominerals, including Ca, P, and K, declined during hot temperatures (Kume et al., 1987; Kume et al., 1989). Kume et al. (1986) also reported that trace element requirements may increase with elevated environmental temperature. Lactating cows subjected to hot climatic conditions and supplemented with K well above minimum NRC recommendations (NRC, 1989) responded with greater milk yield (Mallonee et al., 1985; Schneider et al., 1984; West et al., 1987). Cows supplemented with 5 5 % Na during hot climatic conditions also demonstrated greater feed DMI and milk yield compared with those receiving .18% Na (Schneider et al., 1986). Potassium and Na are the primary cations involved in the maintenance of acid-base chemistry. The cow subjected to hot climatic conditions can have acid-base disturbances resulting from respiratory alkalosis, subsequent renal compensation by increasing urinary excretion of bicarbonate and Na, and renal conservation of K (Collier, 1982a). Electrolytes are a key element of acid-base chemistry and their supplementation during heat stress may be critical to J. h i m . Sci. Vol. 77, Suppl. WJ. Dairy Sci. Vol. 82, Suppl. 211999 MANAGING HEAT-STRESSED DAIRY COWS homeostatic mechanisms. Block ( 1984) pioneered the use of dietary anions and cations to alter the acid-base chemistry of the periparturient cow, creating a metabolic acidosis and altering Ca metabolism. However, during extreme heat stress, cows exhibited a respiratory alkalosis with elevated blood pH and reduced concentrations of blood C02 and bicarbonate (Schneider et al., 1984). Although the respiratory alkalosis, which was indicated by elevated blood pH, suggested an excess of bicarbonate, the elevated pH actually was a result of a carbonic acid deficit created by C02 expiration via hyperventilation (Benjamin, 198lj. Escobosa et al. (1984) found that providing diets that were high in Na and K and normal for C1 resulted in greater DMI and milk yield, compared with diets high in C1 and normal for Na and K. Calculated cation-anion difference (mEq Na + K - C1) was 35.0 mEq/100 g of feed DM for the high cation diet and -14.4 meq/100 g of feed for the high C1 diet. A high C1 diet depressed DMI and was associated with low blood pH and reduced blood buffering. Others have reported similar results, with greater DMI and milk yield from higher cation-anion balance (alkaline) diets (Tucker et al., 1988; West et al., 1991). In work designed to compare efficacy of increasing cationanion difference using either K or Na to increase cation content of the diet, DMI increased linearly with increasing cation content, and Na or K was equally effective as a cation source (West et al., 1992). The improved intake or milk yield observed when more alkaline diets are fed to lactating cows may be a result of improved blood buffering or correction of mineral deficiencies; however, separating the two effects is very difficult. Perhaps acid-base chemistry for the cow is optimized by providing the correct balance or ratio of electrolytes. Baldwin et al. (1980) stated that futile cycles contribute substantially to maintenance energy expenditures and that Na and K transport may account for 20 to 30% of those costs. Cows subjected to heat stress in climatic chambers and fed a high mineral diet (1.93, .68, and 1.85% of K, Na, and C1, respectively) or basal diet (.97, .19, and .2% of K, Na, and C1) responded with more rapid respiratory rates at night for the high mineral diet, as well as higher rectal temperatures. Dry matter intake and milk yields were similar among treatments (Schneider et al., 1988a). This was in the presence of lower blood pH, probably resulting from the high C1 content of the high mineral diet. Greater urinary NH3 was observed, and NH3 was probably serving as a companion ion for hydrogen ion excretion. One of the difficult issues associated with the electrolytes researched revolves around the com- 29 pounds used to alter mineral content of diets. In some studies, salts such as NaCl or KC1 were used, whereas in others Na or K bicarbonates or carbonates were used. The cation-anion difference approach to formulation considers all elements that are critical to the equation. In studies in which the equation was not considered in choosing mineral supplements, results may be clouded by unknown interactions. Florida researchers studied the cation-anion difference approach to electrolyte formulation through empirical modeling of numerous macromineral research studies from several locations (Sanchez et al., 1994). Data were presented for individual macrominerals and for cation-anion difference ( N a + K - C1, mEq/100 g DM) during summer and winter seasons. Dietary K and Na elicited increased DMI response at levels well above NRC ( 1989) recommendations, whereas increasing dietary C1 depressed DMI, especially during hot weather. The combination of respiratory alkalosis and increased renal bicarbonate filtering probably reduced blood buffering capacity sufficiently to make the dairy cow more susceptible to the effects of an acidifying agent such as C1. The DMI response to increasing cation-anion difference was similar during cool and hot weather, which suggests that the effect is consistent across these environments (Sanchez et al., 1994). This is