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[1] Camelus DromedariusCamels belong to the family Camelidae and thereby to the suborder Tylopoda. The tylopoda themselves belong to the order Artiodactyla or cloven-footed animals. The family of Camelidae contains the genera Camelus (old world camel) and Lama (new world camel). The Camelidae originated in North America where the earliest fossil remains of Camelidae have been found. The genus Camelus migrated from North America in the late Tertiary across the then existing land- bridge to Asia and Africa. The Lamas on the other hand reached South America in the ice age across the Central American land-bridge. Included in the genus Camelus are the one-humped dromedary (Camelus dromedarius [1]) and the two-humped bactrian (Camelus bactrianus [2]). The term dromedary is derived from the Greek word "dromados" (run) and in the strict sense is used for riding camels. The name "Bactrian" for the two-humped camel refers to the area "Baktria" in North Afghanistan where this type of camel is thought to have originated . The dromedary is slim, long-legged, short-haired and has its habitat in warm arid and semi-arid areas. The bactrian is stockier, short-legged and has a thicker and longer coat than the dromedary. It mainly occurs in cold and mountainous areas.
[2] Camelus BactrianusThis Website is concerned entirely with the one-humped camel (Camelus dromedarius), the term "camel" should therefore be taken to refer to Camelus dromedarius unless specifically stated otherwise
For more read: An introduction to the camel
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[1] Camelus DromedariusLes chameaux font partie de la famille des Camelidae et de l'ordre des Tylopoda, eux-mêmes des Artiodactyles. La famille des Camelidae contient le genre Camelus (chameau du vieux monde) et Lama (chameau du nouveau monde).
Les Camelidae sont originalement de l'Amérique du Nord où les plus anciens fossiles ont été trouvés. Au Tertaire, le genre Camelus a migré de l'Amérique du Nord en croisant le pont terrestre existant vers l'Asie et l'Afrique. Tandis que les lamas ont migré pendant la glaciation vers l'Amérique latine.
Le genre de Camelus comprend le le dromadaire à une bosse (Camelus dromedarius [1]) et le chameau à deux bosses (Camelus bactrianus [2]). Le terme dromadaire est dérivé du mot grec "dromados" (courir) et est strictement parlant le terme pour des chameaux à monter. Le nom "Bactrianus" pour le chameau à deux bosses fait référence à une région "Baktria" au Nord de l'Afghanistan d'où il est probable que ce type de chameau soit d'origine. Le dromadaire est mince, aux jambes longues et pelage court. Son habitat est dans des zones chaudes et semi-arides. Le chameau est plus trapu aux jambes courtes et son pelage est plus long et plus dense. Il se trouve dans des zones froides et montagnardes.
[2] Camelus BactrianusCe site web ne traite que le chameau à une bosse (Camelus dromedarius). Par conséquent, le terme "chameau" se réfère au Camelus dromedarius.
Pour de plus amples informations veuillez lire la présentation des chameaux (en anglais)
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Camelgate. About Camel Milk. The platform for consulting and knowlege in camel dairy production
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Fresh camel milkIn many arid areas, camels play a central role as milk suppliers. The comparative advantage of the camel as a dairy animal over the other species in the same environment is difficult to quantify; however, it is widely recognised that in absolute terms the camel produces more milk and for a longer period of time than other species maintained in the same environment.
In East Africa, where 60 % of the world camel population is kept, the consumption of camel milk is not limited only to the pastoral nomads, but camel milk is commercialised and sold in the urban areas.
For more read: PDF Milk
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Lait frais de chamelleDans bien des zones arides, les chamelles jouent un rôle essentiel de par le lait qu'elles fournissent. Il est difficile quantifier l'avantage des chamelles face aux autres espèces fournissant du lait dans un même environnement; cependant, il est reconnu qu'en termes absolus la chamelle fournit une plus grande quantité de lait pendant une plus longue période que toute autre espèce gardée dans le même environnement.
En Afrique de l'Est où vivent 60 % des chameaux du monde, la consommation de lait de chamelle n'est pas limitée aux nomades. Au contraire, le lait de chamelle est commercialisé dans les secteurs urbains.
Pour plus d'amples informations veuillez lire le chapitre sur le lait
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Camelgate. Camel Milk Products. The platform for consulting and knowlege in camel dairy production
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1. Fermented Milk
In pastoral societies, milk is traditionally consumed predominantly in the form of fermented milk. Fermentation is the only means of preserving milk under warm condition. In eastern Africa, however, where 60% of the world camel population are held, there is a long tradition in preparing fermented camel milk camel. The milk is either home-consumed or sold.
To prepare fermented camel milk, containers of calabas, cly potes , plant fibre vessels or hollowed wood vessels are smoked by burning chips of Olea africana or Acacia busia. The daily residual fresh milk is poured into the milk container. No starters are used and acidification develops after e few days, either from natural flora of milk when it is not boiled, or from the bacteria growing on the sides of the vessel.
The milk is left in a quite place, often in a covered container sheltered from dust for usually 24-48 hours until it becomes sour. The ambient temperature is normally between 25 and 35 °C. Due to spontaneous nature of the fermentation, this traditional method results in a product with varying taste and flavor and often of poor hygienic quality.
To improve this spontaneous traditional fermentation, controlled fermentation using mesophilic lactic acid bacteria starter culture have been developed and successfully introduced in camel milk processing plants in different eastern African countries.
Journal of dairy research (1990), 57 281-283
2. Cheese
Camel feta cheese immersed in vegetable oil and added herbs
Camel fresh cheese
Most attempts to make cheese from camel milk have revealed major difficulties in getting the milk to coagulate. With the same amount of calf rennet, the coagulation time of camel milk is two to three-folds longer than in cow milk. The action of rennet on camel milk leads to coagulation in the form of flocks, with no firm coagulation. There are some reports in the literature showing that clotting enzyme from one species is more effective and specific with milk from the same species. Chymosin from lamb where found to be more effective with lamb milk than with cow milk. Pig chymosin and pig pepsin have shown higher milk clotting activity against porcine milk than against bovine milk. These findings suggest an adaptation between the proteolytic specificities of the gastric proteases and the structure of the caseins. Accordingly, it can be expected that camel chymosin would be more effective in camel milk than calf chymosin. Following this, our Laboratory developed recombined camel chymosin, from mRNA, obtained from the stomach of a young camel. The process has been patented.
Dried and sweetend camel cheese
After the preliminary tests in laboratory showed the effectiveness of the Camel chymosin field studies for making cheese from camel milk have been successfully run in different east African and Sahel countries.
Camel chymosin can be obtained from Dr. Zakaria Farah function js_43974ce3a9e72(){ string1='mailt'; string2='o:'; string3='farah@camelgate.com'; document.location.href=string1+string2+string3;}farah@camelgate.com
3. Butter
Churning
Camel butter
Like cheese, butter is also not a traditional camel milk product as is it difficult to obtain camel milk butter following the same preparation procedures as for cow’s milk. This is due to the lack of agglutinin, a protein which promotes clustering of fat globules and formation of cream layer in cold milk. Also the high melting point (41-42C°) of camel milk fat makes difficult churning camel milk cream in temperatures commonly used for churning cow milk.
Our laboratory developed a simple method for manufacturing butter from camel milk fat. According to the method, butter was obtained by churning camel milk cream at temperatures between 20 and 25C°. This temperature is considerably higher than that of cow milk which normally varies between 8 and 12C°.
Milchwissenschaft. (1989)44 (7), 412-414
Milchwissenschaft. (1991)46 (6), 361-362
4. Heat treated product
There are very few studies on the effect of heat treatment on the proteins of camel milk. Research work in our Laboratory indicates that the whey proteins in camel milk are more heat resistant than in cow milk. Under the selected experimental conditions the rate of heat denaturation of camel milk whey proteins was approximately twofold lower than cow milk whey proteins. This indicates that camel milk can be easily pasteurised, and there are commercial small and middle scale camel milk processing plants for production of pasteurised milk in Mauritania, Kenya and Somalia.
Milchwissenschaft (1986) 41(12), 763-765
Journal of Dairy Research (1992) 59 229-231
Ongoing investigation to study the ability of camel milk to withstand ultra-high processing temperatures UHT showed heat instability of camel milk. Bulk camel milk collected from camels in Kenya were UHT heat treated applying both direct (150C°/ 2 sec.) and indirect (138C°/ 8-10 sec.) method. After processing the milk was stored at 5, 10, 25 and 30C°for five weeks. After 3 weeks, milk stored at 25 and 30 °C separated forming fine deposit which was more in milk processed by indirect method then the direct. No deposit formation was observed in milk stored at 5 and 10 °C even after 5 weeks storage. This heat instability of camel milk at high processing temperatures can be due to the low content of K-casein and the total absence of â- Lactoglobulin in camel milk. Both proteins play an important role in the heat stability of bovine milk.
Our conclusion so far is that camel milk can not be UHT treated following the same procedure as in cow milk. UHT methods adapted to camel milk are now under investigation.
More on camel milk products, recipes, processing techniques as well as methods for quality control and equipments for small scale camel milk processing plants are in
“Milk and meat from camel: Hand book on products and processing”
Z. Farah and A. Fischer (Editors) . flyer.pdf
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1. Lait fermenté
Selon les traditions des sociétés pastorales, le lait est surtout consommé fermenté. La fermentation est le seul moyen de conserver le lait dans un climat chaud. Cependant, en Afrique orientale où vivent 60% des chameaux du monde, il y a une longue tradition de fermentation de lait de chamelle.
Pour faire fermenter le lait de chamelle, des récipients de calebasse, de fibres de plantes ou de bois creux ou encore des pots d'argile sont fumés en brûlant des coupures d'Olea africana ou d'acacia busia. Chaque jour, le lait frais résiduel est versé dans le récipient de lait. Aucun agent de fermentation n'est utilisé et l'acidification prend son cours après peu de jours, soit du fait de la flore naturelle du lait qui n'est pas bouilli, soit suite aux bactéries présentes aux récipients.
Le lait est conservé à un endroit isolé, souvent dans un récipient couvert abrité de la poussière pendant environ 24-48 heures jusqu'à ce qu'il soit acidulé. La température ambiante est normalement entre le 25 et 35 °C . En raison de la fermentation spontanée, le résultat de cette méthode traditionnelle est un produit aux goûts et saveurs variables et ne correspondant souvent pas aux standards hygiéniques minimaux.
Pour améliorer cette fermentation spontanée traditionnelle, une fermentation contrôlée se basant sur une culture d'amorçage mésophile de bactéries d'acide lactique a été développée et introduite dans des établissements de traitement de lait de chamelle dans différents pays d'Afrique orientale.
Journal of dairy research (1990), 57 281-283
2. Fromage
Feta de lait de chamelle aux herbes dans l'huile végétale
Fromage frais de chamelle
Lors de la plupart des tentatives de faire du fromage à partir du lait de chamelle des difficultés se sont présentées quant à la coagulation du lait. Avec la même quantité de présure de veau, le temps de coagulation du lait de chamelle est deux à trois fois aussi long que celui du lait de vache. La présure fait coaguler en flocons le lait de chamelle, sans qu'une coagulation ferme ne se produise. Il y a des mentions dans la littérature qui indiqueraient que l'enzyme de coagulation d'une espèce est plus efficace et spécifique avec du lait de la même espèce. La chymosine d'agneau a prouvé être plus efficace avec du lait de brebis qu'avec du lait de vache. La chymosine de porc et la pepsine de porc ont déployé une meilleure activité de coagulation dans le lait de truie que dans le lait de vache. Ces résultats suggèrent une adaptation entre les spécificités protéolytiques des protéases gastriques et la structure des caséines. Par conséquent, il peut être assumé que dans le lait de chamelle la chymosine de chameau serait plus efficace que la chymosine de veau. Pour cette raison, notre laboratoire a développé la chymosine de chameau sur la base du mRNA, prélevé de l'estomac d'un chamelon. Le processus est breveté.
Fromage séché et sucré de chameau
Du fait que l'efficacité de la chymosine de chameau a été prouvée par des essais préliminaires dans le laboratoire, des essais pour faire du fromage de lait de chamelle ont été effectués avec succès dans plusieurs pays de l'Afrique orientale et du Sahel.La chymosine de chameau peut être obtenue chez Dr. Zakaria Farah farah@camelgate.com
3. Beurre
Barattage
Beurre de lait de chamelle
Comme le fromage, le beurre n'est pas non plus un produit laitier traditionnel, étant donné qu'il est difficile d'obtenir du beurre du lait de chamelle de la même manière que du lait de vache. Ceci est dû au manque d'agglutinine, une protéine qui favorise l'agglomération des globules de graisse et la formation d'une couche de crème dans le lait froid. En outre, le point de fusion élevé (41 à 42°C) de la matière grasse du lait de chamelle rend celui-ci difficile à baratter aux températures normales du barattage de lait de vache.
Notre laboratoire a développé une méthode simple pour produire du beurre de la matière grasse du lait de chamelle. Selon cette méthode, du beurre est obtenu en battant la crème du lait de chamelle à des températures entre 20 et 25°C. Ces températures sont considérablement plus élevées que celles du lait de vache qui se situent normalement entre 8 et 12°C.
Milchwissenschaft. (1989)44 (7), 412-414
Milchwissenschaft. (1991)46 (6), 361-362
4. Produit réchauffé
Il y a très peu d'études quant à l'effet du traitement thermique sur les protéines du lait de chamelle. Les recherches de notre laboratoire montrent que les protéines de petit-lait du lait de chamelle résistent mieux à la chaleur que ceux du lait de vache. Dans les conditions expérimentales choisies, le taux de dénaturation par la chaleur des protéines de petit-lait du lait de chamelle était environ de la moitié inférieur à celui des protéines de petit-lait du lait de vache. Ceci montre que du lait de chamelle peut facilement être pasteurisé et il y a de petites et moyennes installations commerciales de traitement de lait de chamelle produisant du lait pasteurisé de chamelle en Mauritanie, au Kenya et en Somalie.
Milchwissenschaft (1986) 41(12), 763-765
Journal of dairy Research (1992) 59 229-231
La recherche au sujet de la résistance à ultra haute température (UHT) du lait de chamelle a montré son instabilité à la chaleur. Le lait de chamelle en gros collectionné de chamelles au Kenya était soumis à un traitement à ultra haute température, appliquant aussi bien la méthode directe (150°C/2 sec.) que l'indirecte (138°C/8-10 sec.). Suite au traitement, le lait a été conservé à 5, 10, 25 et 30 °C pendant cinq semaines. Après 3 semaines, le lait conservé à 25 et 30 °C a caillé en former un dépôt fin qui se trouvait davantage dans le lait traité par la méthode indirecte plutôt que dans celle traité par la méthode directe. Aucune formation de dépôt n'a été notée dans le lait conservé à 5 et 10 °C même après 5 semaines. Il est possible que cette instabilité à la chaleur du lait de chamelle aux ultra hautes températures soit due à la basse teneur de K-caséine et à l'absence totale d'alpha-lactoglobuline dans le lait de chamelle. Les deux protéines jouent un rôle important dans la stabilité à la chaleur du lait de vache.
Notre conclusion en est que le lait de chamelle ne peut être traité UHT selon le même procédé que le lait de vache. A présent, des méthodes UHT adaptées au lait de chamelle sont à l'étude.
Vous trouverez des informations détaillées sur des produits laitiers de chamelle, des recettes, des techniques de traitement et des méthodes de contrôle de qualité ainsi que de petites installations de traitement du lait de chamelle dans
“Milk and meat from camel: Hand book on products and processing” (en anglais)
Z. Farah and A. Fischer (rédacteurs). . flyer.pdf
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Camelgate. Camel Meat. The platform for consulting and knowlege in camel dairy production
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Skinning the camel on the skinning trestle
Camel meat is an important by-product mainly as a source of income. Sale
of live camels, usually males and unproductive females for slaughter, is very
common in East Africa and there are now increasing numbers of camel butcheries in many urban centers. There is also a growing export trade of slaughter camels to the Arabian Peninsula. In the following, special knowledge gained in slaughtering, deboning and processing of camels on-site over several years of long-term stays on the Ol Maisor Farm in Kenya is presented. During the development of camel meat products the authors set a great value on the keeping quality of the products as well as creating a wide range of product groups with low and high cost for the local market and a good shelf life.
For more read: PDF: Traditional slaughter, carcass dressing and processing of camels.
or read the following book: “Milk and meat from camel: Hand book on products and processing”
Z. Farah and A. Fischer (Editors) flyer.pdf
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Peler le chameau sur un chevalet
La viande de chameau est un sous-produit important surtout comme source de revenu. La vente de chameaux vivants, habituellement des mâles et des femelles non-productives pour l'abattage, est très commun en Afrique orientale et il y en a un nombre croissant de boucheries de chameaux dans bien des centres urbains. Il y a également une exportation croissante des chameaux d'abattage vers la péninsule arabique. Par la suite sont exposées les connaissances spéciales quant à l'abattage, le désossement et le traitement des chameaux sur place comme elles ont été gagnées au cours de plusieurs années de séjours prolongées à la ferme d'Ol Maisor au Kenya. Lors du développement des produits de viande de chameau, les auteurs ont accordé une grande importance à la possibilité de conservation des produits aussi bien qu'à la création d'un large éventail de groupes de produits aux coûts modérés et élevés pour le marché local et une bonne durée de conservation.
Veuillez lire à ce sujet: PDF: Traditional slaughter, carcass dressing and processing of camels.
ou le livre: “Milk and meat from camel: Hand book on products and processing”
Z. Farah and A. Fischer (rédacteurs) flyer.pdf
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Camelgate. Our Team. The platform for consulting and knowlege in camel dairy production
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We are a group of academics with extensive knowledge on the use of camels (Camelus dromedaries). This knowledge is based on results of many years research work carried out in partnership with universities, camel farmers and pastoralists in arid regions of Africa. Our research covers both basic and practical oriented research, aiming to increase camel productivity through development of market oriented milk and meat products, of high hygienic safety and prolonged shelf life, adding value to the existing traditional camel products. This can improve the income generating opportunities for camel keeping pastoralists and SME engaged in processing and marketing of camel products.
Milk Science and Technology
Dr. Zakaria Farah. Team Leader
Dr. Zakaria Farah is since 1982 senior lecturer for food technology in developing countries at Institute of Food Science and Nutrition, ETHZ. He is a leader of a group within the institue dealing with development oriented research programmes in the area of food security in developing countries.
For the last 15 years camel milk has been on of the main research areas for the group of Dr. Farah. This research, which has been done in collaboration with different research institutions in Kenya, Somalia and Ethiopia, has resulted in many publications in international papers and two textbooks on camel milk and several PhD theses.
Dr. Stefan Kappeler
Dr. Stefan Kappeler, has studied the amounts and characteristics of individual proteins in camel milk in the course of a PhD thesis. One of the projects in a following PostDoc was the recombinant production of camel chymosin for biochemical analysis and evaluation of its effectiveness as a milk coagulant. He presently works as a quality assurance manager in the pharmaceutical industry.
Dr. John Wangoh
Dr. John Wangoh obtained his PhD on chemical and technological properties of camel milk from Swiss Federal Institute of Technology ETH Zurich and is a Lecturer for Dairy technology at Nairobi University. Currently he is EU-consultant for Planning, Installation, Operation and Training for a Camel Milk Plant in Gardo District, Puntland State, Somalia.
Anne Bruntse
Danish Agronomist, worked for Danish extension service, FAO, Privat farming in Kenya, Developed own mini dairy using appropriate technology and developed milk processing recipes suitable for local conditions. Currently assisting KARI (Kenya Agriculture Research Institute) start mini entreprise using appropriate technolgy in pastoral areas, including developing recipes for sale and storage of camel milk.
Animal Health
Dr. Mario Younan
Dr. Mario Khaled Younan is a German veterinarian based in Kenya. He has worked on camel mastitis and on diseases of came calves. Currently he is employed by the Kenya Camel Association under the CTZ/CIM Integrated Experts Program. Besides hunting down bacterical agents in camels and in their milk he is also involved in the hygene monitoring of a camel milk dairy in Somalia
Dr. Omar Abdurahman
Dr. Omar Abdurahman has graduated from the faculty of veterinary medicine of the Somali National University in 1981. He has worked as a pathologist at the central veterinary laboratory in Mogadishu over 10 years and was involved as a researcher in the Somali camel research Project since 1986. During 1987/88 he worked as livestock research officer with NGO in Sablale district development programme. Dr. Abdurahman is a fellow of the Royal Veterinary College (FRCVS, Sweden) and obtained MSc and PhD from the Swedish University of Agricultural Sciences (SLU). He is currently working as a researcher at the Department of Obstetrics and Gynaecology, SLU Uppsala, Sweden.
Meat Science and Technology
Prof. Dr. Albert Fischer
Albert Fischer, Prof. Dr. med. vet., born 1939, study of Veterinary Medicine at the University of Munich, State Examination 1968, Dr. med. vet. 1970, Master Butcher 1972; since 1976 Professor for Meat Technology at the Institute of Food Technology, University of Hohenheim, Stuttgart, Germany.
Kathrin Stephanie Ulmer
Kathrin Stephanie Ulmer, Food Technologist, born 1969, trained as cook from 1989 to 1992, study of Food Technology at the University of Hohenheim, Stuttgart, Germany; since 1999 PhD student at the Department of Meat Technology at the Institute of Food Technology, University of Hohenheim, Stuttgart, Germany.
Kurt Hermann
Kurt Herrmann, born 1960, Master Butcher 1985; since 1986 responsible for the pilot plant of Meat Technology at the Institute of Food Technology, University of Hohenheim, Stuttgart, Germany. From 1996–2001 he stayed several times at the Ol Maisor Farm in Kenya. He took part in developing the described slaughter process, the deboning and cutting of camels and several camel meat products.
