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Ratites: Biology, Housing, and Management Dominique Blache Graeme B Martin Irek Malecki The University of Western Australia, Crawley, Australia INTRODUCTION The large flightless ostrich, emu, cassowary, and rhea, and the small flightless kiwi, compose the ratite family The emu, ostrich, and rhea have been used in farming systems in which their biology influences management and housing Ratite farming is in its infancy and requires further research and development to overcome inherent constraints before each species can reach its full productive potential BIOLOGY OF RATITES The flat, raftlike (ratis) sternum provided the name for the family There is no keel and the pectoral muscles are absent or vestigial In all except rheas, the body feathers lack barbicels, so the plumage is loose and fluffy The feathers of the emu have two shafts The rhea and the ostrich have longer wings than the emu and they use them during elaborate displays (Fig 1) Female emus and rheas are larger than males, but the male ostrich is the largest.[1] All extant ratites are endemic to the Southern Hemisphere, whereas their ancestors were found in both hemispheres.[1] The ostrich, emu, and rhea are found in temperate and Mediterranean regions, but can survive in a wide range of climates.[1,2] The ratites have very strong legs and their muscles have a specific distribution and physiology due to the mechanic constraints of bipedal locomotion Ratites walk most of the day and can run at considerable speed (Table 1) They are nomadic and follow food availability, but are territorial during the breeding season Ratites can also crouch, a posture between standing and sitting (Fig 1) The females lay eggs in this position.[1,2] Vigilant ratites stand with the neck stretched upward; their heads are very mobile and can turn almost 360 degrees Vision is believed to be very efficient because of the elevated position of the eyes and also because of acuity Thus, they are able to see and detect from long distances (few kilometers) Ratites are not active at night and spend most of the dark phase lying Encyclopedia of Animal Science DOI: 10.1081/E EAS 120019782 Copyright D 2005 by Marcel Dekker, Inc All rights reserved Rheas, emus, and ostriches can be found in groups of 30 or more, but also in smaller family groups Ratites are very defensive when eggs or young chicks are present Agonistic behaviors include vocalizations, body postures, and eventual charging The reproductive biology of ratites presents some unique features The males have a large penile organ that erects from the cloaca and penetrates the female’s cloaca during copulation In females, sperm storage tubules in the reproductive tract allow the female to remain fertile for several days after copulation.[3] The mating system varies between species In the wild, male ostriches and rheas form harems, but also copulate with females from other groups, whereas emus form pairs that are stable during the mating period.[1,4] Courtship is based on vocalizations and displays or postures from both sexes (Fig 1) Ostriches reproduce during summer, but rheas and emus reproduce mainly during winter Photoperiod is essential for the emu,[5] but is not that critical for the other ratites because, when nutrition is not limited, they breed at anytime.[1] Ratites nest on the ground in very simple nests (Fig 1) Females lay large eggs at day intervals The total number of eggs laid by one female varies between species and individuals (Table 1) The number of eggs laid over a season seems to be strongly influenced by the level of fat reserves and nutrition of the female before the start of the laying period With the exception of the ostrich, male ratites are solely responsible for incubating the clutch and raising the young (Fig 1) Female emus leave their partner during incubation and mate with other males Ratite chicks are precocious and the nest is usually abandoned within 48 h after hatching.[1] The digestive system is simple and in most respects similar to that of other plant-eating birds, but ratites also consume insects and small animals.[1,2,6] The esophagus is mobile and expandable and ratites swallow their food whole The crop is absent in all ratites, but the structure of the stomach varies among species.[6] Their appetite varies dramatically between the breeding and the nonbreeding seasons, leading to large variations in body weight, mostly due to variation in fat reserves 763 764 Ratites: Biology, Housing, and Management Fig Clockwise from the top left corner Male ostrich with his harem; one female is incubating Male ostrich displaying courtship Male emu displaying courtship (View this art in color at www.dekker.com.) HOUSING AND MANAGEMENT OF RATITES The allocation of space to farmed birds varies with their age and reproductive status (Table 2) Only young chicks need access to an indoor pen Feeding recommendations for ratites are not as precise as those for commercial poultry Nutrient requirements are based on restricted data for ostriches and emus,[2,6] and there are no published data for rheas (Table 2) Feedstuffs of plant origin are the main constituent of the diet and, because of the requirement of Table Ratite biology Species Emu a Number of subspecies Origin Farming Sexual dimorphism Size (m) Weight (kg) Number of digits per foot Running speed (km/h) Breeding season Cluch sizec (eggs) Egg production per female Duration of incubationd (days) Incubating sex a Ostrich Rhea Australia Yes Not obvious 1.5 1.8 45 67 45 Autumn winter (SD) 45 22 ± 56 Male Africa Yes Yes 1.8 2.5 80 140 60 Spring summer (LD)b 36 50 ± 20 42 Male and female Southeastern America Yes Not obvious 1.5 15 40 45 Winter (SD)e 56 16 30 30 44 Male Only present subspecies Breeding season in ostriches varies regionally in Africa and is influenced by rain and food availability c Number of eggs found in one nest d Incubation under natural conditions e SD short day breeder, LD long day breeder (Data from author’s observations and Ref 1.) b Ratites: Biology, Housing, and Management 765 these large birds, a large amount of protein has to be included Most farmed ratites are fed a pelleted diet of crushed grain and other nutrients, formulated according to age and reproductive status.