consistent with Georgia work in which Na and K were equally effective for increasing the cationanion difference (West et al., 1992). Intake was improved by 5.5 and 5.2 kgld in cool and hot weather when the cation-anion difference was increased in the range of -79 to 324 mEq/kg of feed DM (West et al., 1991). Heat stress is cyclic in nature, with cows generally being at the peak of their stress by midafternoon and cooling somewhat in the evening and early morning hours. Cows may exhibit a respiratory alkalosis in the afternoon but physiologic responses may actually overcorrect so that when cows become cool a metabolic acidosis occurs. During the day, cows in an unshaded environment had higher rectal temperatures and respiratory rates than shaded cows, but at night both measures were lower for cows with no shade (Schneider et al., 1988b). Blood bicarbonate was consistently lower for the no-shade cows, and urine pH was elevated during the day and low at night, following trends for blood pH. Ammonium concentration was greater during the cool evening period for cows housed in climatic chambers, reflecting greater H+ ion excretion in response to acidosis (Schneider et al., 1988b). Cows in early lactation during hot weather exhibited a marked decline in plasma Na, K, and C1 concentrations during the day, with concentra- J. h i m . Sci. Vol. 77, Suppl. 215. Dairy Sci. Val. 82, Suppl. 2/1999 30 WEST tions returning to normal at night (Maltz, 1994). The authors speculated that active absorption of Na from the rumen, which is dependent on VFA concentration in the rumen, was limited by reduced VFA production associated with lower DMI, and that C1 absorption is coupled with Na absorption. In addition, K and C1 losses in sweat are significant. The greater losses of K and C1 ions, coupled with potential for reduced absorption and intake of Na, may account for the deficit that occurred during the day. During the cooler hours of the day, DMI was greater, and at night concentrations of elements returned to normal (Maltz et al., 1994). Wide swings in the acid-base chemistry and blood electrolyte content of the cow occur within a short time span. Sanchez et al. (1994) suggested that feeding programs targeted at physiologic needs of the cow at specific times might address the differing physiologic conditions of the cow that occur throughout the day. They also theorized that, because the heat-stressed cow eats more feed during the cooler evening hours and less during the day, ruminal acid production would be greater at the time when the cow is experiencing a compensated metabolic acidosis. Feeding strategies designed around the current physiological status of the cow might help alleviate the metabolic extremes caused by heat stress and improve performance. More work is needed to determine the effectiveness of altering electrolytes and cation-anion difference in the lactating cow, especially during hot climatic conditions, and to determine if targeted feeding is a viable option. Water Utilization and Metabolism Water is the most critical nutrient for cattle, and the need is intensified for those subjected to hot climatic conditions. Classical work demonstrated that water losses through the lungs increased gradually and losses from the skin increased sharply as ambient temperature increased from about 16°C to 35°C (Kibler and Brody, 1950). Cows acclimated to 21.1"C and then exposed to 32.2"C for 2 wk increased water consumption by 110%, and water losses from the respiratory tract and from the skin surface increased by 55% and 177% at the higher temperature (McDowell and Weldy, 1960). Such changes increase the volume of water needed. Water requirements have not been extensively investigated because water is usually supplied in abundance. Water consumption is closely tied to feed DMI; Murphy et al. (1983) reported a highly significant ( P < .0001) correlation of .626 between DMI and water intake. Water consumption increased from 35.6 L/d at 24 d prepartum to 65.2 L/d at 42 d postpartum as DMI increased from 9.6 to 16.2 kgld and milk production increased to 25.7 kgld (Woodford et al., 1984). Quality of water may affect consumption. Water with 4,100 mgL of total dissolved salts from an added complex salt mixture increased water consumption compared with fresh water with less than 450 mgl L total dissolved salts, but persistency of milk yield was reduced by 25 and 27% in winter and summer by the high salt water (Wegner and Schuh, 1988). In Kuwait, brackish well water, which contained 3,574 mg/L of total dissolved solids had little effect on water Consumption when fed under hot, arid conditions and resulted in a slower rate of decline in milk yield during the course of the experiment when compared with fresh water (Bahman et al., 1993). Intake of DM increases with Na and K supplementation during hot weather and with more alkaline diets. A positive correlation between sodium intake and water consumption has been demonstrated (Murphy et al., 19831, and supplementation of diets with high Na and K content will probably increase water intake during hot conditions. Maltz et al. (1994) indicated that water turnover is related to DMI, and milk yield and respiratory-cutaneous water loss by the cow are major factors increasing ion outflow