Why Outcrossing is Not a Panacea

This post is Part 4 in a series of articles on genetic diversity, breed conservation, and genetic tests. For links to all the articles, click here.

This is Saffi. When Saffi was a puppy, she liked to sit on the windowsill. She still tries to sometimes, although she doesn't really fit any more.

Saffi is the daughter of my Indi and her sire is a red poodle dog whom I chose to use on her because, along with other reasons, his pedigree was very different to Indi's. Saffi has a 15-generation COI of just under 5%. To put that in human timescales, a 15-generation pedigree would be all a person's ancestors going back to the 1600s. COI is a probability-based estimate of something called heterozygosity, the similarity between the two 'halves' of the genome inherited from each parent.

Before I get into this, let me just say that COI/heterosis isn't the only factor that should be taken into account when making breeding decisions. It should not just be a race to the bottom. However, the reason Saffi's DNA analysis looks the way I'm about to show here did not happen by accident.

The following graph is an analysis of Saffi's DNA by Genoscoper. Genoscoper's analysis looks at genetic markers in the DNA and estimates heterozygosity from this. Saffi is more heterozygous than the majority of poodles. In fact, she is within the same range as most mixed-breed dogs that Genoscoper has analysed.

Genoscoper is one of two tests I'm aware of that are currently available that analyse DNA to estimate genetic diversity. The other one is VGL. What's cool about VGL is that it gives you a PDF with 35 pairs of DNA markers on and you can put these numbers into third-party software such as SPD and number-crunch them to give estimates of heterosis and other measures of genetic diversity in potential breedings. On SPD, these are expressed as IR (heterosis, more negative numbers are more heterozygous and more positive ones are more homozygous) and OI (higher numbers are more unusual compared to the breed as a whole. VGL has also published some example certificates for different breeds on its website. I thought it might be interesting to compare these dogs to poodles, so I entered some of the data into the SPD.

I recently announced on my blog that I'm hoping to mate Hobsey when she next comes in heat. Obviously I am going to mate Hobsey to a poodle. But let's suppose I wanted to mate Hobsey to something that would be really different from her, so I mated her to my sister's Dobermann. Let's see how that would work:

Nota bene: This is an example Dobermann from VGL, and not my sister's Dobermann, who is in actuality castrated and we're not even sure if he is a real Dobermann.

The IR has a positive value of 0.02. That doesn't look awful, but the zero point is set for poodles, and negative numbers indicate more heterozygosity. There are many poodles that Hobsey could be mated with that would give a number in this range or better. The OI is 0.32, which is again OK, but not the fantastic result one might expect from such a radical step, and there are again many poodles alive and well with higher numbers than this.

How is it that two dogs of different breeds can produce pups of the same ballpark heterozygosity as two of the same breed? Well, they are both DOGS. The oldest bottleneck event was domestication. So some genetic markers are going to be ubiquitous throughout all domestic dogs.

Let's return to the subject of Saffi again. Her VGL analysis gives her a OI of 0.42 and an IR of -0.24. Which is better on both counts than the result here. I can have a more diverse dog by just breeding two poodles together. In fact, one of the conclusions of the original VGL study on poodles was that there is a reasonable amount of genetic diversity in poodles, but that it's unbalanced, and that breeders making breeding decisions to rebalance it would benefit the breed. I'm the first to admit that Saffi is not perfect, but I'm sure she's a damn sight better example of the breed in every single respect than this hypothetical puppy from Hobsey and a Dobermann.

Now, let's try something else. Let's pretend the gamekeeper down the road has a Flatcoat Retriever who is gorgeous and has a lovely temperament (and the dog isn't bad either). As there is a fad for novelty mix-breed dogs, and the price you can claim to them seems to be proportional to how stupid a portmanteau marketing name you can invent, let's say Saffi's mother Indi mated with this Flatcoat and I put an advert on a second-hand website for 'Poo Coat Retrievers'. And I kept a boy puppy... let's call him Matt. His coat is not very pleasant... kind of smelly and greasy, and sheds and gets tangled up, but he has that poodle trait of wanting to be on your lap all the time, and he's kind of a hyperactive lunatic, but he's never bitten me or done anything really horrible so I claim he has a wonderful temperament.