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Nous sommes un groupe d'académiciens aux vastes connaissances quant à l'utilisation des chameaux (dromadaires de Camelus). Cette connaissance est basée sur des résultats de recherches de nombreuses années menées à bien dans de concert avec des universités, des fermiers de chameaux et des pasteurs de régions arides d'Afrique. Nos recherches couvrent la recherche pratique et théorique, visant à augmenter la productivité des chameaux par le développement de produits laitiers et de viande adaptés au marché et d'une excellente sécurité hygiénique ainsi qu'à la durée de conservation prolongée, augmentant ainsi la valeur des produits traditionnels existants. Ceci est sensible de générer des possibilités de revenu pour des pasteurs gardant des chameaux et des PME s'occupant du traitement et du marketing de produits de chameau.
Science et technologie du lait
Dr. Zakaria Farah. Chef de l'équipe
Depuis 1982, le Dr Zakaria Farah est professeur-assistant en technologie alimentaire des pays en voie de développement à l'institut de la science de l'alimentation et de la nutrition de l'ETHZ. Il est le chef d'un groupe de l'institut chargé de programmes de recherche de développement dans le secteur de la sécurité de l'alimentation dans les pays en voie de développement.
Pendant les 15 dernières années, le lait de chamelle a été l'un des domaines principaux de recherche du groupe du Dr Farah. De cette recherche en collaboration avec différents instituts de recherche au Kenya, en Somalie et en Ethiopie est issu un grand nombre de de publications dans des journaux internationaux, deux guides sur le lait de chamelle et plusieurs thèses de PhD.
Dr. Stefan Kappeler
Le Dr Stefan Kappeler a étudié le contenu et les caractéristiques de différentes protéines du lait de chamelle au cours d'une thèse de PhD. Un des projets au cours d'un PostDoc était la recombinaison de la chymosine de chameau pour l'analyse biochimique et l'évaluation de son efficacité comme coagulant de lait. A présent, il est directeur d'assurance de qualité dans l'industrie pharmaceutique.
Dr. John Wangoh
Le Dr John Wangoh a obtenu son PhD sur les propriétés chimiques et technologiques du lait de chamelle à l'Ecole Polytechnique Fédérale à Zurich et est un assistant de technologie laitière à l'université de Nairobi. Actuellement, il conseille l'UE en matière de planification, d'installation, d'opération et de formation en faveur d'un centre de production de lait de chamelle au District de Gardo, état de Puntland, Somalie.
Anne Bruntse
L'agronome danoise a travaillé pour l'office de développement danois, la FAO, l'agriculture privée au Kenya et a établi sa propre petite laiterie en utilisant la technologie appropriée et a développé des recettes pour le traitement du lait adapté aux conditions locales. Actuellement, elle assiste le KARI (Kenya Agriculture Research Institute) lors de l'établissement d'une petite entreprise utilisant la technologie adaptée aux régions pastorales, y compris la commercialisation et la conservation de lait de chamelle.
Santé des animaux
Dr. Mario Younan
Le Dr Mario Khaled Younan est un vétérinaire allemand basé au Kenya. Il a fait des recherches sur la mastite de chamelles et sur les maladies de chamelons. A présent, il est employé par l'association du chameau du Kenya dans le cadre du CTZ/CIM Integrated Experts Program. Outre sa lutte contre des bactéries des chameaux et du lait de chamelle, il est également engagé dans la surveillance de l'hygiène d'une entreprise de transformation de lait de chamelle en Somalie.
Dr Omar Abdurahman
Le Dr Omar Abdurahman a obtenu un diplôme de la faculté de médecine vétérinaire de l'université nationale de Somalie en 1981. Il a travaillé en tant que pathologiste au laboratoire vétérinaire central à Mogadiscio pendant plus de 10 ans. Depuis 1986 il était engagé dans le projet de recherche somali sur les chameaux. Au cours des années 1987/88 il était directeur de recherche sur le bétail d'une organisation non-gouvernementale au sein du programme de développement du district de Sablale. Le Dr Abdurahman est un professeur de l'université vétérinaire royale (FRCVS, Suède) et a obtenu un MSc et un PhD de l'université suédoise des sciences agronomiques (SLU). A présent, il est chercheur au département de l'obstétrique et de la gynécologie, SLU Uppsala, Suède.
Science et technologie de la viande
Prof. Dr. Albert Fischer
Né en 1939, Albert Fischer, Prof. Dr med. vet., a étudié la médecine vétérinaire à l'université de Munich. Diplômé en 1968, doctorat en 1970, maître-boucher en 1972; depuis 1976 professeur de technologie de viande à l'Institut de technologie alimentaire, Université de Hohenheim, Stuttgart, Allemagne.
Kathrin Stephanie Ulmer
Kathrin Stephanie Ulmer, ingénieur d'alimentation, née en 1969, qualifiée comme cuisinière de 1989 à 1992, études de technologie alimentaire à l'Université de Hohenheim, Stuttgart, Allemagne. Depuis 1999 étudiante de PhD au département de la technologie de viande à l'institut de la technologie alimentaire, Université de Hohenheim, Stuttgart, Allemagne.
Kurt Hermann
Kurt Herrmann, né en 1960, maître-boucher en 1985; Depuis 1986 responsable de l'installation pilote de technologie de viande à l'Institut de technologie alimentaire, Université de Hohenheim, Stuttgart, Allemagne. De 1996 à 2001 il a fait plusieurs séjours à la ferme d'Ol Maisor au Kenya. Il a participé au développement des procédés d'abattage, de désossement et de coupe de chameaux décrits et de plusieurs produits à base de viande de chameau.
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We offer a variety of services
We offer the following consulting services to Research institutions, private enterprises, development agencies and government bodies.
Practical oriented services
Market studies to identify needs on camel milk and meat products and opportunities for technology change for sustainable improved production
Assessment of potential consumer health hazard and risk factors of milk and meat products
Planning, supplying, installing and monitoring turnkey camel milk and meat processing plants driven by solar or conventional energy for small und middle enterprises.
Training and capacity building in the area of camel husbandry, veterinary, quality assurance, milk and meat hygiene, meat and milk related processing techniques
Establish contacts to camel breeders for camel acquisition
Research and policy oriented services
Collaborative scientific projects with research institutions
Strategy development and policy Guidelines to improve infrastructure service for the pastoral populations
Feasibility studies on rural livelihood in arid and semi-arid regions
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Nous offrons de multiples services
Nous offrons les services de conseil suivants à des établissements de recherche, à des entreprises privées, à des agences de développement et à des corps de gouvernement.
Services pratiques
Etudes du marché pour identifier les besoins quant à des produits laitiers et de viande de chameau ainsi que des occasions au changement de technologie pour une production durable
Evaluation du risque pour la santé des consommateurs et des risques potentiels de produits laitiers et de viande
Planification, livraison, installation et surveillance de petites et moyennes installations "clés en main" pour la transformation de lait et de viande de chameau alimentées par énergie solaire ou conventionnelle
Promotion de la formation et des connaissances dans le domaine de la garde de chameaux, la médecine vétérinaire, la garantie de la qualité, l'hygiène du lait et de la viande ainsi que les techniques de transformation de la viande et du lait
Etablissement de contacts avec des éleveurs de chameaux pour l'acquisition de chameaux
Services en faveur de la recherche et de la politique
Projets de collaboration scientifiques avec des établissements de recherche
Développement de stratégie et directives de politique afin d'améliorer les services d'infrastructure pour la population pastorale
Etudes de faisabilité sur la subsistance rurale dans des régions arides et semi-arides
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The commercialisation of camel milk
Selling the milk
Collecting the milk
In Eastern African countries (Kenya, Somalia, Sudan and Ethiopia) where most of the world’s camel populations are kept camel milk plays a central role in food security. In the context of advancing urbanisation camel milk, raw or traditionally fermented is increasingly commercialised and consumed in urban areas. Due to increasing demand there is growing interest in processing camel for the urban market. However, good quality dairy products can be obtained only from milk of high hygienic quality. This makes the improvement of milk hygiene the pre-requisite for any future camel dairy development.
The production areas are often located far from markets. Distances to provincial markets range from 20 to 90 km and may be up to 400 km for distant urban markets. During periods of milk surplus (rainy season) transport on dirt roads is unreliable resulting in breakdowns and delays in milk delivery. Storage in unhygienic containers, pooling of milk from different suppliers, prolonged transport times and high environmental temperatures all increase contamination and spoilage of milk.
This results in considerable losses in marketed camel milk caused by frequent spoilage due to the absence of appropriate conservation and storage methods. In addition the presence of zoonotic infectious diseases in camels results in a potentially high health risk to the public.
More on that read: Milk Hygiene and Udder health
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La commercialisation du lait de chamelle
Vente du lait
Collection du lait
Dans les pays d'Afrique orientale (Kenya, Somalie, Soudan et Ethiopie) où le plus grand nombre de chameaux vivent, le lait de chamelle joue un rôle essentiel pour la nourriture quotidienne. Du fait de l'urbanisation en cours, le lait de chamelle - frais ou fermenté - est de plus en plus commercialisé et consommé dans des secteurs urbains. La demande croissante, est à l'origine de l'intérêt de traiter le lait de chamelle pour le marché urbain. Cependant, des produits laitiers de bonne qualité ne peuvent être obtenus qu'à partir de lait d'une excellente qualité hygiénique. L'amélioration de l'hygiène laitière est donc essentielle pour tout développement de produits laitiers à base de lait de chamelle.
Les secteurs de production sont souvent situés loin des marchés. Les distances aux marchés régionaux varient entre 20 à 90 km et peuvent aller jusqu'à 400 km pour les marchés urbains éloignés. Pendant des périodes de surplus de lait (saison des pluies) le transport sur les pistes est incertain ayant pour résultat des pannes et des retards de livraison du lait. Le stockage dans des récipients peu hygiéniques, la collection de lait de différents fournisseurs, des durées de transport prolongées et des températures élevées augmentent le risque de contamination et de détérioration du lait.
Il en résultent des pertes considérables de lait de chamelle en vente sur le marché suite à la détérioration fréquente due à l'absence de méthodes appropriées de conservation. En outre, des maladies infectieuses zoonotiques des chameaux constituent un risque élevé pour la santé du public.
Veuillez lire à ce sujet: Milk Hygiene and Udder health
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Camélidéscamelides.cirad.fr
Cassavawww.cassava.ethz.ch
Centre Suissewww.csrs.ch
ETH-Life (Deutsch)www.eth-life.ethz.ch/articles/kamelbuch.html
Lait Sain pour le Sahelwww.laitsain.com
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Camélidéscamelides.cirad.fr
Cassavawww.cassava.ethz.ch
Centre Suissewww.csrs.ch
ETH-Life (Deutsch)www.eth-life.ethz.ch/articles/kamelbuch.html
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Swiss Tropical Institutewww.sti.ch
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Centre Suissewww.csrs.ch
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Swiss Tropical Institutewww.sti.ch
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ETH-Life (Deutsch)www.eth-life.ethz.ch/articles/kamelbuch.html
Lait Sain pour le Sahelwww.laitsain.com
Swiss Tropical Institutewww.sti.ch
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Our publications
2004 Journal Of Dairy Science 87 2004 2660
2003 Journal Of Dairy Science 86 2003 498
2001 Journal Of Dairy Research 68 2001 1
1999 International Dairy Journal 9 1999 481
1998 Journal Of Dairy Research 65 1998 209
1998 International Dairy Journal 8 1998 617
1998 Dissertation Kappeler 1998 Eth12947
1994 Journal For Vitamin And Nutrition Research 62 1994 30
1993 Milchwissenschaft 48 1993 322
1992 Journal Of Dairy Research 59 1992 229
1991 Milchwissenschaft 46 1991 361
1991 Journal Of Dairy Science 74 1991
1990 Journal Of Dairy Research 57 1990 281
1989 Milchwissenschaft 44 1989 412
1989 Food Microstructure 8 1989 211
1987 Milchwissenschaft 42 1987 689
1986 Milchwissenschaft 41 1986 763
1985 Milchwissenschaft 40 1985 669
Excerpts from the Book "Milk and meat from camel: Hand book on products and processing"
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Milk Hygiene And Udder Health
Milk
Introduction
WhereToOrderTheBook
Weltwoche48 04
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2004 Journal Of Dairy Science 87 2004 2660
2003 Journal Of Dairy Science 86 2003 498
2001 Journal Of Dairy Research 68 2001 1
1999 International Dairy Journal 9 1999 481
1998 Journal Of Dairy Research 65 1998 209
1998 International Dairy Journal 8 1998 617
1998 Dissertation Kappeler 1998 Eth12947
1994 Journal For Vitamin And Nutrition Research 62 1994 30
1993 Milchwissenschaft 48 1993 322
1992 Journal Of Dairy Research 59 1992 229
1991 Milchwissenschaft 46 1991 361
1991 Journal Of Dairy Science 74 1991
1990 Journal Of Dairy Research 57 1990 281
1989 Milchwissenschaft 44 1989 412
1989 Food Microstructure 8 1989 211
1987 Milchwissenschaft 42 1987 689
1986 Milchwissenschaft 41 1986 763
1985 Milchwissenschaft 40 1985 669
Extraits du livre "Milk and meat from camel: Hand book on products and processing"
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Milk Hygiene And Udder Health
Milk
Introduction
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Weltwoche48 04
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2004 Journal Of Dairy Science 87 2004 2660
2003 Journal Of Dairy Science 86 2003 498
2001 Journal Of Dairy Research 68 2001 1
1999 International Dairy Journal 9 1999 481
1998 Journal Of Dairy Research 65 1998 209
1998 International Dairy Journal 8 1998 617
1998 Dissertation Kappeler 1998 Eth12947
1994 Journal For Vitamin And Nutrition Research 62 1994 30
1993 Milchwissenschaft 48 1993 322
1992 Journal Of Dairy Research 59 1992 229
1991 Milchwissenschaft 46 1991 361
1991 Journal Of Dairy Science 74 1991
1990 Journal Of Dairy Research 57 1990 281
1989 Milchwissenschaft 44 1989 412
1989 Food Microstructure 8 1989 211
1987 Milchwissenschaft 42 1987 689
1986 Milchwissenschaft 41 1986 763
1985 Milchwissenschaft 40 1985 669
Slaughter
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Milk Hygiene And Udder Health
Milk
Introduction
WhereToOrderTheBook
Weltwoche48 04
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2004 Journal Of Dairy Science 87 2004 2660
2003 Journal Of Dairy Science 86 2003 498
2001 Journal Of Dairy Research 68 2001 1
1999 International Dairy Journal 9 1999 481
1998 Journal Of Dairy Research 65 1998 209
1998 International Dairy Journal 8 1998 617
1998 Dissertation Kappeler 1998 Eth12947
1994 Journal For Vitamin And Nutrition Research 62 1994 30
1993 Milchwissenschaft 48 1993 322
1992 Journal Of Dairy Research 59 1992 229
1991 Milchwissenschaft 46 1991 361
1991 Journal Of Dairy Science 74 1991
1990 Journal Of Dairy Research 57 1990 281
1989 Milchwissenschaft 44 1989 412
1989 Food Microstructure 8 1989 211
1987 Milchwissenschaft 42 1987 689
1986 Milchwissenschaft 41 1986 763
1985 Milchwissenschaft 40 1985 669
Slaughter
NewsPdf1
Milk Hygiene And Udder Health
Milk
Introduction
WhereToOrderTheBook
Weltwoche48 04
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Important titles and authors of camel related books
Ilse Kohler Rollefson et al.A Field Manual of Camel Diseases: Traditional and Modern Veterinary Care for the DromedaryITDG Publishing 2001
Huelsebusch, C. & Kaufmann, B.Camel breeds and breeding Northern Kenya. Kenya Agricultural research institute [http://www. kari.org] 2002
Evans, J. & Etkin, D.Camel keeping in Kenya, Range Management Handbook of KenyaEnglish Press Limited, Nairobi. 1994
Farah, ZakariaCamel milk: Properties and productsSKAT-Verlag, St. Gallen, Switzerland. 1996
Yagil, R.Camels and camel milkAnimal Production and Health Paper, No. 26, FAO, Rome, pp. 1982
Wangoh, J.Chemical and Technological Properties of Camel MilkDiss. ETH Nr. 12295, Zurich 1997
Kappeler, St.Compositional and structural analysis of camel milk proteins with emphasis on protective proteins. Diss. ETH No. 12947, Zurich, Switzerland. 1998
Bonnet, P. (Editor)Dromedaries and camels, milking animals. Actes du Colloque, 24–26 Octobre 1994, Nouakchott, Mauritanie CIRAD Montpellier France 1998
Ramet, J.P.La technologie des fromages au lait de dromadaire. Etude FAO Production et Santé animals 113 1993
Launois M., Laveissière G., Faye B., Kriska M. A.Le dromadaire pédagogiqueCIRAD Montpellier France 2002 2-87614-528-6
Wilson, R.T.The camelLongman House, Burntmill, Harlow, Essex, UK 1984
Bulliet, R.W.The camel and the wheel. Harvard University Press, Cambridge Mass. 1975
Gautier-Pilters, H. & Dagg, A.I.The camel. Its evaluation, ecology, behaviour and relationship to man.The University of Chicago Press, Chicago. 1981
Schwartz, H. J. & Dioli, M.The one humped camel in eastern Africa – a pictorial guide to diseases, health care and managementVerlag Josef Margraf, Scientific books 1992
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Des livres importants en relation avec les chameaux
Ilse Kohler Rollefson et al.A Field Manual of Camel Diseases: Traditional and Modern Veterinary Care for the DromedaryITDG Publishing 2001
Huelsebusch, C. & Kaufmann, B.Camel breeds and breeding Northern Kenya. Kenya Agricultural research institute [http://www. kari.org] 2002
Evans, J. & Etkin, D.Camel keeping in Kenya, Range Management Handbook of KenyaEnglish Press Limited, Nairobi. 1994
Farah, ZakariaCamel milk: Properties and productsSKAT-Verlag, St. Gallen, Switzerland. 1996
Yagil, R.Camels and camel milkAnimal Production and Health Paper, No. 26, FAO, Rome, pp. 1982
Wangoh, J.Chemical and Technological Properties of Camel MilkDiss. ETH Nr. 12295, Zurich 1997
Kappeler, St.Compositional and structural analysis of camel milk proteins with emphasis on protective proteins. Diss. ETH No. 12947, Zurich, Switzerland. 1998
Bonnet, P. (Editor)Dromedaries and camels, milking animals. Actes du Colloque, 24–26 Octobre 1994, Nouakchott, Mauritanie CIRAD Montpellier France 1998
Ramet, J.P.La technologie des fromages au lait de dromadaire. Etude FAO Production et Santé animals 113 1993
Launois M., Laveissière G., Faye B., Kriska M. A.Le dromadaire pédagogiqueCIRAD Montpellier France 2002 2-87614-528-6
Wilson, R.T.The camelLongman House, Burntmill, Harlow, Essex, UK 1984
Bulliet, R.W.The camel and the wheel. Harvard University Press, Cambridge Mass. 1975
Gautier-Pilters, H. & Dagg, A.I.The camel. Its evaluation, ecology, behaviour and relationship to man.The University of Chicago Press, Chicago. 1981
Schwartz, H. J. & Dioli, M.The one humped camel in eastern Africa – a pictorial guide to diseases, health care and managementVerlag Josef Margraf, Scientific books 1992
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Ilse Kohler Rollefson et al.A Field Manual of Camel Diseases: Traditional and Modern Veterinary Care for the DromedaryITDG Publishing 2001
Huelsebusch, C. & Kaufmann, B.Camel breeds and breeding Northern Kenya. Kenya Agricultural research institute [http://www. kari.org] 2002
Evans, J. & Etkin, D.Camel keeping in Kenya, Range Management Handbook of KenyaEnglish Press Limited, Nairobi. 1994
Farah, ZakariaCamel milk: Properties and productsSKAT-Verlag, St. Gallen, Switzerland. 1996
Yagil, R.Camels and camel milkAnimal Production and Health Paper, No. 26, FAO, Rome, pp. 1982
Wangoh, J.Chemical and Technological Properties of Camel MilkDiss. ETH Nr. 12295, Zurich 1997
Kappeler, St.Compositional and structural analysis of camel milk proteins with emphasis on protective proteins. Diss. ETH No. 12947, Zurich, Switzerland. 1998
Bonnet, P. (Editor)Dromedaries and camels, milking animals. Actes du Colloque, 24–26 Octobre 1994, Nouakchott, Mauritanie CIRAD Montpellier France 1998
Ramet, J.P.La technologie des fromages au lait de dromadaire. Etude FAO Production et Santé animals 113 1993
Launois M., Laveissière G., Faye B., Kriska M. A.Le dromadaire pédagogiqueCIRAD Montpellier France 2002 2-87614-528-6
Wilson, R.T.The camelLongman House, Burntmill, Harlow, Essex, UK 1984
Bulliet, R.W.The camel and the wheel. Harvard University Press, Cambridge Mass. 1975
Gautier-Pilters, H. & Dagg, A.I.The camel. Its evaluation, ecology, behaviour and relationship to man.The University of Chicago Press, Chicago. 1981
Schwartz, H. J. & Dioli, M.The one humped camel in eastern Africa – a pictorial guide to diseases, health care and managementVerlag Josef Margraf, Scientific books 1992
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Ilse Kohler Rollefson et al.A Field Manual of Camel Diseases: Traditional and Modern Veterinary Care for the DromedaryITDG Publishing 2001
Huelsebusch, C. & Kaufmann, B.Camel breeds and breeding Northern Kenya. Kenya Agricultural research institute [http://www. kari.org] 2002
Evans, J. & Etkin, D.Camel keeping in Kenya, Range Management Handbook of KenyaEnglish Press Limited, Nairobi. 1994
Farah, ZakariaCamel milk: Properties and productsSKAT-Verlag, St. Gallen, Switzerland. 1996
Yagil, R.Camels and camel milkAnimal Production and Health Paper, No. 26, FAO, Rome, pp. 1982
Wangoh, J.Chemical and Technological Properties of Camel MilkDiss. ETH Nr. 12295, Zurich 1997
Kappeler, St.Compositional and structural analysis of camel milk proteins with emphasis on protective proteins. Diss. ETH No. 12947, Zurich, Switzerland. 1998
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J. Dairy Sci. 86:498508 American Dairy Science Association, 2003.