under heat stress. Greater water content in the rumen tends to accelerate ruminal turnover (Silanikove, 19921, which may be beneficial to cows during hot weather because the rate of passage of digesta is slower and may contribute to gut fill. Consumed water may have a direct cooling effect via the reticulorumen (Beede and Collier, 1986) in addition to the cooling effects of panting and sweating. Cows offered drinking water at 10°C or 28 to 30.2% consumed more DM (Baker et al., 1988; Milam et al., 1986) and yielded more milk for the cool water treatment (Milam et al., 1986). Cows offered chilled drinking water generally consumed less of the chilled water, but the amount of heat absorbed by the chilled water was still significantly greater than for the warm water treatment. Chilled water cooled the cow, as evidenced by reduced respiratory rate (Lanham et al., 1986). Initial water temperature is critical and the control water temperature for the Texas studies just discussed was relatively high. Economic benefit for the practice must be considered, but the benefits of the consumption of large quantities of cool water on the comfort of the cow and ultimately DMI and milk yield is apparent and should be considered when devising a water supply system for the dairy herd. J. h i m . Sci. Vol. 77, Suppl. 215. Dairy Sci. Vol. 82, Suppl. 211999 31 MANAGING HEAT-STRESSED DAIRY COWS Vifamins The topic of heat stress and vitamin requirements is addressed briefly in regards to niacin use (NRC, 1989), and regarding vitamin A for poultry (NRC, 1981). It was noted that the requirement and the absorption of vitamin A for poultry was unchanged by heat-stress conditions. It becomes apparent that there is a dearth of information in the scientific literature regarding the effects of heat stress on requirements or metabolism of vitamins, although an early study showed that heat stress caused a decline in vitamin A stores in liver of steers (Page et al., 1959). The effect on gain or nutrient utilization has not been elucidated. In practical terms, it appears that adjustments in vitamin content to account for reduced DMI are justified. Further investigation into the effects of high ambient temperature and subsequent heat stress on the requirements and metabolism of specific vitamins is needed. Only then can the impact of heat stress on individual vitamins be ascertained and recommendations established for use in hot climatic conditions. Level of production may determine how effective some nutritional practices are during heat stress conditions. Cattle supplemented with niacin ( 6 g/d) during summer months increased milk yield by .9 kg/d compared with controls (Muller et al., 1986). However, with cows yielding greater than 34 kg of milk per day, yield improved 2.4 kg/d for niacin. Niacin can prevent ketosis and is involved with lipid metabolism. Energy intake for early lactation cows is often inadequate and fat mobilization is excessive. Muller et al. (1986) speculated that niacin improved milk yield by affecting lipid and energy metabolism, by stimulating protein synthesis by ruminal microorganisms or by causing other effects on ruminal microorganisms. The greater response for high producing cows indicates that these cows are more subject to stress, have greater nutritional and metabolic demands, and may derive greater benefit from nutritional modifications than lower producers. It also indicates the need to conduct research with high producing cows so that those demands can be identified. Di Costanzo et al. (1997) reported that supplementation with niacin at three levels ( 12, 24, 36 g/ c0w.d-l) resulted in skin temperatures at the rump and tail of about 0.3% lower with niacin supplementation, an indication that vasodilation and greater blood to flow to the peripheral tissues associated with niacin use improved heat loss by the cow. Rectal temperature and milk yield were unchanged. Cattle fed endophyte-infected tall fescue seed to induce fescue toxicosis exhibited reduced herbage intake per grazing session as ambient temperature increased (Dougherty et al., 1991). Fescue toxicosis increases susceptibility t o heat stress, and cattle ingesting the endophyte often exhibit greater rectal temperature and seek out shade and mud holes during the day. Supplemental thiamine improved grazing intake by extending grazing time during hot weather. The authors theorized that the expression of tall fescue toxicosis in beef cattle subject to high ambient temperatures may involve a thiamin deficiency, which was partially alleviated by the supplemental thiamine. Implications The modern dairy cow with high genetic potential consumes and metabolizes a large quantity of nutrients, and it is apparent that if the cow is to produce to her potential that some heat abatement procedures are necessary to dissipate the large amount of metabolic heat produced. By minimizing the increase in body temperature that occurs with hot environmental conditions, greater DMI is encouraged and the gross efficiency with which dietary nutrients are used by the cow is improved. A reduction in body temperature reduces sweating and panting. The loss of electrolytes via skin secretions is minimized and disturbance of acid-base chemistry from hyperventilation is