[2,6,7] Usually, the diet of females is richer in energy and protein during the breeding season than during the nonbreeding season However, this strategy might not be best considering that, naturally, the birds decrease their intake during the breeding season and replenish their reserves during the nonbreeding season In emus, feed intake is controlled by photoperiod and increases dramatically, by at least 150% (up to kg/ day/bird), when the breeding season ends, allowing them to recover, within two weeks, most of the weight lost over the breeding season.[8] Breeding management differs between countries and farms, but the relative advantages of the different strategies have not been compared scientifically Ostriches are kept in pairs, trios (one male for two females), or colonies.[2] Emus can breed in pairs or in groups Rheas are bred in groups because of their need for harem formation Breeding birds are given more space because of the possibility of fighting Reproductive failure can be due to behavioural problems, such as lack of pair formation Eggs are usually collected and artificially incubated to avoid the assembly of a clutch because the males become sexually inactive as they incubate Eggs need to be cleaned and dried before being set into incubators Damaged, under- and oversized eggs should be discarded Storage of eggs before incubation simplifies hatchery management because it allows batch hatching Optimal conditions for artificial incubation are known (Table 2).[2,6,7] Candling of ratite eggs is possible using commercially available devices Recommendations for hatching conditions are not scientifically proven (Table 2), but the eggs are usually transferred to the hatcher a few days before hatch date because pipping starts 36 hours prior to hatching Sexual maturity is reached at 18 20 months for ostriches and emus and after 24 months for rheas (Table 2) Vent sexing can be successfully carried out within days after hatching with an accuracy of around 85% RATITE PRODUCTION Table Housing conditions and management of farmed emus and ostriches Species Space allocation (m2/bird)a Chicks (0 12 weeks) indoor pen Chicks (0 12 weeks) outdoor pen Chicks (3 months) Young (6 12 months) Yearling (12 24 months) Breeders (>24 months) Free range Breeding pairs Maintenance requirement Energy (kJ/kg0.75/day) Nitrogenb (mg/kg0.75/day) Incubation Dry bulb temperature (°C) Relative humidity (%) Air flow (m3/min/40 eggs) Air quality (% O2, % CO2) Rotation of the eggs Amplitude (degree) Frequency (h) Total egg weight lost (%) Hatching Dry bulb temperature (°C) Relative humidity (%) Emu Ostrich 0.15 0.20 0.30 20 20 40 60 100 0.25 0.30 0.50 10 40 50 250 100 330 625 1000 400 1200 500 2000 600 2500 284 320 440 320 35.3 40 0.71 21, 0.05 36.4 25 30 1.42 21, 0.05 180 15 90 15 35 50 34 25 35 Farming of ratites has great potential that has been exploited most often in the countries of origin of the species Slaughter age is around 12 16 months for emus, 10 12 months for ostriches, and 18 20 months for rheas Methods and regulations for slaughter are already in place in each country, but more development is needed to decrease the cost of slaughtering, especially plucking methods The products are all high quality: soft leather, meat with low-fat and high-iron content, and a fine oil that can be used as a cosmetic base for the administration of topical medicines and as an anti-inflammatory agent, a claim already supported by clinical trials for emu oil Ostrich feathers have been successfully marketed in the past as a fashion item but this market has virtually disappeared The meat market is still small and in need of more marketing for further expansion There is also a need for more scientific input into management and genetic selection to improve productivity and product quality Recent developments of sperm collection and preservation, and artificial insemination techniques specific to both ostrich and emus, provide the industry with major tools for modern methods of selection.[9] CONCLUSION Space allocations vary according to feeding methods a Based on regulations and practices used by the industry b Based on nitrogen requirement of poultry Ratites are scientifically interesting because of their unique biological characteristics Those same characteristics offer a unique opportunity to develop an alternative industry that might have less environmental impact than 766 traditional, imported animal industries These industries exist, but still need a large amount of research and development before they will be successful because ratites are not simply bigger versions of common poultry and cannot be treated as such.[10] REFERENCES Davies, S.J.J.F Ratites and Tinamous: Tinamidae, Rhe idae, Dromaiidae, Casuariidae, Apterygidae, Struthioni dae; Oxford University Press: Oxford, 2002 Deeming, D.C The Ostrich: Biology, Production and Health; CAB International: Wallingford, UK, 1999 Malecki, I.; Martin, G.B Fertile period and clutch size in the Emu (Dromaius novaehollandiae) Emu 2002, 102, 165 170 Blache, D.; Barrett, C.D.; Martin, G.B Social mating system and sexual behaviour in the emu, Dromaius novaehollandiae Emu 2000, 100, 161 168 Ratites: Biology, Housing, and Management Blache, D.; Talbot, R.T.; Blackberry, M.A.; Williams, K.M.; Martin, G.B.; Sharp, P.J Photoperiodic control of the secretion of luteinizing hormone, prolactin and testosterone in the male emu (Dromaius novaehollandiae), a bird that breeds on short days J Neuroendocrinol 2001, 13, 998 1006 Tully, T.N.; Shane, S.M Ratite: Management, Medicine and Surgery; Krieger: Malabar, 1996 Deeming, D.C Improving Our Understanding of Ratites in a Farming Environment; Ratite Conference: Manchester, UK, 1996 Blache, D.; Martin, G.B Day length affects feeding behaviour and food intake in adult male emus (Dromaius novaehollandiae) Br Poult Sci 1999, 40, 573 578 Malecki, I.A.; Martin, G.B.; Lindsay, D.R Semen production by the male emu (Dromaius novohollandiae) Methods for collection of semen Poult Sci 1996, 76, 615 621 10 Malecki, I.; Blache, D.; Martin, G Emu biology and farming Developing management strategies for a valu able resource Land Management October 2001, 20 21 Ratites: Nutrition Management James Sales University of Maryland, College Park, Maryland, U.S.A INTRODUCTION NUTRIENT REQUIREMENTS Ratites (order Struthioniformes) are flightless birds with a raftlike breastbone devoid of a keel, and can be classified into the families Struthionidae (ostriches), Dromiceiidae (emus), Rheidae (rheas), Casuariidae (cassowaries), and Apterygidae (kiwis) The inaccuracy of earlier extrapolation of nutrient requirement specifications for poultry to ostriches and emus soon became evident from various nutrition-related problems encountered by commercial ratite farmers.