Encyclopedia Of Animal Science - W doc

872 Water Water Mineral Composition Fig Typical body water compartments (% of total body water) rumen; the remaining proportion is presumably directed to the omasum Within the remainder of the ruminant gut, quantitative net water absorption is greatest in the proximal small intestine, followed by the omasum and large intestine.[5] Recent data[6] indicate that the lower net water absorption in the large intestine by cattle than by sheep results from a reduced ability to retain absorbed water because more absorbed solvent/solute is drawn back into the lumen through the larger paracellular pores between colonic cells in cattle Drinking water is a source of various minerals that are generally readily available for absorption unless complexed by an interfering nutrient Minerals ingested in water and feed are a variable mix of positively and negatively charged ions that contribute to the dietary cation anion difference of consumed material (DCAD) and have a direct influence on fluid and acid base balance The DCAD is calculated as the milliequivalents (mEq) of Na+, K+, Ca+ +, and Mg+ + minus the mEq of ClÀ, S=, and P=.[13] As anion consumption and concentration in the body increase, cellular acidosis can occur As the DCAD increases from negative to positive (e.g., À 20 to + 100 mEq/kg), feed intake and performance are generally increased However, the prepartum dairy cow is one exception to the generalization Inducing mild metabolic acidosis by feeding anionic diets before calving has been an effective means of preventing milk fever by potentiating calcium resorption from bone before the dramatic calcium needs at parturition arise.[13] Few data are available on the contribution of water minerals to overall DCAD Socha et al.[14] reported average mineral profiles of more than 3600 drinking water TRANSPORTATION-INDUCED DEHYDRATION Transport of animals on semitrailer trucks from the site of birth to the site of growing and finishing can involve periods of up to 24 hours or more without access to water, and variable magnitudes of dehydration can occur Feeder calves seem to lose approximately 3.3% of body weight during the loading and unloading process and can lose an additional 0.3 to 0.4% of body weight/hour of transport.[7,8] Weight losses of feeder pigs during transport can be up to 0.6% of body weight/hour.[9] Loss of gastrointestinal tract contents and carcass weight has accounted for 48 and 32%, respectively, of transport shrink by feeder steers[8] and has accounted for 62 and 27%, respectively, of transport weight loss by feeder pigs Feces, urine, and respiration accounted for 12.6, 26, and 60% of the water loss.[10] Water accounted for 80% of weight lost by wethers during 48 hours of feed and water deprivation.[11] Of total body water loss, 57% was from the intracellular compartment and 29% was from the gastrointestinal tract In steers deprived of water for days, thiocyanate space (assumed to be extracellular space) accounted for 47% of the weight lost (total loss = 16% of body weight).[12] Thiocyanate space decreased 23% and plasma volume decreased 28% during the 4-day period without water The exchange of water within the body in response to dehydration is depicted in Fig Fig Water change between body compartments during dehydration Water is osmotically drawn from transcellular and intracellular compartments to interstitial and plasma compart ments during dehydration, in response to losses by urinary and fecal excretion and insensible routes The magnitude of reduction in compartmental volumes is dependent on the degree of dehydration Water samples collected across the United States Assuming that a growing feedlot steer weighing approximately 300 kg and consuming kg of a mixed diet (>85% dry matter) meeting mineral requirements would drink 30 L of water/ day,[2] this steer would consume twice as much weight in water compared to the weight of feed consumed, and approximately to 7% of calcium, sodium, and sulfur consumed would be derived from water However, approximately 20% of chloride consumed would be derived from drinking water in this example The DCAD calculated for the surveyed samples[14] was approximately 0.4 mEq/kg Estimates of the contribution of drinking water minerals to overall DCAD are needed 873 CONCLUSION The polarity and ability of water to facilitate hydration of polar and ionic molecules are central to the flow of water and metabolites within the body Saliva appears to be a greater proportion of ruminal fluid than previously thought, considering recent observations that some water consumed by drinking in nonsuckling cattle bypasses the rumen, but more intensive study is needed The ability of sheep to form drier feces than cattle results from tighter junctions between colonic cells and a greater ability to establish an osmotic gradient to retain absorbed water Cattle may lose approximately 3% of body weight during loading and unloading for transport, plus an additional 0.3 to 0.4% of body weight per hour of transport Indirect data suggest that water may constitute up to 80% of this weight loss Estimates of the contribution of drinking water minerals to overall cation anion difference and of the influence of water cation anion difference on animal performance are needed 10 11 12 13 REFERENCES 14 Bohinsky, R.C Modern Concepts in Biochemistry, 5th Ed.; Allyn and Bacon, Inc.: Boston, MA, 1987 Parker, D.B.; Brown, M.S Water Consumption for Livestock and Poultry Production In Encyclopedia of Water Science, 1st Ed.; Stewart, B.A., Howell, T.A., Eds.; Marcel Dekker, Inc.: New York, NY, 2003 Christopherson, R.J.; Webster, A.J.F Changes during eating in oxygen consumption, cardiac function and body fluids of sheep J Physiol 1972, 221, 441 457 Zorrilla Rios, J.J.; Garza, D.; Owens, F.N Fate of Drinking Water in Ruminants: Simultaneous Comparison of Two Methods to Estimate Ruminal Evasion; Animal Science Research Report MP 129; Oklahoma Agricultural Experiment Station: Stillwater, OK, 1990; 167 169 Sklan, D.; Hurwitz, S Movement and absorption of major minerals and water in ovine gastrointestinal tract J Dairy Sci 1985, 68, 1659 1666 McKie, A.T.; Goecke, I.A.; Naftalin, R.J Comparison of fluid absorption by bovine and ovine descending colon in vitro Am J Physiol 1991, 261, G433 G442 Bartle, S.J.; Preston, R.L Feedlot Cattle Receiving Experiments, 1988 89; Animal Science Research Report # T 263; Texas Tech University: Lubbock, TX, 1989; 28 30 Self, H.L.; Gay, N Shrink during shipment of feeder cattle J Anim Sci 1972, 35, 489 494 Jesse, G.W.; Weiss, C.N.; Mayes, H.F.; Zinn, G.M Effect of marketing treatments and transportation on feeder pig performance J Anim Sci 1990, 68, 611 617 Mayes, H.F.; Hahn, G.L.; Becker, B.A.; Anderson, M.E.; Nienaber, J.A A report on the effect of fasting and transportation on liveweight losses, carcass weight losses and heat production measures of slaughter hogs Appl Eng Agric 1988, 4, 254 258 Cole, N.A Influence of a three day feed and water deprivation period on gut fill, tissue weights, and tissue composition in mature wethers J Anim Sci 1995, 73, 2548 2557 Weeth, H.J.; Sawhney, D.S.; Lesperance, A.L Changes in body fluids, excreta and kidney function of cattle deprived of water J Anim Sci 1967, 26, 418 423 Goff, J Factors to Concentrate on to Prevent Periparturient Disease in the Dairy Cow, Proceedings of the Mid South Ruminant Nutrition Conference, Texas Agricultural Ex tension Service: College Station, TX, 1998; 63 Socha, M.T.; Ensley, S.M.; Tomlinson, D.J.; Ward, T Water composition variability may affect performance Feedstuffs 2003, 75 (24), 10 Water Buffalo Nguyen van Thu Cantho University, Can Tho City, Vietnam INTRODUCTION The water buffalo is considered to be a very useful animal in many countries, supplying draft power, meat, milk, and other by-products such as hides, horn, etc The water buffalo is closely associated with water or mud, and with smallholder farmers in the rice fields In recent years, buffalo production has developed well, not only in Asia, but also in Europe, South America, and other continents where the buffalo has been introduced This article aims to introduce some basic knowledge of the water buffalo, with an emphasis on its great contribution to our living standards and improved productivity that could be better exploited for a more sustainable agriculture development in the 21st century The water buffalo can be classified into two breed types, the River type (2n = 50) and the Swamp type (2n = 48) River breeds consist of: 1) Asian breeds such as those in India and Pakistan (including Murrah, Nili Ravi, Surti, etc.; and 2) Mediterranean breeds found in Italy, Romania, and the Middle East The skin of River buffaloes is black, but some specimens have a dark slatecolored skin The horns of the River buffalo grow downward and backward, then curve upward in a spiral The Swamp type is found mainly in China and Southeast Asia The skin of the Swamp buffaloes is gray at birth, but becomes slate blue later Albinoid Swamp buffaloes are quite common in some areas, for example, in the north of Thailand Normally, the horns of Swamp buffaloes are longer than those of the River buffaloes, grow outward, and curve in a semicircle More than 70% of the buffaloes in the world belong to the River type.