moderated. Modification of the environment by shading and cooling enhances the quantity of nutrients consumed by the cow but can also affect nutrient requirements. With the onset of hot weather, cows will adapt to the environmental conditions and DMI and milk yield will usually stabilize at a level below that during cool weather. Reformulation of diets is necessary to achieve sufficient nutrient density to maintain nutrient intake. However, some nutrients are required at still higher levels during heat stress ( K and Na, for example), and supplementation above NRC minimums (NRC, 1989) results in improved production during hot weather. There is evidence that a ratio or proportion of mineral elements benefits the heatstressed cow, and the concept of dietary cation-anion difference may have merit. If so, not only is dietary electrolyte content important, but so is the source of electrolytes as well; KC1 or NaCl are neutral in the cation-anion difference equation, whereas Na or K carbonates or bicarbonates contribute to alkalinity of the diet. Apparently, an excess of acidogenic ions such as C1 reduces DMI in the cow, especially during hot weather; this could result from the reduced blood J. h i m . Sci. Vol. 77, Suppl. 215. Dairy Sci. Vol. 82, Suppl. 2/1999 32 WEST buffering capacity that occurs in the presence of a respiratory alkalosis. Although inadequate nutrient intake is of particular importance during hot weather, excesses of nutrients such as protein can contribute t o lowered efficiency in the cow. The practice of boosting nutrients such as CP to high levels to minimize potential deficiencies can have negative consequences because nitrogen in excess of requirement must be metabolized and excreted, requiring energy. Recent research demonstrates that protein content, ruminally undegradable protein value and quality of amino acids delivered to the lower digestive tract play a complex role in the nutrition of the heat-stressed cow. Additional research is needed in this area to establish minimum and maximum protein content of the diet and to determine the effects of protein content on the efficiency of nutrient use by the cow. Heat increment of the dietary ingredients plays a role in both the efficiency with which nutrients are utilized and heat generation potential of the diet. There is a potential for formulation of diets that are particularly favorable for use during hot weather. High fiber diets result in production of large quantities of ruminal acetate, which is used with less efficiency than propionate. The efficiency of use for ME from high fiber diets is lower than for diets with high levels of rapidly fermentable carbohydrate, providing a scientific basis for the practice of feeding low fiber diets during hot weather. Low fiber diets must be implemented cautiously because of the necessity for sufficient fiber to promote good rumen function and t o maintain animal health and wellbeing. Use of very high quality forages for hot weather feeding may ensure both adequate content of digestible fiber and improved content of fermentable sugars to enhance production of rumen VFA. Dietary fats have a relatively low heat increment because of a high efficiency of use by the cow. Fats may be of particular value during hot weather and may be undervalued by current energy equations because of the lower heat increment. Results from research with fat feeding during hot conditions have been variable. Rumen inert fat allows inclusion of relatively high levels of fat in the diet, and experiments t o elucidate the effects of feeding these diets during hot weather should be conducted. Because high levels of dietary fat reduces the amount of fermentable energy in the rumen, the quantity of protein and ruminal escape value for that protein may be particularly critical when feeding high fat diets. Practical management considerations such as frequent feeding and ready water availability are logical. Water should be available in holding pens, travel alleys, and near feed bunks. Provision of fresh feeds through multiple feedings encourages frequent feeding bouts by cows, and making the feeding area as comfortable as possible through shading and(or) cooling should enhance feeding frequency and total intake. Animal behavior during hot weather indicates that the animal will consume more feed during cooler evening hours and feeding quantities and schedules should be adjusted to accommodate changes in behavior because of season of feeding. Research evaluating animal facilities and environmental conditions including temperature, humidity, and photoperiod for their effect on feeding behavior may be beneficial in designing feeding and management programs for the lactating cow during heat-stress conditions. Literature Cited Al-Haidary, A. A,, D. E. Spiers, G. E. Rottinghaus. and G. B. Garner. 1995. Effect of administration of ergovaline on the thermoregulatory functions of beef calves under heat stress. J. Anim. Sci. 73(Suppl. 1):132. Al-Haidary, A,, D. E. Spiers, Q. Zhang, G. E. Rottinghaus, and G. B. Garner. 1994. Shifts in thermoregulatory ability of beef calves following intake of endophyte-infected tall fescue during heat stress. J. Anim. Sci. 72!Suppl. 1):155. Alvarez, M. B., and H. D. Johnson. 