Here is an example of how Matt might look by VGL analysis. I just took the top alleles from a Flatcoat example and the bottom ones from Indi. This outcross works better than the last one, and Matt's done pretty well diversity-wise. He has a good IR in the negative range, and a OI of greater than 0.6, which is something hard to find in the poodle gene pool.

So, I try to recover the type by putting Matt over Hobsey. Let's see how that might come out:

Even though Matt worked quite well as an outcross from a genetic diversity standpoint, it's clear his benefits are much diminished another generation in. The IR is still negative, but only just, and the OI is better than average but not by a great deal, being in a range equal to many full-blood poodles. It's certainly not the quantum leap that might be hoped for in doing such a breeding. Saffi, a real poodle from two poodle parents, still has a result that is better on both counts than this, and Matt and Hobsey's puppy that I kept might have a more poodle-like temperament than her sire, but her coat is still horrible and she doesn't look very poodly.

As the above explains, putting dogs of two different breeds together is not a guarantee of great heterozygosity. More unrelated dogs are more likely to produce more heterozygous offspring, but it's not a given. What outcrossing breeds generally does do is introduce genes that are 'different' but as the Indi and Matt example shows, that difference is subject to diminishing returns the further in generations you progress from the original outbreeding.

Unfortunately there are currently no examples VGL has for poodle-related breeds from the same 'curly water dog' landrace, which would have been interesting to see. Serious outcross programmes (i.e. not the silly examples above) generally use related breeds to get the benefit of genetic difference but minimise the risk of outbreeding depression, however, if these are not managed properly there can also be problems. Unless several breeders are working to the same master plan, there is a risk that outcrosses will be over- or under-bred and harm the gene pool. If I breed English Bog Dogs and I mate my English Bog Dog bitch to a French Bog Dog from the same landrace stock, and other English Bog Dog breeders shun me because of this and my bloodline comes to be seen as something contaminated and unusable, then when I get too decrepit to breed any more Bog Dogs and shuffle off my mortal coil, my bloodline dies with me, and whatever genetic diversity that was in my original line and in the outcross is lost. On the other hand, if the offspring of my French/English Bog Dog breeding is seen as successful by other English Bog Dog breeders, other people might use my dogs and French Bog Dogs as well, and may well even use the same sire because his owner is accommodating of him being used on foreign breeds, and the French outcross dogs become popular sires and overused bloodlines, with all the problems inherent to that. What has to happen in this case is that breeders agree one person will use a German Bog Dog, another a French, another an Irish, and so on, and that they will plan subsequent matings carefully to make sure all the new genetics are incorporated evenly into the breed.

If outcrossing is done carelessly and purely for the sake of heterosis, like the silly examples, without any consideration given to the function and form of the breed we are trying to preserve, outbreeding depression could occur. Outbreeding depression is when the offspring of two very different parents exhibit poor fitness and unwanted traits because of poor compatibility between different genes and the genetic environment. For example, if I were to breed my English Bog Dog to a Himalayan Hairless Handbag Dog using artificial insemination because it's the most 'different' dog I can think of, I doubt the results would be very good for the Bog Dog's original purpose. If the Bog Dog is dolichocephalic, and the Handbag Dog is brachycephalic, there might well be dentition problems and very abnormal bites. The Bog Dog and related breeds are fictitious creations for the purpose of this blog post, as are the Himalayan Hairless Handbag Dogs. In the example of Matt, the poor coat and hyperactive temperament are possibly mild manifestations of this sort of issue.

I'm not lying about outbreeding depression. It is a real phenomenon that's been documented in nature. On another subject close to my heart, alpacas produced some of the finest fibre in the world until war and destruction in their native land caused them to interbreed with llamas. Now there are thought to be hardly any alpacas left alive that do not contain llama DNA, and the fleece has deteriorated accordingly, and is only now after much selective breeding starting to approach what it once was. There's no reason to think that alpacas prior to this had a compromised gene pool, and it has had no benefit to the alpacas. The only antidote for outbreeding depression is brutally aggressive selection over many generations to try to get rid of the deleterious genes that were introduced. Because aggressive selection always results in loss of diversity, there is a real risk that as breeders try to select back for the correct type and inadvertently breed away from an injudicious outcross, diversity will be lost, both the diversity introduced with the outcross as well as in the original breed parent.