5-Flanking Regions of Camel Milk Genes Are Highly Similar to Homologue Regions of Other Species and Can be Divided into Two Distinct Groups
S. R. Kappeler, Z. Farah, and Z. Puhan
Laboratory of Dairy Science, Institute of Food Science, Swiss Federal Institute of Technology, CH-8092 Zurich, Switzerland
ABSTRACT The concentrations of individual casein and whey proteins in camel milk differ markedly to respective protein concentrations in bovine milk. The ratio of casein to -casein is considerably higher in camel milk. -Lactoglobulin is absent, but whey acidic protein and peptidoglycan recognition protein have been detected. Genomic sequences upstream to milk-protein genes, which are known to regulate the expression of milk proteins to a great extent, were determined for 10 camel milk-protein genes and compared to respective sequences in other mammals. Multiple sequence alignment showed closest relationships to homologous sequences from other mammals. Comparison of milk protein regulative regions revealed two distantly related groups with pronouncedly different transcription factor site probabilities. The GC-content in sequences of the first group was considerably higher than in sequences of the second group and combined occurrence of CAAT and TATAA boxes was rare, suggesting that the first group represented mostly the housekeeping gene type, probably regulated by cellular signal transduction pathways, whereas the second group helped to regulate genes specifically expressed in terminally differentiated cells of the lactating alveolar epithelium. A core region of the composite response element, which primarily controls milk protein gene activity, was found by a search for elements conserved within all 5-flanking sequences analyzed, and it is assumed, that the presence of this element determines gene expression in the lactating mammary gland, and binding sites for general activator and repressor factors, surrounding the milk protein gene specific element, are important for regulation of gene activity. (Key words: camel, gene expression, milk protein, transcription factor binding site)
Abbreviation key: 5-flanking region = gene sequence 5-flanking to the transcriptional start site of the milk-protein genes examined, TF = transcription factor. Abbreviations for transcription factors follow the TRANSFAC entries (Wingender et al., 2000). INTRODUCTION Milk proteins may basically be divided into watersoluble proteins, which often have a function in protecting the lactating udder and the newborn against environmental stress, and into amphipatic proteins, which help to preserve the suspension- and emulsionlike character of milk. The quantitative distribution of individual proteins greatly differs between species, and the distribution in camel milk is unique in particular. Some important proteins of bovine milk, such as -lactoglobulin and lysozyme C, are not found in camel milk, whereas other proteins are present, such as the whey acidic protein and the peptidoglycan recognition protein, which were not detected in bovine milk. These proteins most probably function as protective or immunomodulating factors, and some of them are also found as a part of the body immune system. The different distribution of these factors in bovine and camel milk is probably a result of the harsh conditions in the natural habitat of the camels. The immunological situation, which faces the lactating camel and its offspring, requires modifications in the response to environmental stimuli. Our interest in this study was to understand the factors that regulate the expression of the genes that correspond to milk proteins and lead to the observed variation in the distribution of these proteins between species. In particular, we intended to find out whether the mechanisms described to regulate the milk protein gene expression in other species also apply to the homologous camel genes. During the past decade, a new level in understanding of the processes that regulate the tissue-specific expression of milk-protein genes has arisen, with molecular and cellular investigations in the regulation of the mammary gland during pregnancy, lactation,
Received January 31, 2002. Accepted March 26, 2002. Corresponding author: S. R. Kappeler; e-mail: stefan.kappeler@ alumni.ethz.ch.
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and involution. Expression of milk-specific genes in the lobuloalveolar epithelium of the lactating mammary gland was reported to be regulated by hormonal and environmental stimuli, which are transmitted through a set of transcription factors that bind to enhancer elements located on the proximal or distal 5flanking region to the transcriptional start site (Rosen et al., 1998). The minimal requirements to elicit a sufficient lactogenic response include prolactin, glucocorticoids, and insulin (Nagaiah et al., 1981), which may be substituted by the insulin-like growth factor. Progesterone, which is involved in mammogenesis, was shown to repress casein gene expression during pregnancy through a DNA-binding factor of 65 kDa (Lee and Oka, 1992). Epidermal growth factor (Teng, 1999), which is by itself secreted into milk and stimulates the gastrointestinal development of the newborn (Brown et al., 1989), also contributes to mammogenesis (Gallego et al., 2000), counteracting transforming growth factor . On the morphological level, it was found that signals from the basement membrane cooperate with hormonal factors mentioned to maintain the lobuloalveolar structure and the basal-apical polarization of the epithelial cells, which is required for expression and secretion of milk proteins (Aggeler et al., 1991; Close et al., 1997). In particular, the basement membrane protein laminin was shown to be required for activation of the prolactin-receptor by prolactin, acting via membrane anchored 1-integrins (Edwards et al., 1998), as well as factors from the extracellular membrane suppressing histone deacetylases (Rosen et al., 1999). In studies, where stage- or tissue-specific levels of milk proteins and corresponding mRNA were compared, good correlations were found, indicating that quantitative control of expression occurs on the transcriptional level (McClenaghan et al., 1995). Our finding that the protein composition of camel milk is markedly different from the composition in bovine milk over the course of the lactational cycle, prompted us to investigate whether this differential expression pattern can be followed back to differences between the 5-flanking regions of camel and bovine milk-protein genes. The data gained allowed us to start a comparative statistical analysis of putative transacting factor binding sites in 5-flanking regions of homologous milk genes from different species, and to examine, which of the proposed regulative elements is likely involved in the regulation of the examined camel milk-protein genes.
MATERIALS AND METHODS Camel Milk Collection and Quantitative Analysis of Milk Proteins Milk from different breeds and individuals was frozen at -20°C for transport and stored at -70°C until analysis. Caseins and whey proteins were separated, identified and characterized as described (Kappeler et al., 1999b, 1999a). DNA Sequence Analysis Coding sequences for camel milk proteins were determined by using a cDNA library constructed from a Somali breed camel, as described in Kappeler et al (1998). Genomic DNA was extracted from white blood cells of an Arabian camel by conventional phenol-ethanol purification. An average length of about 40 kDa was found after 0.5%-agarose gel separation. Five Genome Walker libraries were created as recommended by the manufacturer (K1807-1; Clontech, Palo Alto, CA). Sequence specific primers were designed, based on information from exon I and intron I regions sequenced previously. PCR-amplification products were sequenced on an ABI Prism 310 Genetic Analyzer using BigDye chemistry. The 5 flanking sequences were entered in the EMBL/GenBank database under the accession numbers AJ409277 (S1-CN), AJ409278 (S2CN), AJ409279 (-CN), AJ409280 (-CN), AJ409281 (-LA), AJ409282 (lactoferrin), AJ409283 (lactophorin), AJ409284 (lactoperoxidase), AJ409286 (peptidoglycan recognition protein), and AJ409285 (whey acidic protein). Computational Sequence Analysis The GCG version 10 software package (Genetics Computer Group, Madison, WI) was used to create a tree illustrating the relatedness of the 5-upstream sequences examined. First, 50 proximal, 5-flanking regions (1120 bp) of milk-protein genes were aligned with the pileup function, using gap weight 1 and gap length weight 0. The penalties for gap creation and extension were set to a low value, since sequences 5upstream to vertebrate genes usually exhibit a high variability in regard to insertion and deletion of DNA fragments. A Tamura distance-corrected matrix of the aligned sequences was created with the distances function. Third, a phylogenetic tree was created out of the matrix according to the unweighted pair group method using arithmetic averages. Additional sequences used for the analyses (see Figure 1) included DNA regions from the following EMBL/GenBank acJournal of Dairy Science Vol. 86, No. 2, 2003
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KAPPELER ET AL.
Figure 1. Phylogenetic tree of 5-flanking regions adjacent to the transcriptional start site of milk-protein genes. Length of the branches corresponds to the number of substitutions. The distance between human mucin 1 and rat gamma-casein was 54.70 substitutions per 100 base pairs.
Journal of Dairy Science Vol. 86, No. 2, 2003
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cession numbers. The information in brackets indicates the corresponding protein and the number of base pairs in front of the transcriptional start site included into analysis. AC005962 (human lactoperoxidase, 2000), AC007785 (human peptidoglycan recognition protein, 2000), AC063956 (human alpha s2-casein, 2000), AF027807 (human beta-casein, 1194), AF044256 (porcine lactoferrin, 1366), AF072711 (human carboxyester lipase, 2000), AF107201 (equine beta-lactoglobulin, 2000), AL121936 (human butyrophilin 1A, 2000), D16108 (murine lactophorin (GlyCAM-1, PP3, 2000), D85424 (human alpha s1-casein, 1180), E12614 (porcine beta-casein, 2000), L11749 (murine mammary tumor virus, LTR, 1361), M10936 (rat beta-casein, 783), M55158 (bovine beta-casein, 1722), M61170 (human mucin 1, gene product of the human polymorphic epithelial mucin gene (PEM), 2000), M74778 (murine lactoferrin, 2000), M75887 (bovine kappa-casein, 2000), M90645 (bovine alphalactalbumin, 1951), M94327 (bovine alpha s2 casein type A (CASAS2), 1509), U02884 (murine fatty acid binding protein, 1515), U16175 (murine mucin 1, 2000), U25810 (bovine lysozyme C, 2000), U28757 (porcine lysozyme C, 2000), U38816 (murine whey acidic protein, 2000), U57623 (human fatty acid binding protein, 1247), U67065 (murine butyrophilin 1A, 2000), U69534 (murine epidermal growth factor, 2000), X01153 (rat whey acidic protein, 1190), X03584 (rat alpha s1-casein, 682), X03589 (rat gamma-casein, 679), X12817 (ovine beta-lactoglobulin, 801), X13484 (murine beta-casein, 2000), X15735 (rabbit beta-casein, 2000), X59856 (bovine alpha s1-casein, 2000), X83391 (bovine lactophorin (GlyCAM-1, PP3), 1014), X98558 (porcine fatty acid binding protein, 1607), Y00726 (guinea pig alpha-lactalbumin, 1195), Y12088 (murine peptidoglycan recognition protein, 1532), Z33882 (caprine kappa-casein, 2000), Z48305 (bovine beta-lactoglobulin, 2000). Repetitive elements were detected in the examined 5-sequences using RepeatMasker version 2, based on the Repbase database (Smit and Green, 2000). Potential binding sites for transcription factors, socalled TF-sites, were detected in the 50 5-flanking regions examined, according to the method of (Schug and Overton, 1997). The TRANSFAC (transcription factor) database (Wingender et al., 2000) version 3.3 was used and the following parameters applied: no allowable mismatch, a minimum element length of six bases and a minimum lg-likelihood of 6. Where possible, the 2000-bp upstream to the transcriptional start site were examined, to be able also to detect more distantly located binding sites. The reliability of TFsites found was 31%, calculated as follows: (nP - nR)/nP with nP as the number of sites found per
base pair of 5-flanking sequence (the total number of base pairs analyzed was 78,598), and nR as the number of sites found per base pair in 200 kbp of randomly generated DNA sequence. Because factors with a large number of TRANSFAC-compiled binding sites were overrepresented by these string-based searches, redundancies were filtered out. The frequencies of occurrence within 1000 bp of the 5-flanking regions were calculated for all TF sites. The same was done with randomly generated DNA sequences. The values obtained from the randomly generated sequences, considered as a background resulting from weakly defined binding sites, were subtracted from the former. The resulting binding site profiles contained information about the grinding potential of known transcription factors to the analyzed promoter regions, under relinquishment, of positional information. A correlation factor for every pair of promoter regions was calculated as the sum of products of the number of background-subtracted, nonredundant binding sites of known transcription factors, according to the formula (nxA-nxR)(nxB-nxR) for x = 1 to z, where A represents a 5-sequence, B another 5-sequence, R a random sequence, n the nonredundant number of binding sites for a certain transcription factor and z the total number of transcription factors compiled in the TRANSFAC library (Wingender et al., 2000). A search for conserved motifs within the 5-upstream sequences was done using MEME (multiple expectation maximization for motif elicitation) Version 3.0 with analysis of both strands and a limiting range for width variability between 6 and 50 bp, expecting at least one occurrence of every motif in each sequence (Bailey and Gribskov, 1998). A set of the following 26 5-upstream sequences was chosen, of which the gene products are abundant in milk: Bos taurus: S1-CN, S2-CN, -CN, -CN, -LA, -LG, lactoferrin, PP3 component (lactophorin). Camelus dromedaries: S1-CN, S2-CN, -CN, -CN, -LA, lactoferrin, PP3 component (lactophorin), peptidoglycan recognition protein, whey acidic protein. Capra hircus: -CN. Equs caballus: -LG. Homo sapiens: S1-CN, S2-CN, -CN. Ovies aries -LG. Sus scrofa: -CN, fatty acid binding protein, lactoferrin. Where possible, the 2000 bp upstream to the transcriptional start site were examined, to be able also to detect more distantly located binding sites. Rodent sequences were not included, due to the more distant evolutionary relationship, and presumably modified TF binding site preferences. In the following, each motif resulting from MEME analysis was searched for TF binding sites, mainly by TESS analysis (Schug and Overton, 1997).
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Table 1. Average amount [mg 1-1] of some casein and whey proteins in mature milk from different species. nd = not detected. indicates a downregulation of gene expression between colostral and mid-lactational milk. indicates an upregulation in case of mastitis. Data from (Aguirre et al., 1998; Cals et al., 1994; Cuilliere et al., 1997; Hennighausen et al., 1994; Kappeler et al., 1999b; Ragona et al., 2000). Milk protein S1-Casein S2-Casein -Casein -Casein -Lactalbumin -Lactoglobulin Whey acidic protein Lactophorin (PP3 component) Lactoferrin Lactoperoxidase Peptidoglycan recognition protein Lysozyme C nd nd Camel 5000 2200 15,000 800 3500 nd 157 950 95 107 nd 100 Cow 12,000 3000 10,000 3500 1260 3500 nd 300 140 30 nd 274 Human minute minute 4670 minute 3400 nd nd nd 565 6 Rodents 1600 nd 4500 nd nd nd 1500 nd nd 465 nd nd Function Formation of casein micelle Formation of casein micelle Formation of casein micelle Formation and rennet coagulation of casein micelles Regulatory subunit of lactose synthetase Binding of fatty acids and retinol Probably an epithelial growth regulator, similar to WDNM1 Lipolysis inhibition Anti-inflammational, nutritive, iron uptake, regulative Anti-inflammational, bacteriolytic activity Anti-inflammational Bacteriolytic activity, N-acetylmuramidase
RESULTS The protein composition of camel and cow milk differed in regard to both casein and whey proteins. Camel milk contained significantly higher concentrations of -CN and lower amounts of -CN (Table 1). -LG, lysozyme C and lactoperoxidase were not detected in mid- to late-lactational camel milk, and the corresponding cDNA of the former two proteins was not found by screening of a lactating mammary gland cDNA library. On the other hand, whey acidic protein, generally described as a major constituent of rodent milk, and peptidoglycan recognition protein, an intracellular protein binding to gram-positive bacteria, and presently not known to be a milk constituent, were detected in major amounts in camel whey, both on the cDNA and protein level. Lactophorin, the proteose peptone 3 component of whey, was detected in higher concentrations than in bovine milk (Kappeler et al., 1999b). A comparative analysis was started for elements regulating genes, which express camel milk proteins. Fewer than or equal to 2 kb of the regions upstream to the transcriptional start sites of 10 camel genes, for which we found the corresponding mRNA in the lactating udder, were sequenced, and compared to homologous regions from other species. Percent identity to homologous bovine sequences were between 56 and 74%, to rodent sequences between 45 and 61%, and to human sequences between 50 and 76%. The overall GC-content of the camel sequences analyzed was about 44%, similar to homologous sequences from other species. Multiple alignment between the 5-flanking regions of milk-protein genes from several species revealed
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two groups of distantly related sequences (Figure 1). The nucleic acid composition of the two groups, thereafter designated as group I and group II, differed markedly, with group I having an average GC content of 54%, and group II having an average GC content of 38%. Interspersed elements and long terminal repeats covered about 16.5% of group I sequences, and 20.5% of group II sequences. Correlation analysis of transcription-factor binding site profiles of the different 5-upstream sequences produced a relational matrix with similar relationships between binding site preferences as the relationships found with multiple sequence alignment (Figure 2). The most abundant potential binding sites found in the 5-flanking regions examined were those for the glucocorticoid receptor, transcription factors of the ets proto-oncogene family, especially MAF (mammary activating factor, predominantly in group I sequences) and PEA3, furthermore those for NF-1 (nuclear factor 1), YY1 (yin yang 1), and the progesterone receptor (Table 2). Sites for TBP/TFIID, proteins of the octamer binding family, among them Pit-1a (a pituitary gland tissue-specific activator), GATA-binding proteins, F2F (repressor of the prolactin promoter), C/EBP (CCAAT enhancer binding protein ) and, to a minor extent, C/EBP were predominantly found in group II promoter sequences, whereas binding sites for the ubiquitous activators Sp1, AP-2 and PU.1, as well as MAF and PPAR (peroxisome proliferator-activated receptor) were mostly found in the GC-rich promoter sequences of group I. MEME analysis of the examined promoter regions revealed six motifs, which occurred in all promoter sequences at least once (Figure 3). A positional over-
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Figure 2. Simplified correlation matrix of 5-flanking regions 2.0 kbp adjacent to the transcriptional start site of milk-protein genes. The average number of binding sites between the examined homologous promoter regions from different species was used for the calculation of the correlation strength between non-homologous promoter regions, as described in Materials and Methods. Strong correlation (Sum of products > 200) is indicated as "++", values > 100 with "+", negative values with "-".
view of these motifs on the sequences is presented in Figure 4. They were in the following compared to the consensus sequences of known TF binding sites, especially to those reported to be involved in the regulation of milk protein genes. The first and the fifth motif contained elements exclusively reported for the regulation of milk protein genes. The mammary activating factor was reported as a regulative factor in the mouse mammary tumor virus long terminal repeat (Reuss and Coffin, 2000), and the milk protein binding factor was described in the regulation of the sheep -LG gene (Watson et al., 1991). The second motif found by MEME analysis corresponded to the core of a composite response element, which was described to be a main regulator of milk protein gene activity (Rosen et al., 1999). The motif contained the STAT5 consensus sequence and putative binding sites for C/EBP and the yin yang 1 repressor, as well as a glucocorticoid receptor half site. The third motif corresponded to an RNA polymerase II promoter sequence. This motif was not clearly noticeable in front of the transcriptional start sites of group I sequences (Figure 1). The fourth motif contained a binding site for the estrogen receptor
and the sixth motif was detected in repeated arrangements in group I sequences. The detection of glucocorticoid receptor half sites (Lechner et al., 1997) was difficult by the method chosen, probably due to the variability is glucocorticoid response elements. NF-1 sites were not found within the detected motifs, most likely because of differences in the local arrangements of this ubiquitous factor in composite response elements of group 1 and group II 5-upstream sequences. DISCUSSION The protein composition of camel and cow milk differs in some fundamental aspects. We assume that the immunological challenges, which face these two ruminating species in their respective natural environment, has led to adaptations in the milk composition. We were not able to isolate -LG, the primary constituent of bovine whey, from camel milk, nor lysozyme C, and did not detect the respective sequences in a lactating mammary gland cDNA library by plaque screening or the PCR method. We also were not able to isolate lactoperoxidase from midlactational camel
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Figure 3. Conserved motifs, i.e., multilevel consensus sequences, detected by MEME analysis of 26 5-flanking regions to genes abundantly expressed in the lactating mammary gland and sorted according to the output file. Transcription factor binding sites were attributed to the motifs, which have previously been reported to be involved in the regulation of gene expression in the course of lactation.