1973. Environmental heat exposure on cattle plasma catecholamine and glucocorticoids. J. Dairy Sci. 56:189-194. Araki, C. T., R. M. Nakamura, L.W.G. Kam, and N. Clarke. 1984. Effect of lactation on diurnal temperature patterns of dairy cattle in hot environments. J. Dairy Sci. 67:1752-1760. Armstrong, D. G., and K. L. Blaxter. 1957. The heat increment of steam-volatile fatty acids in fasting sheep. Brit. J. Nutr. 11: 247-271. Bahman, A. M., J. A. Rooke, a n d J . H. Topps. 1993. The performance of dairy cows offered drinking water of low or high salinity in a hot arid climate. Anim. Prod. 57:23-28. Baker, C. C., C. E. Coppock, J. K. Lanham, D. H. Nave, J. M. LaBore, C. F. Brasington, and R. A. Stermer. 1988. Chilled drinking water effects on lactating Holstein cows in summer. J. Dairy Sci. 71:2699-2708. Baldwin, B. R., N. E. Forsberg, and C. Y. Hu. 1985. Potential for altering energy partition in the lactating cow. J. Dairy Sci. 68: 3394-3402. Baldwin, R. L., N. E. Smith, J. Taylor, and M. Sharp. 1980. Manipulating metabolic parameters t o improve growth rate and milk secretion. J. Anim. Sci. 51:1416-1428. Beede, D. K., and R. J. Collier. 1986. Potential nutritional strategies for intensively managed cattle during thermal stress. J. Anim. Sci. 62543-554. Belibasakis, N. G., P. Ambatzidis, P. Aktsali, and D. Tsirgogianni. 1995. Effects of degradability of dietary protein on milk production and blood components of dairy cows in hot weather. World Rev. Anim. Prod. 30:21-26. Benjamin, M. M. 1981. Fluid and electrolytes. In: Outline of Veterinary Clinical Pathology. pp 213-228. Iowa State UniT.. Press, Ames. J. Anim. Sci, Vol. 77, Suppl. 2/.J. Dairy Sci. Vol. 82, Suppl. 2/1999 MANAGING HEAT-STRESSED DAIRY COWS Blackshaw, J . K., and A. W. Blackshaw. 1994. Heat stress in cattle and the effect of shade on production and behaviour: A review. Aust. J . Exp. Agric. 34:285-295. Block, E. 1984. Manipulating dietary anions and cations for prepartum cows to reduce incidence of milk fever. J. Dairy Sci. 67: 2939-2948. Brokken, R. F. 1971. Formulating beef rations with varying levels of heat increment. J. Anim. Sci. 32692-703. Bucklin, R. A,, L. W. Turner, D. K. Beede, D. R. Bray, and R. W. Hemken. 1991. Methods to relieve heat stress for dairy cows in hot, humid climates. App. Eng. In Agnc. 7:241-247. Chan, S. C., J. T. Huber, K. H. Chen, J. M. Simas, and Z. Wu. 1997. Effects of ruminally inert fat and evaporative cooling on dairy cows in hot environmental temperatures. J. Dairy Sci. 80: 1172-1178. Chen, K. H., J. T. Huber, C. B. Theurer, D. V. Armstrong, R. C. Wanderly, J . M. Simas, S. C. Chan, and J. L. Sullivan. 1993. Effect of protein quality and evaporative cooling on lactational performance of Holstein cows in hot weather. J . Dairy Sci. 76: 819-825. Collier, R. J., D. K. Beede, W. W. Thatcher, L. A. Israel, and C. J. Wilcox. 1982a. Influences of environment and its modification on dairy animal health and production. J. Dairy Sci. 65: 2213-2227. Collier, R. J. , S. G. Doelger, H. H. Head, W. W. Thatcher, and C. J. Wilcox. 1982b. Effects of heat stress during pregnancy on maternal hormone concentrations, calf birth weight and postpartum milk yield of Holstein cows. J . h i m . Sci. 54:309-319. Coppock, C. E. 1985. Energy nutrition and metabolism of the lactating dairy cow. J. Dairy Sci. 68:3403-3410. Coppock, C. E., W. P. Flatt, L. A. Moore, and W. E. Stewart. 1964. Effect of hay to grain ratio on utilization of metabolizable energy for milk production by dairy cows. J. Dairy Sci. 47: 1330-1338. Cummins, K. A. 1992. Effect of dietary acid detergent fiber on responses to high environmental temperature. J. Dairy Sci. 75: 1465-147 1. Danfaer, A,, I. Thysen, and V. Ostergaard. 1980. The effect of the level of dietary protein on milk production. 1. Milk yield, liveweight gain and health. Beret. Statens Husdyrbrugsfors. 492. Di Costanzo, A,, J. N. Spain, and D. E. Spiers. 1997. Supplementation of nicotinic acid for lactating Holstein cows under heat stress conditions. J . Dairy Sci. 80:1200-1206. Dougherty, C. T., L. M. Lauriault, N. W. Bradley, N. Gay, and P. L. Cornelius. 1991. Induction of tall fescue toxicosis in heatstressed cattle and its alleviation with thiamin. J . Anim. Sci. 69:1008-1018. Escobosa, A,, C. E. Coppock, L. D. Rowe, Jr., W. L. Jenkins, and C. E. Gates. 1984. Effects of dietary sodium bicarbonate and calcium chloride on physiological responses of lactating dairy cows in hot weather. J. Dairy Sci. 67:574-584. Hassan, A,, and J . D. Roussel. 1975. Effect of protein concentration in the diet on blood composition and productivity of lactating Holstein cows under thermal stress. J . Agric. Sci. (Camb.) 85: 409-415. Higginbotham, G. E., J . T. Huber, M. V. Wallentine, N. P. Johnston, and D. Andrus. 1989a. Influence of protein percentage and degradability on performance of lactating cows during moderate temperature. J . Dairy Sci. 72:1818-1823. Higginbotham, G. E., M. Torabi, and J. T. Huber. 198913. Influence of dietary protein concentration and degradability on performance of lactating cows during hot environmental temperatures. J. Dairy Sci. 72:2554-2564. 33 Holter, J. B., H. H. Hayes, W. E. Urban, Jr., and A. H. Duthie. 1992. Energy balance and lactation response in Holstein cows supplemented with cottonseed with or without calcium soap. J. Dairy Sci. 75:1480-1494. Holter, J . B., J. W. West, and M. L. McGilliard. 