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 Заголовок сообщения: Ссылки на интересные кинологические сайты и форумы:
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Ссылки на интересные кинологические сайты и форумы:


Переводчик с Собачьего с Цезарем Милланом
Dog Whisperer with Cesar Millan (Animal Planet)

http://perevodchik-s-s.do.am/


PesIQ - Один из самых больших и популярный Форумов кинологов:
http://pesiq.ru/forum/index.php

(я редко общаюсь на форумах и не очень люблю, но на этом форуме можно найти много нужной информации по выставкам, разведению, воспитанию.... если уметь отфильтровывать всякую ерунду, склоки и ругань) :D

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kennel `COLISTO`S'
Чирнеко дель Этна
Россия, Москва
www.cirneco.ru


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 Заголовок сообщения: Re: Ссылки на интересные кинологические сайты и форумы:
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Ооооооооо))) это моя самая любимая передача на Animal Planet :) всегда её сморю, Цезарь Миллан-гений!!!! 8-)


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 Заголовок сообщения: Re: Ссылки на интересные кинологические сайты и форумы:
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Сразу скажу - я лично эти передачи не смотрю, но слышала довольно много. И не давно у нас, на латвийском форуме собачников была бурная и обширная дискуссия про этого дрессировщика и про механизмы собачьего поведения вообще. В рамках дискуссии давалась ссылка на вот такой ролик.
http://www.youtube.com/watch?v=RFCGtatp ... r_embedded
Я свои выводы сделала.
К стати, никто из москвичей не был на только что проходившим семинаре Gerard O'Shea? Это человек, который на меня произвёл в своё время огромнейшее впечатление.

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Дети, собаки, мужья - это все иногда так неудобно. А без них, любимых так невозможно. ((c)К.М.)


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 Заголовок сообщения: Re: Ссылки на интересные кинологические сайты и форумы:
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Sanita, на видео в основном фрагменты его передачи, т.к. вы не смотрите её, конечно ролик мог вызвать некоторые вопросы). А на самом деле к Цезарю обращаются хозяева совершенно неуправляемых собак, которые не могут существовать в обществе, например недавно я смотрела как он помогал хозяевам собаки, которая СЪЕЛА соседского чихуахуа :!: этого пса, который ненавидел маленьких собачек, хотели усыпить, а Цезарь помог и в конце программы показали как эта же собака играет в обществе миников и проявляет дружелюбие. И таких примеров множество, просто стоит посмотреть ;) . Я почти все методы воспитания Артика взяла из програмы Цезаря, Артик у нас, кто не знает, с 4,5 месяцев ходит в туалет только на улице и 2 раза в день, то есть терпит по 12 часов. Так же, мы научили его не просить еду со стола, сидеть на месте когда нужно и мн. другое). Мне эта программа очень нравится. И ещё, я вижу, что собаки, которых перевоспитывает Миллан, любят его и уважают, а не боятся, на видео же, просто нарезка) на самом деле этот человек очень любит животных и его методы просты и гениальны. В другом случае, он просто не бы широко известен во всём мире)


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 Заголовок сообщения: Re: Ссылки на интересные кинологические сайты и форумы:
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До того как у нас появилась возможность регулярно смотреть "Переводчик с собачьего", у меня было просто навязчивой идеей раздобыть ролики и посмотреть методы воздействия этого легендарного дрессировщика...
Но теперь я могу смотреть это не через интернет, а на "АнималПлэнэт" ... Так вот не смотрю ведь. У меня просто нервы не выдерживают смотреть на людей, у которых не хватило (нужное подчеркнуть) интелекта, интуиции, внимания, деликатности, жажды знаний, ответственности, энтузиазма, любви... Ну не знаю чего им не хватило чтобы все сделать правильно и надежно и что бы не довести животных до точки.Очень часто самый большой враг собаки - это ее владелец. Ведь всего этого вообще можно было избежать. Собаки в принципе не могут стать такими, каких там корректируют, если к ним относиться правильно и любить их по ИХ ПРАВИЛАМ, а не по своим, не очеловечивать, и не пускать все на самотек. Я на людей в этих роликах просто не могу смотреть спокойно. Ведь покупая автомобиль, стиральную машину, тостер все хотя бы читают руководство по эксплуатации (простите за цинизм), приступая к проффессиональной деятельности - учатся годами и десятилетиями... И только собак (ну и в некоторых случаях детей) заводят и воспитывают "на авось" или "на обум"... Причем ждут ГАРАНТИРОВАННО ПОЛОЖИТЕЛЬНОГО РЕЗУЛЬТАТА (!?) , если нужны гарантии - пусть купят тостер.
Не смотря на некоторую жесткость отдельных приемов коррекции поведения, у Цезаря несомненно есть чему поучиться и многое взять на вооружение как саму систему поведения в обществе собаки и по отношению к собаке, т.к. это во многом может посодействовать , что бы избежать ситуаций, в которых уже потребуется серьезная коррекция поведенческих актов, уже просто в целях выживания собаки. Ведь если стоит вопрос о том, что бы избавиться от собаки одним из способов, или изолировать ее от людей или соплемеников, или лишить жизни... То безусловно, из двух зол выбираешь меньшее. Ведь если пара профессиональных пинков в бочину и шкуротрясных хваток на начальном этапе, в конечном результате откроют путь к полноценному и обоюдобезопасному общению владельца со своей собакой, то пусть будет так. Если кто-то может в таких совершенно диких и запущенных ситуациях сделать что-нибудь вообще не подавляя личность собак психически и не применяя грубой силы, то пусть он поскорее найдется...