milk, although this enzyme can be isolated from bovine milk throughout lactation. Nevertheless, we were able to isolate and sequence a corresponding clone from a cDNA library produced from udder tissue four
Table 2. Most prominent TF sites with possible involvement in the regulation of milk protein genes after subtraction of background and sorting out of redundancies. The numbers represent the percentage of a particular site, as compared to the total number of sites detected. Factor Group I Group II All sequences
ETS (MAF; PEA3) MAF GR GR half sites TFIID TBP C/EBPbeta C/EBPdelta NF-1 YY1 PU.1 AP-2 Oct Sp1 PPAR Pit-1a PR F2F GATA C/EBPalpha MPBF NF-kappaB Pit-1 COUP STAT5
5.49 1.00 6.89 0.27 0.94 1.38 1.39 0.37 2.38 0.91 1.97 12.44 0.00 12.54 2.07 0.00 0.88 0.26 0.00 0.30 0.61 0.61 0.00 0.26 0.02
(% of sites detected) 5.55 5.97 0.11 0.43 5.47 6.40 1.66 1.31 7.87 6.10 6.41 5.18 5.15 4.26 3.72 2.86 2.04 2.32 2.58 2.21 0.85 1.31 0.10 4.39 8.24 5.68 0.00 1.83 0.30 0.94 5.55 3.99 1.84 1.65 3.24 2.46 2.88 1.33 1.49 1.19 0.00 0.21 0.00 0.20 0.48 0.30 0.07 0.14 0.06 0.03
weeks after parturition. This enzyme is probably rapidly downregulated in the camel mammary gland, as it was also observed in human tissue. On the other hand, high concentrations of lactophorin (the PP3 component), whey acidic protein and the peptidoglycan recognition protein were found in camel whey. Furthermore, we found a considerably higher ratio of -CN to -CN in camel milk from different breeds that reported for cow milk (Table 1). In the course of this study, we sequenced the proximal (2 kbp) regions 5-upstream to the transcriptional start of 10 camel milk protein genes, to find a rationale to the different expression pattern of milk protein genes in the lactating mammary gland of camels and cows. We supposed these regions to be sufficient to direct the stage- and tissue-specific expression of the respective genes, as it was shown for corresponding sequences in other mammals (Faerman et al., 1995; Lee et al., 1998). Alignment of the 50 5-upstream sequences analyzed (Figure 1) showed in a first step, that all 5flanking regions of camel milk-protein genes were most closely related to their homologous counterparts from other species, independently from the gene expression level in the mammary gland of the other species (Figure 1). Interestingly, the camel 5-upstream sequences also showed the observed relatedness in those cases, where the respective genes were reported to be expressed to a very different level in the lactating mammary glands of the different species, e.g., in the case of camel and bovine -CN (Kappeler et al., 1998). This sequence relatedness gave indication that the
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Figure 4. A summary of the motifs from Figure 3 showing an optimized (nonoverlapping) tiling of all of the motif occurrences with a positional P-value (the probability of a single random subsequence of the length of the motif scoring at least as well as the observed match) 0.0001 for each of the sequences in the training set. The product of motif probabilities on a sequence is given as "Expect" value. Motif occurrences are indicated by numbers, motif lengths on a sequence as empty spaces, and the motif direction with plus or minus.
regulation of gene expression has to be restricted to small conserved areas within the 5-upstream sequences, and that mutations in these areas will initiate, abolish, or modulate the tissue- and stage-specific expression of milk protein genes. Additionally, two very distantly related groups of 5flanking regions were discerned, designated as group I and group II (Figure 1). Group I sequences enclosed some of the whey and the milk-fat globule membrane protein gene sequences, group II sequences encompassed casein and some whey and butyrophilin gene sequences. The higher GC-content of group I sequences, and the scarcity in combined CAAT and TATAA boxes towards the transcriptional start sites let us assume that group I 5-flanking regions predominantly were regulating housekeeping genes, which became highly expressed in the lactating mammary gland of some species, whereas group II genes were specifically expressed in the end-differentiated cells of the lobuloalveolar epithelium. Our interest at this point was to see if binding site probabilities for the different transcription factors were similar for the 50 sequences examined. A position
independent pattern search strategy was developed, as described in Materials and Methods, because similar regulative elements, especially with regard to hormone responsive elements, have been localized on different positions relative to the transcriptional start site (Rosen et al., 1998). A correlation matrix was generated out of the background corrected binding site probabilities. A simplified view of this matrix is shown in Figure 2 combining the results of homologous sequences. The resulting matrix showed the same division into two weakly related groups as the multiple sequence alignment before, with the exception of binding site probabilities for wap 5-flanking sequences, which also exhibited good TF-site correlation results with some group II sequences. Figure 2 shows, nonetheless, that the majority of the different 5-flanking regions weakly correlate with each other, indicating that most sequences contain some common regulative elements. Some TF-sites were detected on nearly all sequences with a background-corrected frequency of more than one per 1000 bp. Most abundantly, binding sites for the glucocorticoid receptor and for transcription factors of the Ets
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family were found on sequences of both groups (Table 2). Ets factors, especially MAF and PEA3, were localized on regulative sequences of milk-protein genes and probably mediate signals of the epidermal growth factor and of insulin to the prolactin gene (Jacob et al., 1999). Glucocorticoid receptor sites in the first group were often mere half sites, as described to exist particularly on milk-protein gene promoter regions (Lechner et al., 1997), whereas sites in the second group preferably consisted of near-palindromic sequences. Binding sites for the peroxisome proliferator activated receptor (PPAR) were detected on most group I sequences. This fatty-acid activated nuclear receptor controls genes involved in the lipid metabolism and is enhanced by prolactin during adipogenic conversion of cells (Nanbu et al., 2000). The 2-isoform was reported to be downregulated (Gimble et al., 1998) or upregulated (Jain et al., 1998) during pregnancy and lactation, and a role in response to physiologic and pathologic stimuli, which alter lipid metabolism, was suggested. This indicates that the factor may help to regulate group I gene expression in differentiating and involuting mammary tissue. The ubiquitous transcription factors of the octamer binding family were reported to bind to elements in the proximal 5 sequence of casein genes (Groenen et al., 1992) and in the long terminal repeat of the mouse mammary tumor virus (Brueggemeier et al., 1991), enhancing the activity of the mediators of hormonal and local signals. Octamer binding sites were abundantly detected in all group II sequences, but only weakly in group I sequences. There are a number of observations on the histological level that genes belonging to either of the two groups show strikingly different stage- and cell-specific expression patterns. A mutually exclusive histological localization of lactoferrin mRNA on one hand, -LA and S1-CN mRNA on the other hand was reported in the alveolar epithelium (Molenaar et al., 1992), depending on the lactational status of the cells. Whereas -LA and S1-CN mRNA have been detected in terminally differentiated, lactating alveoli, lactoferrin mRNA was almost exclusively found in emerging and regressing alveoli. Lactoferrin expression also did not require basement membrane signals and its expression was even repressed in epithelial cells with basal-apical orientation, but stimulated in nonpolarized cells (Close et al., 1997). This finding was in contrast to the expression of genes that belong to typical milk proteins, such as -LA, caseins and also the whey acidic protein, although we found the 5-flanking sequences of wap genes to belong to group I (Figure 1). Fatty acid binding protein, also known as mammaryderived growth inhibitor, was localized in the vacuolar, nonlactating cell type (Erdmann and Breter,
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1993), whereas butyrophilin mRNA was detected in lactating cells (Molenaar et al., 1995). Most genes of the second group are solely expressed in alveolar epithelial cells of the late-pregnant and lactating mammary gland, whereas the first group, with the exception of the whey acidic protein, was made up of genes, which are known to be expressed in a broader range of tissues. Fatty acid binding protein, for example, participates in the intracellular transport of fatty acids in different tissues, and peptidoglycan recognition protein is a protein of the innate immune system of vertebrates and invertebrates. Additionally, many proteins of the first group are likely involved in feedback regulation of protein and fat secretion into the alveoli and thus need to be expressed in a manner different to caseins and related proteins, to execute these functions. Altogether, a majority of the proteins, which help to preserve the structural properties of milk, seem to be expressed in terminally differentiated cells under control of a TATA-like promoter. Proteins, which protect against environmental stress, on the other hand, rather seem to be constitutively expressed and merely upregulated in the lactating mammary gland. Established models describing the regulation of milk-protein gene transcriptional control are mainly based on studies with genes for rodent whey acidic protein and rodent and bovine -CN. Additionally, the long terminal repeat of the murine mammary tumor virus, which directs expression to lactating mammary epithelial cells, is well investigated. Information about the regulation of other genes expressing milk proteins is rare. DNase I hypersensitivity and footprinting experiments, as well as electrophoresis mobility shift assays and transgenic studies, helped to identify binding sites for transcription factors and hormone responsive elements (Rosen et al., 1999). A crucial role in the regulation of milk genes plays a pathway, by which the prolactin signal is effected via phosphorylation of the membrane-bound prolactin-receptor, inducing tyrosine phosphorylation of STAT5 A and B isoforms by Janus kinase, dimerization, nuclear translocation, and DNA binding (Rosen et al., 1999). Short forms of the prolactin-receptor and of STAT5 have been reported, which were found in nonlactating mammary tissue and act as dominant negative regulators, repressing expression of target genes (Edwards et al., 1998). STAT5 binding sites were shown to reside within a composite response element, which also included sites for NF-1 and GR in the rodent whey acidic 5-flanking region, and for GR, C/EBP and isoforms, and YY1 in the bovine -CN 5-flanking region (Doppler et al., 1995). The core region of this element was
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retrieved as the second motif from a MEME analysis of 26 regions 5-upstream to genes highly expressed in the lactating mammary gland of different species (Figure 3). The motif was lost when sequences were included in the MEME training set, which are not highly expressed in the lactating mammary gland, supporting the idea of a crucial role for this response element in the control of milk protein gene expression. We conclude by comparing our results from TF-site analysis and from MEME detection of conserved elements, that genes lacking the composite response element, which combines the STAT5 binding site with TF-sites for other activating and repressing factors, will not be expressed in the lactating mammary gland of a particular species. However we did not succeed to associate the variances in milk protein levels between species to a structural disparity in the corresponding 5-flanking regions. We presume that the fine-tuning of milk protein gene expression will probably reside in the arrangement of binding sites for ubiquitous factors, such as NF-1 or the Octamer family, nearby the conserved elements. Additionally, superior regulative regions and mRNA stability are likely involved in the control of milk protein gene expression. ACKNOWLEDGMENTS The authors thank the laboratory group of the central veterinary laboratory of Dubai for providing genomic camel DNA. REFERENCES
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KAPPELER ET AL. mammary gland cells share functional elements. J. Virol. 74:81838187. Rosen, J. M., C. Zahnow, A. Kazansky, and B. Raught. 1998. Composite response elements mediate hormonal and developmental regulation of milk protein gene expression. Biochem. Soc. Symp. 63:101113. Rosen, J. M., S. L. Wyszomierski, and D. Hadsell. 1999. Regulation of milk protein gene expression. Annu. Rev. Nutr. 19:407436. Schug, J., and G. C. Overton. 1997. Subject: TESS (Transcription Element Search Software on the WWW). http://www.cbil. upenn.edu/tess. Accessed Dec. 28, 2001. Smit, A. F. A., and P. Green. 2000. Subject: RepeatMasker. http:// ftp.genome.washington.edu/RM/RepeatMasker.html. Accessed Dec. 28, 2001. Teng, C. T. 1999. Regulation of lactoferrin gene expression by estrogen and epidermal growth factor: molecular mechanism. Cell Biochem. Biophys. 31:4964. Watson, C. J., K. E. Gordon, M. Robertson, and A. J. Clark. 1991. Interaction of DNA-binding proteins with a milk protein gene promoter in vitro: identification of a mammary gland-specific factor. Nucleic Acids Res. 19:66036610. Wingender, E., X. Chen, R. Hehl, H. Karas, I. Liebich, V. Matys, T. Meinhardt, M. Pruess, I. Reuter, and F. Schacherer. 2000. TRANSFAC: An integrated system for gene expression regulation. Nucleic Acids Res. 28:316319.
Molenaar, A. J., S. R. Davis, L.-J. W. Jack, and R. J. Wilkins. 1995. Expression of the butyrophilin gene, a milk fat globule membrane protein, is associated with the expression of the alpha-S1 casein gene. Histochem. J. 27:388394. Molenaar, A. J., S. R. Davis, and R. J. Wilkins. 1992. Expression of alpha-lactalbumin, alpha-S1-casein, and lactoferrin genes is heterogeneous in sheep and cattle mammary tissue. J. Histochem. Cytochem. 40:611618. Nagaiah, K., F. F. Bolander, Jr., K. R. Nicholas. T. Takemoto, and Y. J. Topper. 1981. Prolactin-induced accumulation of casein messenger RNA in mouse mammary explants: A selective role of glucocorticoid. Biochem. Biophys. Res. Commun. 98:380387. Nanbu, W. R., Y. Fujitani, Y. Mashuho, M. Muramatu, and H. Wakao. 2000. Prolactin enhances CCAAT enhancer-binding protein-beta (C/EBP beta) and peroxisome proliferator-activated receptor gamma (PPAR gamma) messenger RNA expression and stimulates adipogenic conversion of NIH-3T3 cells. Mol. Endocrinol. 14:307316. Ragona, L., F. Fogolari, L. Zetta, D. M. Perez, P. Puyol, K. De Kruif, F. Loehr, H. Rueterjans, and H. Molinari. 2000. Bovine betalactoglobulin: Interaction studies with palmitic acid. Prot. Sci. 9:13471356. Reuss, F. U., and J. M. Coffin. 2000. The mouse mammary tumor virus transcription enhancers for hematopoietic progenitor and
Journal of Dairy Science Vol. 86, No. 2, 2003
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Milk and Meat from the Camel
Handbook on Products and Processing The book is based on results of research work carried out in partnership with universities, camel farmers and pastoralists in Eastern African countries. The first part deals with chemical composition, technological properties, hygienic safetyandqualitycontrolofcamelmilk.Thisisfollowedbythepresentationofprocessing methods for different dairy products which should allow smallholder producers to process milk on farm or in centralised small scale dairy unit. In the second part special knowledge gained in slaughtering, deboning and processing of camels on-site over several years of long-term stays on the Ol Maisor Farm in Kenya is presented. During the development of camel meat products the authors set a great value on the keeping quality of the products as well as creating a wide range of product groups with low and high cost for the local market and a good shelf life. The book presents background information and recipes on the manufacture of camel milk and meat products and it is intended as a practical guide to professionals, government bodies and development agencies interested in building up small scale processing units for camel meat and milk products. It is hoped that it may also serve as a reference for extension personal working with camel herders, food scientists and students. For further information please contact the author: zakaria.farah@ilw.agrl.ethz.ch published: February 2004 232 p., 17 x 24 cm, hardcover, many tables, recipes and 4-coloured figures US/Europe: CHF 47./EUR 32. (D)/USD 33.50, plus freight and taxes Rest of the world: USD 25., plus freight and taxes ISBN 3 7281 2527 X
Zakaria Farah, Albert Fischer (eds.)
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TABLE OF CONTENTS Dedication Acknowledgement Preface An introduction to the camel Part A: Camel as a milk animal 1. Milk 1.1 Milk production 1.2 Milk composition 2. Milk products 2.1 Introduction 2.2 Cheese 2.3 Fermented milk 2.4 Butter 3. Methods for quality control 3.1 Introduction 3.2 Determination of milk freshness and hygienic quality 3.3 Milk adulteration 4. Milk Hygiene and Udder Health 4.1 Milk Hygiene 4.2 Udder Health 5. Equipment for small scale milk plants 5.1 Introduction 5.2 Milk collection 5.3 Small-scale milk processing systems 5.4 Equipment requirement and specification Part B: Camel as a meat animal 6. Traditional slaughter, carcass dressing and processing of camels 6.1 Locations for slaughter of camels
6.2 The traditional method of slaughter for camels 6.3 Traditional dressing of the carcass 6.4 Traditional meat products in Africa ands Asia
7. Method for hygienic slaughter of camels 7.1 Requirementstobesatisfiedbyslaughterhouses 7.2 Slaughter procedure 7.3 The dressing percentage of camels 7.4 By-products from slaughter of camels
8.Dressing of the camel carcass 8.1 Requirements to be satisfied by deboning rooms and equipment 8.2 Cutting of the camel into primal cuts 8.3 Deboning of the camel 8.4 Cutting of the camel into retail cuts 8.5 Standardization of processing material 8.6 Percentageofretailcuts,processingmaterial and remaining parts 8.7 Camel meat composition
9.Meat products from camel meat 9.1 Product groups 9.2 Equipment needed for meat products 9.3 Additives and ingredients for meat products 9.4 Casings for meat products 9.5 Recipes for camel meat products
Recommended references About the Authors
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Journal of Dairy Research (1990), 57, 281-283 Printed in Great Britain
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SHORT COMMUNICATION Preparation and consumer acceptability tests of fermented camel milk in Kenya
By ZAKARIA FARAH, THOMAS STREIFF AND MARC R. BACHMANN Laboratory of Dairy
Science, Swiss Federal Institute of Technology, ETH-Zentrum, CH-8092 Zürich, Switzerland
(Received 3 April 1989 and accepted for publication 16 October 1989)
There are estimated to be 600000 camels (Camelus dromedarius) in Kenya (Wandera, 1985). Almost 80% of these are kept by pastoral tribes living in arid areas in eastern and north-eastern parts of the country. In these regions, camels are important dairy animals. A camel in north-east Kenya can be expected to yield about 4 kg milk daily as compared with 0.5-1.5 kg for a cow in the same area. Most of the camel milk is consumed in the form of fermented milk. The milk is allowed to ferment naturally at ambient temperature and without prior heat treatment until it turns sour. The resulting fermented camel milk is known as Susa. Due to the spontaneous nature of the fermentation, this traditional method results in a product with varying taste and flavour and is often of poor hygienic quality. In addition, because of the limited scale of production, the product can be sold only in the immediate vicinity of the herd. For production of fermented milk under controlled conditions, thermophilic or mesophilic lactic acid cultures are normally used. In warm countries, mesophilic lactic cultured milk offers some advantages, as it can be incubated at ambient temperature (20-30 °C) and the fermentation stops at 1-1.2 % lactic acid, eliminating the need for cooling to stop further souring as occurs in the case of yoghurt (Kurwijila, 1980). Considering these advantages, Kurwijila (1980) developed, in Kenya, fermented cows' milk using mesophilic lactic cultures. Tests conducted with adult Kenyans showed high consumer acceptability. The present investigation was undertaken in Garissa, a north-eastern province town of Kenya, where a majority of the population subsists almost entirely on camel milk. The objective of this study was to develop fermented camel milk using mesophilic lactic culture and to test the acceptability of the product in comparison with the traditional fermented camel milk.
EXPERIMENTAL Milk samples Fresh camel milk was obtained from herds owned by nomads around the town of Garissa.
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Z. FARAH AND OTHERS
Cultures Multiple strain, mixed type, lyophilized, mesophilic lactic cultures, O-CH:143 (homofermentative) and B-CH:40 (heterofermentative) were obtained from Chr. Hansen's Laboratorium, Denmark. Preparation of fermented milk Two 10-litre churns of fresh whole camel milk were placed in a container filled with water and heated until the milk attained 85°C. This temperature was maintained for 30 min. After cooling in a water bath to ambient temperature, each milk churn was inoculated with 2 % of a 24 h culture and incubated at a room temperature of 27°C for 24 h. Chemical analysis For each milk the following parameters were determined; acidity by titration expressed in Soxhlet-Henkel degrees (° SH), fat by the Gerber method, total solids by calculation after Fleischmann (1896) from the values of fat content, and specific gravity using a lactometer. The same determinations were also made with the traditionally fermented camel milk, Susa, from the local market. Sensory evaluation In the sensory tests, the two fermented camel milk samples were compared with a traditionally fermented camel milk, Susa. Two groups of people were selected for the sensory evaluation. Group A consisted of 13 Somali nomads with no formal education. All claimed to consume Susa regularly. Group B consisted of nine Somalis and three Canadians. They all worked as senior officers in the Provincial and District Administrations. Seven of the group reported consuming Susa regularly and five only occasionally. As the panellists had no previous experience of testing products, the rating test was simplified and limited in respect of consumer preference. Each person was asked to taste the three coded samples and score each product for preference on a threepoint scale ranging from' most preferred' (preference score = 1) to 'least preferred' (preference score = 3). The instructions were given orally in both the Somali and English languages. The panellists were told that they were testing camel milk, but the identities of the individual samples were revealed only after the tests were completed.