1997. Predicting ad libitum dry matter intake and yield of Holstein cows. J. Dairy Sci. 80:2188-2199. Holter, J . B., J. W. West, M. L. McGilliard, and A. N. Pell. 1996. Predicting ad libitum dry matter intake and yields of Jersey cows. J. Dairy Sci. 79:912-921. Huber. J. T., G. Higginbotham, R. A Gomez-Alarcon, R. B. Taylor, K. H. Chen, S. C. Chan, and Z . Wu. 1994. Heat stress interactions with protein, supplemental fat, and fungal cultures. J. Dairy Sci. 77:2080-2090. Ibrahim, I. F., S.M. Seif, and H. D. Johnson. 1970. Plasma insulin and free amino acid changes with heat exposure in bovine. J. Dairy Sci. 53:651 (Abstr.). Igono, M. O., and H. D. Johnson. 1990. Physiological stress index of lactating dairy cows based on diurnal pattern of rectal temperature. J. Interdiscipl. Cycle Res. 21:303-320. Igono, M. O., H. D. Johnson, B. J. Steevens, W. A. Hainen, and M. D. Shanklin. 1988. Effect of season on milk temperature, milk growth hormone, prolactin, and somatic cell counts of lactating cattle. Intl. J. Biometeor. 32:194-200. Igono, M. O., H. D. Johnson, B. J. Steevens, G. F. Krause, and M. D. Shanklin. 1987. Physiological, productive, and economic benefits of shade, spray, and fan system versus shade for Holstein cows during summer heat. J. Dairy Sci. 70:1069-1079. Igono, M. O., B. J. Steevens, M. D. Shanklin, and H. D. Johnson. 1985. Spray cooling effects on milk production, milk and rectal temperatures of cows during a moderate temperature summer season. J . Dairy Sci. 68:979-985. Jenkinson, D. M., and R. M. Mabon. 1973. The effects of temperature and humidity on skin surface pH and the ionic composition of skin secretions in Ayrshire cattle. Brit. Vet. J. 129:282-295. Johnson, H. D. 1967. Climate effects on physiology and productivity of cattle. In: R. H. Shaw ( E d . ) Ground Level Climatology. Amer. Assoc. Adv. Sci. Pub. 86. Washington, DC. Johnson, H. D., L. Hahn, H. H. Kibler, M. K. Shanklin, and J. E. Edmondson. 1967. Heat and acclimation influences on lactation of Holstein cattle. Missouri Agric. Exp. Sta. Res. Bull. No. 916. Columbia. Johnson, H. D., P. S. Katti, L. Hahn, and M. D. Shanklin. 1988. Short-term heat acclimation effects on hormonal profile of lactating cows. Univ. of Missouri Rsch. Bull. No. 1061. Columbia. Johnson, H. D., A. C. Ragsdale, I. L. Berry, and M. D. Shanklin. 1963. Temperature-humidity effects including influence OF acclimation in feed and water consumption of Holstein cattle. Univ. of Missouri Res. Bull. No. 846. Johnson, H. D., and W. J. Vanjonack. 1976. Effects of environmental and other stressors on blood hormone patterns in lactating animals. J . Dairy Sci. 59:1603-1647. Kabuga, J . D., and K. Sarpong. 1991. Influence of weather conditions on milk production and rectal temperature of Holsteins fed two levels of concentrate. Int. J . Biometeorol. 34:226-230. Kamal, T. H., and H. D. Johnson. 1970. Whole body 4oK loss as a predictor of heat tolerance in cattle. J. Dairy Sci. 53:1734-1738. Kelly, R. O., F. A. Martz, and H. D. Johnson. 1967. Effect of environmental temperature on ruminal volatile fatty acid levels with controlled feed intake. J. Dairy Sci. 50:531-533. Kibler, H. H. and S. Brody. 1950. Influence of temperature, 5" t o 95"F, on evaporative cooling from the respiratory and exterior body surfaces in Jersey and Holstein cows, Missouri Agric. Exp. Sta. Bull. No. 461. Columbia. J. Anim. Sci. Vol. 77, Suppl. 215. Dairy Sci. Val. 82, Suppl. 2/1999 34 WEST Kibler, H. H., and S. Brody. 1953. Influence of humidity on heat exchange and body temperature regulation in Jersey, Holstein, Brahman and Brown Swiss cattle. Missouri Agric. Exp. Sta. Bull. No. 522. Columbia. Knapp, D. M., and R. R. Grummer. 1991. Response of lactating dairy cows to fat supplementation during heat stress. J . Dairy Sci. 74:2573-2579. Kronfeld, D. S., S.Donoghue, J . M. Naylor, K. Johnson, and C. A. Bradley. 1980. Metabolic effects of feeding protected tallow to dairy cows. J . Dairy Sci. 63:545-552. Kume, S., M. Kurihara, S.Takahashi, M. Shibata, and T. Aii. 1987. Effect of hot environmental temperature on major mineral balance in lactating cows. Jpn. J. Zootech. Sci. 58:764-770. Kume, S., M. Shibata, M. Kurihara, and T. Aii. 1986. Effect of environmental temperature on trace element metabolism of cows during feeding and fasting. Jpn. J. Zootech. Sci. 57: 687-693. Kume, S., S. Takahashi, M. Kurihara, and T. A i . 1989. Effect of hot environment on Ca and P metabolism in dairy cow. Asian-Aust. J . Anim. Sci. 2:259-260. Kurihara, M. 1996. Energy requirements and feed of dairy cows under high temperature conditions. Japan Agric. Rsch. Quarterly. 30:107-112. Lanham, J. K., C. E. Coppock, K. Z. Milam, J . M. Labore, D. H. Nave, R. A. Stermer, and C. F. Brasington. 1986. Effects of drinking water temperature on physiological responses of lactating Holstein cows in summer. J. Dairy Sci. 69:1004-1012. Leighton, R. E., and I. W. Rupel. 1956. Effects of fiber content of the diet on milk production and hot weather discomfort in producing dairy cows. J . Dairy Sci. 39:937 (Abstr.). Lough, D. S., D. K. Beede, and C. J. Wilcox. 