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 Заголовок сообщения: Re: Ссылки на интересные кинологические сайты и форумы:
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^^ :twisted: :(

Cesar Milan :twisted: :twisted:


http://www.wikihow.com/Control-Your-Dog ... ack-Leader :D

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Erja & Necku Iosono Verona "Nala" & Iosono Fiamma "Jemma"

Iosono Cirnechi & Italian Greyhounds


Последний раз редактировалось Erja Luomanmäki 20 май 2011, 18:31, всего редактировалось 2 раз(а).

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http://www.youtube.com/watch?v=G31N_gmUkE0


Early Neurological Stimulation

By Dr. Carmen L. Battaglia

Surprising as it may seem, it isn't capacity that explains the differences that exist between individuals because most seem to have far more capacity than they will ever use. The differences that exist between individuals seem to be related to something else. The ones who achieve and outperform others seem to have within themselves the ability to use hidden resources. In other words, it's what they are able to do with what they have that makes the difference.

In many animal breeding programs, the entire process of selection and management is founded on the belief that performance is inherited. Attempts to analyze the genetics of performance in a systematic way have involved some distinguished names such as Charles Darwin and Francis Galton. But it has only been in recent decades that good estimates of heritability of performance have been based on adequate data. Cunningham, (1991) in his study of horses, found that only by using Timeform data, and measuring groups of half brothers and half sisters could good estimates of performance be determined. His data shows that performance for speed is about 35% heritable. In other words, only about 35% of all the variation that is observed in track performance is controlled by heritable factors, the remaining 65% is attributable to other influences, such as training, management and nutrition. Cunningham's work while limited to horses, provides a good basis for understanding how much breeders can attribute to the genetics and the pedigrees.

Researchers have studied these phenomena and have looked for new ways to stimulate individuals in order to improve their natural abilities. Some of the methods discovered have produced life long lasting effects. Today many of the differences between individuals can now be explained by the use of early stimulation methods.

Man for centuries has tried various methods to improve performance. Some of the methods have stood the test of time, others have not. Those who first conducted research on this topic believed that the period of early age was a most important time for stimulation because of its rapid growth and development. Today, we know that early life is a time when the physical immaturity of an organism is susceptible and responsive to a restricted but important class of stimuli. Because of its importance many studies have focused their efforts on the first few months of life.

Newborn pups are uniquely different from adults in several respects. When born, their eyes are closed and their digestive system has a limited capacity requiring periodic stimulation by their dam who routinely licks them in order to promote digestion. At this age they are only able to smell, suck, and crawl. Body temperature is maintained by snuggling close to their mother or by crawling into piles with other littermates. During these first few weeks of immobility, researchers noted that these immature and under-developed canines are sensitive to a restricted class of stimuli which includes thermal and tactile stimulation, motion and locomotion.

Other mammals such as mice and rats are also born with limitations, and they also have been found to demonstrate a similar sensitivity to the effects of early stimulation. Studies show that removing them from their nest for three minutes each day during the first five to ten days of life causes body temperatures to fall below normal. This mild form of stress is sufficient to stimulate hormonal, adrenal and pituitary systems. When tested later as adults, these same animals were better able to withstand stress than littermates who were not exposed to the same early stress exercises. As adults, they responded to stress in "a graded" fashion, while their non-stressed littermates responded in an "all or nothing way."

Data involving laboratory mice and rats also shows that stress in small amounts can produce adults who respond maximally. On the other hand, the results gathered from non-stressed littermate show that they become easily exhausted and are near death if exposed to intense prolonged stress. When tied down so they were unable to move for twenty-four hours, rats developed severe stomach ulcers, but litter mates exposed to early stress handling were found to be more resistant to stress tests and did not show evidence of ulcers. A secondary affect was also noticed.

Sexual maturity was attained sooner in the littermates given early stress exercises. When tested for differences in health and disease, the stressed animals were found to be more resistant to certain forms of cancer and infectious diseases and could withstand terminal starvation and exposure to cold for longer periods than their non-stressed littermates.

Other studies involving early stimulation exercises have been successfully performed on both cats and dogs. In these studies, the Electrical Encephalogram (EEG) was found to be ideal for measuring the electrical activity in the brain because of its extreme sensitivity to changes in excitement, emotional stress, muscle tension, changes in oxygen and breathing. EEG measures show that pups and kittens when given early stimulation exercises mature at faster rates and perform better in certain problem solving tests than non-stimulated mates.

In the higher level animals the effect of early stimulation exercises have also been studied. The use of surrogate mothers and familiar objects were tested by both of the Kelloggs and Dr. Yearkes using young chimpanzees. Their pioneer research shows that the more primates were deprived of stimulation and interaction during early development, the less able they were to cope, adjust and later adapt to situations as adults.

While experiments have not yet produced specific information about the optimal amounts of stress needed to make young animals psychologically or physiologically superior, researchers agree that stress has value. What also is known is that a certain amount of stress for one may be too intense for another, and that too much stress can retard development. The results show that early stimulation exercises can have positive results but must be used with caution. In other words, too much stress can cause pathological adversities rather than physical or psychological superiority.