[6] Studies by Cilliers[7] and O’Malley[4] revealed significant information on the energy and amino acid requirements of these two species (Tables and 3) It is evident that different diets, each with different nutrient concentrations, have to be fed at different stages of the life cycle; for example, a starter diet up to three months of age, a grower diet till slaughter age, and a breeder diet for breeder birds DIGESTIVE PHYSIOLOGY Despite their similarities to other birds, ratites have developed unique characteristics, such as modifications in the gastrointestinal tract, in order to survive in their natural habitat.[1] Ratites not have teeth or a crop (the feed storage organ in other avian species) Ostriches, emus, and rheas could be considered monogastric herbivores, which means they are simple-stomached animals that have developed the ability to utilize forage Whereas fiber fermentation appears to take place in the large intestine (colon) of the ostrich, the distal ileum serves as a fermentation organ in the emu The most distinctive characteristic of the gastrointestinal tract of the rhea is the relatively large cecum (Table 1) RATITE DIETS Many different diets have been utilized in commercial ostrich production, varying from single ingredients such as alfalfa, to compound diets with several ingredients including vitamin/mineral mixtures, since the domestication of the ostrich as a farm animal around 1865 in South Africa.[2] The first book on ostrich feeds and feeding was already published in 1913 by Dowsley and Gardner.[3] Reliance on compound, commercial, manufactured diets, mostly in a pelleted form, has become the norm since the spread of ostrich farming to countries outside South Africa around 1990 and the recognition of emu farming as being technically feasible in Australia in 1987.[4] At the few pilot operations for the domestication of the rhea as a commercial farm animal in South America, a variety of compound pelleted diets, consisting mainly of alfalfa and corn meal, are fed.[5] Encyclopedia of Animal Science DOI: 10.1081/E EAS 120019784 Copyright D 2005 by Marcel Dekker, Inc All rights reserved Mineral Requirements Currently, dietary mineral, as vitamin, specifications for ratites are based on suggestions A major problem in ostrich feeding is that calcium is very often overfed, with the result of depressed uptake of zinc and manganese Although a total dietary calcium concentration of 2.0 to 2.5% is recommended for ostrich layers in intensive production systems, excellent laying and fertility results have been achieved with dietary calcium levels as low as 1.6% on a dry matter basis.[8] Under intensive farming conditions, leg problems seldom occur in young ostrich chicks fed a diet with calcium levels around 1.5 to 1.6%.[9] NUTRITION OF CHICKS Although the rearing of young ostriches is a wellestablished practice, high mortalities are often encountered.[9] Ostrich feed and water should be available from day one after hatch A chopped fresh alfalfa or grass topping on feed will stimulate chicks to start eating It was also found in rhea chicks[10] that the first few chicks required frequent stimulation, for example, by poking with a finger or pencil at the food, to induce proper feeding Many ostrich producers supplement the starter diet or water of the newly hatched ostrich with a booster pack containing: 1) electrolytes that will ensure that the correct 767 768 Ratites: Nutrition Management Table Comparison of the digestive tract of ostriches, emus, and rheas Length (cm) Region Relative length (% of total) Ostrich Rhea Ostrich Emu Rhea 512 94 800 Small intestine Cecum Colon Emu 51 28 140 48 40 36 57 90 61 21 17 (From Ref 1.) supplementation of any product, for example, yogurt, that might stimulate immunity is highly recommended ratio of sodium to potassium will be consumed and that the absorption of moisture will be normal during these early stages of life; 2) acidification substances that will lower the pH of the digestive tract and enhance its adaptation to high-protein starter diets; 3) amylase, protease, and cellulase enzymes to ensure more efficient digestion of starch, protein, and fiber; and 4) vitamins A, D, E, and B complex to ensure immunity against infections and other diseases.[8] It is well known that ostrich chicks have poor resistance against infectious and other diseases The CONCLUSION Ratites are unique in that they resemble the characteristics of avian species with nutritional adaptations similar to that of ruminants Despite studies on ratites that enable the modeling of energy and amino acid requirements, dietary Table Estimated dry matter intake (DMI),a energy (TMEn), and protein and amino acid requirements for maintenance and growth of African black ostriches AGE (Days) LW (kg) ADG (g/b/d) DMI (g/b/d) TMEn (MJ/kg DMI) Prot (g/kg DMI) Amino acids (g/kg DMI) Lys Meth Cys Arg Thr Val Isoleu Leu His Phe Tyr 30 60 90 120 150 180 210 240 270 300 330 360 390 420 450 480 510 540 570 600 4.0 11.0 19.5 28.5 39.5 52.1 63.4 73.3 82.4 91.0 96.3 99.9 103.5 107.0 110.0 112.3 114.2 116.0 118.6 120.3 105 233 283 300 367 420 375 330 305 287 177 120 120 117 100 77 63 60 87 57 220 440 680 820 