Outcrossing to other breeds can potentially deflect attention from the vital work of preserving the diversity that already is in the breed's gene pool. Notice how Saffi is more diverse than Matt's offspring? If I only had room to keep one puppy from Indi, and I did this mating and used Matt instead and was eager to retire him and move forward with his offspring because of his lack of type, Saffi would never have been born and her contribution to the gene pool would not be there. Outcrossing is often seen as an easier path to newcomers to a breed rather than familiarising oneself with bloodlines and doing a lot of pedigree research, and trying to track down rare lines. Some of these rare lines have ended up in the hands of not very salubrious breeders, and accessing others might involve trips to foreign countries. Personally I have a rare British breed of chicken, and doing an outcross to a breed that looks phenotypically similar that I can buy eggs from off Ebay is looking a lot easier than trying to track down elderly, technologically illiterate, and hard to find breeders who have the genetic diversity that already exists in the breed and working out how related these remnants might be to each other. Slogging through a few generations of chickens to recover the type looks easier than travelling all over the country to obtain birds that might keel over before they manage to breed because of issues probably caused by inbreeding depression in the hope that they might be different enough to each other that the offspring will thrive and breed normally. But it's better for the breed in the long run to try to save what already exists first rather than letting it die out in pursuit of bringing in new material. Outcrossing with other breeds needs to be a last resort, and every other possibility for keeping the gene pool viable and hanging on to the unique genetics of the breed has to be pursued as a first priority. Once the original diversity of a breed is lost, it is lost forever.

Outcrossing can be a lifeline for a breed that is in dire genetic straits, and has been used successfully in many breeds, in Dalmatians, Setters, and Barbets, and in countless poultry breeds. But it's not a panacea for everything, and in rare breeds and endangered species that have sustainable genetic diversity, it can introduce more problems than it resolves, waste resources that would be better used for more important genetic conservation within the population, and even contribute to loss of diversity. It shouldn't be used without a great deal of caution and research to make sure that firstly it is necessary on a case-by-case basis, and lastly that when it is needed, the diversity it introduces isn't inadvertently squandered in subsequent generations.

Hobsey pups summer/autumn 2018(?)

Teenage Hobsey

I am getting a lot of enquiries at the moment for pups. Unfortunately I have none available at the moment, but I am hoping to try to have a litter later in the year and will explain what I'm doing later in the post.

If you are interested in a puppy, I operate a waiting list system. There are already people on the waiting list for the next litter. I (ideally and hopefully) don't have pups available 'now', and in order to be on the waiting list, I need to have your questionnaire filled in to tell me who you are and some information so I can see what you want the puppy for and the sort of poodle personality who would be the best fit for your family. If I don't have your questionnaire, unfortunately I will likely just lose your information as I'm disorganised like that and I get a lot of enquiries from people who aren't sure or are just looking to find out more (which is fine) and it will get buried with all the other bits of paper and emails.

After I have your questionnaire, you need to come and visit, where you can meet my poodles and ask any questions, and I can also make sure you're a suitable person. By this I don't mean I want to give people nasty interviews, but a couple of people when they've come to visit turned out to be allergic to the adult poodles or realised a different breed was what they should be looking for. I am always happy to give advice, and if you or your home aren't a suitable match for one of my pups, I will try to be helpful and suggest alternatives and where you can find out about them. The order of priority for the waiting list is the order in which I have met people.

All of my puppies are endorsed and sold on a contract that forbids them from being used for breeding, unless they're going to someone I know well and trust. They are suitable as companions for active people or for doing dog sports with. Occasionally I get enquiries from people who think they can get around this, and if you're such a person you need to stop wasting your own time reading this and look elsewhere.