The U.S. Military in their canine program developed a method that still serves as a guide to what works. In an effort to improve the performance of dogs used for military purposes, a program called "Bio Sensor" was developed. Later, it became known to the public as the "Super Dog" Program. Based on years of research, the military learned that early neurological stimulation exercises could have important and lasting effects. Their studies confirmed that there are specific time periods early in life when neurological stimulation has optimum results. The first period involves a window of time that begins at the third day of life and lasts until the sixteenth day. It is believed that because this interval of time is a period of rapid neurological growth and development, and therefore is of great importance to the individual.

The "Bio Sensor" program was also concerned with early neurological stimulation in order to give the dog a superior advantage. Its development utilized six exercises which were designed to stimulate the neurological system. Each workout involved handling puppies once each day. The workouts required handling them one at a time while performing a series of five exercises. Listed in order of preference, the handler starts with one pup and stimulates it using each of the five exercises. The handler completes the series from beginning to end before starting with the next pup.

1. The handling of each pup once per day involves the following exercises: Tactical stimulation (between toes): Holding the pup in one hand, the handler gently stimulates (tickles) the pup between the toes on any one foot using a Q-tip. It is not necessary to see that the pup is feeling the tickle. Time of stimulation 3 - 5 seconds.(Figure 1)
2. Head held erect: Uusing both hands, the pup is held perpendicular to the ground, (straight up), so that its head is directly above its tail. This is an upwards position. Time of stimulation 3 - 5 seconds (Figure 2)
3. Head pointed down: Holding the pup firmly with both hands the head is reversed and is pointed downward so that it is pointing towards the ground. Time of stimulation 3 - 5 seconds(Figure 3)
4. Supine position: Hold the pup so that its back is resting in the palm of both hands with its muzzle facing the ceiling. The pup while on its back is allowed to sleep. Time of stimulation 3-5 seconds.(Figure 4)
5. Thermal stimulation: Use a damp towel that has been cooled in a refrigerator for at least five minutes. Place the pup on the towel, feet down. Do not restrain it from moving. Time of stimulation 3-5 seconds. (Figure 5)

These five exercises will produce neurological stimulations, none of which naturally occur during this early period of life. Experience shows that sometimes pups will resist these exercises, others will appear unconcerned. In either case a caution is offered to those who plan to use them.
Do not repeat them more than once per day and do not extend the time beyond that recommended for each exercise. Over stimulation of the neurological system can have adverse and detrimental results. These exercises impact the neurological system by kicking it into action earlier than would be normally expected, the result being an increased capacity that later will help to make the difference in its performance. Those who play with their pups and routinely handle them should continue to do so because the neurological exercises are not substitutions for routine handling, play socialization or bonding.
Benefits of Stimulation

Five benefits have been observed in canines that were exposed to the Bio Sensor stimulation exercises.

1. The benefits noted were: Improved cardio vascular performance (heart rate)
2. Stronger heart beats
3. Stronger adrenal glands
4. More tolerance to stress
5. Greater resistance to disease.

In tests of learning, stimulated pups were found to be more active and were more exploratory than their non- stimulated littermates over which they were dominant in competitive situations.

Secondary effects were also noted regarding test performance. In simple problem solving tests using detours in a maze, the non-stimulated pups became extremely aroused, whined a great deal, and made many errors. Their stimulated littermates were less disturbed or upset by test conditions and when comparisons were made, the stimulated littermates were more calm in the test environment, made fewer errors and gave only an occasional distress sound when stressed.

As each animal grows and develops, three kinds of stimulation have been identified that impact and influence how it will develop and be shaped as an individual. The first stage is called early neurological stimulation and the second stage is called socialization. The first two (early neurological stimulation and socialization) have in common a window of limited time. When Lorenz, (1935) first wrote about the importance of the stimulation process, he wrote about imprinting during early life and its influence on the later development of the individual. He states that it was different from conditioning in that it occurred early in life and took place very rapidly producing results which seemed to be permanent. One of the first and perhaps the most noted research effort involving the larger animals was achieved by Kellogg & Kellogg (1933). As a student of Dr. Kellogg's, I found him and his wife to have an uncanny interest in children and young animals and the changes and the differences that occurred during early development. Their history-making study involved raising their own newborn child with a newborn primate. Both infants were raised together as if they were twins. This study, like others that followed attempted to demonstrate that among the mammals, there are great differences in their speed of physical and mental development. Some are born relatively mature and quickly capable of motion and locomotion, while others are very immature, immobile and slow to develop. For example, the Rhesus monkey shows rapid and precocious development at birth, while the chimpanzee and the other "great apes" take much longer. Last and slowest is the human infant.

One of the earliest efforts to investigate and look for the existence of socialization in canines was undertaken by Scott-Fuller (1965). In their early studies, they were able to demonstrate that the basic technique for testing the existence of socialization was to show how readily adult animals would foster young animals, or accept one from another species. They observed that, with the higher level animals, it is easiest done by hand rearing. When the foster animal transfers its social relationships to the new species, researchers conclude that socialization has taken place. Most researchers agree that among all species, a lack of adequate socialization generally results in unacceptable behavior and often times produces undesirable aggression, excessiveness, fearfulness, sexual inadequacy and indifference toward partners.

Socialization studies confirm that one of the critical periods for humans (infant) to be stimulated are generally between three weeks and twelve months of age. For canines the period is shorter, between the fourth and sixteenth weeks of age. The lack of adequate social stimulation, such as handling, mothering and contact with others, adversely affects social and psychological development in both humans and animals. In humans, the absence of love and cuddling increases the risk of an aloof, distant, asocial or sociopathic individual. Over-mothering also has its detrimental effects by preventing sufficient exposure to other individuals and situations that have an important influence on growth and development. It occurs when a parent insulates the child from outside contacts or keeps the apron strings tight, thus limiting opportunities to explore and interact with the outside world. In the end, over-mothering generally produces a dependent, socially maladjusted and sometimes emotionally disturbed individual.