1990. Effects of feed intake and thermal stress on mammary blood flow and other physiological measurements in lactating dairy cows. J . Dairy Sci. 73:325-332. MacRae, J. C., and G. E. Lobley. 1982. Some factors which influence thermal energy losses during the metabolism of ruminants. Livestk. Prod. Sci. 9:447-456. Magdub, A,, H. D. Johnson, and R. L. Belyea. 1982. Effect of environmental heat and dietary fiber on thyroid physiology of lactating cows. J . Dairy Sci. 65:2323-2331. Mallonee, P. G., D. K. Beede, R. J. Collier, and C. J . Wilcox. 1985. Production and physiological responses of dairy cows to varying dietary potassium during heat stress. J . Dairy Sci. 68: 1479- 1487. Maltz, E., N. Silanikove, U. Shalit, and A. Berman. 1994. Diurnal fluctuations in plasma ions and water intake of dairy cows as affected by lactation in warm weather. J. Dairy Sci. 77: 2630-2639. LMaust,L. E., R. E. McDowell, and N. W. Hooven. 1972. Effect of summer weather on performance of Holstein cows in three stages of lactation. J. Dairy Sci. 55:1133-1139. McDowell, R. E. 1972. The animal body in warm environments. In: Improvement of Livestock Production in Warm Climates. p 65. W. H. Freeman and Co., San Francisco, CA. McDowell, R. E., E. G. Moody, P. J. Van Soest, R. P. Lehmann, and G. L. Ford. 1969. Effect of heat stress on energy and water utilization of lactating cows. J. Dairy Sci. 52:188-194. McDowell, R. E., and J. R. Weldy. 1960. Water exchange of cattle under heat stress. Proc. Third Intl. Biometerological Cong., London. pp 414-424. Pergamon Press, New York. McGuire, M. A,, D. K. Beede, R. J . Collier, F. C. Buonomo, M. A. DeLorenzo, C. J. Wilcox, G. B. Huntington, and C. K. Reynolds. 1991. Effects of acute thermal stress and amount of feed intake on concentrations of somatotropin, insulin-like growth factor (1GF)-I and IGF-11, and thyroid hormones in plasma of lactating Holstein cows. J . Anim. Sci. 69:2050-2056. McGuire, M. A,, D. K. Beede, M. A. DeLorenzo, C. J. Wilcox, G. B. Huntington, C. K. Reynolds, and R. J . Collier. 1989. Effects of thermal stress and level of feed intake on portal plasma flow and net fluxes of metabolites in lactating Holstein cows. J. Anim. Sci. 67:1050-1060. Milam, K. Z., C. E. Coppock, J. W. West, J . K. Lanham, D. H. Nave, J. M. LaBore, R. A. Stermer, and C. F. Brasington. 1986. Effects of drinking water temperature on production responses in lactating Holstein cows in summer. J . Dairy Sci. 69: 1013-1019. Mitra, R., G. I. Christison, and H. D. Johnson. 1972. Effect of prolonged thermal exposure on growth hormone ( G H ) secretion in cattle. J. h i m . Sci. 34:776-779. Moe, P. W. 1981. Energy metabolism of dairy cattle. J . Dairy Sci. 64: 1120-1 139. Moody, E. G., P. J . Van Soest, R. E. McDowell, and G. L. Ford. 1967. Effect of high temperature and dietary fat on performance of lactating cows. J. Dairy Sci. 50:1909-1915. Muller, L. D., A. J . Heinrichs, J. B. Cooper, and Y. H. Atkin. 1986. Supplemental niacin for lactating cows during summer feeding. J. Dairy Sci. 69:1416-1420. Murphy, M. R., C. L. Davis, and G. C. McCoy. 1983. Factors affecting water consumption by Holstein cows in early lactation. J . Dairy Sci. 66:35-38. NRC. 1981. Effect of environment on nutrient requirements of domestic animals. National Academy Press, Washington, DC. NRC. 1989. Nutrient Requirements of Dairy Cattle. 6th rev. ed. Natl. Acad. Sci., Washington, DC. Nianogo, A. J.,H. E. A m o s , M. A. Froetschel, and C. M. Keery. 1991. Dietary fat, protein degradability, and calving season: effects on nutrient use and performance of early lactation cows. J . Dairy Sci. 74:2243-2255. O’Kelly, J. C. 1987. Influence of dietary fat on some metabolic responses of cattle to hyperthermia induced by heat exposure. Comp. Biochem. Physiol. 87A:677-682. Oldham, J. D. 1984. Protein-energy interrelationships in dairy cows. J . Dairy Sci. 67:1090-1114. Page, H. M., E. S. Erwin, and G. E. Nelms. 1959. Effect of heat and solar radiation on vitamin A utilization by the bovine animal. Am. J. Physiol. 196:917-918. Palmquist, D. L., and T. C. Jenkins. 1980. Fat in lactation rations: A review. J . Dairy Sci. 63:l-14. Parker, D. S. 1984. Metabolic limitations to milk production in the tropics. Trop. Anim. Prod. 9:251-256. Purwanto, B. P., Y. Abo, R. Sakamoto, F. Furumoto, and S. Yamamoto. 1990. Diurnal patterns of heat production and heart rate under thermoneutral conditions in Holstein F:-iesian cows differing in milk production. J . Agric. Sci. (Camb.) 114: 139-142. Reynolds, C. K., H. F. Tyrrell, and P. J. Reynolds. 1991. Effects of diet forage-to-concentrate ratio and intake on energy metabolism in growing beef heifers: Whole body energy and nitrogen balance and visceral heat production. J. Nutr. 121: 994-1003. Robinson, J. B., D. R. Ames, and G. A. Milliken. 1986. Heat production of cattle acclimated to cold, thermoneutrality, and heat when exposed to thermoneutrality and heat stress. J. Anim. Sci. 62:1434-1440. Sanchez, W. K., M. A. McGuire, and D. K. Beede. 1994. Macromineral nutrition by heat stress interactions in dairy cattle: review and original research. J . Dairy Sci. 77:2051-2079. J . Anim. Sci. Vol. 77, Suppl. 215. Dairy Sci. Vol. 82, Suppl. 2/1999 MANAGING HEAT-STRESSED DAIRY COWS Saunders, R. G., C. E. Coppock, D. L. Wilks, A. G. Lane, and C. E. Gates. 