[2] TAXONOMY AND TYPES MEAT, MILK, AND DRAFT ATTRIBUTES The world’s buffaloes are classified into two groups, the African and the Asian, with genus names Syncerus and Bubalus, respectively According to the zoological classification,[1] buffaloes belong to the class Mammalia, subclass Ungulata, order Artiodactila, suborder Ruminantia, family Bovidae, subfamily Bovinae, tribe Bovini The tribe Bovini includes three groups: Bovina (cattle), Bubalina (the Asian buffalo), and Syncerina (the African buffalo) The Asian and African buffaloes are generally similar, but there are some anatomic differences The African buffalo includes only one species, Syncerus caffer, while the Asian buffalo comprises three species: Anoa (Bubalus depressicornis) from the Island of Celebes, Tamarao (Bubalus mindorensis) from the Island of Mindoro, and Arni (Bubalus Arnee), or the Indian wild buffalo Of these four species of African and Asian buffalo, only the India Arni buffalo has been domesticated and given the species name bubalis Therefore, the domestic buffalo currently reared with the name of water buffalo is classified as bubalus bubalis It is believed that the domestication of the buffalo occurred about 5000 years ago on the Indian subcontinent, and the domestication of the Swamp buffalo took place in China about 1000 years later 874 In general, the River types are mainly used for milk in South Asian countries, while the Swamp types are used for draft power in Southeast Asian countries and China (Table 1) However, both the River and Swamp types have been used for multiple purposes such as work, milk, meat, manure, fuel, etc by small farmers in different crop livestock farming systems In addition, crossbreeding programs of the River and Swamp buffaloes have shown great potential for improving meat, work, and milk outputs Recently, the U.S Department of Agriculture (USDA) estimated the nutritional value of water buffalo meat and compared it to beef and chicken The findings showed that water buffalo meat has 41% less cholesterol, 92% less fat, and 56% fewer calories than traditional beef Furthermore, there are as yet no reports on the occurrence of bovine spongiform encephalopathy (BSE), also known as mad cow disease, in buffaloes in any part of Asia.[4] The milk yield of the buffalo is lower than that of cattle, and average milk production is 1500 kg per lactation However, some individuals can produce 3500 kg per lactation Buffalo milk has high nutritive value and is excellent for the preparation of dairy products Using Encyclopedia of Animal Science DOI: 10.1081/E EAS 120019837 Copyright D 2005 by Marcel Dekker, Inc All rights reserved Water Buffalo 875 Table Plowing and harrowing performance of swamp buffaloes in the Mekong delta, Vietnam Sex Female (na = 24) Mean ± std Criteria Plowing timeb (hrs/day) Plowed area (ha/pair/day) Harrowing timeb (hrs/day) Harrowed area (ha/pair/day) Male (na = 24) Mean ± std 5.35 ± 0.58 5.39 ± 0.31 0.29 ± 0.025 0.31 ± 0.035 5.05 ± 0.17 5.28 ± 0.30 0.73 ± 0.167 0.77 ± 0.170 a In pair With a break (From Ref 3.) b buffaloes for a single purpose makes them less competitive with cattle and tractors This is believed to be an important reason for the serious decline of the buffalo population in a number of Southeast Asian countries, including some parts of Vietnam Alexiev reported that the Swamp-type Wenzhou buffalo in China can give an average milk yield of 1030 kg per 280-day lactation.[2] Thus, milk production of the Swamp buffalo is sufficient for family consumption In addition, the Swamp buffalo provides draft power, and thus has potential in rural areas of China and Southeast Asian countries In Europe and the Near East, the main purpose of raising buffaloes is for milk Milk can be used for liquid consumption and making different cheeses or yogurt, particularly in Italy, where most of the buffalo milk is used for making a well-known cheese called Italian Mozarella, which retails at a very high price.[5,6] The total number of buffalo in the World in 2002 was about 167,126,000 and it is increasing, particularly in India, China, Brazil, etc., where the River buffaloes are raised However, there is a serious reduction of the Swamp buffalo population in some countries such as Thailand, Malaysia, and Cambodia due to mechanization, overslaughtering for meat, and other reasons (Table 2) In many cases, knowledge from studies on cattle can also be applied to buffalo research and practices However, differences in anatomy, physiology, feeding behavior, reproductive characteristics, and productivity between the species have been reported.[8] The water buffalo is a ruminant, and the rumen reticulum of buffaloes is similar to that of cattle However, it is heavier than in cattle and 10% more capacious.[9] Studies comparing buffaloes to cattle have suggested a higher feed intake, longer retention time of feed in the digestive tract, longer rumination, less depression of cellulose digestion by soluble carbohydrates, a wider range of plant preferences, and a higher population of cellulolytic bacteria.[9] However, some authors have found no significant difference in feed digestibility between the two species It was suggested that the better performance of buffaloes fed coarse fodder may not be related to a superior capacity for fiber digestion, but rather that they are less discriminating against plants not readily eaten by cattle In Colombia, cattle are sometimes first used to graze pasture, whereafter buffaloes are allowed to graze the remaining and less desirable parts of the sward.[10] Recently, in a comparative study on cattle and Swamp buffaloes raised under the same village conditions, some authors reported higher bacteria, lower protozoa, and higher fungal zoospore counts in Swamp buffaloes.[11] It was also found that the Swamp buffalo can adapt better in the acid sulphate soil areas compared to the cattle and goats in the Mekong delta of Vietnam Based on results of a number of studies, buffalo might utilize protein more efficiently than cattle.[9] An ability of buffaloes to utilize endogenous urea more efficiently than cattle may explain in part their apparent superiority in utilizing high-fiber and low-nitrogen feed resources It is concluded that there have been contradictory results for fiber digestion abilities of buffaloes compared to cattle Buffaloes, however, seem to have a superior ability to consume coarse roughage, perhaps as a result of a better rumination capacity There is evidence that urea recycling and purine excretion in buffaloes are different from those in cattle, but more comprehensive studies are lacking Table Buffalo population (head) in the world and in selected countries (1970 2000) 1970 World India China Brazil Italy (From Ref 7.) 1980 1990 2000 107,437,984 56,118,000 15,713,063 118,000 48,600 121,757,733 66,070,000 18,439,152 495,000 88,900 148,184,210 80,570,000 21,421,975 1,397,097 112,400 164,339,658 93,772,000 22,596,439 1,102,551 201,000 876 Water Buffalo CONCLUSION It may be concluded that the water buffalo has a great potential to develop in the future A number of promising buffalo farming models have been developed in Brazil, Australia, Italy, Philippines, Colombia, etc Valuable products of water buffaloes, such as milk, meat, draft power, and manure, are relevant for the people and our living environment, particularly with respect to the trend toward organic agriculture in many parts of the world REFERENCES Alexiev, A The Water Buffalo; St Kliment Ohridski University Press: Sofia, 1998 Chantalakhana, C Long term breeding strategies for ge netic improvement of buffaloes in developing countries Asian Aust J Anim Sci 1999, 12, 1152 1161 Thu, N.V A Study of Performance, Physiological Param eters and Economic Efficiency of Working Buffaloes in the Mekong Delta of Vietnam In Working Animals in Agriculture and Transport; Pearson, R.A., Lhoste, P., Saastamoinen, M., Martin Rosset, W., Eds.; EAAP Tech nical Series, Wageningen Academic Publisher, 2003; Vol 6, 165 171 Ranjhan, S.K A Vision of buffalo production with special 10 11 reference to milk and meat production Proc Symp Series of the 8th World Conf Anim Prod., Seoul, Korea, June 28 July 4, 1998; 263 270 Borghese, A.; Moioli, B.; Tripadi, C Processing and Product Development in Mediterranean Countries In Proceedings of the Third Asian Buffalo Congress, Kandy, Sri Lanka, 2000; 37 46 Chantalakhana, C Long term breeding strategies for genetic improvement of buffaloes in developing countries Asian Aust J Anim Sci 1999, 12 (7), 1152 1161 FAO Live Animals FAOstat Agriculture Data; 2003 http://apps.fao.org/page/collections?subset = agriculture Cockrill, W.R The Husbandry and Health of Domestic Buffalo; FAO: Rome, 1974 Khajarern, S.; Khajarern, J.M Feeding Swamp Buffalo for Milk Production In Feeding Dairy Cows in the Tropics; FAO Animal Production and Health Paper, Wageningen Academic Publishers: The Netherlands, 1991; Vol 86, 115 125 Thu, N.V A Study of the Use of Female Cattle and Buffalo Crusing Sugar Cane in Colombia M.Sc Thesis; Swedish University of Agricultural Sciences: Uppsala, Sweden, Food and Agriculture of the United Nations, 1994 Wanapat, M.; Ngarmsang, A.; Korkhuntot, S.; Nontaso, N.; Wachirapakorn, C.; Keakes, G.; Rowlinson, P A comparative study on the rumen microbial population of cattle and Swamp buffalo raised under traditional village conditions in the Northeast of Thailand Asian Aust J Anim Sci 2000, 13 (7), 918 921 Well-Being and Handling Temple Grandin Colorado State University, Fort Collins, Colorado, U.S.A INTRODUCTION Reducing stress during handling for procedures such as vaccinations, milking, and herding will improve both animal welfare and productivity Pigs and dairy cows that are afraid of people have reduced productivity Pigs have lower weight gains and fewer piglets and dairy cows produce less milk Fearfulness was assessed by measuring the animal’s willingness to approach people Cows on dairies where the employees had received training in stockmanship and animal behavior had a smaller flight zone and gave more milk.[1] The trained employees engaged in fewer negative interactions with the cows, such as hitting or yelling Further studies have shown that wild, excitable cattle that become highly agitated in the squeeze chute had lower weight gains,[2] poor beef quality, and tougher meat squeeze chute, sweating in horses when there is little physical exertion, flapping in caged layers, and a horse rearing when he is suddenly startled Isolation is a strong stressor, and a single cow or lamb may run into a fence or try to jump it when it is separated from its herdmates Physiological measures such as cortisol in the blood can also be used as indicators of fear stress that occurs during nonpainful restraint in a squeeze chute.