Protected youngsters who grow up in an insulated environment often become sickly, despondent,lacking in flexibility and unable to make simple social adjustments. Generally, they are unable to function productively or to interact successfully when they become adults.

Owners who have busy life styles with long and tiring work and social schedules often cause pets to be neglected. Left to themselves with only an occasional trip out of the house or off of the property they seldom see other canines or strangers and generally suffer from poor stimulation and socialization. For many, the side effects of loneliness and boredom set-in. The resulting behavior manifests itself in the form of chewing, digging, and hard- to-control behavior (Battaglia).

It seems clear that small amounts of stress followed by early socialization can produce beneficial results. The danger seems to be in not knowing where the thresholds are for over and under stimulation. Many improperly socialized youngsters develop into older individuals unprepared for adult life, unable to cope with its challenges, and interactions. Attempts to re-socialize them as adults have only produced small gains. These failures confirm the notion that the window of time open for early neurological and social stimulation only comes once. After it passes, little or nothing can be done to overcome the negative effects of too much or too little stimulation.

The third and final stage in the process of growth and development is called enrichment. Unlike the first two stages it has no time limit, and by comparison, covers a very long period of time. Enrichment is a term which has come to mean the positive sum of experiences which have a cumulative effect upon the individual. Enrichment experiences typically involve exposure to a wide variety of interesting, novel, and exciting experiences with regular opportunities to freely investigate, manipulate, and interact with them. When measured in later life, the results show that those reared in an enriched environment tend to be more inquisitive and are more able to perform difficult tasks. The educational TV program called “Sesame Street” is perhaps the best known example of a children's enrichment program. The results show that when tested, children who regularly watched this program performed better than playmates who did not. Follow-up studies show that those who regularly watch “Sesame Street” tend to seek a college education and when enrolled, performed better than playmates who were not regular watchers of the “Sesame Street” program.

There are numerous children’s studies that show the benefits of enrichment techniques and programs. Most focus on improving self-esteem and self-talk. Follow-up studies show that the enriched “Sesame Street” students, when later tested were brighter and scored above average, and most often were found to be the products of environments that contributed to their superior test scores. On the other hand, those whose test scores were generally below average, (labeled as dull) and the products of underprivileged or non- enriched environments, often had little or only small amounts of stimulation during early childhood and only minimal amounts of enrichment during their developmental and formative years. Many were characterized as children who grew up with little interaction with others, poor parenting, few toys, no books and a steady diet of TV soap operas.

A similar analogy can be found among canines. All the time they are growing they are learning because their nervous systems are developing and storing information that may be of inestimable use at a later date. Studies by Scott and Fuller confirm that non-enriched pups, when given free choice, preferred to stay in their kennels. Other litter mates who were given only small amounts of outside stimulation between five and eight weeks of age were found to be very inquisitive and very active. When kennel doors were left open, the enriched pups would come bounding out while littermates who were not exposed to enrichment would remain behind. The non-stimulated pups would typically be fearful of unfamiliar objects and generally preferred to withdraw rather than investigate. Even well-bred pups of superior pedigrees would not explore or leave their kennels, and many were found difficult to train as adults. These pups, in many respects, were similar to the deprived children. They acted as if they had become institutionalized, preferring the routine and safe environment of their kennel to the stimulating world outside their immediate place of residence.

Regular trips to the park, shopping centers and obedience and agility classes serve as good examples of enrichment activities. Chasing and retrieving a ball on the surface seems to be enriching because it provides exercise and includes rewards. While repeated attempts to retrieve a ball provide much physical activity, it should not be confused with enrichment exercises. Such playful activities should be used for exercise and play or as a reward after returning from a trip or training session. Road work and chasing balls are not substitutes for trips to the shopping mall, outings or obedience classes most of which provide many opportunities for interaction and investigation.

Finally, it seems clear that stress early in life can produce beneficial results. The danger seems to be in not knowing where the thresholds are for over and under stimulation. The absence or the lack of adequate amounts of stimulation generally will produce negative and undesirable results. Based on the above, it is fair to say that the performance of most individuals can be improved, including the techniques described above. Each contributes in a cumulative way and supports the next stage of development.
Conclusion

Breeders can now take advantage of the information available to improve and enhance performance. Generally, genetics account for about 35% of the performance, but the remaining 65‰ (management, training, nutrition) can make the difference. In the management category, it has been shown that breeders should be guided by the rule that it is generally considered prudent to guard against under and over stimulation. Short of ignoring pups during their first two months of life, a conservative approach would be to expose them to children, people, toys and other animals on a regular basis. Handling and touching all parts of their anatomy is also a necessary part of their learning which can be started as early as the third day of life. Pups that are handled early and on a regular basis generally do not become hand-shy as adults.

Because of the risks involved in under-stimulation, a conservative approach to using the benefits of the three stages has been suggested based primarily on the works of Arskeusky, Kellogg, Yearkes and the "Bio Sensor" program (later known as the "Super Dog Program").

Both experience and research have dominated the beneficial effects that can be achieved via early neurological stimulation, socialization and enrichment experiences. Each has been used to improve performance and to explain the differences that occur between individuals, their trainability, health and potential. The cumulative effects of the three stages have been well documented. They best serve the interests of owners who seek high levels of performance when properly used. Each has a cumulative effect and contributes to the development and the potential for individual performance.