1220 1490 1630 1710 1760 1800 2160 2210 2250 2250 2250 2250 2250 2250 2250 2250 15.2b 17.5b 15.3b 14.9b 12.5b 12.2b 11.3 10.8 10.7 10.8 8.0 7.4 7.4 7.5 7.5 7.3 7.3 7.3 7.7 7.5 239 272 224 207 174 168 148 135 130 128 85 74 74 75 73 69 67 67 74 68 10.6 12.5 10.8 10.6 9.1 9.0 8.5 8.2 8.3 8.4 6.3 5.9 5.9 6.1 6.1 6.0 6.0 6.0 6.4 6.2 3.1 3.6 3.2 3.2 2.7 2.7 2.6 2.5 2.6 2.6 2.0 1.9 1.9 2.0 2.0 1.9 1.9 2.0 2.1 2.0 2.8 3.3 2.8 2.7 2.3 2.3 2.1 2.0 2.0 2.0 1.5 1.3 1.4 1.4 1.4 1.3 1.3 1.3 1.4 1.4 9.8 11.5 10.1 9.9 8.5 8.5 8.0 7.8 7.9 8.1 6.1 5.7 5.8 5.9 5.9 5.9 5.9 5.9 6.2 6.1 6.5 7.6 6.6 6.4 5.5 5.5 5.1 5.0 5.0 5.1 3.8 3.5 3.6 3.7 3.7 3.6 3.6 3.6 3.8 3.7 7.9 9.3 8.2 8.1 7.0 6.9 6.6 6.4 6.5 6.7 5.1 4.8 4.8 4.9 5.0 4.9 4.9 5.0 5.2 5.1 8.7 10.3 9.0 8.8 7.6 7.6 7.2 7.0 7.1 7.2 5.4 5.1 5.2 5.3 5.3 5.2 5.2 5.3 5.6 5.4 14.5 17.0 14.7 14.3 12.3 12.2 11.4 11.0 11.1 11.2 8.4 7.8 7.9 8.1 8.0 7.9 7.9 8.0 8.4 8.2 3.6 4.3 3.8 3.8 3.3 3.3 3.1 3.1 3.1 3.2 2.4 2.3 2.3 2.4 2.4 2.4 2.4 2.4 2.5 2.5 8.5 10.0 8.7 8.5 7.3 7.2 6.8 6.6 6.6 6.7 5.0 4.7 4.7 4.8 4.8 4.8 4.7 4.8 5.1 4.9 4.4 5.1 4.5 4.5 3.9 3.9 3.7 3.6 3.7 3.8 2.9 2.7 2.7 2.8 2.8 2.8 2.8 2.8 3.0 2.9 LW live weight; ADG average daily gain; DMI dry matter intake; TMEn true metabolizable energy corrected for nitrogen retention; Prot protein; Lys lysine; Met methionine; Cys cystein; Arg arginine; Thr threonine; Val valine; Isoleu isoleucine; Leu leucine; His histidine; Phe phe nylalanine; Tyr tyrosine a Based on a diet with a TMEn (ostrich) content of 11.25 MJ/kg b In calculating TMEn requirements from results obtained for seven month old birds, similar energy contents were assumed for younger birds This assuption is incorrect, resulting in an overestimation of dietary energy requirements (From Ref 7.) Ratites: Nutrition Management 769 Table Estimated dry matter intake (DMI),a and protein and amino acid requirements for maintenance and growth of emus Amino acids (g/kg DMI) AGE (Weeks) LW (kg) ADG (g/b/d) DMI (g/b/d) Prot (g/kg DMI) Lys Met Met + Cys Thr Isoleu Leu 2 3 4 6 8 10 10 12 12 16 16 20 20 24 24 28 28 32 32 36 36 40 40 44 44 48 48 52 52 56 56 60 60 62 62 63 0.5 0.8 1.3 2.3 3.9 5.9 7.8 10.7 14.6 18.2 23.8 23.8 26.1 28.0 30.0 32.5 35.7 38.8 41.5 43.1 44.0 14 59 80 106 124 153 121 145 134 125 92 95 71 58 80 110 117 104 87 72 98 35 88 140 220 259 368 374 561 603 630 597 545 544 614 604 820 851 829 1,051 1,026 1,175 119 170 151 137 146 133 116 94 90 89 91 114 116 110 134 113 112 114 88 88 84 6.5 9.8 8.7 7.9 7.7 7.1 6.4 5.3 5.0 5.0 5.2 6.7 7.0 6.9 8.7 7.5 7.3 7.6 5.8 5.8 5.6 1.8 2.8 2.5 2.2 2.3 1.8 1.8 1.5 1.5 1.6 1.6 2.1 2.1 2.1 2.5 2.2 2.2 2.2 1.7 1.6 1.6 5.9 6.7 5.7 4.8 5.0 4.1 4.1 3.4 3.6 3.8 4.1 5.1 5.3 5.1 6.0 5.0 5.0 5.1 3.9 3.9 3.6 5.8 7.9 6.9 6.2 6.7 6.1 5.4 4.4 4.2 4.2 4.4 5.5 5.7 5.4 6.6 5.5 5.4 5.6 4.3 4.3 4.1 5.0 6.7 5.9 5.4 5.8 5.3 4.7 3.8 3.7 3.7 3.8 4.8 4.9 4.7 5.6 4.7 4.7 4.8 3.7 3.8 3.6 13.1 17.4 15.2 13.7 14.7 13.3 11.8 9.7 9.4 9.4 9.8 12.3 12.6 12.0 14.6 12.3 12.1 12.4 9.6 9.6 9.2 LW live weight; ADG average daily gain; DMI dry matter intake; TMEn true metabolizable energy corrected for nitrogen retention; Prot protein; Lys lysine; Met methionine; Cys cystein; Thr threonine; Isoleu isoleucine; Leu leucine a Based on a diet with gross energy content of 11.5 MJ (From Ref 4.) recommendations on minerals and other nutrients are still based on data from other avian species Different dietary nutrient concentrations are needed through the successive stages of the life cycle Low immunity in the digestive system of the ratite chick until the age of three months is one of the reasons for high mortalities Of the commercial ratite species (ostriches, emus, rheas), nutritional research has mainly concentrated on the ostrich Due to similarities in the digestive system, information obtained with ostriches could probably be extrapolated to the rhea, the least studied species REFERENCES Angel, C.R A review of ratite nutrition Anim Feed Sci Technol 1996, 60, 241 246 Drenowatz, C.; Sales, J.; Sarasqueta, D.V.; Weilbrenner, A History & Geography In Ratite Encyclopedia; Drenowatz, C., Ed.; Ratite Records, Inc.: San Antonio, TX, USA, 1995; 29 Dowsley, W.G.; Gardner, C Ostrich Foods and Feed ing; Crocott & Sherry: Grahamstown, South Africa, 1913 O’Malley, P.J An Estimate of the Nutritional Require ments of Emus In Improving Our Understanding of Ratites in a Farming Environment; Deeming, D.C., Ed.; Ratite Conference: Oxfordshire, UK, 1996; 92 108 Sales, J.; Navarro, J.L.; Bellis, L.; Manero, A.; Lizurume, M.; Martella, M.B Carcass and component yields of rheas Br Poult Sci 1997, 38, 378 380 Cilliers, S.C.; Angel, C.R Basic Concepts and Recent Advances in Digestion and Nutrition In The Ostrich: Biology, Production and Health; Deeming, D.C., Ed.; CAB International: Wallingford, Oxon, U.K., 1999; 105 128 Cilliers, S.C Feedstuffs Evaluation in Ostriches (Struthio camelus) Ph.D Thesis; University of Stellenbosch: South Africa, 1995 Smith, W.A.; Sales, J Feeding and Feed Management In Practical Guide for Ostrich Management and Ostrich Products; Smith, W.A., Ed.; An Alltech Inc Publica tion, University of Stellenbosch Publishers: Stellen bosch, South Africa, 1995; 19 Verwoerd, D.J.; Deeming, D.C.; Angel, C.R.; Perelman, B Rearing Environments Around the World In The Ostrich: Biology, Production and Health; Deeming, D.C., Ed.; CAB International: Wallingford, Oxon, U.K., 1999; 191 216 10 Kruczek, R Breeding Darwin’s rheas at Brookfield Zoo Chicago Int Zoo Yearb 1968, 8, 150 153 Religious Foods: Jewish and Muslim Laws for Animal Slaughter/Welfare Joe M Regenstein Cornell University, Ithaca, New York, U.S.A Carrie E Regenstein University of Wisconsin, Madison, Wisconsin, U.S.A Muhammad M Chaudry Islamic Food and Nutrition Council, Chicago, Illinois, U.S.A INTRODUCTION Prohibition of Mixing Milk and Meat The kosher dietary laws determine which foods are fit or proper for consumption by Jewish consumers who observe these laws The halal dietary laws determine which foods are lawful or permitted for Muslims The kosher and halal dietary laws both deal extensively with animal issues More details about these laws and the additional requirements not covered in this article can be found in other sources.