[4] Cortisol is a time-dependent measure and it takes 10 to 20 minutes for it to reach peak levels It is important to differentiate between fear and pain stress Cortisol levels can also rise in response to pain from procedures such as hot iron branding The variable of the handling stress needs to be separated from the variable of pain caused by a procedure such as castration Handling stress is mostly fear, and stress from castration is caused by pain and injury to tissues BIOLOGICAL BASIS OF FEAR VARIATIONS IN HANDLING STRESS Fear is a strong stressor and it can be detrimental to both productivity and welfare People working with animals should take steps to reduce the animal’s fear Other stressors such as weather extremes often cannot be avoided, but livestock producers can easily reduce fear Fear is a basic emotion and it motivates animals to avoid predators The amygdala is the brain’s fear center.[3] If the amygdala is destroyed, the animal will no longer become fearful of things that would normally cause fear, such as sudden loud noise It also loses learned fear responses An example of a learned fear response is refusing to enter a squeeze chute for vaccinations because the cow was accidentally hit on the head by the headgate In wild animals that are not accustomed to handling, destruction of the amygdala will make them act tame Fear stress during handling can vary from almost none to extreme Extensively raised cattle that were not accustomed to close contact with people had much higher cortisol levels when they were restrained in a squeeze chute compared to hand-reared dairy cattle.[5] Taming of an animal may reduce physiological reactivity of the nervous system Hand-reared deer that were raised in close contact with people had significantly lower cortisol levels after restraint than free-range deer.[6] There are three basic variables that will affect both the intensity of fear stress during handling and the size of the animal’s flight zone They are: 1) genetic factors; 2) amount of contact with people; and 3) previous experiences with handling that can be either aversive or nonaversive INDICATORS OF FEARFULNESS GENETIC FACTORS One indicator of fearfulness in grazing animals is the size of the flight zone Animals with larger flight zones are more fearful Another indicator is the startle response to a sudden stimulus such as a firecracker Some other behavioral indicators of fear are a cow struggling in a The domestic phenotype has reduced responses to changes in its environment.[7] Several studies have shown that there are differences in how different breeds of cattle react to handling Brahman cattle had higher cortisol levels after restraint than crosses of the English breeds such as Encyclopedia of Animal Science DOI: 10.1081/E EAS 120019847 Copyright D 2005 by Marcel Dekker, Inc All rights reserved 877 878 Hereford or Angus Some genetic lines of cattle, pigs, or chickens are more likely to be extremely agitated during handling Animals that have flighty, excitable, high-fear genetics are more likely to become highly agitated when they are suddenly placed in a new situation, compared to animals with a calmer temperament Flighty animals have to be introduced more gradually to new things to avoid agitation and panic, compared to animals with a calmer temperament An experiment by Ted Friend showed that measurements of epinephrine (adrenalin) showed that some pigs habituated to a novel, nonpainful swimming task where they were suddenly placed in a pool of water The task was repeated over a series of days In some of the pigs, the elevated epinephrine levels returned to normal and in other individuals, the epinephrine levels remained high Some of the pigs lost their fear of swimming and others remained scared Genetic factors may have accounted for these differences EFFECT OF PREVIOUS EXPERIENCES An animal’s previous experiences with handling will affect how it will react in the future Cattle that had been accidentally bumped on the head in a squeeze chute were more reluctant to reenter the chute a month later Sheep that had been turned upside down in a restraint device were more reluctant to reenter the facility the following year compared to sheep that were restrained in an upright position.[8] It is important that an animal’s first experience with a new person or new place be a good one Progressive ranchers walk cows and calves through the corrals prior to doing procedures so that they will associate corrals with being fed Sometimes painful procedures have to be done, but it is recommended that they not be associated with the animal’s first experience with either a new person or a new place A rat experiment indicated that if a rat was shocked severely the first time it entered a new arm on a maze, it would never enter that arm again However, if the rat was fed the first time he went into the new arm and then subjected to gradually increasing shocks, he would keep entering the arm to get the food.[9] FEAR MEMORIES If an animal is subjected to either a frightening or a painful experience, it may form a permanent fear memory that cannot be erased.[3] This memory is formed in the lower subcortical pathway in the brain, and extinguishing the Well-Being and Handling conditioned fear is difficult because it has to be suppressed by an active learning process that requires input from higher parts of the cortex The fear memory is suppressed by the cortex, but it can sometimes reappear Careful, quiet handling of animals will help prevent the formation of fear memories that may compromise welfare, lower productivity, or cause behavior problems, as in horses Animals can associate certain types of clothing or a person’s voice with either a frightening or a painful experience Animals also have the ability to recognize the voice of a familiar safe person who can calm them down FEAR OF NOVELTY New experiences and new things are both scary and attractive to animals They are attractive when the animal is allowed to voluntarily approach, but frightening when suddenly introduced.[7] If a flag is placed in the middle of a large field, cattle and horses will approach it and investigate However, if the same flag is suddenly waved next to a horse, he may become highly agitated.[7] Animals can be trained to tolerate new things if they are gradually introduced Cattle should become accustomed to being handled and fed by different people in different vehicles This will help reduce stress when they are moved to a new place Training animals to tolerate new experiences will help keep them calmer It is important to train cattle on being moved by both people on foot and people on horses Cattle appear to perceive a person riding a horse and a person walking on foot as two different things TRAIN FOR HANDLING Training calves and pigs to handling procedures helps to produce calmer adult animals Pigs differentiate between a person in the aisle and a person in their pens Pigs will move more easily in and out of trucks and through chutes at a meat plant if the producer trained them by walking through their pens several times each week Animals will have the lowest amount of fear stress when they voluntarily cooperate with being restrained and handled Zoos and aquariums are training animals, such as apes, lions, and dolphins, to cooperate with blood testing and veterinary procedures Highly excitable Bongo antelope were trained to enter a box and allow blood samples to be taken when they were fed treats Almost baseline cortisol (stress hormone) levels were obtained The levels of glucose in the blood of trained animals was significantly lower compared to the same animal immobilized with a dart.[10] Well-Being and Handling CONCLUSIONS Reducing fear during handling will improve animal productivity.[1] There are many different stressors that animals encounter such as stimuli that evoke fear, heat stress, cold stress, pain, or fatigue Fear is a strong stressor and it is one stressor that is easy to reduce Fearful animals have lower productivity Animals remember frightening or painful events and producers should be careful to avoid creation of fear memories An animal’s first experience with a new corral or person should be low stress Training animals to handling procedures will help reduce fear stress Both animal welfare and productivity will be improved by reducing fear stress 879 ARTICLE OF FURTHER INTEREST Animal Handling-Behavior, p 22 REFERENCES Hemsworth, P.H.; Coleman, G.J.; Barnett, J.C.; Berg, S.; Dowling, S The effect of cognitive behavioral interven tions on the attitude and behavior of stock persons and the behavior and productivity of commercial dairy cows J Anim Sci 2002, 80, 68 78 10 Voisinet, B.D.; Grandin, T.; Tatum, J.D.; O’Connor, S.F.; Struthers, J.J Feedlot cattle with calm temperaments have higher daily weight gains than cattle with excitable temperaments J Anim Sci 75, 892 896 LeDoux, J The Emotional Brain; Simon and Schuster: New York, New York, 1996 Grandin, T Assessment of stress during handling and transport J Anim Sci 1997, 75, 249 257 Lay, D.C.; Friend, T.H.; Bowers, C.C.; Grissom, K.K.; Jenkins, O.C A comparative physiological and behavioral study of freeze and hot iron branding using dairy cows J Anim Sci 1992, 70, 1121 1125 Hastings, B.E.; Abott, D.E.; George, L.M.; Staler, S.G Stress Factors influencing plasma cortisol levels and adrenal weights in Chinese water deer Res Vet Sci 1992, 53, 375 380 Grandin, T.; Deesing, M.J Behavioral Genetics and Animal Science In Genetics and the Behavior of Domestic Animals; Grandin, T., Ed.; Academic Press: San Diego, CA, 1998; 30 Hutson, G.D The influence of barley food rewards on sheep movement through a handling system Appl Anim Behav Sci 1985, 14, 263 273 Miller, N.E Learning resistance to pain and fear, effects of over learning exposure and rewarded exposure in context J Exp Psych 1960, 60, 137 142 Phillips, M.; Grandin, T.; Graffam, W.; Irlbeck, N.A.; Cambre, R.C Crate conditioning of Bongo (Trage laptous eurycerus) for