* References: Battaglia, C.L., "Loneliness and Boredom" Doberman Quarterly, 1982
* Kellogg, W.N. & Kellogg The Ape and the Child, New York: McGraw Hill
* Scott & Fuller, (1965) Dog Behavior -The Genetic Basics, University Chicago Press
* Scott, J.P., Ross, S., A.E. and King D.K. (1959) The Effects of Early Enforced Weaning Behavior of Puppies, J. Genetics Psychologist, p 5: 261-81.

About The Author

Carmen L Battaglia holds a Ph.D. and Masters Degree from Florida State University. As an AKC judge, researcher and writer, he has been a leader in promotion of breeding better dogs and has written many articles and several books. Dr. Battaglia is also a popular TV and radio talk show speaker. His seminars on breeding dogs, selecting sires and choosing puppies have been well received by the breed clubs all over the country. Those interested in learning more about his seminars should contact him directly.
Early Stimulation Exercises

* Figure # 1 Tactical stimulation
Изображение

* Figure # 2 Head held erect
Изображение

* Figure # 3 Head pointed down

Изображение

* Figure # 4 Figure Supine position

Изображение

* Figure # 5 Thermal stimulation

Изображение

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Erja & Necku Iosono Verona "Nala" & Iosono Fiamma "Jemma"

Iosono Cirnechi & Italian Greyhounds


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Новый кинологический портал:

ИНФОДОГС - Кинологический Информационный Портал

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Тань при регистрации там есть случайный вопрос , как ты ответила?)). У меня всё пишет неправильный ответ) Не могу зарегистрироваться.


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о собаках

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Пишет не правильно-я уже и о психологии и т.д. писала, о собаках тоже-но пишет неверное)) не хочет меня пускать)


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попробуй о собаке (посмотри значение шрифта (крупный у тебя или мелкий). пиши мелким, без заглавных букв. Вчера, меня с первого раза зарегистрировали, проблем не было. Я ответила - о собаках -

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Спасибо Тань, получилось..надо было "о" с маленькой буквы. Уф.


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Переводчик с собачьего -Цезарь Миллан
http://cirneco.spb.ru/news/cesar-millan/


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На Бельгийском форуме меня спросили, откуда такая информация, что чирнеко 6000 лет? Я просканировала книгу.
Тогда меня послали в архив американского журнала. Статья 2004 года. Он там мне откопировал все и про фараонов, андалузов и родственные породы
, что генотип не принадлежит волку, а выведен человеком.
Мне стало интересно, я послала мэил, теперь я там член... Но теперь все материалы могу только с ссылкой помещать.
http://www.sciencemag.org/cgi/content/full/304/5674/1160/DC1
Science and AAAS журнал...
Если кому интересно можете попереводить, я только переведу красный абзац.

Следует отметить, что несколько пород обычно счита:тся древнего происхождения, таких как собаки фараона и Поденко Ибиценко, не включены в эту группу/древних пород/. Они часто думают, самый старый из всех пород собак, произошедших непосредственно из древних египетских собак, нарисованных на стенах гробниц фараонов более, чем 5000 лет назад. Наши результаты показали, однако, что эти две породы воссозданные в последнее время из комбинаций других пород. Таким образом, хотя их облик совпадает с древнеегипетскими собаками, их геномы этого не показывают.