RESULTS AND DISCUSSION Compared with cows' milk, the consistency of fermented camel milk is thin. After fermentation a precipitate in the form of flocs was formed rather than a coagulum. In a preliminary experiment an attempt was made to improve the consistency by the addition of cows' milk powder. However, this had to be omitted in the final experiments, as sensory evaluation and conversation with consumers of camel milk revealed that mixing camel with cows' milk affected the typical camel milk taste and was undesired. Results of chemical and sensory analysis arc presented in Table 1. The values of total solids and fat content were the same in all the samples. Homofermentative culture O-CH:143 showed less titratable acidity. Fermented milk made with mesophilic lactic cultures was clearly preferred by both groups. Within the two
Fermented camel milk
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Table 1. Test panel preference scores of three fermented samples of camel milk
Fermented samples of camel milk Mesophilic lactic cultures Sensory evaluation Group A, n = 13 Mean score s.d. Group B, n = 12 Mean score s.d. Chemical analysis Total solids, % Fat content, % Acidity °SH O-CH:143 B-CH:40 Susa
2.08 0.76 1.58 0.67 12.7 4.1 36.2
1.39 0.77 1.50 0.52 12.7 4.0 40.0
2.31 0.86 2·92 0.29 12.5 4.0 40.0
starter cultures used, the homofermentative culture O-CH:143 was least preferred. Both groups described the two prepared cultured milks as particularly good Susa with uniform fresh taste. The study shows that the traditional Susa can be improved by using selected mesophilic lactic acid culture. Seasonal variations in camel milk production are great in north-eastern Kenya, and' much surplus milk is wasted during the rainy season (R. Muriuki, pers. comm.). The method described here for fermented milk production can be introduced in rural areas. It allows small-holder producers to process surplus milk on-farm or in centralized small-scale units. Obtaining starter culture could be a limiting factor for large-scale production of fermented milk. However, simple commercial systems for producing frozen starter cultures which maintain their activity for years are in operation in Kenya (Kurwijila, 1983; Schulthess, 1988). We thank the staff of Garissa Community Service Centre in Kenya for providing us with all the facilities during our field work. REFERENCES
FLEISCHMANN, W. 1896 The Book of the Dairy, pp. 71-72. KURWIJILA, R. L. N. 1980 Low cost optimization of the flavour, consistency and keeping quality of fermented milk with particular reference to consumer acceptability in Kenya. M.Sc. Thesis, University of Nairobi KURWIJILA, R. L. N. 1983 Maintenance of reactivated, lyophilised mixed type lactic cultures by subculturing and by cold storage. Indian Journal of Dairy Science 36 338-343 SCHULTHESS, W. 1988 [Appropriate milk product development-an example in Kenya.] LebensmittelTechnologie 21 226-228 W ANDERA, J. G. 1985 Camel pastoralism in Kenya. In Significance and Prospects of Camel Pastoralism in Kenya pp. 61-62 (Ed. S. E. Migot-Adholla), Nairobi, Kenya: Institute for Development Studies
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International Dairy Journal 9 (1999) 481}486
Sequence analysis of camel (Camelus dromedarius) lactoferrin
Stefan R. Kappeler*, Manfred Ackermann, Zakaria Farah, Zdenko Puhan
Laboratory of Dairy Science, Institute of Food Science, Swiss Federal Institute of Technology, CH-8092 Zurich, Switzerland Received 25 January 1999; accepted 5 May 1999
Abstract The aim of this study was to characterise camel lactoferrin in terms of primary structure and molecular weight. The protein was eluted from a heparin-sepharose column at a sodium chloride concentration of 0.5 M, and corresponded to bovine lactoferrin in terms of N-terminal sequence and the molecular weight of 80.16}80.73 kDa. Lactoferrin cDNA was PCR ampli"ed, using a cDNA library from lactating mammary gland of a Somali camel. The sequenced clone had a length of 2337 bp and an open reading frame of 2124 bp, which coded for a protein of 708 amino acid residues. The mature protein had a length of 689 amino acid residues, a calculated molecular weight of 75.250 kDa and a calculated isoelectric point at pH 8.14. Primary structure identity to bovine lactoferrin was 74.9%. Concentration of lactoferrin in whole, late-lactational milk was 220 mg l\, which was higher than the lactoferrin concentration in comparable bovine milk, which was 140 mg l\. 1999 Elsevier Science Ltd. All rights reserved.
Keywords: Camelus dromedarius; Lactoferrin
1. Introduction Lactoferrin is a protein of the innate immune system, which is also expressed in the lactating mammary gland. It is found in milk and di!erent other body secretions, and in neutrophil leukocytes (Masson, 1970). The concentration in milk strongly depends on the species and the stage of lactation. Industrial scale puri"cation from whey is carried out by cation exchange, and use as a preserving agent in food, drugs and cosmetics has been proposed (Saito, Takase, Tamura, Shimamura & Tomita, 1994). Lactoferrin belongs to the family of transferrins, together with blood serotransferrin (siderophilin), egg white ovotransferrin (conalbumin), melanotransferrin of malignant melanoms, the porcine inhibitor of carbonic anhydrase, and other proteins. The common property of this protein family is the binding of two metal cations, preferably Fe>, at structurally closely related binding sites. Most proteins of the transferrin type are needed for storage or transport of iron. It can be assumed, that lactoferrin in colostral milk acts as an iron scavenger, which depletes the milk from free iron and thereby slows down microbial growth.
* Corresponding author.
Brock (1997) proposed, that the in vivo function of apolactoferrin is the prevention of iron-mediated lipid peroxidation, a property, which was already demonstrated with monocytes. This function is based on the ability of lactoferrin, to bind to cell membranes. The higher a$nity for iron, as compared to other transferrins, would enable it to function at the reduced pH found in the stomach and upper intestine. The high resistance of apolactoferrin to proteolysis, compared with other apotransferrins, would enable it to maintain its iron-binding potential in the face of proteolytic activity in the gut. Since diferric lactoferrin was reported to be even more resistant to proteolysis, it was supposed, that the iron}lactoferrin complex would resist degradation, and was sequestered by hepatocytes, or was excreted from the gut. A higher lactoferrin concentration also could help to prevent lipid peroxidation by free radicals in an infected udder, which has an elevated iron content. Iron-saturated lactoferrin, which is found in milk from the second week to the end of the lactational period, may primarily prevent microbial growth in the gut. This would help the new-born, which is easily infected, to survive the "rst weeks, until its own immune system becomes developed, and the gut becomes adapted to food digestion. Iron-saturated lactoferrin could also be a source of iron for the suckling, once the protein is degraded in the gut.
0958-6946/99/$ - see front matter 1999 Elsevier Science Ltd. All rights reserved. PII: S 0 9 5 8 - 6 9 4 6 ( 9 9 ) 0 0 1 1 7 - X
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Camel milk was frequently reported to have a high antimicrobial activity, and was shown to slow down the growth of pathogenic bacteria more than bovine milk (Elagamy, Ruppanner, Ismail, Champagne & Assaf, 1992). Antimicrobial properties were partially attributed to well characterised proteins, such as lactoferrin, lactoperoxidase, lysozyme and immunoglobulin A. These proteins were shown to have higher concentrations or higher activity in camel milk, as compared to bovine milk. In this study we determined the basic physico-chemical and structural parameters of lactoferrin, of which the relationship between structure and function is well characterised in human and bovine counterparts.
Molecular masses of proteins were measured by matrix assisted laser desorption/ionization mass spectrometry (Kappeler et al., 1998). 2.3. Quantixcation Lactoferrin eluted from the Heparin-Sepharose column was collected, tenfold diluted in distilled water and the absorbency measured at 280 nm. For calculation of the protein concentration, an extinction coe$cient of 84540 M\ cm\ was used for camel lactoferrin and of 102890 M\ cm\ for bovine lactoferrin (Gill & Von Hippel, 1989). One litre whole milk was estimated to yield 0.8 l whey. 2.4. cDNA Sequence analysis PolyA-mRNA isolation and construction of a cDNA library was done as described in Kappeler et al. (1998). Overlapping fragments of the lactoferrin cDNA, which together covered the complete sequence, were produced by the polymerase chain reaction (PCR). The following protocol was applied to most of the reactions: 2 l of the -cDNA library were taken as templates in 50 l PCR assays with 2.5 units Taq Polymerase (Amersham Pharmacia), which was blended with 0.05 units Pfu Polymerase (Stratagene, La Jolla, CA), and 5 l 10; TaqPlus Precision incubation bu!er (Stratagene), 20 nmol of each dNTP (Amersham Pharmacia) and 50 pmol of speci"c primers. A series of 30 cycles was run with initial 2 min denaturation at 943C, followed by 10 s denaturation at 943C, 30 s annealing at 553C and 2 min 30 s elongation at 683C. Elongation prolongation was 20 s per cycle. A "nal 10 min incubation step at 723C was added to increase the concentration of full-length products. Each PCR product was generated twice and ligated into a pGEM -T Easy vector (Promega, Madison, WI) according to the manufacturer's instructions. In case of base reading ambiguities, a third PCR product was generated. Two -gt11 vector speci"c general primers were constructed to cover the 5- and 3-ends of lactoferrin cDNA: -gt11 forward: 5-GACGACTCCTGGAGCCCGTCAGTA-3, -gt11 reverse: 5-CACCAGACCAACTGGTAATGGTAG-3. The following PCR products were generated, mostly with the help of highly conserved regions in the cDNA sequences of other species (mixed base sites according to IUB code): A 0.4 kbp PCR product was generated with 5-CTGTCCCATAGACCTCTGCCGCTA-3, and gt11 reverse.
2. Materials and methods 2.1. Isolation of lactoferrin from whey Pooled milk of Arabian camels was stored at !703C until analysis. After thawing, the milk, which had a pH of about 6.6, was skimmed at 1000 g, 43C for 15 min. Whey was obtained by acid precipitation of casein at pH 4.6 and 373C for 20 min, using 0.1% acetic acid, followed by addition of 10 mM sodium acetate for neutralisation, and centrifugation at 4000 g for 5 min. The supernatant was dialysed twice against double distilled water for 5 h at 43C, and once against 10 mM sodium phosphate bu!er at pH 7.4 for 14 h at 43C, using an autoclaved SPECTRA/POR membrane tubing with a molecular cuto! of 6}8 kDa (Spectrum Medical Industries, Inc., Los Angeles, CA). Prior to chromatography, samples were "ltered through a hydrophilic 0.45 m membrane (ME25; Schleicher and Schuell, Dassel, Germany). A Heparin-Sepharose HiTrap column (1 ml; Amersham Pharmacia, Uppsala, Sweden) was loaded with 40 ml whey. The column was washed with 10 ml PBS (10 mM sodium phosphate, 20 mM sodium chloride, pH 7.4). Elution was performed at ambient temperature by a linear gradient from 0.02 to 1 M sodium chloride over 40 min. The column e%uent was monitored with an UV detector (L-7300; Merck, Darmstadt, Germany) at 280 nm. Proteins eluted were collected manually and lyophilised. Fractions were further puri"ed, prior to micro-sequencing and molecular mass determination, by reversed-phase C HPLC (Kappeler, Farah & Puhan, 1998). Elution was performed by a linear gradient from 0.1% TFA in double distilled, nano"ltered water, to 0.1% TFA in acetonitrile, over 60 min. 2.2. Physico-chemical characterisation Proteins collected from the e%uent of the C -column were used directly for N-terminal sequencing (Kappeler et al., 1998).
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483
A 0.8 kbp PCR product was generated with 5-GTTCRRTGGTGTRCCRTMTCCMMA-3, and 5-GTCTTTGAACAGCAGGTCCTTCTG-3. A 1 kbp PCR product was generated with 5-TTCCAGCTCTTTGGCTCYCC-3, and 5-TTGAACAGAAGGTTTTTGGT-3. A 0.4 kbp PCR product was generated with 5-CCAGGCAAGTTTTGCTTGTTCCAG-3, and gt11 reverse. The ligation products were dialysed and transformed into E. coli XL1-Blue (Stratagene) by electroporation with a Gene-Pulser (BioRad, Hercules, CA) at 2.5 kV, 25 FD, and 200 in 0.2 cm cuvettes. The transformed bacteria were plated overnight at 373C on IPTG/XGal/Ampicillin-selective agar. White colonies were picked and grown overnight in 20 ml LB-Ampicillin 100 (Maniatis, Sambrook & Fritsch, 1989). Plasmid DNA was puri"ed for #uorescent sequencing with the Wizard Plus SV Minipreps DNA Puri"cation System (Promega). Fluorescent sequencing was carried out using an ALF automated device (Amersham Pharmacia) with standard operating procedures. Sequencing samples were prepared, using the Cy52+-dATP labelled, vector speci"c primers: Cy5-SP6: 5-TACTCAAGCTATGCATCCAACGCG-3, and Cy5-T7: 5-ACTCACTATAGGGCGAATTGGGCC-3 and the Thermo Sequenase cycle sequencing kit RPN 2438 (Amersham Pharmacia) according to the manufacturer's instructions. The following 25 cycles were applied: 953C, 30 s, 503C, 30 s, 723C, 50 s. Where sequencing results di!ered, a third PCR product was sequenced. Overlapping sequences were detected using the FASTA module of the gcg/egcg programme package (Genetics Computer Group, Madison, WI). Consecutive sequences were joined and vector speci"c sequences removed. In this way, complete cDNA sequences were obtained. 2.5. Computational sequence analysis Alignments of DNA and protein sequences and DNA similarity searches were performed using the gcg/egcg programme package (Genetics Computer Group). Protein sequence similarity searches against the Swissprot database (Swiss Institute of Bioinformatics, Geneva, Switzerland) were made using a Smith and Waterman algorithm with default values (Barton, 1997). A low-resolution model of the tertiary structure of camel milk lactoferrin was obtained by comparative modelling (Guex & Peitsch, 1997). The primary structure,
which was revealed by amino acid and cDNA sequencing, was threaded over resolved tertiary structures of di!eric lactoferrins from other species. Multiple sequence alignments were made for improvement of modelling reliability. Energy minimisation of the model was done with force "eld computation by GROMOS96. 3. Results 3.1. Primary structure PCR ampli"cation products of a full-length cDNA clone of camel lactoferrin were sequenced (EMBL/GenBank2+ accession number AJ131674). The clone was 2337 bp long, and contained a 5-untranslated region of 22 bp and a 3-untranslated region of 191 bp. The 5untranslated region contained a partial Kozak-box (Kozak, 1989) in front of the translational start site ATG, with a purine at !3 bp, and cytosines at !1 bp, !5 bp and !8 bp. The 3-untranslated region contained a polyadenylation signal AATAAA. The open reading frame ranged from A to G, and coded for a polypeptide of 707 aa residues. The start site of the mature protein was determined by similarity as Ala. The 19 aa signal peptide was to 94.7, 84.2 and 78.9% identical to the respective signal sequences of bovine, porcine and human lactoferrin. The mature protein was 689 aa residues long, and had a molecular weight of 75.250 kDa (Table 1), without postranslational modi"cations. The isoelectric point of the unmodi"ed peptide was at pH 8.14. The protein shared 74.9% sequence identity with bovine, 74.5% with porcine, and 74.0% with human lactoferrin. The pronounced homology gave indication for a nearly identical biological function of the proteins. 3.2. Glycosylation N-linked glycans contribute about 4}11% (3}9 kDa) to the total mass of bovine lactoferrin, which is about
Table 1 Physicochemical characteristics of camel and bovine lactoferrin Camel Amino acid residues Molecular mass (kDa) based on mass spectrometry Molecular mass (kDa) based on amino acid sequence Isoelectric point Concentration in milk (mg l\) Sequence identity 689 80.16}80.73 75.250 8.14 220 74.9% Bovine 689 84.0 76.143 8.18 140
Data on bovine lactoferrin after Schanbacher, Goodman and Talhouk (1993), Yip and Hutchens (1997). Calculated with the gcg programme (Genetics Computer Group, Madison, WI 53711, USA).
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84.0 kDa (Spik, Coddeville, Mazurier, Bourne, Cambillaut & Montreuil, 1994); Yip & Hutchens, 1997). Glycosylation enhances the solubility of the secreted protein and may help to bind at speci"c cell types, such as liver parenchymal cells (Ziere, Bijsterbosch & Van Berkel, 1993). Camel milk lactoferrin was found to contain 6.2% carbohydrates in colostral milk and 5.6% in milk collected 15}30 days after parturition (Mahfouz, ElSayed, Abd El-Gawad, El-Etriby & Abd El-Salam, 1997). The content of N-acetyl-glucosamine in camel milk lactoferrin was markedly higher than in ruminants' milk lactoferrins (3.35% in colostral camel milk compared to about 1.75% in colostral ruminants' milk, Mahfouz et al., 1997). In our study, the carbohydrate content of lactoferrin from end-lactational milk was 6.2}6.8% of total protein mass, calculated as a di!erence between the protein mass measured by MALDI-MS and the protein mass of the amino acid sequence (Table 1). Possible glycosylation sites, based on pattern analysis (Gavel & Von Heijne, 1990), are Asn, Asn, Asn and Asn. In bovine lactoferrin, four of "ve sites with N-glycosylation potential, Asn, Asn, Asn, and Asn, are glycosylated (Spik et al., 1994), and contribute to an overall carbohydrate content of 11.2%. Degree of glycosylation in human lactoferrin is about 6.40% (Spik et al., 1994), and thus similar to camel lactoferrin. Human lactoferrin contains 2 glycosylated sites, Asn and Asn, with glycans of the N-acetyllactosaminic type, which were also found in camel lactoferrin. By comparison with bovine and human lactoferrin, glycosylation of two of the four possible sites of camel lactoferrin is proposed (Fig. 1). 3.3. Concentration in camel milk Colostral camel milk was reported to have an extremely high lactoferrin content of 5.10 g l\ on the second day after parturition, compared to about 0.50 g l\ in bovine colostral milk. After 30 days of milking, the lactoferrin level in camel milk went down to 0.34 g l\, whereas in bovine milk, only about 0.06 g l\ were found (Abd El-Gawad, El-Sayed, Mahfouz & Abd El-Salam, 1996). In our studies, we used an extinction coe$cient of 84540 M\ cm\ at 280 nm to calculate a lactoferrin concentration of 0.22 g l\ in a milk sample, which was taken at the end of the lactation period, 360 days after parturition. In a sample of pooled cow milk, lactoferrin concentration was 0.14 g l\. If it is assumed, that the main function of lactoferrin in milk is the inhibition of bacterial growth, a di!erently composed micro#ora in the gut of the new-born could be a reason for the apparently higher lactoferrin concentration in camel milk. Nevertheless, it has to be taken into account, that the concentration of protective proteins in milk also depends on the milk yield, which was about 5 l d\ for the camels, and about 15 l d\ for the cattle studied. It also has to be considered that the
immunological situation, with regard to the placenta type, the colostrum, the development and stimulation of the immune system in the calf, and the nutritional properties in general, will strongly di!er between both species, as a result of the di!erent habitats, in which the animals live, adaptation of camels to a sub-optimal food supply and quality, di!erences in the way, the o!spring is raised, and the more distant paleontological relationship. 3.4. Tertiary structure and ligand binding The polypeptide chain of transferrins consists of about 700 amino acids and is folded into two, tandemly arranged, asymmetrical metal binding sites, designated as N- and C-lobes, which probably evolved by gene duplication. The sequence of the camel lactoferrin N-lobe, which extended from Val to Arg, shared 39.8% sequence identity with the sequence of the C-lobe, which ranged from Val to Arg. Under physiological conditions, transferrins bind one Fe> cation in each lobe with a low dissociation constant of about 10\ (Brock, 1997). Cation binding requires synergistic binding of a bicarbonate anion, probably for charge compensation. In bovine and probably also in camel lactoferrin, the side chains of Asp, Tyr, Tyr and His are involved in binding of the cation in the N-lobe (Baker et al., 1998; Fig. 1). Two oxygens from the bidentate CO\ anion are suggested to complete a dis torted octahedral geometry (Anderson, Baker, Norris, Rice & Baker, 1989). In the N-lobe, the side chains of Thr, Arg, and Tyr, and two backbone hydrogens of Ala and Gly, are involved in binding of the bicarbonate anion. Lactoferrin retains its iron binding potential at pH values below pH 5.5, and even in the presence of citrate, in contrast to the other known transferrins (Brock, 1997). The primary structure and a modelled tertiary structure of the binding sites of bovine and camel lactoferrin were found to be nearly identical. We therefore assume that cations are bound by both lobes of the camel protein with similar a$nities as in bovine lactoferrin. 3.5. Bacteriostatic activity of the N-terminal end A high amount of Arg and Lys are clustered at the N-terminal end of lactoferrin, near and between a loop, which is formed by disulphide bonding of Cys and Cys (Fig. 1). The N-terminus of human and bovine lactoferrin was found to have strong bacteriostatic activity on gram-negative bacteria, as a result of non-speci"c binding to the negatively charged outer bacterial membrane, and subsequent release of lipopolysaccharides, thereby altering the permeability properties (Ellison, Giehl & LaForce, 1988). The N-terminal part of camel lactoferrin contained 13 basic residues (Fig. 1), at sites more similar to bovine lactoferrin than to human
S.R. Kappeler et al. / International Dairy Journal 9 (1999) 481}486
485
Fig. 1. Schematic drawing of the relationship between structure and function in camel and bovine lactoferrin. Signal sequences in italics. Functional residues in bold. -Helical regions designated as & ', -pleated regions as & '. E indicate probably glycosylated asparagine residues (in bold).
lactoferrin. The isoelectric point of the N-terminal fragment from Ala to Ala was at pH 10.98, compared to pH 11.57 for the bovine fragment at the same position, which had one more basic residue. As in bovine lactoferrin, most residues were found in or near the loop. Based
on the high homology between camel and bovine lactoferrin in general, and particularly on the similar grouping of the N-terminal basic residues, we suggest a similar e!ect of camel lactoferrin as observed of bovine lactoferrin. The bacteriostatic activity of camel lactoferrin on
486
S.R. Kappeler et al. / International Dairy Journal 9 (1999) 481}486 Barton, G. J. (1997). SCANPS Version 2.3.1, User Guide. UK: University of Oxford. Brock, J. H. (1997). Lactoferrin Structure*Function Relationships. In T. W. Hutchens, & B. Lonnerdal, Lactoferrin: Interactions and K Biological Functions (pp. 3}23). Totowa, NJ: Humana Press. Elagamy, E. I., Ruppanner, R., Ismail, A., Champagne, C. P., & Assaf, R. (1992). Antibacterial and antiviral activity of camel milk protective proteins. Journal of Dairy Research, 59, 169}175. Ellison, R. T., Giehl, T. J., & LaForce, F. M. (1988). Damage of the outer membrane of enteric gram-negative bacteria by lactoferrin and transferrin. Infection and Immunity, 56, 2774}2781. Gavel, Y., & von Heijne, G. (1990). Sequence di!erences between glycosylated and non-glycosylated Asn-X-Thr/Ser acceptor sites: implications for protein engineering. Protein Engineering, 3, 433}442. Gill, S. C., & Von Hippel, P. H. (1989). Calculation of protein extinction coe$cients from amino acid sequence data. Analytical Biochemistry, 182, 319}326. Guex, N., & Peitsch, M. C. (1997). SWISS-MODEL and the SwissPdbViewer: an environment for comparative protein modelling. Electrophoresis, 18, 2714}2723. Kappeler, S., Farah, Z., & Puhan, Z. (1998). Sequence analysis of Camelus dromedarius milk caseins. Journal of Dairy Research, 65, 209}222. Kozak, M. (1989). The scanning model for translation: an update. Journal of Cell Biology, 108, 229}241. Mahfouz, M. B., El-Sayed, E. M., Abd El-Gawad, I. A., El-Etriby, H., & Abd El-Salam, A. M. (1997). Structural studies on colostrum and milk lactoferrins from di!erent species. Egyptian Journal of Dairy Science, 25, 41}53. Maniatis, T., Sambrook, J., & Fritsch, E. F. (1989). Molecular Cloning (2nd ed.). New York: Cold Spring Harbor Laboratory Press. Masson, P. L. (1970). La lactoferrine, proteine des secretions externes et des leucocytes neutrophiles. Brussels: Editions Arsca S.A. Saito, H., Takase, M., Tamura, Y., Shimamura, S., & Tomita, M. (1994). Physicochemical and antibacterial properties of lactoferrin and its hydrolysate produced by heat treatment at acidic pH. In T. W. Hutchens, S. V. Rumball, & B. Lonnerdal, Lactoferrin Structure and K Function (pp. 21}31). New York: Plenum Press. Schanbacher, F. L., Goodman, R. E., & Talhouk, R. S. (1993). Bovine mammary lactoferrin: implications from messenger ribonucleic acid (mRNA) sequence and regulation contrary to other milk proteins. Journal of Dairy Science, 76, 3812}3831. Spik, G., Coddeville, B., Mazurier, J., Bourne, Y., Cambillaut, C., & Montreuil, J. (1994). Primary and three-dimensional structure of lactotransferrin (lactoferrin) glycans. In T. W. Hutchens, S. V. Rumball, & B. Lonnerdal, Lactoferrin Structure and Function (pp. 21}31). K New York: Plenum Press. Yip, T. T., & Hutchens, T. W. (1997). A$nity Mass Spectrometry. In T. W. Hutchens, & B. Lonnerdal, Lactoferrin: Interactions and BioloK gical Functions (pp. 39}58). Totowa, NJ: Humana Press. Ziere, G. J., Bijsterbosch, M. K., & Van Berkel, T. J. (1993). Removal of 14 N-terminal amino acids of lactoferrin enhances its a$nity for parenchymal liver cells and potentiates the inhibition of beta-very low density lipoprotein binding. Journal of Biological Chemistry, 268, 27069}27075.
di!erent bacterial strains was found to have equal strength as the activity of bovine lactoferrin (Elagamy et al., 1992). 3.6. Food preservation Lactoferrin was discussed to be a promising choice for preservation in food and cosmetics, since it is highly stable towards heat treatment and at low pH conditions (Saito et al., 1994). It helps to establish a favourable micro#ora, promotes growth of bi"dobacteria, and may therefore "nd attraction for use in functional food products. The antimicrobial peptides formed upon gastric digestion of lactoferrin are also promising candidates as additives for food preservation. Primary structures of peptides formed from camel lactoferrin should be studied and activity of such peptides on inhibition of bacterial growth tested, to get better understanding of the action of lactoferrin in camel milk. The higher amounts of lactoferrin in camel milk are of advantage for natural preservation of the milk in arid regions, where technology for milk preservation is often not available.