[1–7] ‘‘Thou shalt not seeth the kid in its mother’s milk’’ appears three times in the Torah (the first five books of the Holy Scriptures) and is therefore considered a very serious admonition Meat has been rabbinically extended to include poultry Dairy includes all milk derivatives To keep meat and milk separate requires that the processing and handling of all food products and production equipment that are kosher fall into one of three categories: meat, dairy, or pareve (neutral) Pareve includes all plant products plus eggs, fish, honey, and lac resin (shellac) Pareve foods can be used with either meat or dairy, except that fish cannot be mixed directly with meat Some kosher supervision agencies permit products without meat but made on meat equipment to be listed as ‘‘meat equipment (M.E.).’’ KOSHER DIETARY LAWS Allowed Animals and the Prohibition of Blood Ruminants with split hoofs that chew their cud, the traditional domestic birds, fish with fins and removable scales, and a few grasshoppers are generally permitted Everything else is prohibited Ruminants and fowl must be slaughtered according to Jewish law by a specially trained religious slaughterer using a special knife that is very straight, very sharp, and at least twice the neck diameter in length These animals are subsequently inspected for various defects In the United States, a stricter inspection requirement requires smooth lungs (Glatt), i.e., less than two perforations or adhesions The meat and poultry must be further prepared by properly removing certain veins, arteries, prohibited fats, blood, and the sciatic nerve Therefore, only the front quarter cuts of red meat are generally used To remove more blood, red meat and poultry are soaked and salted within a specified time period All animal ingredients for kosher production must come from kosher-slaughtered animals Thus, fats or oils used for kosher products are mostly obtained from plant sources 770 Equipment Koshering There are three ways to make equipment kosher and/or to change its status Which procedure is required depends on the equipment’s prior production history Converting pareve equipment to use for meat or dairy does not require kosherization The first and simplest equipment kosherization occurs with equipment made from materials that have only been handled cold These require a good caustic/soap cleaning However, materials such as ceramics, rubber, earthenware, and porcelain cannot be koshered Heating above 120°F is usually defined rabbinically as cooking To kosher these items, the second form of equipment kosherization requires that the equipment be thoroughly cleaned with caustic/soap The equipment must be left idle for 24 hours and then flooded with boiling water in the presence of a kosher supervisor For ovens or other equipment that use fire, the third form of Encyclopedia of Animal Science DOI: 10.1081/E EAS 120021146 Copyright D 2005 by Marcel Dekker, Inc All rights reserved Religious Foods: Jewish and Muslim Laws for Animal Slaughter/Welfare equipment kosherization involves heating the metal until it glows with the rabbi present HALAL DIETARY LAWS Prohibited and Permitted Animals; Prohibition of Blood Meat of pigs is strictly prohibited, and so are carnivorous animals and birds of prey Some of the animal and birds are permitted only under special circumstances, e.g., horsemeat may be allowed under certain distressing conditions Animals fed unclean or filthy feed, e.g., sewage or tankage protein, must be fed clean feed for three to 40 days before slaughter Eggs and milk must come from permitted animals According to Quran, blood that pours forth is prohibited from being consumed whether from permitted or nonpermitted animals and any derivatives For seafood, some groups accept only fish with scales as halal, while others consider everything that lives in water, all or some of the time, as halal Animals that live both in water and on land (e.g., amphibians) are not consumed by most Muslims The status of insects is unclear, except that locust is specifically mentioned as halal The use of honey was very highly recommended by Prophet Muhammad Other insect products are generally acceptable; however, some consider shellac and carmine makrooh offensive to their psyche 771 animal may only be dismembered after the blood is drained completely and the animal is lifeless Animalderived food ingredients must be made from Muslimslaughtered halal animals Hunting of wild halal animals is permitted for the purpose of eating, but not for pleasure Allah’s name should be pronounced when ejecting the tool rather than when catching the hunt On catching, the animal must immediately be bled by slitting the throat If the blessing is made at the time of pulling the trigger or shooting an arrow and the hunted animal dies before the hunter reaches it, it would still be halal as long as slaughter is performed and some blood comes out Fish and seafood may be hunted or caught by any reasonable means available as long as it is done humanely The requirements of proper slaughtering and bleeding are applicable to land animals and birds Fish and other water creatures need not be ritually slaughtered Similarly, there is no special method of killing locust The meat of animals that die of natural causes, diseases, from being gored by other animals, by being strangled, by falling from a height, through beating, or killed by wild beasts, is unlawful to be eaten, unless such animals are slaughtered before they become lifeless Fish that dies of itself, if floating on water or lying on shore, is halal as long as it shows no signs of decay or deterioration An animal must not be slaughtered in dedication to other than Allah, or immolated to anyone other than Allah under any circumstances GELATIN Proper Slaughtering of Permitted Animals There are special requirements for slaughtering the animal It must be a halal species slaughter by a sane, adult Muslim with the name of Allah pronounced