veterinary and husbandry proce dures at Denver Zoological Garden Zoo Bio 1998, 17, 25 32 Well-Being Assessment: Behavioral Indicators J C Swanson M Rassette Kansas State University, Manhattan, Kansas, U.S.A INTRODUCTION Animal well-being can be characterized as the harmony an animal is experiencing mentally and physically with its environment Animal well-being is often used interchangeably with the term animal welfare Domestic livestock and poultry are raised under a variety of environmental conditions that are vastly different from those of their wild ancestors The scientific assessment of the well-being of livestock and poultry has become important to the sustainability of raising them for food The best scientific approach and criteria to assess animal well-being have yet to achieve a scientific consensus, but it is generally accepted that behavior, physiology, health, productivity, cognition, and system ecology are indicators of animal well-being BEHAVIORAL INDICATORS The repertoire of behavior expressed by a domestic animal reflects a living history of its natural and artificial selection Generally, behavior is used to identify and assess animal needs, preferences, state of health, ability to adapt and cope with its social and physical environment, emotional state, and to gain insight into what an animal may comprehend or feel about its environment Several behavioral indicators are commonly cited as useful to understanding and assessing animal well-being including abnormal behavior, posture, vocalization, responsiveness, grooming and displacement behavior, preferences animals express toward features of their living environment, and the presence/absence of stereotypies Abnormal Behavior The use of abnormal behavior as an indicator of wellbeing requires a clear knowledge of what constitutes normal behavior for a species Species behavior is sequenced, measured, described, and recorded to construct an ethogram The ethogram characterizes both instinctive and learned behavior displayed throughout a species’ life cycle Ethograms of wild ancestors, close relatives, or feral members of the same species are useful in studying 880 the behavioral similarities and differences induced by domestication An example of abnormal behavior is an outbreak of tail biting in pigs The interpretation of behavior elicited under domestic conditions is complicated and requires that we understand the cause, developmental aspects, and function of the behavior within the construct of the evolutionary and domestic history of the species Posture The posture of an animal represents a coping response to a stimulus Posture is often coupled with other behavioral indicators such as vocalization and locomotion to assess well-being Researchers have studied the usefulness of posture to correctly assess the amount of pain and distress an animal may experience after being subjected to common animal management procedures For example, a behavioral method using posture was validated to assess acute pain associated with different castration procedures used on lambs.[1] Each procedure was ranked according to an established index of expected pain Physiologic and behavioral data (including posture) were then collected for a period of 60 minutes postprocedure The data were analyzed according to the ability to place a lamb into the correct procedure group A combination of behavior and posture data correctly placed 79% of the lambs into their respective treatment groups.[1] As technology advances, so too does the sophistication of using an animal’s posture or movement for assessing well-being For example, computer image analysis has been used to measure the severity of head movements of cattle undergoing various types of branding to measure their aversion to the procedure,[2] and to evaluate the thermal comfort of pigs based on their proximity to one another.[3] While assessments must be validated for other species and for different types of practices, postural measures appear to be useful behavioral indicators of well-being Vocalization Animals convey a range of emotional states through various types of vocalizations Vocalizations are contextEncyclopedia of Animal Science DOI: 10.1081/E EAS 120019844 Copyright D 2005 by Marcel Dekker, Inc All rights reserved Well-Being Assessment: Behavioral Indicators specific, and the circumstances under which vocalizations are emitted must be carefully considered For example, a recent study compared the vocalizing of cattle in slaughter plants before and after modifications were made in animal handling procedures.[4] The data were used to evaluate the effectiveness of the plant modifications Indeed, a reduction in observable aversive events (prod use, slippage, excessive restraint pressure) decreased the amount of vocalization behavior.[4] Other researchers have found similar uses for vocalization in different species One study measured the occurrence and frequency of calls in piglets being castrated, and found a significantly greater rate of high-frequency calls (> 1000 Hz) compared with controls who were handled similarly but not actually castrated.[5] The researchers were able to isolate the most painful part of the procedure itself, and the effect these vocalizations had on other piglets, both of which have important implications for well-being Responsiveness The degree of an animal’s responsiveness to stimuli also acts as an indicator of well-being For example, the attitudes and behavior of dairy stockpersons toward cows have been researched and a correlation found between the stockperson’s behaviors and the avoidance distance of cows.[6] Avoidance behavior can shed light on an animal’s past relationship with humans and reflect the well-being of individuals or groups Another example of responsiveness as an indicator of well-being comes from a study using tonic immobility.[7] Tonic immobility is a state of petrification induced by positioning a bird on its back or side consequently, no movement is detected for a given period of time The time until the bird recovers head movement, stands, and walks is measured Shorter latencies to recovery indicate a better coping response by the bird Reduced or absent responsiveness of an animal has been recognized as an indicator of poor well-being Grooming and Displacement Behaviors Grooming as a social and self-maintenance behavior can reflect the relative well-being of an individual or an entire group Disruption or abnormal manifestations of grooming are measurable events The lack of grooming, indicated by poor hair/fur coat or feather condition, is often used as an indicator of sickness or depression for individual animals Abnormal pulling of hair/fur or feathers or obsessive grooming activities may occur in individuals or within groups Both are considered abnormal A displacement behavior is the result of frustration or behavioral disinhibition, or is performed when an animal is in conflict with how to behave in a given set of 881 circumstances For example, abnormal feather pecking in laying hens may be the displaced behavior of natural foraging or dustbathing and has been used to assess different housing conditions of egg-laying hens.[8] Feather pecking in hens can lead to significant feather loss or even skin damage Thus, the occurrence of displacement behavior and abnormal forms of grooming can be measured and used to assess well-being Preferences Preference tests are valuable tools to evaluate stimuli or conditions by appealing to the desires of the animal For example, such tests can be used to assess the effects on well-being of different enrichment devices or housing conditions In one study, researchers tested the preferences of dairy cattle for different kinds of flooring sand, straw, or a soft rubber mat.[9] The cows avoided sand and preferred either the mat or straw The researchers then tested whether a preference existed between the mat and straw They found that cattle preferred straw in winter, but in summer, cows showed no special preference for one system over the other Preference testing of this type allows for better design of housing systems However, extreme care must be taken when designing and drawing conclusions from such tests For example, exposure to resource cues can affect the performance of an animal in preference tests.[10] Cues such as odors can be undetectable to humans, but obvious to animals Carefully controlled preference tests are useful in validating the needs and choices of animals Stereotypies Stereotypy is a common abnormal behavior observed in intensively farmed species and thought to be the product of impoverished environments Stereotypies are behavior patterns repeated without variation and appear to have no obvious goal or function Examples include bar-biting; fur, hair, or wool chewing; sham chewing; tongue lolling; and a variety of locomotion patterns such as headweaving Once developed, stereotypies can be difficult to extinguish, even when animals are moved into more enriched environments This indicates an addictive quality to the behavior that requires an understanding of its neurophysiological development Performance of stereotypic behavior is often cited as an indicator of poor well-being Researchers have studied stereotypies in nearly all farmed species, including those farmed for fur, such as mink raised in cages.[11] Potential remedies such as environmental enrichment are often explored to provide relief However, the view that all stereotypies indicate poor well-being is controversial.[12–14] Performance of 882 Well-Being Assessment: Behavioral Indicators stereotypy could also indicate excitement or anticipation of a resource Thus, stereotypic behaviors are complex and must be fully examined to determine the effect on well-being Although the motivation to stereotype in domestic species has been researched, the neurophysiological implications are only beginning to be elucidated For example, recent studies have linked altered brain functioning and enhanced frustration to stereotypies found in caged birds.