Acknowledgements The authors thank Mr. and Mrs. Breitling of the Kamelfarm Fatamorgana in Rotfelden, Germany, for providing milk samples, and the laboratory group of Prof. M. Teuber for giving the opportunity to do some molecular-biological work in their laboratory.
References
Abd El-Gawad, I. A., El-Sayed, E. M., Mahfouz, M. B., & Abd ElSalam, A. M. (1996). Changes of lactoferrin concentration in colostrum and milk from di!erent species. Egyptian Journal of Dairy Science, 24, 297}308. Anderson, B. F., Baker, H. M., Norris, G. E., Rice, D. W., & Baker, E. N. (1989). Structure of human lactoferrin: crystallographic structure analysis and re"nement at 2.8 A resolution. Journal of Molecular s Biology, 209, 711}734. Baker, E. N., Anderson, B. F., Baker, H. M., MacGillivray, R. T., Moore, S. A., Peterson, N. A., Shewry, S. C., & Tweedie, J. W. (1998). Three-dimensional structure of lactoferrin. Implications for function, including comparisons with transferrin. In G. Spik, D. Legrand, J. Mazurier, A. Pierce, & J.-P. Perraudin, Advances in Lactoferrin Research (pp. 1}14). New York: Plenum Press.
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15
An introduction to the camel
Z. Farah
Present distribution and economic potential
According to FAO statistics there are about 19 million camels in the world, of which 15 million are found in Africa and 4 million in Asia. Of this estimated world population, 17 million are believed to be one-humped dromedary camels (Camelus dromedarius) and 2 millions two-humped (Camelus bactrianus). Approximately 11 million dromedaries, representing two thirds of the world's camel population, are in the arid areas of Africa, particularly in North East Africa, i.e. Somalia, Sudan, Ethiopia and Kenya.
Table 1: Estimated camel populations of Africa and the world (FAOSTAT Database 2001)
Country Africa: Algeria Chad Djibouti Egypt Ethiopia Kenya Libya Mali Mauritania Other regions: China India Iraq 326 1030 76 Mongolia Pakistan Saudi Arabia 360 800 400 240 725 70 120 1070 830 72 467 1230 Morocco Niger Nigeria Senegal Somalia Sudan Tunisia West Sahara 36 415 18 4 6200 3200 231 105 Camel population (in 1000) Country Camel population (in 1000)
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An introduction to the camel
Most of these animals are kept by pastoralists in subsistence production systems. They are very reliable milk producers during dry seasons and drought years when milk from cattle sheep and goats is scarce. At such times camel can contribute up to 50% of the nutrient intake of the pastoralists. In recent years the picture of "moving" nomads has changed to some extent. With growing urbanisation the demand for milk among the urban population has been increasing. On the other hand the demand for a number of goods such as grain, oil, sugar and clothes increased among the pastoralists and the milk sales became the most important part of cash income for many camel owning pastoralists. Camel meat is also an important by-product mainly as a source of income. Sale of live camels, usually males and unproductive females for slaughter, is very common in East Africa and there are now increasing numbers of camel butcheries in many urban centres. There is also a growing export trade of slaughter camels to the Arabian Peninsula. The camel is also a means for transportation and for domestic use as drawing water from wells, rivers and dams. From a global perspective, the economic significance of camel production is minimal in comparison with that of other domestic animals. Nevertheless, in Africa, especially in East Africa and Sahel countries, the camel population makes a significant contribution to national economies. However, it is difficult to evaluate this economic contribution as most of the camel products are traded in the informal sector. Owing to the increasing human population and declining per capita production of food in Africa, there is an urgent need to develop marginal resources, such as arid land, and optimise their utilisation through appropriate livestock production systems of which camel production is the most suitable without doubt.
Traditional husbandry and management
Camels are held by nomads in arid regions. The pastoral land is mainly covered with annual grass, acacias, euphorbias and dwarf bushes. The annual rain fall varies between 100 and 400 mm, the amount of rain varying from year to year and the rains being restricted to widely separated areas. This type of pasture permits only extensive types of animal production. Because of its high mobility, its modest fodder requirement and its water regulation perfectly adapted to the environment, the camel is better suited than any other domestic animal to use this type of pasture. According to the nomads, camels can survive in times of extreme need for up to 30 days without water. This depends, however, on the grazing and prevailing temperatures.
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17
The salt requirement of camels is very high and is six to eight times higher than that of other domestic animals. The salt requirement is only partly satisfied by grazing. When the herdsmen observe that the camels are restless, with reduced appetite and milking performance, they take this as a sure sign of salt deficiency. The camels are then driven to salty water sources and watered repeatedly. Alternatively, salt-containing earth collected from other areas is given to the animals. Studies of many nomadic people in several countries show that female animals constitute 70% to 80% of a camel herd. The high number of female animals is needed to satisfy the large milk requirement of the nomad economy. The number of male animals in the herd is reduced in two ways and at two points in the camel's life cycle. A percentage of male calves are slaughtered at birth or within few weeks of it. This allows more milk for female calves and for family consumption or sales. Males not slaughtered at birth are allowed to grow until they are about 4 years old. At this age the majority are castrated, fattened, sold and slaughtered for meat. Reproduction of the herds is achieved by selection of suitable male camels. According to the traditional husbandry, these should have the following characteristics: · The bull or its father should have had predominately female progeny with good milk performance; · It should be fully grown and strong; · It should be a good fighter able to overcome other males. It would be difficult to evaluate to what extent these selection criteria influence the quality of the progeny. One restriction arises from the fact that only the characteristics of the father, and not the characteristics inherited from mother, are taken into account in the selection. In general, breeds of camels are not as differentiated and classified as breeds of other domestic species. In most camel rearing societies, breed classifications are based on names of the ethnic group, clan as well as on the geographical localities where these camels are raised, rather than upon phenotypical characteristics. In Kenya for example there are three main types of camel classified as Somali breed, Rendille/Gabbra breed and Turkana breed. The Somali breed camels are primarily owned by Somali people of North-Eastern province of Kenya and are generally much larger than the other breeds found in the country. Adult females average 500600 kg and males 600800 kg. Average milk daily yield is 5 kg to 8 kg during a lactation of 10 to 12 months. The Rendille/Gabbra breed is found mainly in Marsabit district amongst the
18
An introduction to the camel
Rendille and Gabbra tribes. It is generally smaller than Somali breed camels. Live weights average 350450 kg and 400500 kg for females and males respectively. Milk yields average 3 to 4 kg, over a lactation of 12 months. The Turkana breed is the smallest camel found in Kenya averaging 350 kg for females and 400450 kg for males. Milk yields are much lower than from other breeds and are in average 2 kg to 3 kg per day over a lactation of 9 to 10 months. The small body size and small feet make Turkana camels very swift and able to climb steep lava hills.
Fig. 1: Somali breed bull
In recent years dromedaries from Pakistan have been introduced in some camel farms in Kenya in order to improve milk production through cross breeding. Camels are slow reproducers. A female camel is sexually mature at the age of 45 years. Pregnancy is just over 12 months and the calving interval in pastoral production systems is normally 24 months or more. Female camels can remain fertile up to the age of 25 years and it is often reported that they produce 810 calves during a lifetime. In pastoral production systems, however, only a small proportion of the breeding females can reach this production performance. A major problem in camel productivity is the high mortality rate of camel calves in the first 3 months. The causes for mortality are mainly poor management practice and infectious diseases. The new-born calf has no natural protection against diseases, as there is no antibody transfer from the mother during foetal development. The calf can
Z. Farah
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obtain immediate immunisation soon after birth only through the colostrum, which has a very high concentration of antibodies. Therefore, it is vital for the calf to suckle as soon and as much as possible. Unfortunately there is a common belief among many pastoralists that colostrum causes diarrhoea and, consequently, is unsuitable for the new-born calf. This wide spread practice of withholding the colostrum from the new-born calves, depriving them of essential antibodies, is certainly a crucial factor in the frequently reported high calf mortality in pastoral production systems. The milking of camels is a process that varies according to the different pastoral groups. Camels may be milked once or several times a day. In general, it is normal practice among most nomadic tribes to milk their camels in the early morning before animals are taken to grazing and at night when they return from grazing. Before milking, the calf is allowed to suckle until the milk is flowing and then the camel can be milked. Without this stimulation, the dam cannot be milked. The milker stands on one leg, puts the milk pot on the upper part of the other leg, and milks with one or two hands. Sometimes, milking may be done by two persons, each milking two teats. To prevent calves from suckling while at pasture, it is a usual practice to tie up one or more teats with special strings.
Fig. 2: Somali breed female with calf
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An introduction to the camel
Fig. 3: Milking of camels
Fig. 4: Adult female Somali breed. (Piers Simpkin)
Z. Farah
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Fig. 5: Adult female Turkana breed. (Piers Simpkin)
Fig. 6: Adult female Somali/Turkana crossbreed. (Piers Simpkin)
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An introduction to the camel
Fig. 7: Adult female Pakistan breed. (Piers Simpkin)
Fig. 8: Pregnant Somali heifer (L) and pregnant mature Rendille/Gabbra female (R). (Chris Field)
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Farah, Camel milk butter
Manufacture and characterization of camel milk butter
By Z. FARAH, T. STREIFF and M.R. BACHMANN Laboratory of Dairy Science, SWISS Federal Institute of Technology, Zurich. Switzerland 1. Introduction Camels (Camelus dromedarius) are Important as daily animals in certain regions of the world. Most of the camel milk is drunk fresh or when it has just turned sour. Among many camel rearing societies, there is a common belief that butter cannot be made from camel milk. This belief has been supported by some authors (1) while others (2, 3) claim that butter can be made from camel milk. The present study was undertaken in order to gain a clear picture of this rather contradictory situation. Experiments on butter from camel milk were made in rural areas in North Eastern Kenya, where a majority of the population subsists almost entirely on camel milk. The butter obtained, along with camel milk samples, was analyzed at the Dairy Laboratory of the Swiss Federal Institute of Technology (ETH) Zurich. 2.5 Chromatography of fatty acids Milk from 2 different camel herds was used for the analysis. For each herd 10 samples were collected from 10 Individual camels Samples were kept refrigerated and transported to our laboratory within 36 h. Upon arrival, fat of each sample was extracted by a mixture of chloroform and methanol (2:1 v/v). The solvent was removed in a stream of nitrogen and the fat stored in sealed tubes at -20°C until analyzed. Fatty acids analysis of milk was carried out as described by PRABUCKI (6). The methyl esters were prepared by the procedure of CHRISTOPHERSON (7). The analysis was performed on a Hewlett Packard 5830 gas chromatograph with a flame ionization detector. The columns (2 m x 1/8") were packed with Chromosorb W-AW, 100/120 mesh containing 3 % SP-231 0 and 2 % SP-2300. The samples were injected at an oven temperature of 75°C; then the oven temperature was raised to 220°C at a rate of 5 °C/min. The detector temperature was 250°C. 3. Results Centrifugation and churning Due to the low fat content of cream after the first pass of centrifugation, the cream had to be separated a second time. As a result the separation temperature decreased from 65°C at the first to 55°C at the second centrifugation pass. In 5 milk samples with a fat content between 2.5 and 3.8 %, the butter fat content of the skim milk obtained after the second centrifugation varied between 0.2-0.9 %. Cow milk separated in the same centrifuge Yielded skim milk with a butter fat content of 0.1 -0.2 %. After centrifugation the cream samples of varying fat content were churned immediately at different
Camel milk cream samples churned under varying cream fat content and churning temperature Cream
2. Materials and methods 2.1 Milk samples Camel milk was obtained from the following sources: (a) Camel herd of the University of Nairobi, (b) Ngare Ndare ranch, Laikipla District, Kenya, (c) individual herds owned by nomads around the town of Garissa, North Eastern Province, Kenya. 2.3 Starter culture As starter culture for sour cream freeze-dried mesophilic lactic culture O-CH-143 obtained from Chr. Hansen's Laboratory, Denmark, was used 2.3 Cream separation and churning Milk was heated to 65°C and separated with an AlfaLaval hand centrifuge, type 24 S. After the first pass, the cream was immediately separated a second time. The cream was divided into small portions of 1 I on average, and the fat content was adjusted to 20.5, 25 and 30 %. Some cream samples were inoculated with 2 % starter culture and incubated at room temperature (27°C). Churning took place in a domestic glass hand churn of a 2 I capacity. The filling level was approximately 1 I each time. The churning temperatures were varied between 15 and 36°C. The butter grains were washed twice with 500 ml drinking water at ambient temperature (27°C). The butter samples were frozen for further analysis. 2.4 Butter analysis Cream and butter fat contents were determined by the Gerber method (4). The following physical and chemical constants of the butter were determined according to the Swiss Manual for Food Analysis (5): Melting point (capillary method), refractive index (Zeiss Butyro-Refractometer), iodine value, saponification value, Reichert-Meissl value and Polenske value.
Table 1:
Sample No
fat
%
Churning temperature °C
Churning time
Butler fat yield
%
mln 13 18 16 11 10 35 38 6 50 10 5 5
Sweet cream:
1 2 3 4 5 6 7 8
9 Sour cream:
20.0 22.5 22.5 22.5 25.0 25.0 25.0 30.0 30.0 20.0 25.0 30.0
25 15 20 25 25 32 36 25 30 25 25 25
81.1 80.8 84.0 85.3 78.2 60.7 65.6 64.8 58.7 59.6 55.0 55.2
10 11 12
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churning temperatures. Results of the churning trials are shown in Table 1 and Fig. 1. Increasing of the cream fat content decreased the churning time in both sour and sweet cream (Fig. 1). At the same cream fat content sour cream yielded a lower butter fat content than sweet cream (Fig. 1). The highest fat yields In butter (8085.3%) with cream of 20-25% fat were obtained at churning temperatures between 15 and 20°C. The corresponding values for cow milk cream vary between 8-12 °C, being on average 10°C lower (8).
higher than the corresponding values in cow milk butter. Due to the high melting point, reading with the butyrorefractometer had to be made at 46°C and corrected to 40 °C. The mean value of the reading was 49.1. Compared with cow milk butter, camel milk butter was found to have low Reichert-Meissl, Polenske and saponification values, but a higher iodine value. Fatty acid analysis The molar percentages of fatty acid composition of milk fat from 2 different camel herds along with those of cow milk (10) are presented in Table 3. Each herd consisted of 10 camels. Herd A was fed exclusively by free grazing, whereas the camels of herd B grazed during the day and upon return from pastures in the evening, they were fed a supplement consisting of a mixture of maize and barley. In both groups milk samples were taken in September during the dry season. Apart from the dietary supplement to herd B, management and pasture quality was similar for both herds. Compared to cow milk fat (Table 3), camel milk fat contains less short chain fatty acids, but relatively high concentrations of C14:0 and C16:0 acids. Long chain unsaturated fatty acids occur to about the same extent as in cow milk fat. Compared to herd A camel milk fat from herd B which received supplement showed an Increased concentration in C18:0 and C18:1 and a decreased concentration in the yield of fatty acids C16:0 and C14:0, This seems to follow the effect of dietary supplements on the yields of fatty acids generally observed In bovine milk fat (11).
11 min
Churning time (min)
_ Sweet cream ~ Sour cream 6min
I
22.5
25
30
Cream fat content (%)
Fig 1 Effect 01 cream fat content on fat yield (churning temperature 25°C)
Butter constants Some chemical and physical constants of the butter obtained were determined. Mean values of 10 butter samples are presented In Table 2. Values for cow milk fat are also included for comparison (9). The mean melting point is at 41.4 °C and is on average 8°C
Table 2: Physical and chemical constants of 10 camel butter samples Sample No 1 2 3 4 5 6 7 8 9 10 Mean Cow milk butter Melting point 41.5 40.6 40.7 40.9 41.3 40.6 41.5 41.5 42.5 42.5 41.4 28-38 Refractometer readings 49.5 48.5 49.0 49.5 49.5 49.0 49.5 49.5 48.5 48.5 49.1 39-46
4. Discussion and conclusions Although the experiments were carried out in rural areas under difficult working conditions, a reasonable
Saponification value 201 201 202 200 200 200 200 200 200 200 200.4 220-233
Iodine value 51.3 51.0 50.5 47.8 47.3 47.8 48.0 50.1 47.8 48.0 48.96 25-38
Reichert-Meissl value 2.27 2.25 214 2.16 2.09 2.12 2.00 2.22 2.00 2.00 2.12 24-34
Polenske value 0.68 0.60 0.60 0.62 0.60 0.58 0.60 0.67 0.62 0.62 0.62 1.5-5
Table 3: Milk fat From Camel herd A Camel herd B Cow
Fatly acid composition (mole %) of milk fats from 2 camel herds along with cow milk fat for comparison Molar percentage mean values 01 tatty acids from 10 individual animals for each camel herd C120 C140 C141 C15:0 C15:1 C160 C161 C170 C171 C180 C181 0.26 1.03 0.80 3.50 14.20 9.70 10.00 1.29 0.80 1.29 1.98 0.50 0.26 0.20 31.50 28.50 24.60 1030 720 220 0.78 1.10 052 083 11.40 17.60 14.90 24.60 12.20 25.10
C40 1.81 1.10 8.85
C60 0.78 0.20 3.30
C8:0 0.52 1.10 2.60
C100 C10:1 1.29 0.20 4.07
C182 1.10 4.40 2.20
C183 1.29 1.38 0.70
C200 0.78 1.38 -
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Farah. Camel milk butter
efficiency of separation has been achieved. In order to find out optimum churning conditions, cream was churned at different temperatures. At low cream temperature (10 to 12°C) no butter grams could be recovered above 36 °C butter yield began to decrease. Hence, the temperature trials were made between 15 and 36°C. The highest butterfat yield was obtained at a churning temperature of 25°C from cream with 22.5 % fat content. The time taken to churn at this temperature was 11 min. The reason for this different churning behaviour of camel milk fat in comparison with cow milk fat can partly be attributed to the high melting point of camel milk fat. This seems to shift the ideal ratio of solid to liquid fat in globules at a given temperature towards a point higher than that of cow milk fat. The different churnability can also be attributed to the reported small size of camel milk fat globules (12). Small globules have a larger surface in relation to their mass which tends to increase their resistance. Compared to cow milk butter, camel milk butter is white in colour and is more sticky and greasy in consistency. The fatty acid analysis by gas chromatography confirmed the results of the butterfat constants. As indicated the analysis represents mean values of the fatty acids from 10 individual animals for each camel herd. Fatty acid composition is influenced to some degree by environmental and physiological factors such as diet, stage of lactation and genetic differences within the species. Within this limitation the general pattern of the camel milk fatty acids found in the present investigation are comparable to those of GLASS et al. (13) obtained by gas chromatography technique, but differ from the data reported by OHINGRA (14) who analyzed milk fat of not specified Indian camels using a fat fractionation method. From the present investigation it can be concluded, that butter can be prepared from camel milk. Owing to the nature of its milk fat, camel cream has different churning properties compared to cream from cow milk.