1990. Effects of supplemental fats on intake, production, and heat stress in lactating Holstein cows in summer. J. Dairy Sci. 73(Suppl. 1):193 (Abstr.). Schneider, P. L., D. K. Beede, and C. J. Wilcox. 1986. Responses of lactating cows to dietary sodium source and potassium quantity during heat stress. J. Dairy Sci. 69:99-110. Schneider, P. L., D. K. Beede, and C. J. Wilcox. 1988a. Effects of supplemental potassium and sodium chloride salts on ruminal turnover rates, acid-base and mineral status of lactating dairy cows during heat stress. J. Anim. Sci. 66:126-135. Schneider, P. L., D. K. Beede, and C. J. Wilcox. 1988b. Nyctohemeral patterns of acid-base status, mineral concentrations and digestive function of lactating cows in natural or chamber heat stress environments. J. Anim. Sci. 66:112-125. Schneider, P. L., D. K. Beede, C. J. Wilcox, and R. J. Collier. 1984. Influence of dietary sodium and potassium bicarbonate and total potassium on heat-stressed lactating dairy cows. J. Dairy Sci. 67:2546-2553. Seath, D. M., and G. D. Miller. 1948. Effect of water sprinkling with and without air movement on cooling dairy cows. J. Dairy Sci. 5:361-366. Silanikove, N. 1992. Effects of water scarcity and hot environment on appetite and digestion in ruminants: A review. Livestock Prod. Sci. 30:175-194. Skaar, T. C., R. R. Grummer, M. R. Dentine, and R. H. Stauffacher. 1989. Seasonal effects of prepartum and postpartum fat and niacin feeding on lactation performance and lipid metabolism. J. Dairy Sci. 72:2028-2038. Stott, G. H. 1981. What is animal stress and how is it measured? J. Anim. Sci. 52:150-153. Stott, G. H., and E. G. Moody. 1960. Tolerance of dairy cows to high climatic temperatures on low roughage ration. J. Dairy Sci. 43: 871 (Abstr.). Taylor, R. B., J. T. Huber, R. A. Gomez-Alarcon, F. Wiersma, and X. Pang. 1991. Influence of protein degradability and evaporative cooling on performance of dairy cows during hot environmental temperatures. J. Dairy Sci. 74:243-249. Tsai, Y. C., L. S. Castillo, W. A. Hardison, and W. J. A. Payne. 1967. Effect of dietary fiber level on dairy cows in the humid tropics. J. Dairy Sci. 50:1126-1129. Tucker, W. B., G. A. Harrison, and R. W. Hemken. 1988. Influence of dietary cation-anion balance on milk, blood, urine, and rumen fluid in lactating dairy cattle. J. Dairy Sci. 71:346-354. Tyrrell, H. F., P. W. Moe, and W. P. Flatt. 1970. Influence of excess protein intake on energy metabolism of the dairy cow. Fifth Symp. Energy Metab. Farm Anim. pp 69-71. 35 Tyrrell, H. F., P. J. Reynolds, and P. W. Moe. 1979. Effect of diet on partial efficiency of acetate use for body tissue synthesis by mature cattle. J. h i m . Sci. 48:598-606. Valtorta, S. E., M. R. Gallardo, H. C. Castro, and M E. Castelli. 1996. Artificial shade and supplementation effects on grazing dairy cows in Argentina. Trans. Amer. SOC.Agric. Eng. 39: 233-236. Van Soest, P. J. 1982. Limitations of ruminants. In: Nutritional Ecology of the Ruminant. pp 325-344. 0 & B Books, Inc., Corvallis, OR. Warren, W. P., F. A. Martz, K. H. Asay, E. S. Hilderbrand, C. G. Payne, and J. R. Vogt. 1974. Digestibility and rate of passage by steers fed tall fescue, alfalfa and orchardgrass hay in 18 and 32°C ambient temperature. J. Anim. Sci. 39:93-96. Wayman, O., H. D. Johnson, C. P. Merilan, and I. L. Berry. 1962. Effect of ad libitum or force-feeding of two rations on lactating dairy cows subject t o temperature stress. J. Dairy Sci. 45: 1472-1478. Webster, A. J. F., P. 0. Osuji, F. White, and J. F. Ingram. 1975. The influence of food intake on portal blood flow and heat production in the digestive tract of sheep. Brit. J. Nutr. 34:125-139. Wegner, T. N., and J. D. Schuh. 1988. Effect of water quality and season on milk production and water metabolism in Holstein cows. J. Dairy Sci. 71(Suppl. 1):185 (Abstr.). West, J. W., C. E. Coppock, K. Z . Milam, D. H. Nave, J. M. LaBore, and L. D. Rowe, J r . 1987. Potassium carbonate as a potassium source and dietary buffer for lactating Holstein cows during hot weather. J. Dairy Sci. 70:309-320. West, J. W., K. D. Haydon, B. G. Mullinix, and T. G. Sandifer. 1992. Dietary cation-anion balance and cation source effects on production and acid-base status of heat-stressed cows. J. Dairy Sci. 75:2776-2786. West, J. W., G. M. Hill, P. Mandebvu, J. M. Fernandez, and B. G. Mullinix. 1998. Effects of dietary fiber on intake, milk yield, and digestion by lactating dairy cows during cool or hot, humid weather. J. Dairy Sci. ( I n press). West, J. W., B. G. Mullinix, and T. G. Sandifer. 1991. Changing dietary electrolyte balance for dairy cows in cool and hot environments. J. Dairy Sci. 74:1662-1674. Wolfenson, D., I. Flamenbaum, and A. Berman. 1988. Dry period heat stress relief effects on prepartum progesterone, calf birth weight, and milk production. J. Dairy Sci. 71:809-818. Woodford, S. T., M. R. Murphy, and C. L. Davis. 1984. Water dynamics of dairy cattle as affected by initiation of lactation and feed intake. J . Dairy Sci. 67:2336-2343. Zook, A. B. 1982. The effects of two levels of dietary protein solubility on lactating cows exposed to heat stress and in thermal neutral environments. Diss. Abstr. Intl. 43:1760B. J. Anim. Sci. Vol. 77, Suppl. 215. Dairy Sci. Vol. 82, Suppl. 211999
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