Genetic Structure of the Purebred Domestic Dog
1. Heidi G. Parker1,2,3,
2. Lisa V. Kim1,2,4,
3. Nathan B. Sutter1,2,
4. Scott Carlson1,
5. Travis D. Lorentzen1,2,
6. Tiffany B. Malek1,3,
7. Gary S. Johnson5,
8. Hawkins B. DeFrance1,2,
9. Elaine A. Ostrander1,2,3,4,*,
10. Leonid Kruglyak1,3,4,6
+ Author Affiliations
1. 1 Division of Human Biology, Fred Hutchinson Cancer Research Center, Post Office Box 19024, 1100 Fairview Avenue North, D4-100, Seattle, WA 98109–1024, USA.
2. 2 Division of Clinical Research, Fred Hutchinson Cancer Research Center, Post Office Box 19024, 1100 Fairview Avenue North, D4-100, Seattle, WA 98109–1024, USA.
3. 3 Molecular and Cellular Biology Program, University of Washington, Box 357275, Seattle, WA 98195–7275, USA.
4. 4 Department of Genome Sciences, University of Washington, Box 351800, Seattle, WA 98195–7275, USA.
5. 5 Department of Veterinary Pathobiology, College of Veterinary Medicine, University of Missouri, Columbia, MO 65211, USA.
6. 6 Howard Hughes Medical Institute, 1100 Fairview Avenue North, D4-100, Seattle, WA 98109–1024, USA.
1. ↵* To whom correspondence should be addressed. E-mail: eostrand@fhcrc.org
Abstract
We used molecular markers to study genetic relationships in a diverse collection of 85 domestic dog breeds. Differences among breeds accounted for ∼30% of genetic variation. Microsatellite genotypes were used to correctly assign 99% of individual dogs to breeds. Phylogenetic analysis separated several breeds with ancient origins from the remaining breeds with modern European origins. We identified four genetic clusters, which predominantly contained breeds with similar geographic origin, morphology, or role in human activities. These results provide a genetic classification of dog breeds and will aid studies of the genetics of phenotypic breed differences.
The domestic dog is a genetic enterprise unique in human history. No other mammal has enjoyed such a close association with humans over so many centuries, nor been so substantially shaped as a result. A variety of dog morphologies have existed for millennia, and reproductive isolation between them was formalized with the advent of breed clubs and breed standards in the mid–19th century. Since that time, the promulgation of the “breed barrier” rule— no dog may become a registered member of a breed unless both its dam and sire are registered members—has ensured a relatively closed genetic pool among dogs of each breed. At present, there are more than 400 described breeds, 152 of which are recognized by the American Kennel Club (AKC) in the United States (1). Over 350 inherited disorders have been described in the purebred dog population (2). Many of these mimic common human disorders and are restricted to particular breeds or groups of breeds as a result of aggressive inbreeding programs used to generate specific morphologies. We have previously argued that mapping genes associated with common diseases, including cancer, heart disease, epilepsy, blindness, and deafness, as well as genes underlying the striking diversity among breeds in morphology and behavior, will best be accomplished through elucidating and taking advantage of the population structure of modern breeds (3). Understanding the genetic relationships among breeds will also provide insight into the directed evolution of our closest animal companions.
Mitochondrial DNA analyses have been used to elucidate the relationship between the domestic dog and the wolf (4–6), but the evolution of mitochondrial DNA is too slow to allow inferences about relationships among modern dog breeds, most of which have existed for fewer than 400 years (1, 7, 8). One previous study showed that nuclear microsatellite loci could be used to assign dogs from five breeds to their breed of origin, demonstrating large genetic distances among these breeds (9). Another study used micro-satellites to detect the relatedness of two breed pairs in a collection of 28 breeds but could not establish broader phylogenetic relationships among the breeds (10). The failure to find such relationships could reflect the properties of microsatellite loci (10), the limited number of breeds examined, or the analytical methods used in the study. Alternatively, it may reflect the complex structure in purebred dog populations, resulting from the recent origin of most breeds and the mixing of ancestral types in their creation. Here, we show that microsatellite typing of a diverse collection of 85 breeds, combined with phylogenetic analysis and modern genetic clustering methods (11, 12), allows the definition of related groups of breeds and that genetic relatedness among breeds often correlates with morphological similarity and shared geographic origin.
To assess the amount of sequence variation in purebred dogs, we first resequenced 19,867 base pairs of noncontiguous genomic sequence in 120 dogs representing 60 breeds. We identified 75 single nucleotide polymorphisms (SNPs), with minor allele frequencies ranging from 0.4 to 48% (table S1). Fourteen of the SNPs were breed specific. When all dogs were considered as a single population, the observed nucleotide heterozygosity (13) was 8 × 10–4, essentially the same as that found for the human population (14, 15).
To further characterize genetic variation within and among breeds, we genotyped 96 microsatellite loci in 414 purebred dogs representing 85 breeds (five unrelated dogs that lacked any common grandparents were sampled from most breeds; table S2). We predicted that, because of the existence of breed barriers, dogs from the same breed would be more similar genetically than dogs from different breeds. To test this prediction, we estimated the proportion of genetic variation among individual dogs that could be attributed to breed membership. An analysis of molecular variance (16) in the microsatellite data showed that variation among breeds accounts for more than 27% of total genetic variation. Similarly, the average genetic distance between breeds calculated from the SNP data is FST = 0.33. These observations are consistent with previous reports that analyzed fewer dog breeds (9, 10), confirming the prediction that breed barriers have led to strong genetic isolation among breeds, and are in marked contrast to the much lower genetic differentiation (typically in the range of 5 to 10%) found among human populations (17, 18). Variation among breeds in dogs is on the high end of the range reported for domestic livestock populations (19, 20).
Strong genetic differentiation among dog breeds suggests that breed membership could be determined from individual dog genotypes (9). To test this hypothesis, we first applied a Bayesian model–based clustering algorithm, implemented in the program structure (11, 12, 21), to the microsatellite data. The algorithm attempts to identify genetically distinct subpopulations on the basis of patterns of allele frequencies. We applied structure to overlapping subsets of 20 to 22 breeds at a time (22) and observed that most breeds formed distinct clusters consisting solely of all the dogs from that breed (Fig. 1A). Dogs in only four breeds failed to consistently cluster with others of the same breed: Perro de Presa Canario, German Shorthaired Pointer, Australian Shepherd, and Chihuahua. In addition, six pairs of breeds clustered together in the majority of runs. These pairings—Alaskan Malamute and Siberian Husky, Belgian Sheepdog and Belgian Tervuren, Collie and Shetland Sheepdog, Greyhound and Whippet, Bernese Mountain Dog and Greater Swiss Mountain Dog, and Bullmastiff and Mastiff—are all expected on the basis of known breed history. To test whether these closely related breed pairs were nonetheless genetically distinct, we applied structure to each of these clusters. In all but one case, the clusters separated into two populations corresponding to the individual breeds (Fig. 1B). The single exception was the cluster containing Belgian Sheepdogs and Belgian Tervurens. The European and Japanese Kennel Clubs classify these as coat color and length varieties of a single breed (23, 24), and although the AKC recognizes them as distinct breeds, the breed barrier is apparently too recent or insufficiently strict to have resulted in genetic differentiation.