(10) FOX, PF.: Development in Dairy Chemistry - 2. Appl. Sci. Publishers, London, New York, p. 15 (1983) (11) FOX, P.F.: Development In Dairy Chemistry - 2. Appl. SCI Publishers, London, New York. p. 60 (1983) (12) Y AGIL, R.: FAO Animal Production and Health Paper, Rome 26 17 (1982) (13) GLASS, RL et al.: Camp. biochem. physiol. 22 415-425 (1967) (14) OHINGRA, DR Biochem. J. 28 73-78 (1934)
6. Summary
FARAH, Z., STREIFF, T., BACHMANN, M.R: Manufacture and characterization of camel milk butter. Milchwissenschaft 44 (7) 412-414 (1989). 44 Butter composition (camel milk) Experiments on butter manufacture from camel milk were carried out in rural areas In North Eastern Kenya. The milk was heated to 65°C and separated with an Alfa-Laval hand centrifuge. In order to find optimum churning conditions. cream of varying fat content was churned at different temperatures. Under the selected experimental conditions the highest butter fat yield (85.3 %) was found at 25°C from cream with a fat content of 22.5 %. The churning time at this temperature was 11 min. Chemical and physical constants of the butter obtained as well as the fatty acid composition of milk fat tram 2 different camel herds were also studied. Compared with cow milk butter, camel milk butter was found to have low Reichert-Meissl, Polenske, and saponification values, but a higher melting point, refractive index and Iodine value. The fatty acids of camel milk differ markedly from those at cow milk fat in the lower content of short chain fatty acids. FARAH, Z., STREIFF. T., BACHMANN. M.R.: Herstellung und Charakterisierung van Kamelmilch-Butter. Milchwissenschaft 44 (7) 412-414 (1989) 44 Butterzusammensetzung (Kamelmilch) Es wurden Versuche zur Herstellung von Butter aus Kamelmilch in einer ländlichen Region im Nordosten Kenias durchgeführt. Die Milch wurde auf 65°C erhitzt und mit einer Handzentrifuge der Marke Alfa-Laval separiert. Um die idealen Butterungsbedingungen zu ermitteln, wurde der Rahm bei verschiedenen Temperaturen und unterschiedlichem Rahmfettgehalt verbuttert. Unter den gewählten experimentellen Bedingungen wurde die höchste Ausbeute an Butterfett (85,3%) bei einer Butterungstemperatur von 25°C und einem Rahmfettgehalt von 22.5% erreicht. Bei dieser Temperatur betrug die Butterungszeit 11 min. Einige chemische und physikalische Konstanten der gewonnenen Butter sowie die Zusammensetzung der Fettsäuren in Milch von zwei verschiedenen Kamelherden wurden ebenfalls untersucht. Verglichen mit Kuhmilchbutter weist Kamelmilchbutter Polenskeund niedrigere Reichert-Meissl-, Verseifungszahlen, aber höheren Schmelzpunkt, Brechungsindex sowie eine höhere Jodzahl auf. Kamelmilchfett unterscheidet sich van Kuhmilchfett im niedrigeren Gehalt an kurzkettigen Fettsäuren. FARAH. Z., STREIFF, T, BACHMANN'. M.R.: Fabrication
et caracterisation de beurre du lait de chamelle. Milchwissenschaft
Acknowledgements The authors express their thanks to Prot. A. Prabucki for his valuable technical assistance and Prof. Z. Puhan for stimulating discussion.
5. References
(1) DICKSON, H.R.P, in The Arab at the Desert. London, George Allan and Unwin Ltd., p. 409 (1951) (2) KNOSS, K.H.: World Animal Rev. 57 11-21 (1986) (3) YAGIL. R.: FAO Animal Production and Health Paper, Rome, 2622-24 (1982) (4) SCHNEIDER, K., ROEDER, H.: Ole praktische Milchprüfung und die Kontrolle van Molkereiprodukten, 12. Aufl. Verlag Wyss, Bern (1979) (5) Schweiz. Lebensmittelbuch, 2. Band-spezieller Teil, Kapitel Rahm und Butter, Eidg. Drucksachen- und Materialzentrale, Bern (1967) (6) PRABUCKI, A.: Anleitung zur gaschromatographlschen Bestimmung van Fettsäuren-Methylestern. ETH Zürich (unveröffentlicht) (7) CHRISTOPHERSON, FW, GLASS. RL J. Dairy Science 52 1289-1290 (1969) (8) HUNZIKER. O.R., In: The Butter Industry. La Grange, Illinois, p. 286 (1927) (9) TOPEL, A.: Chemie und Physik der Milch, 2. Aufl. VEBVerlag, Leipzig, p. 90 (1981)
44 (7)412-414 (1989). 44 Composition du beurre (Iait de chamelle) FARAH, Z., STREIFF, T., BACHMANN, M.R.: Producción y caracterización de manteca de leche de camella. Milchwissenschaft 44 (7) 412-414 (1989). 44 Composición de manteca (Ieche de camella)
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Journal of Dairy Research (1992) 59 229-231 Printed in Great Britain
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Heat coagulation of camel milk
By ZAKARIA FARAH AND DEBORAH ATKINS*
Laboratorium für Milchwissenschaft, Institut für Lebensmittelwissenschaft, ETH-Zentrum, CH-8092 Zürich, Schweiz
* Ol Maisor Farm, PO Box 9, Rumuruti, Kenya
(Received: 30 July 1991 and accepted for publication 28 October 1991)
Camel milk is an important component of the human diet in many parts of the world. It contains all essential nutrients and the composition is similar to that of cows' milk (Yagil, 1982). Present knowledge about the milk production potential of camels (Oamelu8 dromedariu8) is very limited. Data available show, however, that a camel on good feed can produce 2000 I milk per lactation (Yagil, 1982), and even higher milk yields have been recorded (Knoess, 1980). Camel milk is drunk fresh or in the form of fermented milk. Heat processing, such as pasteurization and sterilization, as a means of preserving camel milk is unknown. Information on the heat stability of camel milk is therefore scarce. In an earlier study (Farah, 1986), camel milk was heated to 63, 80 and 90°C for 3o min and the distribution of N between the total protein, non-casein N and non-protein N fractions was determined. The whey proteins were also examined by PAGE. The camel milk whey protein showed generally higher heat stability than that from cows' mille In order to study the ability of camel milk to withstand higher processing temperatures, the heat coagulation time (HCT) was determined in the range 100-130°C and pH 6-:~-7'1, and compared with measurements on cows' milk.
EXPERIMENTAL Milk samples Camel milk samples were taken at Ol Maisor Camel Farm, which is situated just north of the equator in Kenya's Laikipia District at an altitude of between 1767 and 1889 m above sea level. The animals were of indigenous breed and were fed throughout the year exclusively by grazing. The milk samples were collected from ten individual camels. The pH of the ten samples and that of a pooled sample were determined. The milks were then kept refrigerated at 4°C and transported to our laboratory within 24 h. Upon arrival, the milk samples were skimmed and analysed. For comparison, bulk cows' milk from our Zurich laboratory was used.
Determination of heat stability Milk was adjusted to various pH in the range 6.3 - 7.1 by adding 0.1 M NaOH or 0·1 MHCI. HCT was determined in a thermostatically controlled oil bath at 100, 120 and 130°C according to the method of Davies & White (1966).
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Fig. I. Heat coagulation time-pH curves for (camel milk at . 100°C; . 120°C and O. 130 °C, and for cows' milk at 130°C (X).
RESULTS AND DISCUSSION The heat stability of milk can be defined in terms of the time required to induce coagulation at a given temperature. For bovine milk, the most widely used temperature for heat coagulation is 130 or 140°C. Preliminary experiments showed that in camel milk the HCT at 140°C was too short (< 1 min) for the present assay. Coagulation times were therefore determined at. 100, 120 and 130 °C. Fig. I shows the HCT-pH curves for pooled camel and cows' milks. All ten individual camel milk samples gave similar HCT-pH curves. The HCT-pH curve of cows' milk is in agreement with findings reported previously (Rose, 1963; Fox, 1982). It. showed a marked maximum around pH 6.7 and a minimum near pH 6.8. The heat stability increased above pH 6.9. The shape of the HCT-pH curve for camel milk at low temperature was different from those at high temperatures. The milks heated at. 130 and 120°C were very unstable at all pH and coagulated in 2-3 min. At 100 °C the HCT initially increased with pH, remained constant between pH 6.4 and 6.7 and then increased progressively with increasing pH. Milks from different species differ in their heat stability. Compositional differences and heat-induced interaction between the caseins and whey proteins, particularly -casein and lactoglobulin, are reported to be responsible for these differences (Haynes & Fox, 1975; Fox & Hoynes, 1976; Ganguli, 1979). Casein fractions homologous with bovine a- and ft-casein were isolated and identified by PAGE and ion-exchange chromatography (Farah & Farah-Riesen, 1985; Larsson- Ranikiewicz & Mohamed, 1986). In these studies no protein fraction corresponding to K-casein could be clearly detected. It is possible that camel casein contained so little -casein that it escaped detection or was obscured by other casein fractions. On the other hand, foul' whey proteins have been isolated from camel milk: two proteins similar to serum albumin and -lactalbumin, and two novel milk proteins of no structural similarity to other milk proteins. The evidence for the presence of -lactoglobulin in camel milk is conflicting (Farah, 1986; Beg et al. 1984, 1987).
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The present study found that the heat stability of camel milk differs markedly from that of cows' milk -Casein and -lactoglobulin play an important role in the stability of bovine milk. Therefore, the absence or deficiency of these two proteins in camel milk might be a cause of its poor stability at high temperatures. However, this remains to be confirmed.
REFERENCES
BEG, O. U. VON BAHR-LINDSTRÖM, H. ZAIDI, Z.H. & JÖRNVALL, H. 1987 Characterisation of a heterogeneous camel milk whey non-casein protein. FEBS Letters 216 270-274 DAVIES. T. & WIIITE, J. C. D. 1 966 The stability of milk protein to heat. I. Subjective measurement of heat stability of milk. Journal of Dairy Research 33 6i-81 FARAH, Z. 1986 Effect of heat treatment on whey proteins of camel milk. Milchwissenschaft 41 763-765 FARAH, Z. & FARAH-RIESEN, M. 1985 Separation and characterization of major components of camel milk casein. Milchwissen8chaft 40 66B-6i 1 Fox, P. F. 1982 Heat-induced coagulation of milk. In Developments in Dairy Chemistry-l Proteins pp. 189-228 (Ed. P. F. Fox). London: Applied Science Publishers Fox, P. F. & Haynes, M. C. T. 1976 Heat stability characteristics of ovine, caprine and equine milks. Journal of Dairy Research 43 433-442 GANGULI, N. C. 1979 Stability of buffalo casein micelles. Journal of Dairy Research 46 401-405 Hoynes,M. C. T. & Fox. P. F. 1975 Some physico-chemical properties of porcine milk. Journal of Dairy Research 42 4:3-.56 KNOESS, K. H. 1H80 Milk production of the dromedary. In Camels pp. 201-214. International Foundation for Science Provisional Report No. (j LARSSON-RAZNIKIEWICZ, M. & MOHAMED, M. A. 1986 (Analysis of the casein content, in camel (Camelus dromedarius) milk. Swedish Journal of Agricultural Research 16 13-18 ROSE, D. 1963 Heat stability of bovine milk: a review. Dairy Science Ab8tracts 25 45-52 Y AGIL, R. 1982 Camels and camel milk. Rome: Food and Agriculture Organization (FA 0 Animal Production and Health Paper No. 26 pp. 14-19) BEG, O. D., Von BAHR-LINDSTROM, H., ZAIDI, Z. H. & JÖRNVALL, H. 1984 A small camel-milk protein rich in cysteine/half-cystine. Bioscience Reports 4 1065-1070
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Farah, Camel milk
Rennet coagulation properties of camel milk
By Z. FARAH and M. R. BACHMANN Labor für Milchwissenschaft, Eidg. Technische Hochschule, Zürich, Switzerland
1.Introduction According to FAO statistics there are 17 million camels in the world, of which 12.2 million are in Africa and 4.8 million in Asia (1). The camel is a potentially important source of milk. Indeed, in some countries hosting large camel populations, camel milk is one of the main components of the human diet. The present knowledge about milk production potential is very limited. However, milk production varying between 1,800 and 12,775 kg during a lactation period between 9 and 18 months has been reported (2). Most camel milk is consumed fresh or when it has just turned sour. Reports on the possibility of obtaining cheese from camel milk are scarce and often contradictory. Some authors report the existence of rennet coagulated cheese made from camel milk while others categorically state that cheese cannot be made from camel milk (3, 4, 5). The present investigation was therefore undertaken to obtain a better insight into the problem of rennet coagulation of camel milk. The study describes the action of rennet on camel milk as well as the effect of pH, temperature, and added calcium chloride on the coagulum development. The release of non protein nitrogen (NPN) by the action of rennet on camel milk has also been studied. 2. Materials and methods 2.1 Milk samples Camel milk samples were taken at Ngare Ndare Camel Farm) which is situated in Kenya's Laikipia District at an altitude of 1,730 to 1,890 m above sea level. The animals of indigenous breed (Camelus dromedarius) were fed year round exclusively by grazing. The milk samples were collected from 10 individual camels. After measuring the pH of the individual samples the milks were mixed to form one batch, skimmed and prepared for coagulation studies in the laboratory. For comparison bulk cow milk was used. 2.2 Milk coagulation activity determination The rennet was commercial powder from Chr. Hansen with an activity of 1:100,000. Rennet solution of 0.4% was prepared and an appropriate amount of this solution was taken to give a visually observed coagulation time of approximately 5 min in cow milk. For measurement of the milk coagulation time the following two methods were used. Visual method 25 ml samples of milk were measured into a 125 ml beaker placed in a water bath at 35°C for 30 min. After the addition of appropriate volume of rennet solution, the coagulation time was measured with the visual method described by HOSTETTLER (6). The coagulation time was defined as the time required for the first appearance of graininess in the moving film of milk on the surface of the glass walls.
Formagraph method The coagulation time for pooled camel and cow's milk samples were measured on Formagraph Type 11700 (Foss Electric, Denmark) according to the procedure of McMAHON and BROWN (7). The milks were adjusted to pH 6.65 and equilibrated at 35 °C. 2.3 Electron microscopic examination For electron microscopic examination freeze dried reconstituted camel and cow milks were used. After adjusting the pH to 6.65 the milk were equilibrated at 35 °C and coagulation monitored by visual observation. Samples were taken before and after adding the rennet at time intervals up to the visually observed coagulation time. Samples were prepared for electron microscopy with the freeze-fracturing technique and without adding any cryoprotectives. The freeze-fracturing was carried out in a modified Batzers BAF 300 unit, at object temperature of -150 °C. Details of the procedure have been described by MÜLLER et al. (8). Electron micrographs were made at 25.000 X magnification. The negatives were enlarged 3-5 times, but magnification at 75,000 was used for the interpretation of the results. 2.4 Effect of temperature The milk samples were adjusted to pH 6.65 by slow addition of 1 M HCI, placed in a water bath, and the coagulation time was measured at temperatures over the range at 25 to 40 °C. 2.5 Effect of pH All samples were equilibrated at 35 °C and the coagulation time determined at pH between 6.25 and 7.00. 2.6 Effect of added calcium All the samples were adjusted to pH 6.65 and placed in a water bath at 35°C. Calcium sensitivity was evaluated by measuring the coagulation time after addition of an appropriate amount of calcium chloride. 2.7 Measurement of NPN released by the action of rennin on milk This method is patterned after that described by NITSCHMANN (9). An aliquote rennet solution low enough to give a coagulation time of around 20 min in cow milk was added to 25 ml portions of camel and cow milk samples incubated in a water bath at 35°C. At pre-determined intervals trichloroacetic acid was added to each portion to obtain a final concentration of 12 % (w/v) in order to stop the reaction and precipitate the milk proteins. The trichloroacetic acid soluble NPN, split-off during the action of rennet as well as the total nitrogen (TN) were determined by the Kjeldahl method (10).
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3. Results and discussion pH values and coagulation times of camel and cow milk are presented in Table 1. The lowest pH measured in the 10 individual camel milks was 6.48 and the highest 6.70. The most frequently observed pH values were 6.55 and 6.65. which agreed with camel milk pH values reported in the literature (11). Coagulation times for each individual camel milk sample as well as those of mixtures were measured and compared with bulk cow milk adjusted to pH 6.65. The coagulum obtained was a precipitate in the form of flocks Table 1: The pH values and coagulation times of 10 Individual camel milk samples at 35 °c pH 6.48 6.55 6.55 6.56 6.60 6.63 6.65 6.65 6.65 6.70 6.65 (adjusted) 6.65 (adjusted) Coagulation time (sec) 540 600 660 600 720 720 780 720 840 1,020 840 300 Fig. 1: Formagramm of duplicate camel (A), and cow (B milk samples. (pH adjusted 6.65 Rcow = 330 sec, Rcamel = 930 sec)
Camel milk sample No. 1 2 3 4 5 6 7 8 9 10 Mixture 1-10 Cow milk
and no clot was formed. In all the samples the visually observed coagulation time of camel milk was 2 to 3 times longer than that of cow milk. Typical tracing by Formagraph of duplicate cow and camel milk samples are shown in Fig. 1. The coagulation time (R) was determined by measuring the distance from the origin to the point where the baseline began to increase in width. Comparison of the coagulation time measured by the 2 methods showed that the coagulation point for the visual method occurred prior to that of the Formagraph. This is due to the requirement of minimal gel formation to induce pendulum movement by the Formagraph before coagulation can be detected (7). However, the coagulation time recorded by the Formagraph is consistent with that of the visual method and shows that under the applied experimental conditions camel milk coagulated approximately 3 times slower than cow milk.
By the Formagraph method the progress of the curd formation can be detected. This is expressed as the time from the start of gel development until a width of 20 mm is reached (7). Following this definition no curd firmness could be measured in camel milk as this width of Formagraph was not reached, due to the failure of curd formation. The primary aim of the electron microscopic examination was to give a visual illustration of the coagulation process and to compare the structural changes of cow and camel casein micelles during milk coagulation. Electron micrographs of the casein micelles of camel and cow milk are presented in Fig. 2. Both camel and cow casein micelles appeared as almost spherically shaped particles, composed of numbers of submicelles. Cow casein micelles are dispersed over the field (Fig. 2, A) whereas camel casein micelles are more aggregated and grouped together (Fig. 2, B). In all the examined fields the size of the camel casein particles appears bigger. In the present investigation the size distribution of the casein micelles has not been studied. Due to this limitation, no comparison can be made here between the size of cow and camel casein micelles. In a study on particle size distribution of casein micelles in camel milk by transmission electron microscopy ALI and ROBINSON (12) reported a range of size of casein micelles between 14 to 560 nm with most of the micelles having diameters ranging between 28 to 240 nm.
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The effects of temperature, pH and calcium chloride on coagulation time are shown in Figs. 3-5. In both camel and cow milk coagulation time was reduced with decreasing pH,
Fig. 2: Electron micrographs (x 75,000) of freeze-fractured casein micelles in camel and cow milk. Scale bar = 100 nm. A - cow milk before rennet addition B - camel milk before rennet addition C - cow milk after rennet addition (100% coagulation time) D - camel milk after rennet addition (120 % coagulation time) The visually obtained coagulation times of cow and camel milk were 5 and 15 min respectively. In the cow milk the onset of aggregation as observed in electron micrographs began approximately at 60% of the coagulation time. At 80% of the coagulation time many casein micelles were already linked together forming the beginning of a network. At the coagulation time the aggregation process was more advanced and a continuous cross-linked network of casein micelle chains could be detected (Fig. 2,C). In camel milk the electron microscopic observation did not show any micellar aggregation until about 120 % of the coagulation time (Fig. 2, D). At that time the formation of flocks was already well advanced. In contrast to cow milk, where at the coagulation point a network of mostly fused micelles is formed, camel casein micelles appear to form a less compact and looser network linked merely by contact with very little change in the original micellar structure (Fig. 2, D).
Fig. 3: Influence of temperatures on the coagulation time of camel and cow milk at pH 6.65
Fig.5: Effect of added calcium on the coagulation time of camel and cow milk at 35°C and pH 6.65
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with increasing temperature and added calcium. This means that the response to changes in pH, temperature and calcium concentration is the same for camel and cow milk, but the difference in the coagulation time still remains. The rate of liberation of NPN from casein by the action of rennet was measured by monitoring the increase in 12 % trichloroacetic acid (TCA) soluble in N-compounds. As Fig. 6 shows, in both camel and cow milk the amount of NPN released by the action of rennet increased at first, reaching a maximum at the coagulation point, and declined then at a slow constant rate. This is consistent with the findings of MEHAIA (13), who reported the release of NPN soluble in 12% trichloroacetic acid in camel milk. It seems reasonable to assume that a primary reaction of the bovine type occurred between rennet and camel milk caseins. The nature of the released fragments have not been examined. However, MEHAIA (13) reported the existence of glyco· and non-glyco--cascin in camel casein. Our earlier attempt to isolate -casein-like protein from camel milk was however unsuccessful (14). Other investigations also reported difficulties in detecting -casein in camel milk (15). The main objective of this work was to examine the rennet coagulation and some of its properties in camel milk. From the obtained results it can be concluded that camel milk casein is accessible to rennin and the effects of variables like pH, temperature and calcium chloride on the coagulation time are similar to that of cow milk but not as pronounced. The action of rennet on camel milk leads to coagulation in the form of flocks with no evidence of gel formation.