[15] Greater understanding of the disruption to brain function could eventually adjudicate the competing views on stereotypic behavior At present, the exhibition of stereotypies in domestic animals should prompt a closer look at other well-being indicators to further assess the possibility of a poor state of well-being CONCLUSION Behavior is one of several indicators used to assess animal well-being There is still much to be learned about the behavior of our domestic livestock and poultry and what constitutes a state of good well-being or contentment Although scientific consensus has not been reached regarding good versus poor well-being, there is general agreement that behavior provides insight into factors that promote or detract from an animal’s quality of life 10 11 12 REFERENCES Molony, V.; Kent, J.E.; McKendrick, I.J Validation of a method for assessment of an acute pain in lambs Appl Anim Behav Sci 2002, 76 (3), 215 238 Schwartzkopf Genswein, K.S.; Stookey, J.M.; Crowe, T.G.; Genswein, B.M Comparison of image analysis, exertion force, and behavior measurements for use in the assessment of beef cattle responses to hot iron and freeze branding J Anim Sci 1998, 76 (4), 972 979 Xin, H Assessing swine thermal comfort by image 13 14 15 analysis of postural behaviors J Anim Sci 1998, 77 (supplement 2), Grandin, T Cattle vocalizations are associated with handling and equipment problems at beef slaughter plants Appl Anim Behav Sci 2001, 71 (3), 191 201 Weary, D.M.; Braithwaite, L.A.; Fraser, D Vocal response to pain in piglets Appl Anim Behav Sci 1998, 56 (2 4), 161 172 Waiblinger, S.; Menke, C.; Coleman, G The relationship between attitudes, personal characteristics and behaviour of stockpeople and subsequent behaviour and production of dairy cows Appl Anim Behav Sci 2002, 79 (3), 195 219 Hocking, P.M.; Maxwell, M.H.; Robertson, G.W.; Mitchell, M.A Welfare assessment of broiler breeders that are food restricted after peak rate of lay British Poultry Science 2002, 43 (1), 15 El Lethey, H.; Aerni, V.; Jungi, T.W.; Wechsler, B Stress and feather pecking in laying hens in relation to housing conditions British Poultry Science 2000, 41 (1), 22 28 Manninen, E.; de Passille, A.M.; Rushen, J.; Norring, M.; ´ Saloniemi, H Preferences of dairy cows kept in unheated buildings for different kinds of flooring Appl Anim Behav Sci 2002, 75 (4), 281 292 Warburton; Mason, G.J Is out of sight out of mind? The effects of resources cues on motivation in mink Anim Behav 2003, 65 (4), 755 762 Nimon, A.J.; Broom, D.M The welfare of farmed mink (Mustela vison) in relation to housing and management: A review Animal Welfare 1999, (3), 205 228 Vinke, C.M Some comments on the review of nimon and broom on the welfare of farmed mink Animal Welfare 2001, 10 (3), 315 324 Mason, G.J.; Mendel, M Do Stereotypies of pigs, chickens, and mink reflect adaptive species differentiation in control of foraging? Appl Anim Behav Sci 1997, 53 (1/2), 45 58 Broom, D.M.; Nimon, A.J Response to Vinke’s short communication: Comments on mink needs and welfare indicators Animal Welfare 2001, 10 (3), 325 326 Garner, J.P.; Mason, G.J.; Smith, R Stereotypic route tracing in experimentally caged songbirds correlates with general behavioural disinhibition Anim Behav 2003, 66 (4), 711 727 Well-Being Assessment: Concepts and Definitions John J McGlone Texas Tech University, Lubbock, Texas, U.S.A INTRODUCTION Animal welfare and animal well-being are more or less interchangeable terms Assessment of animal welfare seems to include some subjective assessments, while the term animal well-being is viewed as more objective in some circles In practice, the two terms have very similar meaning to the public and most scientists Animal welfare/well-being assessment is often criticized by scientists as being anthropomorphic Anthropomorphism is the ascribing of human traits to nonhumans (e.g., animals or inanimate objects) Most scientists have historically not been comfortable with assessing animal happiness or pleasure Still, there is a need to objectively measure and assess animal well-being From this need, the science of farm animal welfare was born Animal cognitive experiences, including their feelings, are included in this science along with measures of physiological status (endocrine and immune status), behavior, growth, and reproduction HISTORICAL PERSPECTIVE Philosophers have examined the relationship between humans and animals from moral and theological views for centuries The modern concept of farm animal wellbeing began with the issuing of the Brambell report in 1965 in the United Kingdom The group of biologists, led by Brambell, concluded that animals have ‘‘Five Freedoms.’’ These freedoms (some would call them ‘‘rights’’ today) include the freedom to get up, lie down, stretch their limbs, turn around, and groom (themselves or others, depending on the species) The assignment of the original ‘‘Five Freedoms’’ is considered more of a moral argument than a scientific argument there was no science to support these basic freedoms in 1965 ANIMAL RIGHTS VS ANIMAL WELFARE/WELL-BEING The public and the media often confuse animal rights and animal welfare/well-being Animals have limited legal Encyclopedia of Animal Science DOI: 10.1081/E EAS 120019846 Copyright D 2005 by Marcel Dekker, Inc All rights reserved rights and few widely agreed-upon moral rights Animals have a legal right to not be abused or neglected Other than that right, animals not have the right to life or liberty Some activist groups attribute rights to animals to the extent that they believe animals should not be eaten, exhibited, or used in research Animal welfare/well-being is the concern of all people who own animals People give animals adequate environments to ensure that they have good welfare/well-being The subject of animal welfare/well-being science is a recognized area of investigation Those who hope to improve the lives of animals will so through careful examination of animal welfare/well-being DEFINING AND ASSESSING ANIMAL WELFARE/WELL-BEING Scientists working in the field of farm animal welfare science have struggled with defining and assessing animal welfare/well-being The most widely-held view is that to properly assess farm animal welfare, a multidisciplinary approach is required Measures should include behavior, physiology, growth, and reproduction All these measures are responsive to stress to varying degrees A sample of other views are provided here Duncan[1] suggested that animal welfare has to with how animals feel their cognitive experiences Moberg[2] suggested that when animals experience stress, their welfare is compromised when they reach a prepathological state as measured by animal physiology and disease state (including infectious and metabolic diseases) In another view, because behavior is adaptive, simply finding a behavioral effect cannot be said to be a negative welfare situation Only when the environment is stressful to the point that physiological changes are invoked can the animal be said to be in a state of reduced welfare, McGlone[3] argued In another model, animal welfare has to with behavioral needs, and when behavioral needs are met, welfare is adequate.[4] The most recent model, proposed by Curtis,[5] includes an assessment of the animals’ state of being its state relative to a continuum from a bad to a good state of being (Fig 1) Many models of animal welfare/well-being overlap 883 884 Well-Being Assessment: Concepts and Definitions Fig The continuum of states of animal welfare/well being THE MULTIDISCIPLINARY APPROACH In the multidisciplinary approach, one measures behavior, physiology, and performance and then uses all of this information to determine whether welfare/well-being is adequate This approach is the safest approach in that several of the other models can be examined if all of these measures are collected This approach was used recently to assess sow welfare in various housing systems using a meta-analysis of selected scientific publications.[6] Measures of performance include, for growing animals, rates of growth and efficiency of nutrient utilization Table Definitions in the field of animal welfare/well being science Name Agonistic Fixed action pattern Rights (12 definitions were given in this source) Rights (animal) Stereotyped Stereotyped behaviorb Stereotypy Welfare Well being a Definition Aggressive, submissive, and threat behaviors Any action pattern typical of a given species or breed that is performed in a very similar way by its individual members In contemporary ethology, the term ‘‘fixed action pattern’’ often is replaced by ‘‘modal action pattern’’ because of inevitable individual variations in behavior Examples: face grooming in mice, egg retrieval in geese Qualities (as adherence to duty or obedience to lawful authority) that together constitute the ideal of moral propriety or merit moral approval; something to which one has a just claim, such as the power or privilege to which one is justly entitled The idea that animals have a just or moral claim or privilege to certain items such as lack of abuse or neglect, life, or freedom Repeated behaviors shown in sequence that vary only slightly in sequence; may be caused by the environment genetics, or a combination Examples: chewing, suckling Behavior repeated in a very constant way The term generally is used to refer to behavior that develops as a consequence of a problem situation such as extended social isolation, low level of environmental complexity, deprivation, etc Stereotypy also may arise from genetic predispositions, or from disease of, or damage to, the brain Stereotyped behavior that serves no apparent function; often associated with disease or adaptation to a stressful environment Example: navel sucking in weaned piglets The state of being of an animal Welfare can range from very good to very bad A term used in the scientific literature to indicate animal welfare Sourcea Hurnik et al.[7] Merriam Webster[8] Hurnik et al.