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Fig. 1.
Clustering assignment of 85 dog breeds. (A) Seventy-four breeds are represented by five unrelated dogs each, and the remaining 11 breeds are represented by four unrelated dogs each. Each individual dog is represented on the graph by a vertical line divided into colored segments corresponding to different genetic clusters. The length of each colored segment is equal to the estimated proportion of the individual's membership in the cluster of corresponding color (designated on the y axis as a percentage). Breeds are labeled below the figure. (B) Six clusters containing two breeds each are subdivided at K = 2, with colors representing the estimated proportion of individual membership in only two possible clusters. Black lines separate individual dogs and the two breeds are labeled below the figures.
We next examined whether a dog could be assigned to its breed on the basis of genotype data alone. Using the direct assignment method (25) with a leave-one-out analysis, we were able to assign 99% of individual dogs to the correct breed. Only 4 dogs out of 414 were assigned incorrectly: one Beagle as a Perro de Presa Canario, one Chihuahua as a Cairn Terrier, and two German Shorthaired Pointers as a Kuvasz and a Standard Poodle. All four errors involved breeds that did not form single-breed clusters in the structure analysis.
Having demonstrated that modern dog breeds are distinct genetic units, we next sought to define broader genetic relationships among the breeds. We first used standard neighbor-joining methods to build a majority-rule consensus tree of breeds (Fig. 2), with distances calculated using the chord distance measure (26), which does not assume a particular mutation model and is thought to perform well for closely related taxa (27). The tree was rooted using wolf samples. The deepest split in the tree separated four Asian spitz-type breeds, and within this branch the Shar-Pei split first, followed by the Shiba Inu, with the Akita and Chow Chow grouping together. The second split separated the Basenji, an ancient African breed. The third split separated two Arctic spitz-type breeds, the Alaskan Malamute and Siberian Husky, and the fourth split separated two Middle Eastern sight hounds, the Afghan and Saluki, from the remaining breeds.

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Fig. 2.
Consensus neighbor-joining tree of 85 dog breeds and the gray wolf. Nine breeds that form branches with statistical support are shown. The remaining 76 breeds show little phylogenetic structure and have been combined into one branch labeled “All other breeds” for simplification. The entire tree is shown in fig. S1. The trees that formed the consensus are based on the chord distance measure. Five hundred bootstrap replicates of the data were carried out, and the fraction of bootstraps supporting each branch is indicated at the corresponding node as a percentage for those branches supported in more than 50% of the replicates. The wolf population at the root of the tree consists of eight individuals, one from each of the following countries: China, Oman, Iran, Sweden, Italy, Mexico, Canada, and the United States. Branch lengths are proportional to bootstrap values.
The first four splits exceeded the majority-rule criterion, appearing in more than half of the bootstrap replicates. In contrast, the remaining breeds showed few consistent phylogenetic relationships, except for close groupings of five breed pairs that also clustered together in the structure analysis, one new pairing of the closely related West Highland White Terrier and Cairn Terrier, and the significant grouping of three Asian companion breeds of similar appearance, the Lhasa Apso, Shih Tzu, and Pekingese (fig. S1). A close relationship among these three breeds was also observed in the structure analysis, with at least two of the three clustering together in a majority of runs. The flat topology of the tree likely reflects a largely common founder stock and occurrence of extensive gene flow between phenotypically dissimilar dogs before the advent of breed clubs and breed barrier rules. In addition, it probably reflects the fact that some historically older breeds that died out during the famines, depressions, and wars of the 19th and 20th centuries have been recreated with the use of stock from phenotypically similar or historically related dogs.
Whereas the phylogenetic analysis showed separation of several breeds with ancient origins from a large group of breeds with presumed modern European origins, additional subgroups may be present within the latter group that are not detected by this approach for at least two reasons (28). First, the true evolutionary history of dog breeds is not well represented by the bifurcating tree model assumed by the method because existing breeds were mixed to create new breeds (a process that continues today). Second, methods based on genetic distance matrices lose information by collapsing all genotype data for pairs of breeds into a single number. The clustering algorithm implemented in structure was explicitly designed to overcome these limitations (11, 12, 28) and has been applied to infer the genetic structure of several species (17, 28, 29). We therefore ran structure on the entire data set using increasing values of K (the number of subpopulations the program attempts to find) to identify ancestral source populations. In this analysis, a modern breed could closely mirror a single ancestral population or represent a mixture of two or more ancestral types.
At K = 2, one cluster was anchored by the first seven breeds to split in the phylogenetic analysis, whereas the other cluster contained the large number of breeds with a flat phylogenetic topology (Fig. 3A). Five runs of the program produced nearly identical results, with a similarity coefficient (17) of 0.99 across runs. Seven other breeds share a sizeable fraction of their ancestry with the first cluster. These fourteen breeds all date to antiquity and trace their ancestry to Asia or Africa. When a diverse set of wolves from eight different countries was included in the analysis, they fell entirely within this cluster (Fig. 3B). The branch leading to the wolf outgroup also fell within this group of breeds in the phylogenetic analysis (Fig. 2).