(5) CHAPMAN, M.).: World Animal Rev. 5514-19 (1985) (6) HOSTETTLER, H., STEIN, J.: Schweiz. Milchztg. 81 Wiss. Beil. 20 24 (1955) (7) McMAHON, D.)., BROWN, R.).: J. Dairy ScL 651639 (1982) (8) MÜLLER, M. et 01.: Microskopie (Wien) 36 129140 (1980) (9) NITSCHMANN, H., BOHREN, H.U.: Helv. Chem. Acta 38 1953-1963 (1955) (10) International Dairy Federation: International Standard No. 20 (1982) (11) SHALASH, M.R.: in: Camels. Intern. Foundation for Science (IFS) Symposium Khartum, Sudan 285-306 (1979) (12) All, M.Z., ROBINSON, R.K.: J. Dairy Res. 52 303--307 (1985) (13) MEHAIA, M.A.: J. Dairy ScL 69 (suppl. 1) 89 (0119) (1986) (14) FARAZ, Z., FARAH-RIESEN, M.: MiJchwissenschaft 40 669671 (1985) (15) LARSSON·RAZNIKIEWICZ, M., MOHAMED, A.M.: Swedish ). Agric. Res. 1613-18 (1986) 5. Summary FARAH, Z., BACHMANN, M.R.: Rennet coagulation properties of camel milk. Milchwissenschaft 42 (11) 689-692 (1987). 51 Camel milk (cheesemaking) The action of rennet on camel milk has been compared with cow milk. The coagulation time was determined and the structural changes of the casein micelles during coagulation were examined by electron microscopy. The effect of pH, temperature, and added calcium chloride on the coagulation time as well as the release of NPN by the action of rennet was also studied. The coagulum obtained from camel milk was a precipitate in the form of flocks, and no clot was formed. With the same amount of rennet the coagulation time of camel milk was two to three fold longer than that of cow milk. In both, camel and cow milk coagulation time was reduced with decreasing pH, with increasing temperature and added calcium. This means, that the response to changes in pH, temperature and calcium concentration is the same for camel and cow milk, but the difference in the coagulation time still remains. Camel and cow milk showed similar NPN liberation patterns. This suggests that a primary phase reaction of bovine type occurs between rennet and camel casein.
Acknowledgement This work was supported in part by the Swiss Directorate for Technical Cooperation and Humanitarian Aid, Berne. The authors express their thanks to Dr. W. Schulthess of the University of Nairobi (Kenya) for providing the camel milk samples and to Dr. E. Wehrli of the Electron Microscopy Unit EHT Zurich for his valuable technical assistance.
4. References (1) FAO: Production Year Book, FAO Rome (1978) (2) YAGIL, R.: FAO Production and Health Paper, Rom 269 (1982) (3) GAST, M.L. et 01.: in: FAO Animal Production and Health Paper, Rom 26 19-24 (1982) (2) (4) RAO, M.B., GUPTA, R.C., DASTUR, N.N.: Ind. ). Dairy ScL 2372-78 (1970)
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FARAH, Z., BACHMANN, M.R.: Labgerinnungseigenschaften der Kamelmilch. Milchwissenschaft 42 (11) 689-692 {19B]}. 51 Kamelmilch (Käseherstellung) Es wurde die Wirkung des Labenzyms auf Kamelmilch mit Kuhmilch verglichen. Die Gerinnungszeit wurde ermittelt und die Strukturveränderung der Caseinmicellen wahrend der Gerinnung untersucht. Der Einfluss des pH-Wertes, der Temperatur und des zugesetzten Calciumchlorids auf die Gerinnungszeit sowie die Spaltung des NichtProteinstickstoffs durch die Einwirkung des Labenzyms wurde ebenfalls verfolgt. Das erhaltene Koagulum der Kamelmilch bestand in einer Ausfallung in Form von Flocken und es bildete sich keine Gallerte. Bei gleichbleibender Labkonzentration war die Gerinnungszeit van Kamelmilch zwei- bis dreimal langer als bei Kuhmilch. In beiden Milchen verminderte sich die Gerinnungszeit mit der Abnahme des pH-Wertes, der Zunahme der Temperatur und der zugesetzten Menge Calciumchlorid. Dies bedeutet, dass die Reaktion auf Veränderung des pH, der Temperatur und der Calciumkonzentration gleich ist, dass jedoch der Unterschied in der Gerinnungszeit der beiden Milchen bestehen bleibt. Kamel- und Kuhmilch zeigten ein ähnliches NichtProtein-Stickstoff-Abspaltungsmuster. Dies lässt vermuten, dass zwischen Labenzym und Kamel-Casein eine dem KuhCasein ähnliche Primärphasenreaktion stattfindet. FARAH, Z., BACHMANN, M.R.: Proprithes de coagulation presure du lait de chamelle. Milchwissenschaft 42 (11) 689-692 (1987). 51 Lait de chamelle (fromagerie) FARAH, Z., BACHMANN, M.R.: Propriedades de coagulacion por el cuajo de leche de camella. Milchwissenschaft 42 (11) 689-692 (1987). 51 Leche de camella (queseda)
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K. Ulmer and A. Fischer
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Traditional slaughter, carcass dressing and processing of camels
K. Ulmer and A. Fischer
The following chapter uses Kenya as an example to describe the traditional way camels are slaughtered, deboned and processed. There is every likelihood that the slaughter, deboning and processing methods described can be adapted to other countries in Africa and Asia.
6.1
Locations for slaughter of camels
In rural areas facilities for slaughter are often non-existent or very simple. In most cases camels are slaughtered in the open air, on bare ground, without any roof to give protection from dust and sun. Simple scaffolds or trees are used to hang the carcasses up for processing. The place where camels are slaughtered in Isiolo, Kenya, can be cited as an example. Here camels are slaughtered in the open air, at a location which is quite separate from the slaughterhouse used for cattle. The smooth concrete floor is surrounded by a six-foot-high fence. The slaughter takes place on the ground, so the meat is soon contaminated with dust and dirt. Hanging racks are available for hanging up the cuts. There is also running water for cleaning, with a hole in the ground for drainage. Even in public slaughterhouses there are usually no technical facilities for carrying out the hygiene measures that are urgently needed. Inadequate energy and water supplies often make it difficult to clean and disinfect the slaughterhouse and equipment and dispose of offal and effluent. This means there is a very high risk of contamination of the meat. Adequate cold storage rooms are rarely available. This is why fresh meat is many times of poor quality and has a short shelf life in most developing countries.
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Traditional slaughter, carcass dressing and processing of camels
Fig. 6.1: Place where camels are slaughtered in Isiolo, Kenya
6.2
The traditional method of slaughter for camels
The traditional slaughter of camels requires the meat to be "halal", in accordance with Islamic custom. In this type of slaughter the animals are not stunned. The camel is first put into a sitting position, the head is secured in a caudal position (i.e. turned towards the tail), and the main blood vessels between the neck and the thorax are severed with a single cut. Death occurs immediately. Although the Koran forbids consumption of blood, in some parts of Kenya the blood is collected and consumed. If the cut in the neck is not expertly executed, however, this traditional method of slaughter can be cruel and causes great suffering to the animal, since death then does not occur immediately which is a strong argument for prior stunning of the animal. The camel's skin is removed, starting from the backbone and going down both sides of the carcass to the belly. The skin is laid on the ground with the flesh side uppermost. The hump is then split lengthwise and removed. The shoulders are separated and the ribs are cut away from the vertebrae. Next, the gastrointestinal tract is removed. The backbone is then cut out, so that the carcass collapses in on itself. The hind legs are split in the pelvis and divided up into smaller cuts in the joints. Camels are usually slaughtered early in the morning, when the outside temperatures are relatively low, about 10ºC. In public slaughterhouses there is usually a post-mortem inspection for camels as well as cattle. After the post-mortem inspection the meat is taken away very quickly by dealers in special meat transport cases on handcarts or donkey-drawn carts.
K. Ulmer and A. Fischer
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Fig. 6.2: Traditional slaughter of the onehumped camel (Field, 1995)
6.3
Traditional dressing of the carcass
In the traditional method of slaughter the carcass is already divided into cuts at the end of the slaughter procedure. As already mentioned in the previous chapter, after skinning the hump is first removed, then the shoulders are detached and the ribs are cut away from the spinal column. The spinal column is then removed, so that the carcass collapses in on itself. The hind legs are split in the pelvis and divided up into smaller cuts in the joints. There is no further dressing of the carcass. The meat is cut directly off the bone. There is no grading into high-quality and lower-quality cuts. This means that standardization is impossible and there is no guarantee of consistent quality by uniform grading of cuts of meat. The only sorting that takes place is into meat with or without bones, fat and offal.
Fig. 6.3: Traditional marketing of camel meat in rural areas of Kenya
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Traditional slaughter, carcass dressing and processing of camels
6.4
Traditional meat products in Africa and Asia
The range of traditional product groups is fairly narrow. The traditional products in Africa and Asia are mainly dried products, which are made by a lowtechnology approach. Because of the climatic conditions and lack of cold storage facilities it is virtually impossible to keep meat or meat products fresh for any length of time. The drying of meat, taking advantage of existing natural factors such as temperature, humidity and air movement, is the oldest method of food preservation. Examples of typical dried meat products are biltong, odka, qwanta, kilishi and pastirma. For the first three of these the meat is cut into strips, then dry-salted or rubbed with a paste of spices and dried in the sun on straw mats. For odka and qwanta the meat is then heated in oil and dried in the sun again. So as to be able to keep the products for up to 12 months, they are covered with oil and stored in a closed container. To make kilishi, muscle meat is cut into slices which are dried for a short time in the sun. The slices are then dipped into a mixture of water, flour and various spices and dried in the sun again with this coating. Dried products are frequently smoked over a fireplace, to improve their flavor and microbiological stability. Another dried product is pastirma. Here the dry-salting process takes several days, since large pieces of meat are used. After the dry-salting, the meat is pressed for several days to remove water and give it an attractive shape. During the subsequent drying process, which usually takes place in the shade and involves air-drying, the pieces of meat are pressed again. The dried muscle meat is then coated with a paste made from water, salt, garlic, fenugreek seeds, paprika and mustard, and dried again. In spite of the simple procedures used to make traditional dried products, the seasonal weather conditions often make it difficult to achieve consistent quality. Thus, the products often vary widely in degree of drying and processing time, and this results in poorer microbiological stability and reduced keeping quality.
For more on: Method for hygienic slaughter of camels, Dressing of the camel carcass, Meat products from camel meat, read : "Milk and meat from camel: Hand book on products and processing" Z. Farah and A. Fischer (Editors)
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Diese Woche Camburger & Co. Mathias Plüss Forschung auf Weltniveau: Die ETH widmet sich der Kamelmilch. Dass die ETH Zürich in Quantenchemie und Turbinentechnik brilliert, hatten wir gewusst. Doch nun ereilt uns die überraschende Nachricht, dass das Polytechnikum auch in der nicht gerade landesüblichen Disziplin der Kamelkunde eine Vorreiterrolle spiele. Das Kamel, sei es in seiner ein- oder zweihöckrigen Form, ist perfekt ans Leben in der Wüste angepasst. Bei Trockenheit kann es bis zu dreissig Tage ohne Wasser auskommen. Im Sandsturm verschliesst es wohlweislich seine etwas hoch getragene Nase. Für den Menschen besonders wichtig ist, dass der Passgänger selbst in Notzeiten, wenn Schafs- und Ziegenzitzen unbarmherzig versiegen, zuverlässig Milch produziert, bis zu acht Liter am Tag. Darum ist das Kamel vor allem in den Trockengebieten Ostafrikas ein zentraler Ernährungs- und Wirtschaftsfaktor. Mit dem Handel ist das allerdings so eine Sache: Oft fehlt es an Verarbeitungstechniken, um Kamelprodukte verkaufstauglich zu machen. Gerade hier leistet die ETH Entwicklungshilfe: Sie liefert Weltpremiere! eine Anleitung zur Produktion von Kamelkäse. Bislang fehlte es nämlich an einem geeigneten Stoff, der Kamelmilch schadlos zur Gerinnung bringt. Verwendet man gewöhnliches Kuhlab, wird der Kamelkäse bitter. Ein Forscherteam vom Institut für Lebensmittelwissenschaft hat es nun geschafft, einem Schimmelpilz die Produktion eines geeigneten Gerinnungsenzyms beizubringen Gentechnik sei Dank. Mit diesem Stoff, so heisst es, gelinge ein perfekter Kamelkäse. Der Kopf des Teams ist Zakaria Farah, ein gebürtiger Somali und promovierter Chemiker, der sich an der ETH seit 15 Jahren mit den Feinheiten der Kamelmilch auseinander setzt. Zusammen mit Dromedarexperten aus der ganzen Welt hat er seine Erkenntnisse nun in einem Buch veröffentlicht (Milk and Meat from the Camel, vdf-Hochschulverlag, 230 S., Fr. 47.) ein Meilenstein der Kamelforschung. Den Höhepunkt des Werks bilden zweifellos die gut vierzig Rezepte, die auszuprobieren uns leider in Ermangelung geeigneter Zutaten nicht vergönnt war: Kamelbierwurst, Camel Irish Stew, Kamelkäse, Corned Camel, Kamelcamembert und unser Liebling: Camburger. Ein kleiner Dämpfer zum Schluss: Die ETH hält keine eigenen Dromedare, wie man uns auf Anfrage versicherte. Die für die Forschung benötigte Kamelmilch werde in gekühltem Zustand aus Kenia eingeflogen. (c) 2004 by Die Weltwoche, Zürich - E-mail: webmaster@weltwoche.ch
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J. Dairy Sci. 87:26602668 American Dairy Science Association, 2004.
Expression of the Peptidoglycan Recognition Protein, PGRP, in the Lactating Mammary Gland
S. R. Kappeler, C. Heuberger, Z. Farah, and Z. Puhan
Laboratory of Dairy Science, Institute of Food Science, Swiss Federal Institute of Technology, CH-8092 Zurich, Switzerland
ABSTRACT The peptidoglycan recognition protein, PGRP, known as an intracellular component of neutrophils, has been isolated from camel (Camelus dromedarius) milk by acid precipitation followed by heparin-sepharose affinity chromatography of the supernatant. The mean concentration in milk was about 120 mg/L. It decreased during lactation by 19% and increased in the event of severe mastitis by 45%. The protein bound to lactic acid bacteria and other gram-positive bacteria with an affinity similar to that reported for the human and murine orthologs, although the isoelectric point of the molecule was distinctly higher at pH 9.02. The N-terminus of mature camel PGRP was determined as NH2ArgGluAspProPro-CO2H. Calculated and measured molecular masses were both 19.1 kDa, excluding the possibility of posttranslational modifcation or binding of cation ligands. The peptide probably builds a homotrimer at high concentration. The corresponding mRNA was isolated from lactating mammary gland tissue, and 5.3 kbp of the corresponding gene was sequenced. Similarities were found to the camel lactoferrin gene with regard to sites of expression and to the region 5 upstream to the gene. (Key words: camel milk, lactating mammary gland, peptidoglycan recognition protein) Abbreviation key: MALDI-MS = matrix-assisted laser desorption/ionization mass spectrometry, pglyrp = gene for the peptidoglycan recognition protein (short pglyrp is synonymous to murine tumor-associated gene 7 and human tnfsf3l), PGRP = peptidoglycan recognition protein (the short variant is synonymous to murine tumor-associated gene 7 and bovine oligosaccharidebinding protein). INTRODUCTION The ability of milk to protect the offspring against various diseases is based on factors that belong primar-
Received October 6, 2003. Accepted March 27, 2004. Corresponding author: S. R. Kappeler; e-mail: stefan.kappeler@ alumni.ethz.ch.
ily to the innate and adaptive immune systems. The cellular and humoral components of the latter system, such as leukocytes, immunoglobulins, and the complement system, are selectively imported from serum by intracellular routing (Praetor et al., 1999). On the other hand, the components of the first line host defense are mainly synthesized in the lactating mammary gland, conferring antimicrobial protection to the newborn. Among these factors, pattern-recognition proteins, such as lactoferrin, which binds to lipopolysaccharides of gram-negative bacteria, and lysozyme C, which binds and hydrolyzes peptidoglycans, preferably of gram-positive bacteria, exhibit antimicrobial, immunomodulating, and antiinflammatory activities, and are synthesized in the lobuloalveolar tissue of the lactating mammary gland. Lysozyme C, a 1,4--N-acetylmuramidase closely related to -lactalbumin, is expressed in the functionally differentiated, lactating epithelium, in contrast to lactoferrin, which is predominantly expressed in developing, resting, and regressing tissue (Molenaar et al., 1992). Short peptidoglycan recognition protein (PGRP) is a soluble, conserved pattern-recognition protein of vertebrates and invertebrates that binds to bacterial peptidoglycan with similar specificity, but with a higher affinity than lysozyme C (Liu et al., 2000). It has been previously detected in different mammalian tissues, such as spleen and lung (Kustikova et al., 1996), but it has not yet been isolated from milk. Expression of the gene for short peptidoglycan recognition protein (pglyrp) has been conversely discussed; the expression of the human, bovine, and murine variants exclusively in polymorphonuclear leukocytes and bone marrow stem cells, accompanied by intracellular storage in cytoplasmic granules, was demonstrated (Liu et al., 2000; Tydell et al., 2002). Low levels of pglyrp expression have been detected in different human and murine tissues (Kang et al., 1998; Liu et al., 2001). The corresponding cDNA has first been cloned from 2 mammary adenocarcinoma cell lines (Kustikova et al., 1996), and a majority of murine expressed sequence tag entries related to pglyrp in GenBank are derived from mammary gland tumors. The 19 kDa peptide is basic, monomeric, and probably aggregates to homo-trimers at higher concen-
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PGRP IN CAMEL MILK
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trations (Fornhem et al., 1996; Kiselev et al., 1998; Liu et al., 2000). Specific binding with nanomolar affinity to polymeric, noncrosslinked peptidoglycans was shown for murine PGRP (Liu et al., 2000). The molecule showed weak or no affinity to other microbial polysaccharides or to peptidoglycans of low molecular weight (Yoshida et al., 1996; Liu et al., 2000). A bacteriostatic activity toward gram-positive bacteria was reported, without any bacteriolytic or otherwise bactericidal effect, and an antiinflammatory modulation of the immune response was observed. However, in a more recent report (Tydell et al., 2002), a microbicidal effect of bovine PGRP was observed, and the protein was also effective against gram-negative bacteria and a fungal pathogen. Our interest was to characterize camel PGRP and its binding to lactic acid bacteria and pathogenic bacteria. The regulation of pglyrp expression in the lactating mammary gland of camels is furthermore correlated with data from 68 individual milk samples, with regard to health status, incidence of bacterial infection, and lactational stage of the animals. MATERIALS AND METHODS Milk Sample Collection A total of 68 samples of camel milk were collected at 2 breeding stations in Isiolo and Rumuruti in Kenya. Milk from the four quarters of the camel udders were collected separately and examined for total SCC (California mastitis test) and for bacterial growth on blood agar and Edwards agar, and in Todd-Hewitt broth. The lactational stage of the camels was between 2 and 13 mo. Additionally, 5 porcine mastitis-negative milk samples were collected on a local farmyard on the day of parturition and 3 wk thereafter. Pooled human and bovine mastitis-free raw milk samples from midlactation were obtained from a women's hospital in Zurich and from a Swiss brown cattle farm near Zurich. The milk was centrifuged at 10,000 × g and at 4°C for 30 min, and filtered through nylon mesh. The skimmed milk was acidified by addition of 1% (vol/vol) acetic acid and incubated at 37°C for 10 min. Following, it was neutralized with 1/10 vol. of 1 M sodium acetate, and centrifuged at 10,000 × g and at 4°C for 5 min. The supernatant, containing soluble whey proteins, was filtered through a 0.45-µm screen. Heparin-Sepharose Affinity Chromatography Soluble whey proteins (10 mL) were loaded on a 1mL Heparin-Sepharose HiTrap column (GE Healthcare Bio-Sciences, Little Chalfont, UK). The column was washed with 20 mL of 0.01 M sodium phosphate and
0.1 M sodium chloride at pH 7.4. Elution was performed at ambient temperature applying a linear gradient from 0.1 to 0.7 M sodium chloride over 37 min. The column effluent was monitored at 280 nm. The collected fractions were separated by 4 to 12% gradient SDS-PAGE, and proteins were detected by Coomassie staining. An additional purification step was carried out, prior to micro-sequencing and molecular mass determination, by reversed-phase HPLC on an analytical silica-coated C18-column (5 µm, 250 × 4.6 mm). Elution was performed at ambient temperature by applying a linear gradient from 0.1% trifluoroacetic acid in double-distilled, nanofiltered water to 0.1% trifluoroacetic acid in acetonitrile, at a flow rate of 1 mL/min over 60 min. Amino Acid N-Terminal Sequence Analysis Protein (0.5 nmol) collected from the effluent of the C18-column was used directly for N-terminal sequencing. Eluted proteins were checked for purity by SDSPAGE, applied on a trifluoroacetic acid-treated polyvinylidene-fluoride cartridge filter and dried under continuous nitrogen flow. Automated Edman degradation over 35 cycles was performed using an ABI 471A sequencer (Applied Biosystems, Foster City, CA) equipped with a 120A HPLC for online reversed-phase C18-HPLC analysis of phenylthiohydantoinyl AA derivatives. Mass Determination of PGRP The mole