at slaughter The throat is cut in a manner that induces rapid and complete bleeding, resulting in quick death Generally, at least three of the four passages, i.e., carotids, jugulars, trachea, and esophagus, must be cut to give zabiha or dhabiha meat (meat acceptable for Muslim consumption) Although kosher meat is similarly slaughtered, a prayer is not said over each animal Thus, most Muslim scholars not accept kosher meat as halal In the absence of halal meats, individual Muslims may choose to purchase kosher meat products Islam places great emphasis on humane treatment of animals, especially before and during slaughter Some conditions include giving the animal proper rest and water, avoiding or reducing stress, not sharpening knives in front of animals, and using a very sharp knife The Gelatin is probably the most controversial kosher and halal ingredient Gelatin can be derived from pork skin, beef bones, or beef skin along with fish skin and bones Currently available gelatins even if called kosher are not acceptable to the mainstream kosher supervision organizations or to halal consumers However, limited kosher hide gelatin is available Similarly, at least two sources of certified halal gelatin are available BIOTECHNOLOGY Rabbis and Islamic scholars currently accept products made by simple genetic engineering, e.g., chymosin (rennin) used in cheese making The production conditions in the fermenters must still be kosher or halal, i.e., the ingredients and the fermenter, and any subsequent processing must use kosher or halal equipment and ingredients of the appropriate status A product produced in a dairy medium would be dairy Mainstream rabbis may 772 Religious Foods: Jewish and Muslim Laws for Animal Slaughter/Welfare approve porcine lipase made through biotechnology when it becomes available, if all the other conditions are kosher The Muslim community is still considering the issue of products with a porcine gene; although a final ruling has not been announced, the leaning seems to be toward rejecting such materials If the gene for a porcine-derived product were synthesized, i.e., it did not come directly from the pig, Muslim leaders are prepared to accept it The religious leaders of both communities have not yet determined the status of more complex genetic manipulations and, therefore, such a discussion is premature CONCLUSION ANIMAL WELFARE In the United States, the Food Marketing Institute (representing the major supermarkets) and the National Council of Chain Restaurants (in conjunction with the production agriculture trade associations) has undertaken to develop a set of minimal animal welfare standards As part of that process, a kosher/halal standard and audit requirements have been developed, based on the American Meat Institute’s requirement for upright slaughter.[8] In addition, the Northeast Sheep and Goat Program at Cornell University has developed a low-cost, upright holding pen for small animals, and has identified a commercial knife appropriate for halal slaughter The Cornell program is currently developing a poster on onfarm humane/halal slaughter that will be available in a number of different languages (e.g., English, Arabic, Persian, Spanish) As consumers continue to refine their food requirements, more companies may well choose to provide kosher and halal food products in the marketplace REFERENCES Chaudry, M.M Islamic food laws: Philosophical basis and practical implications Food Technol 1992, (10), 92 Chaudry, M.M.; Regenstein, J.M Implications of bio technology and genetic engineering for kosher and halal foods Trends Food Sci Technol 1994, 5, 165 168 Chaudry, M.M.; Regenstein, J.M Muslim dietary laws: Food processing and marketing Enc Food Sci 2000, 1682 1684 Regenstein, J.M Health aspects of kosher foods Activ Rep Min Work Groups Sub work Groups R & D Assoc 1994, 46 (1), 77 83 Regenstein, J.M.; Regenstein, C.E An introduction to the kosher (dietary) laws for food scientists and food processors Food Technol 1979, 33 (1), 89 99 Regenstein, J.M.; Regenstein, C.E The kosher dietary laws and their implementation in the food industry Food Technol 1988, 42 (6), 86, 88 94 Regenstein, J.M.; Regenstein, C.E Kosher foods and food processing Enc Food Sci 2000, 1449 1453 Regenstein, J.M.; Grandin, T Animal welfare Kosher and halal Inst Food Technol Relig Ethnic Foods Div Newsl 2002, (1), 16 Rumen Microbiology Todd R Callaway United States Department of Agriculture, Agricultural Research Service, College Station, Texas, U.S.A Scott A Martin University of Georgia, Athens, Georgia, U.S.A R C Anderson Tom S Edrington David J Nisbet Kenneth J Genovese United States Department of Agriculture, Agricultural Research Service, College Station, Texas, U.S.A INTRODUCTION The ruminant animal is able to digest feeds due to a mutually beneficial relationship with microorganisms in the rumen (forestomach) These diverse microorganisms degrade and ferment feedstuffs and in turn, provide the animal with usable nutrients The ruminal fermentation is important to the success of ruminant animals, but is inefficient Therefore, strategies have been sought to improve the efficiency of the ruminal fermentation very large, diverse population of microorganisms (bacteria, protozoa, fungi, and viruses) These microorganisms degrade feeds through the process of fermentation (described subsequently) Feedstuffs in the rumen are continuously broken down into smaller and smaller pieces by microbial activity as well as regurgitation and remastication (a process known variously as ruminantion, or chewing the cud) As feed is broken down to pieces less than mm in size, it passes out of the rumen and then to the abomasum (or true stomach) for further degradation and to the intestine for digestion by mammalian enzymes THE RUMINANT ANIMAL The ability of the ruminant animal to utilize low-quality fibrous feedstuffs (e.g., grasses and forages) to produce a high-quality end-product (i.e., meat, milk, and wool) is the result of a mutually beneficial relationship between the mammalian host and the fermentative microbial population inhabiting the rumen (forestomach).