[7] A source is given when the definition is widely accepted This definition has been functionally divided into ‘‘normal’’ stereotyped behavior and stereotypies among farm animal welfare scientists b Well-Being Assessment: Concepts and Definitions Among adult animals, rates of reproduction are included in animal performance measures Growth and reproduction are suppressed when animals are stressed Measures of behavior include maintenance behaviors (feeding, drinking, standing, moving, laying, and sleeping), social behaviors (agonistic and nonagonistic behaviors), goal-directed behaviors (exploration, food-searching, water-searching), preferences, emotional behaviors (fear, frustration, rage, etc.) and abnormal behaviors Among abnormal behaviors are aberrant behaviors including tail biting, ear chewing, navel sucking, buller-steer mounting, wind sucking, and cribbing in horses, woolpicking in sheep, and a host of others In a gray area of science, certain behaviors are considered abnormal by some authors but other authors simply conclude they have unknown cause Included in this gray area are stereotyped behaviors that develop into stereotypies (Table 1) Examples of behaviors that clearly are stereotyped but may become stereotypies are bar biting in sows, tongue rolling in calves, and pacing among captive wild animals Measures of physiology include both endocrine and immune measures Endocrine measures used in assessment of animal welfare include adrenal cortical and medullary hormones Glucocorticoids (cortisol or corticosterone) and catecholamines are the most commonly measured endocrine measures of stress Measures of immune status are measures of stress in that if the immune system is suppressed and a pathogenic microorganism (or even a normally nonpathogenic microorganism) is present in sufficient quantity, then the animal will become ill Illness is clearly a state of reduced welfare/well-being Stress suppresses the immune system and so an important measure of the animal’s welfare/well-being would be its relative immune status Examples of measures of immunity that are sensitive to stress include natural killer cell activity, neutrophil function (chemotaxis and phagocytosis), and levels of some cytokines Other measures of immunity such as antibody response to a foreign antigen and lymphocyte proliferation in the presence of mitogen have been used in welfare/well-being assessment; however, these measures require very stressful environments to induce changes Two examples of use of the multidisciplinary approach to assessment of animal welfare are given below Hicks et al.[9] examined the effects of heat stress, shipping stress, and social stress on pig behavior, immunity, and endocrine and performance measures Pig behavior was significantly changed by all acute, mild stressors Pig physiology was only slightly changed Pig social stratus (dominant, intermediate, or submissive) interacted with stress treatments Dominant pigs were heavier and less negatively influenced by stressors than were subordinate pigs The authors concluded that behavioral changes were more consistent and reliable 885 measures of the effects of acute stress Stockpeople could use the behavioral responses as early indicators of reduced welfare and as a sign that interventions are required to maintain adequate animal welfare Mitlohner et al.[10] examined the effects of shade on cattle performance, carcass traits, physiology, and behavior while they were experiencing heat stress The provisions of shade increased weight gain of cattle that were in a warm climate Shade also reduced neutrophil numbers and respiratory rates and caused altered cattle behavior Because shade increased cattle weight gain and improved some measures of physiology, one could conclude that the cattle with shade in the summertime had improved welfare/well-being CONCLUSIONS Animal welfare/well-being can be examined as a science; as a legal, moral, or ethical argument; or as a subject for activism Farm animals have the right to not be abused or neglected, but beyond that they have few agreed-upon rights Livestock producers provide environments that are conducive to good animal welfare Several animal welfare models are presented Measuring animal welfare by using a multidisciplinary approach would provide information on animal behavior, physiology, and performance so that decisions about animal welfare/well-being can be made with the most possible information[11] and if possible in context with other society issues.[12] REFERENCES Duncan, I.J.H Animal welfare defined in terms of feelings Acta agric Scand., A Anim Sci 1996, 27, 29 35 Moberg, G.P Suffering from stress: An approach for evaluating the welfare of an animal Acta Agric Scand., A Anim Sci 1996, 27, 46 49 McGlone, J.J What is animal welfare? J Agric Ethics 1993, 6, 26 36 Duncan, I.J.H Behavior and behavioral needs Poultry Sci 1998, 77, 1766 1772 Curtis, S.E Stress: State of being Encycl Anim Sci 2004 (in press) McGlone, J.J.; von Borell, E.H.; Deen, J.; Johnson, A.K.; Levis, D.G.; Meunier Salaun, M.; Morrow, J.; Reeves, D.; ă Salak Johnson, J.L.; Sundberg, P.L Review: Compilation of the scientific literature comparing housing systems for gestating sows and gilts using measures of physiology, behavior, performance, and health Prof Anim Sci 2004, 20, 105 119 Hurnik, J.F.; Webster, A.B.; Siegel, P.B Dictionary of 886 Farm Animal Behavior, 2nd Ed.; Iowa State University Press: Ames, 1995 Merriam Webster Merriam Webster Online Dictionary; 2004 http://www.m w.com/netdict.htm Accessed March 28, 2004 Hicks, T.A.; McGlone, J.J.; Whisnant, C.S.; Kattesh, H.G.; Norman, R.L Behavioral, endocrine, immune, and perfor mance measures for pigs exposed to acute stress J Anim Sci 1998, 76, 474 483 10 Mitlohner, F.M.; Galyean, M.L.; McGlone, J.J Shade ¨ Well-Being Assessment: Concepts and Definitions 11 12 effects on performance, carcass traits, physiology, and behavior of heat stressed feedlot heifers J Anim Sci 2002, 80, 2043 2050 Brambell, F.W.R Report of the Technical Committee to Enquire into the Welfare of Animals Kept Under Intensive Livestock Husbandry Systems; Command Paper, Her Majesty’s Stationery Office: London, 1965; Vol 2836 McGlone, J.J Farm animal welfare in the context of other society issues: Toward sustainable systems Livest Prod Sci 2001, 72, 75 81 Well-Being Assessment: Physiological Criteria Katherine Albro Houpt Cornell University, Ithaca, New York, U.S.A INTRODUCTION There is no single valid measure of stress (or well-being) Nevertheless, we can use physiological variables to assist in validation The hormone most often used for measuring well-being is cortisol, the product of the mammalian adrenal cortex One also can measure the levels of hormones and metabolites that are affected by cortisol The sympathetic nervous system is the other major source of reactions to stress, pain, or fright SYMPATHETIC NERVOUS SYSTEM There are two components to the sympathetic nervous system neural and hormonal The most rapid response is neural Centers in the diencephalon (the hypothalamus, primarily) are stimulated by the frightening event, and therefore, the sympathetic pathways in the spinal cord and then the nerves of the sympathetic chain are stimulated The neurotransmitters released by the sympathetic nerves are norepinephrine and epinephrine (adrenaline and noradrenaline are alternative names) The structures innervated by the sympathetic nerves are the blood vessels, the hair follicles, the heart and lungs, and the gastrointestinal tract The action on the gastrointestinal tract is primarily negative: Secretion and motility are inhibited The actions on the heart are to increase the frequency and strength of contraction and to dilate the bronchioles of the lungs The pupils of the eyes dilate The hair stands on end (piloerection) Any of these reactions can be measured to assess welfare The hormones norepinephrine and epinephrine are very quickly degraded, so blood samples need to be taken quickly and the blood kept cold and processed quickly The hormones can also be measured in saliva, which is a less invasive method, but still involves restraint of the animal For this reason, it is more practical and probably more valid to measure the results of sympathetic stimulation, for example, heart rate There are heart rate monitors that can be attached to the animal with a chest band These can be retrieved later to determine any change in heart rate or in variability of heart rate Encyclopedia of Animal Science DOI: 10.1081/E EAS 120019845 Copyright D 2005 by Marcel Dekker, Inc All rights reserved There can be confounding factors in any measure of stress For example, ceiling effects can make interpretation difficult A ceiling effect occurs when the response is already high and cannot be any higher physiologically Branding is used for identification of beef cattle in the United States Although the modern techniques of microchipping would also make identification possible, what the rancher needs is a symbol, unique to his ranch, that is visible from a distance There are two methods of branding: hot-iron branding and freeze branding Hotiron branding destroys the hair follicles and creates a scar Freeze branding does not destroy the hair follicles, but causes the hair to regrow white rather than pigmented These brands are somewhat harder to read than hot-iron brands, but presumably are more humane When the responses of beef cattle to the two types of branding were compared, the heart rate and catecholamine levels were high following both procedures The explanation is that the restraint necessary to brand the animals was extremely stressful to all the cattle, so their response was maximal In other words, there was a ceiling effect When the comparison of branding methods was repeated using dairy cattle, hot-iron branding caused higher heart rate and more avoidance than freeze branding.[1] Dairy cattle are much more accustomed to the presence of humans, to restraint, and to being handled than are most beef cattle HYPOTHALAMIC-PITUITARY ADRENAL AXIS Stress to the animal leads to stimulation of those hypothalamic neurons that produce corticotropin releasing factor (CRF).