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Fig. 3.
(A) Population structure of 85 domestic dog breeds. Each individual dog is represented by a single vertical line divided into K colors, where K is the number of clusters assumed. Each color represents one cluster, and the length of the colored segment shows the individual's estimated proportion of membership in that cluster. Black lines separate the breeds that are labeled below the figure. Representative breeds pictured above the graph from left to right: Akita, Pekingese, Belgian Sheepdog, Collie, Doberman Pinscher, Basset Hound, American Cocker Spaniel, Bedlington Terrier, Flat-Coated Retriever, Newfoundland, and Mastiff. Results shown are averages over 15 structure runs at each value of K. (B) Population structure, as in (A), but with gray wolves included. Graph shown is averaged over five structure runs at K = 2.
At K = 3, additional structure was detected that was not readily apparent from the phylogenetic tree. The new third cluster consisted primarily of breeds related in heritage and appearance to the Mastiff and is anchored by the Mastiff, Bulldog, and Boxer, along with their close relatives, the Bullmastiff, French Bulldog, Miniature Bull Terrier, and Perro de Presa Canario. Also included in the cluster are the Rottweiler, Newfoundland, and Bernese Mountain Dog, large breeds that are reported to have gained their size from ancient Mastiff-type ancestors. Less expected is the inclusion of the German Shepherd Dog. The exact origins of this breed are unknown, but our results suggest that the years spent as a military and police dog in the presence of working dog types, such as the Boxer, are responsible for shaping the genetic background of this popular breed. Three other breeds showed partial and inconsistent membership in this cluster across structure runs (fig. S2), which lowered the similarity coefficient to 0.84.
At K = 4, a fourth cluster was observed, which included several breeds used as herding dogs: Belgian Sheepdog, Belgian Tervuren, Collie, and Shetland Sheepdog. The Irish Wolfhound, Greyhound, Borzoi, and Saint Bernard were also frequently assigned to this cluster. Although historical records do not suggest that these dogs were ever used to herd livestock, our results suggest that these breeds are either progenitors to or descendants of herding types. The breeds in the remaining cluster are primarily of relatively recent European origins and are mainly different types of hunting dogs: scent hounds, terriers, spaniels, pointers, and retrievers. Clustering at K = 4 showed a similarity coefficient of 0.61, reflecting similar cluster membership assignments for most breeds but variable assignments for other breeds across runs (fig. S2). At K = 5, the similarity coefficient dropped to 0.26 and no additional consistent subpopulations were inferred, suggesting a lack of additional high-level substructure in the sampled purebred dog population.
Our results paint the following picture of the relationships among domestic dog breeds. Different breeds are genetically distinct, and individuals can be readily assigned to breeds on the basis of their genotypes. This level of divergence is surprising given the short time since the origin of most breeds from mixed ancestral stocks and supports strong reproductive isolation within each breed as a result of the breed barrier rule. Our results support at least four distinct breed groupings representing separate “adaptive radiations.” A subset of breeds with ancient Asian and African origins splits off from the rest of the breeds and shows shared patterns of allele frequencies. At first glance, it is surprising that a single genetic cluster includes breeds from Central Africa (Basenji), the Middle East (Saluki and Afghan), Tibet (Tibetan Terrier and Lhasa Apso), China (Chow Chow, Pekingese, SharPei, and Shi Tzu), Japan (Akita and Shiba Inu), and the Arctic (Alaskan Malamute, Siberian Husky, and Samoyed). However, several researchers have hypothesized that early pariah dogs originated in Asia and migrated with nomadic human groups both south to Africa and north to the Arctic, with subsequent migrations occurring throughout Asia (5, 6, 30). This cluster includes Nordic breeds that phenotypically resemble the wolf, such as the Alaskan Malamute and Siberian Husky, and shows the closest genetic relationship to the wolf, which is the direct ancestor of domestic dogs. Thus, dogs from these breeds may be the best living representatives of the ancestral dog gene pool. It is notable that several breeds commonly believed to be of ancient origin, such as the Pharaoh Hound and Ibizan Hound, are not included in this group. These are often thought to be the oldest of all dog breeds, descending directly from the ancient Egyptian dogs drawn on tomb walls more than 5000 years ago. Our results indicate, however, that these two breeds have been recreated in more recent times from combinations of other breeds. Thus, although their appearance matches the ancient Egyptian sight hounds, their genomes do not. Similar conclusions apply to the Norwegian Elkhound, which clusters with modern European breeds rather than with the other Arctic dogs, despite reports of direct descent from Scandinavian origins more than 5000 years ago (1, 24).
The large majority of breeds appears to represent a more recent radiation from shared European stock. Although the individual breeds are genetically differentiated, they appear to have diverged at essentially the same time. This radiation probably reflects the proliferation of distinct breeds from less codified phenotypic varieties after the introduction of the breed concept and the creation of breed clubs in Europe in the 1800s. A more sensitive cluster analysis was able to discern additional genetic structure of three subpopulations within this group. One contains Mastiff-like breeds and appears to reflect shared morphology derived from a common ancestor. Another includes Shetland Sheepdog, the two Belgian Sheepdogs, and Collie, and may reflect shared ancestral herding behavior. The remaining population is dominated by a proliferation of breeds dedicated to various aspects of the hunt. For these breeds, historical and breed club records suggest highly intertwined bloodlines, consistent with our results.
Dog breeds have traditionally been grouped on the basis of their roles in human activities, physical phenotypes, and historical records. Here, we defined an independent classification based on patterns of genetic variation. This classification supports a subset of traditional groupings and also reveals previously unrecognized connections among breeds. An accurate understanding of the genetic relationships among breeds lays the foundation for studies aimed at uncovering the complex genetic basis of breed differences in morphology, behavior, and disease susceptibility.
Supporting Online Material
http://www.sciencemag.org/cgi/content/f ... 4/1160/DC1
Materials and Methods
Figs. S1 and S2
Tables S1 to S5
References
Meld dit bericht aan de moderator Gelogd

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http://cirnecoforum.webnode.nl/


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