[1] Animals equipped with a rumen include cattle, buffalo, sheep, antelope, gazelle, duiker, reindeer, deer, giraffe, and goats; other animals that consume grass (e.g., horses and donkeys) are not considered true ruminants, but rather utilize a postgastric fermentation Mammals not produce enzymes that degrade cellulose (a primary fibrous component of plant materials), but ruminants are able to degrade cellulose via fermentation because of the presence of the rumen and its resident microbial population Ruminant animals are characterized as having teeth on the bottom jaw, and a hard dental pad on the top This arrangement of teeth results in incomplete mastication (chewing) of ingested feed Feed is swallowed and deposited into a large pouch (the rumen) at the end of the esophagus The rumen is a large chamber (can compose up to 30% of the mass of the animal) that is anaerobic (does not contain oxygen) and populated by a Encyclopedia of Animal Science DOI: 10.1081/E EAS 120019789 Copyright D 2005 by Marcel Dekker, Inc All rights reserved FERMENTATION: ANAEROBIC DIGESTION Fermentation is defined as the process of substrate degradation in the absence of oxygen The best known (to humans) fermentations involve the production of beer, wine, or vinegar (acetate), which is also an important endproduct of ruminal and intestinal fermentations Nearly all feed protein and carbohydrates can be degraded by bacteria via ruminal fermentation to produce volatile fatty acids (VFA) and microbial cells (Fig 1) The VFA are absorbed by the host animal and provide the animal with a source of carbon and energy for maintenance and productive functions The most important VFA to the animal are acetate (vinegar), propionate, and butyrate Microbial cells (bodies) are washed out of the rumen along with the small feed particles and are also digested, providing the ruminant animal with an excellent source of high-quality protein (especially essential amino acids), as well as B vitamins as a by-product of fermentation Thus, the ruminant provides the microorganisms a hospitable environment and food in exchange for the microorganisms providing nutrients derived from a low-quality feed to the animal truly, a mutualistic relationship 773 774 Rumen Microbiology Fig Activity of the rumen Although the ruminal fermentation is generally beneficial to the animal, in some cases, the end-products of the ruminal fermentation can be detrimental to the animal, or even to the environment Some bacteria ferment specific amino acids (tryptophan) and produce 3-methylindole, which can be inhaled by the animal, resulting in asphyxiation (bovine emphysema) Other problems that can be traced to production of harmful end-products of the rumen fermentation include bloat (swelling of the rumen caused by gas production) and lactic acidosis (accumulation of strong acid in the rumen, which damages the tissues of the rumen and inhibits the beneficial fermentation) Some of the ammonia produced from ruminal protein fermentation is not utilized by the animal, but is excreted in the urine and directly impacts the environment (environmental nitrogen pollution) Methane is a powerful greenhouse gas produced by ruminal microorganisms that is eructated (belched) by all ruminant animals MICROBIAL ECOLOGY OF THE RUMEN The rumen is one of the most densely populated and diverse microbial ecosystems It is composed of bacteria and protozoa (single-celled microorganisms), fungi (multicellular), and viruses These flora and fauna break down feedstuffs by sequential colonization and synergistic effort (i.e., fungi and bacteria can colonize grass fibers, and break them down to constituent parts, which are further degraded by bacteria to produce VFA and more bacteria) (Fig 1) The most well-understood members of the ruminal ecosystem are the bacteria and, to a lesser degree, the protozoa Well over 200 species of bacteria have been isolated from the rumen However, because the rumen microbial population is very dense, many bacterial species present at very low populations probably have not been isolated The bacterial population is extremely dense and has been estimated to be as high as 1010 cells/ml of ruminal fluid (that is, 10,000,000,000 bacteria/ml) Considering that the ruminal volume of a cow is 75,000 ml or greater, it is no surprise that the rumen has been characterized as the world’s largest fermentation process.[2] Ruminal bacteria can ferment nearly all dietary components, and are often grouped based on their fermentation substrate and/or the end-products of their metabolic activity (Table 1).[3] Some bacteria are generalists and can ferment many substrates fairly well (e.g., Butyrivibrio), while other bacteria are highly selective in what they can utilize (i.e., Anaerovibrio), but can ferment very rapidly Some bacteria specialize in degrading cellulose, others primarily degrade protein in the diet, and still other species produce methane Protozoa are larger, multicellular microorganisms They can ingest and ferment feedstuffs as well as bacteria and smaller protozoa and play a role in nitrogen cycling within the rumen The exact role of protozoa in the rumen is still unclear, although they provide a home for some bacteria to attach to and can share a mutualistic relationship with these bacteria Ruminal fungi are thought to help initiate the degradation of forage and to quickly utilize oxygen ingested with feedstuffs; however, the true significance of ruminal fungi is unclear Whatever their role in the microbial ecosystem, each microbial species has adapted to fill a specific niche in this complex environment USE OF ANTIMICROBIALS TO ENHANCE FERMENTATION EFFICIENCY The microbial fermentation allows ruminant animals to utilize low-quality feedstuffs; however, the process of fermentation is inherently inefficient It can often require more than five pounds of feed to produce one pound of animal gain (meat) or milk This low-feed efficiency makes ruminant production in feedlots and dairy farms quite expensive Therefore, methods to