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The rest of the top 10 dog breeds are: Border Terrier, Brittany, Cairn Terrier, Welsh Springer Spaniel, English Cocker Spaniel, Cocker Spaniel, Papillon, Australian Cattle Dog, and Shetland Sheepdog. The rest of the bottom 10 dog breeds are: Saint Bernard, Great Dane, Mastiff, Bullmastiff, Irish Wolfhound, Bloodhound, Chow Chow, Alaskan Malamute and Boxer.
A selective sweep containing the insulin-like growth factor 1 (IGF1) gene is associated with size variation in domestic dogs.
We present DNA sequence data that confirms the absence of the derived small SNP allele in the intron 2 region of IGF1 in a large sample of grey wolves and further establishes the absence of a small dog associated SINE element in all wild canids and most large dog breeds. The absence of both the SINE element and SNP allele in grey wolves suggests that the mutation for small body size post-dates the domestication of dogs.
This paper notes that though recent genomic work utilizing mtDNA pinpoints the origin of domestic dogs to East Asia, this does not comport with the archaeological evidence. Around PC 1 you can see the relationship between dogs and in particular Middle Eastern wolves. The explanation of how these Middle Eastern derived small dogs arose in a evolutionary genetic sense seems a bit muddled to me, but I think that’s the point. Note: Since I started blogging, my attitudes toward small dogs have changed, and I am more positively inclined toward them. Small dogs can be very good pets, but I like a dog that can go on long, strenuous walks with me. Large dogs are harder to care for than small dogs, because they can’t sit around the house all day.
I used to have the same attitude towards small dogs until I had an opportunity to interact with a number of hearing ear dogs (service dogs for deaf people). PS: I was shocked when data looked like dogs were only domesticated once, and I am shocked again to think that IGF1 mutation was only found and selected for once.
Designer David McCandless made this neat matrix mapping dog breeds according to factors like popularity, intelligence, longevity, or cost. Intelligence comes in many forms, even in dogs, so it's hard to say whether one breed is really "smarter" than another.

The bumper sticker "My border collie is smarter than your honor student" may be an exaggeration, but border collies are considered the smartest breed in training and obedience. In terms of social intelligence -- the ability to communicate and cooperate with others -- dogs are very clever. Experiments show that dogs will purposely fool people and other dogs to get what they want. Not only can old dogs learn new things, but new challenges also can help hold back the mental decline that's sometimes a part of aging.
Intron 2 of IGF1 contains a SINE element and single nucleotide polymorphism (SNP) found in all small dog breeds that is almost entirely absent from large breeds. However, because all small dogs possess these diagnostic mutations, the mutations likely arose early in the history of domestic dogs.
Charles Darwin leaned toward the proposition that modern dog breeds derive from a wide range of canids, but the more recent genetic work seems to imply one domestication from gray wolves, though the details are still to be worked out.
Though the preponderance of evidence seems to lean toward a single-origin model, there may be detailed nuances due to the interfertility of domestic dogs and gray wolves which will always complicate the picture (it is a matter of debate whether dogs are really just a morph of wolves, or a separate species altogether). Small size could have been more desirable in more densely packed agrarian societies where dogs may have lived partly indoors or in confined outdoor spaces…. In some ways small dogs can be thought of as cats with a more servile attitude; terriers originated as ratters. But it is probably funnier and more popular to hate on small dogs, so I’ll continue to strike that pose for the sake of blogging.
Comparing breeds can be hard for the first two types, but there's a wide range in brainpower among breeds in working and obedience intelligence.
But tests show that dogs can count up to four or five and understand the idea of addition and subtraction.
Not only can they interact well with other dogs, but they are very good at communicating with humans.
Dogs raised in a mentally stimulating environment learn faster than dogs raised in a boring one.

This could be due to selective breeding for skills that require intelligence -- such as hunting, guarding, guiding, and shepherding. Canine cognitive dysfunction syndrome in dogs is often compared to Alzheimer's disease in people. But smart dogs can be more demanding; they tend to need more attention and may be more high-strung and quicker to react in both positive and negative ways. In this paper the authors focused on a major phenotypic difference among dogs, size, and its localization in terms of genetic control to a QTL which overlaps with the IGF1 locus (insulin-like growth factor). This I think is a convincing story of why small dogs might have emerged in the Middle East first.
Small dogs also consume less precious protein in a community than large dogs or most cats, so they’re ecologically and economically more sensible to encourage to breed. The others, in order, were poodles, German shepherds, golden retrievers, Dobermans, Shetland sheepdogs, Labrador retrievers, papillons, Rottweilers, and Australian cattle dogs.
They were considered the least "trainable" breed, followed by the basenji, bulldog, chow chow, borzoi, bloodhound, Pekingese, beagle, mastiff, and basset hound. Thus, in concordance with past archeological studies, our molecular analysis is consistent with the early evolution of small size in dogs from the Middle East.
If the subsequent spread of small dogs was purely diffusion-based, as suggested by the genetic variation of IGF1 (there’s only one family of variants which results in a small dog, implying singular origin), that has implications for our ideas of how human cultures evolve. People who bred small dogs over large probably had more kids as they had more protein to share! Instead of local small dog varieties emerging to fill the vermin-controlling niche, small dogs of Middle Eastern provenance seem to have spread their particular genetic architecture across the world.

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