[2] This is carried in the hypothalamic pituitary portal system to the anterior pituitary, where it stimulates release of adrenal corticotropic hormone (ACTH) This, in turn, stimulates release of the adrenal cortical hormones, in particular cortisol (in mammals) and corticosterone (in birds) The mineral corticoids aldosterone may also be released to a lesser degree This hormonal cascade will take some time (minutes to hours), in contrast to the more rapid neural activity of the sympathetic nervous system One important question is how much does an animal’s cortisol level have to rise 887 888 before we should consider the animal stressed Barnett and Hemsworth[3] have suggested that a 40% increase indicates stress One could use any increase above the normal range for the particular laboratory and species There are pitfalls in the use of cortisol (or any other physiological measurement), not because cortisol is not an indicator of stress, but because of confounding circumstances For example, veal calf welfare is frequently questioned, so measuring cortisol was assumed to be a valid measure As expected, when the calves were first placed in veal crates their cortisol was elevated, but several weeks later their cortisol was lower than agematched calves that were housed in pens.[4] The controls had higher cortisol than the confined calves, probably because they had to be chased and caught before the blood samples were taken The method of obtaining the sample is important If blood is taken by direct venipuncture and several attempts have to be made before the vein is punctured, the cortisol may be high for that reason Taking blood from the anterior vena cava of a supine pig is much more likely to be stressful than taking it from the jugular vein of a horse habituated to handling and injections Preplacement of an indwelling vascular catheter avoids some of those problems There is a definite circadian rhythm of cortisol secretion, so that morning cannot be used as a control for afternoon In fact, loss of the rhythmicity is another sign of stress Twenty minutes should be allowed after the stressor for cortisol to rise Cortisol can be measured in other body fluids Salivary cortisol can be collected easily by putting a cotton-tipped applicator in the animal’s mouth Urinary cortisol can be measured, but creatinine must be measured also in order to control for concentration of the urine A low cortisol concentration in dilute urine could represent a higher plasma level than a higher level of cortisol in concentrated urine Fecal cortisol has been measured successfully and is particularly useful when the well-being of free-ranging or wild species is to be evaluated One advantage of measuring fecal cortisol is that cortisol production over a matter of hours is represented, rather than cortisol at a single point in time, as with a blood sample The actions of cortisol on the rest of the body can also be measured and used to evaluate welfare Cortisol has effects on the liver, the fat depots, and the immune system The hormone stimulates gluconeogenesis Gluconeogenesis is the deamination of amino acids, freeing glucose for immediate energy The ammonia produced forms urea, and urea can be measured as a sign of stress In this case cortisol stimulation more urea is produced, but levels may be high because less is excreted Impaired excretion would indicate a kidney problem Therefore, when a high level of urea is detected, renal health should be evaluated before stress is diagnosed Renal function can be Well-Being Assessment: Physiological Criteria measured from the specific gravity of the urine, from the presence or absence of protein in the urine, and by the ratio of urea to creatinine, a compound that rarely varies in plasma concentration Under the influence of cortisol, fatty acids are metabolized rather than forming more adipose tissue These two actions, gluconeogenesis and antilipogenesis, complement the actions of the adrenal medullary hormones that stimulate glycogenolysis and lipolysis One of the major actions of cortisol is the reduction of inflammation, and inflammation is reduced by suppression of the immune system The number and type of white blood cells can be measured There are several types of white blood cells, including neutrophils and lymphocytes The lymphocytes are the antibody-producing cells, and these are the cells suppressed by cortisol The ratio of neutrophils to lymphocytes can therefore be used as a measure of stress The fewer the lymphocytes, the more likely the animal is secreting more cortisol and is stressed One can also measure the activity of white blood cells rather than simply the number of cells Some of these measures are mitogen-induced lymphocytic proliferation and natural killer-cell cytotoxicity These have been used to assess well-being, but the results are often inconsistent.[5] Suppression of the immune system is the most dangerous effect of cortisol Although the swelling and pain of inflammation will be decreased, the white blood cells that cause these signs will not be protecting the body from invasion by bacteria or viruses Antibodies will not form complexes with foreign antigens, and bacteria will not be destroyed by phagocytosis The result of suppression of the immune response is illness The respiratory or gastrointestinal pathology (shipping fever) seen in newly mixed or transported animals is a result of stressinduced immunosuppression The adrenal glands are not the only ones stimulated by stress Thyroid-stimulating hormone is released from the pituitary and stimulates release of thyroxine from the thyroid gland Thyroxine increases metabolic rate and, therefore, calorigenesis Carbohydrate stores will be utilized first, and then fat stores Cortisol is a useful measure of some kinds of stress, but not others For example, cortisol increases when horses are transferred from one environment to another and when they are trailered, but chronic deprivation of water or exercise does not cause cortisol to rise or the response of cortisol to ACTH to change Fortunately, there are other physiological values that can be used Examples include plasma protein, which can be used to assess the effects of furosemide Furosemide is a drug frequently administered to race horses, ostensibly to prevent exercise-induced pulmonary hemorrhage However, it also improves the animal’s performance, because Well-Being Assessment: Physiological Criteria the horse is 10 20 kilograms lighter in weight as a consequence of diuresis If a horse is treated with furosemide, the loss of fluid from the circulation causes an increase in plasma protein If horses are given limited amounts of water, as in mares used for estrogen production, they have normal plasma protein but an elevated osmotic pressure.[6] The most recently used physiological measure of wellbeing is acute phase proteins These are haptoglobins, a glycoprotein of the alpha-2-globulin fraction by haptocytes in response to stress, ACTH, and cortisol They are elevated following castration of piglets and after transporting older pigs for more than hours CONCLUSION The animal whose well-being is compromised responds with a variety of physiological changes These can be used, in combination with behavioral measures, to help us determine the optimum housing, social grouping, and transport of farm animals 889 REFERENCES Lay, D.C., Jr.; Friend, T.H.; Bowers, C.L.; Grissom, K.K.; Jenkins, O.C A comparative physiological and behav ioral study of freeze and hot iron branding using dairy cows J Anim Sci 1992, 70, 1120 Dantzer, R.; Mormede, P Stress in Domestic Animals: A Psychoneuroendocrine Approach In Animal Stress; Moberg, G.P., Ed.; American Physiological Society: Bethesda, MD, 1985; 81 95 Barnett, J.L.; Hemsworth, P.H The validity of physiolog ical and behavioral measures of animal welfare Appl Anim Behav Sci 1990, 20, 177 187 Stull, C.; McDonough, P Multidisciplinary approach to evaluating welfare of veal calves in commercial facilities J Anim Sci 1994, 72, 2518 2524 McGlone, J.J.; Salak, J.L.; Lumpkin, E.A.; Nicholson, R.I.; Gibson, M.; Normal, R.L Shipping stress and social status effects on pig performance, plasma cortisol, natural killer cell activity, and leukocyte numbers J Anim Sci 1993, 71 (4), 888 Houpt, K.A.; Houpt, T.R.; Johnson, J.L.; Erb, H.N.; Yeon, S.C The effect of exercise deprivation on the behaviour and physiology of straight stall confined pregnant mares Anim Welf 2001, 10, 257 267 Wool: Biology and Production A C Schlink N R Adams CSIRO Livestock Industries, Wembley, Western Australia INTRODUCTION Wool is a generic description of hair from various breeds of domesticated sheep (Ovis aries) Wool appears to be the earliest material man used to spin and weave into clothing, with evidence of shears for harvesting wool being used around 1000 B.C Requirement for shearing implies development of sheep with a continuously growing fleece These developments associated with domestication have continued until this day, although wool is no longer a dominant textile fiber BIOLOGY The gross morphology of a wool fiber is shown in Fig 1.