improve the efficiency of the ruminal fermentation have been examined Antibiotics are antimicrobial compounds that kill or stop the growth of bacteria Often, antibiotics are used to treat bacterial diseases in humans or in animals In some cases, antibiotics have been used to try to increase the efficiency of the fermentation or to reduce pathogenic bacteria (both human and animal) in the gastrointestinal tract For example, Tylosin is currently fed to cattle to reduce the incidence of Fusobacteria necrophorum (a bacterium responsible for liver abscesses in cattle); Rumen Microbiology 775 Table Groups of important ruminal bacteria, and the dietary components they are capable of fermenting Dietary component Group of bacteria Forage Cellulose fermenting species Forage Hemicellulose fermenting species Forage Pectin fermenting species Grain Starch fermenting species Any Protein fermenting species Any Fermentation acid utilizing species Any Any Lipid utilizing species Methane producing species Important genera Ruminococcus Fibrobacter Butyrivibrio Butyrivibrio Bacteroides (Prevotella) Ruminococcus Butyrivibrio Bacteroides (Prevotella) Succinovibrio Streptococcus Streptococcus Bacteroides (Prevotella) Succinomonas Lactobacillus Clostridium Peptostreptococcus Bacteroides (Prevotella) Butyrivibrio Megasphaera Megasphaera Selenomonas Anaerovibrio Butyrivibrio Methanobacterium Methanobrevibacter (Adapted from Ref 3, among other sources.) neomycin sulfate has been suggested to be used in feedlot cattle to reduce the human pathogenic bacterium Escherichia coli O157:H7.[4] However, the use of antibiotics as animal growth promotants has come under increased scrutiny due to problems associated with antibiotic resistance (discussed elsewhere in this encyclopedia) In response to this issue, the European Union has recently (2003) enacted a ban on the use of all antimicrobial feed additives in animal rations; it remains to be seen if the United States will follow suit Therefore, the use of antibiotics, especially those used in human medicine, to enhance the efficiency of the rumen fermentation is not widespread or encouraged Ionophores are the most widely used compound that can increase the efficiency of ruminant production.[5] Ionophores are antimicrobials (but not antibiotics) that inhibit Gram-positive bacteria Because the rumen is populated by both Gram-positive and -negative bacteria, the Gram-negative bacteria gain a competitive advantage in the rumen Due to this shift caused by ionophore treatment, ruminal methane, ammonia, and lactic acid production is reduced, and animal growth efficiency is increased This increase in efficiency has led to the widespread use of ionophores in most feedlot cattle in the United States BACTERIAL PATHOGENS IN THE RUMEN Because the rumen is ideally suited for microbial growth, it is no surprise that pathogenic bacteria can also inhabit the rumen E coli O157:H7 and Salmonella (many serotypes) are foodborne pathogenic bacteria that have been isolated from the rumen Both Salmonella and E coli O157:H7 can pose a risk to humans via direct animal contact or through consumption of contaminated meat products Additionally, some Salmonella serotypes can cause severe illness in the host animal Processing plants an excellent job of controlling the spread of these pathogens after slaughter; however, foodborne illnesses that are associated with ruminant-derived food products still occur Therefore, recent research has focused on strategies to reduce these pathogens in animals prior to entry into the food chain CONCLUSION Our knowledge of rumen microbiology has grown immensely over the past 50 years, yet many people still regard the ruminal fermentation processes as a black box 776 Like any other well-developed ecosystem, the rumen is very complex and changes imposed upon the fermentation can have unintended repercussions throughout the ecosystem, which may have a profound effect on the animal Therefore, technologies to improve the efficiency of the ruminal fermentation proposed for the future (including introduction of designer bacteria or super-bugs that can address any perceived shortcomings of the ecosystem) need to be approached with caution Future directions of research into the area of rumen microbiology will certainly include the use of genomics (sequencing of the DNA of ruminal microorganisms) The recent complete sequencing of the genome of predominant cellulose degrading bacteria will surely allow a greater understanding of the complex ruminal ecosystem Rumen Microbiology GI Tract: Anatomical and Functional Comparisons, p 445 GI Tract: Animal/Microbial Symbiosis, p 449 REFERENCES ARTICLES OF FURTHER INTEREST Digesta Processing and Fermentation, p 282 Digestion and Absorption of Nutrients, p 285 Hungate, R.E The Rumen and Its Microbes; Academic Press: New York, NY, 1966 Weimer, P.J Cellulose degradation by ruminal micro organisms Crit Rev Biotechnol 1992, 12, 189 223 Yokoyama, M.G.; Johnson, K.A Microbiology of the Rumen and Intestine In Microbiology of the Rumen and Intestine; Waveland Press: Englewood Cliffs, NJ, 1988; 125 144 Elder, R.O.; Keen, J.E.; Wittum, T.E.; Callaway, T.R.; Edrington, T.S.; Anderson, R.C.; Nisbet, D.J Intervention to reduce fecal shedding of enterohemorrhagic Escherichia coli O157:H7 in naturally infected cattle using neomycin sulfate J Anim Sci 2002, 80 (Suppl 1), 15 Russell, J.B.; Strobel, H.J Effect of ionophores on ruminal fermentation Appl Envir Microbiol 1989, 55,  ... or adhesions The meat and poultry must be further prepared by properly removing certain veins, arteries, prohibited fats, blood, and the sciatic nerve Therefore, only the front quarter cuts of. .. the rabbi present HALAL DIETARY LAWS Prohibited and Permitted Animals; Prohibition of Blood Meat of pigs is strictly prohibited, and so are carnivorous animals and birds of prey Some of the animal. .. indoor pen Feeding recommendations for ratites are not as precise as those for commercial poultry Nutrient requirements are based on restricted data for ostriches and emus,[2,6] and there are no