[1] The fiber is surrounded by cuticle cells that overlap in only one direction, leading to directional frictional characteristics and wool felting The cuticle has four layers with a combined thickness of 0.5 to 0.8 mm, occupying between and 16% of total fiber weight The cortex, composing 90% of fiber weight, consists of two cell types, ortho- (60 to 90%) and paracortex cells (10 to 40%), the latter containing higher quantities of sulphur than the former, resulting in a tougher cell with more cross-linkage Cortex cell-type arrangement changes with increasing fiber diameter In fine-wool Merinos, the cortical cells are arranged in a bilateral manner, and the border between cell types is arranged in helical pattern along the fiber axis This helical pattern results in fiber crimp, with paracortex being situated in the inner part and orthocortex in the outer part of the crimp Cortex cells have a spindlelike shape, being 45 to 95 mm long and to mm wide Ortho-cortex cells rarely contain nuclear remnants and cytoplasmic residues At its widest point, each cortical cell contains to 20 clearly separated macrofibrils embedded in intermacrofibrillar matrix material, in a hexagonal array Macrofibrils are composed of bundles of 500 to 800 microfibrils Microfibrils, or intermediate filaments, are composed of alpha-helical proteins of comparatively low cystine 890 content that are linked by both disulphide and hydrogen bonds A matrix of intermediate filament-associated proteins surrounds microfibrils and is composed of two families of nonhelical proteins, one being cystine-rich and the other, glycine- and tyrosine-rich Wool is almost entirely composed of a family of proteins known as alpha-keratins Merino wool has higher cystine content than coarse wools as a result of having a larger proportion of high-sulphur alpha-keratin proteins Amino acid composition can vary between sheep, with the growth phase of the wool follicle cycle and with the nutritional status of the sheep Wool fiber diameters range 10 to 80 mm and have a density of 1.304 g/cm3, with slightly and imperfectly elliptical cross-sections Wools with higher fiber diameter tend to be hairlike and medulated Proteins in wool have the ability to adsorb water At standard atmosphere of 65% relative humidity and 20°C, water regain ranges from 14 to 18%.[2] Wool fibers are highly elastic, and if not strained by more than 30% of length for longer than one hour, they can return to their original state by soaking in water The intrinsic strength of wool is low, varying between 50 and 300 megapascals Sheep wool follicles have a very long anagen phase, with 2% of follicles inactive at any one time The general morphology of anagen follicles is shown in Fig 2.[3] A connective tissue sheath surrounds the tubular down growth of epithelium, and there is a dermal papilla responsible for cell division Blood vessels are found in the connective tissue sheath and, except in the smaller secondary follicles of Merino sheep, the dermal papilla Primary and secondary follicles are distinguished by their appendages and time of initiation in fetal skin Primary follicles form first, at about 60 days postconception, and secondary follicles start 14 to 20 days later Variable numbers of secondary follicles may form either as separate follicles or as outgrowths of other secondary follicles Sweat glands and arrector pili muscles are appendages of primary follicles Both follicle types have sebaceous glands The ratio of primary to secondary follicles varies between sheep Merino sheep with 19 mm Encyclopedia of Animal Science DOI: 10.1081/E EAS 120023830 Published 2005 by Marcel Dekker, Inc All rights reserved Wool: Biology and Production 891 Fig Gross morphology of a Merino wool fiber (From CSIRO Livestock Industries.) (View this art in color at www.dekker.com.) Fig Diagram of skin and wool follicle groups showing primary follicles, with arrector pili muscles and sweat glands, and secondary follicles (From CSIRO Livestock Industries.) 892 Wool: Biology and Production Table Principal greasy wool producing countries and world production from 1988 to 2002 (million kg) Year Australia New Zealand China Eastern Europe World 1988 1989 1990 1991 1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 916 959 1102 1066 875 869 829 731 725 700 684 678 652 600 565 346 339 311 305 296 256 284 289 275 266 252 253 237 246 253 209 222 237 239 240 238 240 260 298 255 277 283 291 294 293 613 627 623 592 526 456 439 409 263 229 198 188 187 188 189 2,919 2,967 2,970 2,953 2,989 2,913 2,817 2,689 2,541 2,418 2,379 2,348 2,303 2,239 2,217 (From the International Wool Textile Organization.) wool have 80 follicles/mm2 and a secondary to primary follicle ratio of 20:1, whereas Drysdale sheep (43 mm) have 13 follicles/mm2 and a secondary to primary follicle ratio of 5:1 PRODUCTION Wool production normally occurs in areas where the pasture quality is adequate, but insufficient for meat production Seasonal variation in clean wool growth is similar to the variation in availability of digestible dry matter per hectare, or crude protein available per hectare.[4] The amplitude in seasonal wool growth rate ranged from 20.3% in Armidale, New South Wales, to 650% in Huang Cheng, China Peak wool growth occurred in the spring for cold, wet winter areas and in the summer in summer rainfall, tropical areas These changes in wool growth are also seen in follicle bulb diameter, dermal papilla length, skin weight, and the incidence of active follicles in the skin.[5] Wool growth also varies between years, reflecting annual pasture production cycles, with amplitude in fleece weight between years varying from 15.8% to 118.5% Only 20% of the protein synthesized in skin is excreted as wool, with the remaining 80% being degraded in the skin or desquamated as epithelial cells The value of 20% is robust for a wide range of sheep breeds and feeding levels.[6] Thus, genetic selection to increase fleece weight also increases the rate of skin protein synthesis Fleece weight is affected by age, pregnancy, lactation, and sex, with rams producing more wool than wethers or ewes.[7] Fiber diameter increases by 3.9 mm from 18 months to years of age in rams, but for ewes the increase is 0.4 mm over the same period Pregnancy and lactation reduce fleece weight by 30 to 600 grams and fiber diameter by 0.4 to 1.5 mm, and also affect staple strength.[8] Strong seasonal patterns in wool growth occur in many breeds, with annual rhythms synchronized by photoperiod acting through melatonin secreted by the pineal gland Breeds such as the Merino have a reduced response to photoperiod Fiber diameter is the first determinant of wool price, and many wool breeding programs aim to reduce fiber diameter Fiber diameter is changed by nutrient availability between and within years, but is mainly determined by the strain of sheep Seasonal variation in wool growth reduces staple strength, which is the second most important determinant of wool price Staple strength has a strong genetic component, depending on variation in fiber diameter, fiber shedding, and the strength of individual fibers For breeding ewes, the most critical time remains the last two weeks of pregnancy.[9] World greasy wool production peaked in 1990, with a total production of 2970 million kilograms, and declined to 2217 million kilograms in 2002 (Table 1) On a clean wool basis, Australia and New Zealand are the world’s two largest wool producers, although China produces more greasy wool than New Zealand Australia is the dominant producer of Merino sheep in the 18- to 23-mm range for apparel production New Zealand wool production is dominated by Romney sheep in the 30- to 38-mm range, which is suitable for carpets China has increased wool production from 7.2% of world greasy wool in 1988 to 13.2% in 2002 World production statistics are provided on a greasy wool basis, and greasy wool contains from 30 to 70% impurities Wool impurities are wax, suint, dust, and vegetable matter Sheep coats successfully reduce these contaminants Low wool yields in countries such as China are a consequence of overnight corralling of sheep for feeding during cold winters and protection from the elements CONCLUSION The average fine-wool sheep produces some 6000 kilometers of a complex protein fiber each year This fiber is produced from wool follicles that use 20% of the protein turnover in skin Wool is predominantly produced from Wool: Biology and Production grazing lands and is highly seasonal in growth Australia is the largest producer, with 25.5% of the world production of 2217 million kilograms in 2002 REFERENCES Hocker, H Fibre Morphology In Wool: Science and Technology; Simpson, W.S., Crawshaw, G.H., Eds.; CRC Press: Cambridge, England, 2002; 60 79 Heale, J.W.S Physical Properties of Wool In Wool: Science and Technology; Simpson, W.S., Crawshaw, G.H., Eds.; CRC Press: Cambridge, England, 2002; 80 129 Orwin, D.F.G Variation in Wool Follicle Morphology In The Biology of Wool and Hair; Rogers, G.E., Reis, P.J., Ward, K.A., Marshall, R.C., Eds.; Chapman and Hall: New York, 1989; 227 241 Schlink, A.C.; Mata, G.; Lea, J.M.; Ritchie, A.J.M Seasonal variation in fibre diameter and length in wool of grazing 893 Merino sheep with low or high staple strength Aust J Exp Agric 1999, 39, 507 517 Schlink, A.C.; Sanders, M.; Hollis, D.E Seasonal variations in skin and wool follicle morphology of grazing Merino sheep with low or high staple strength Asian Australas J Anim Sci 2000, 13 (Suppl A), 253 256 Adams, N.R.; Liu, S.; Masters, D.G Regulation of Protein Synthesis for Wool Growth In Ruminant Physiology: Digestion, Metabolism, Growth and Reproduction; Cronje, P.B., Ed.; CAB International, 2000; 255 272 Corbett, J.L Variation in Wool Growth with Physiolog ical State In Physiology and Environmental Limitations to Wool Growth; Black, J.L., Reis, P.J., Eds.; The Uni versity of New England Publishing Unit: Australia, 1979; 79 98 Hynd, P.I.; Masters, D.G Nutrition and Wool Growth In Sheep Nutrition; Freer, M., Dove, H., Eds.; CAB Interna tional, 2002; 165 187 Robertson, S.M.; Robards, G.E.; Wofle, E.C The timing of nutritional restriction during reproduction influences staple strength Aust J Agric Res 2000, 51, 125 132  ... of animal welfare/well-being DEFINING AND ASSESSING ANIMAL WELFARE/WELL-BEING Scientists working in the field of farm animal welfare science have struggled with defining and assessing animal welfare/well-being... good welfare/well-being The subject of animal welfare/well-being science is a recognized area of investigation Those who hope to improve the lives of animals will so through careful examination of. .. RIGHTS VS ANIMAL WELFARE/WELL-BEING The public and the media often confuse animal rights and animal welfare/well-being Animals have limited legal Encyclopedia of Animal Science DOI: 10.1081/E EAS