Hobsey age 4

I am hoping to attempt a litter from Hobsey when she next comes in season, likely in June. I haven't finalised yet who will be the sire and whether I might try to have just brown pups or both black and brown. If it works, pups would be born in August and ready to go home around October. Hobsey has previously had one litter in 2016 and all the pups are well. I usually try to avoid having pups in summer as there's so much else going on and the bitches can get really uncomfortable if the weather is hot, but Hobsey coped really well with her previous litter, and I did want to breed her on her last season, but for various reasons it didn't happen and as she will be 5 this year I don't want to leave it until 2019. The alpacas are due to give birth in July so hopefully the constant going outside to check for Stiltlambs will be over by then, and I will get her a special cooling mat to lie on if it works. :-)

Basil, recent picture of a puppy from Hobsey's 2016 litter, sent from his owner

Alpacas at the Turn of Spring

The young alpacas are being trained, which as usual is going down like a lead balloon:

They eat very slowly, so it's easier to encourage them to come to be fed if the older ones aren't there eating it all.

Barbets, Water Dogs, and Grooming

The Barbet is an ancient breed of dog that will soon be recognised as a breed registerable with the Kennel Club. Although the breed is the French variation of the 'curly water dog' landrace and has been in existence for pretty much as long as the other breeds of that ancestral origin (poodles, Lagotto Romagnola, Spanish water dogs, Portuguese water dogs, etc.) it never seems to have caught on in this country until now, and from what I understand was at one point very endangered in its native France and was outcrossed to dogs from other curly water breeds to rescue it from genetic collapse, which is why it has only just developed a large enough population here to be recognised officially as a breed.

I hope water dog breeds continue to enjoy steady growth in interest in this country, and that none of these breeds will become endangered again. As a poodle breeder, I occasionally get a strange enquiry from a potential owner who says they want a water dog but don't like how poodles are groomed, asking if they can have a poodle puppy and can they not clip its face. In effect they want a poodle to be groomed to look like a Barbet or a Lagotto.

There are a couple of problems with this. First of all, if you have done your research properly and have your heart set on the look of a particular breed that is suitable, you should get that breed, not another breed as a compromise. These breeds are still numerically small in the country, and if you want one you are likely to need to get on a breeder's waiting list. I assume these people have either not looked for a Barbet/Lagotto/etc. breeder, or they have looked but can't wait for a litter to be born with space on the waiting list, or have been turned away by the breeder, which brings me to the second point:

Here's a video of a gentleman demonstrating how he grooms his IWS, a beautiful water breed. Note that the dog has to be washed and blowdried straight before scissoring to brush out any knots and achieve a good finish. Afterwards, the IWS is wetted again and left to dry naturally to give it the correct curly appearance for show

A Lagotto being groomed. As the IWS, the Lagotto has to be wetted (with fizzy pop?) after it has been trimmed to achieve the desired finish.

People also may not realise that the traditional show trims of most of the curly water retrievers are likely to involve work. Their standards call for them to be shown in a 'rustic' style. In order to be kept clean and prevent matting, they will likely need to be washed and blow-dried as poodles are to get any knotting out and do any needed trimming, but to achieve the 'rustic' look, the dog needs to be wetted again after it's been brushed out and trimmed, and let dry naturally to stop the coat from looking fluffy like a poodle's. While shaping the coat of one of these breeds might not require quite as much 'sculpting' skill as doing a show trim on a poodle, there's a comparable amount of work involved in its maintenance. Conversely, people are often shocked at how low maintenance poodles actually can be. The clipped face, feet, and base of tail in traditional poodle trims are actually like that for ease of management and cleanliness. An adult poodle in a short enough trim typically doesn't need its coat touched in between a clip, wash, and thorough brushing out of any longer parts every 6 weeks. Additionally, coats vary slightly between water breeds. Although they are all water dogs with curly coats, some coats are better at forming cords than others and some are less tightly curled than others, which means some are easier to manage in different ways. A poodle with a good poodle coat will often mat and be hard to maintain in a style intended for a looser-curled rustic coat breed. Poodles are proportionally tall and more upright compared to most other water dogs, and have leaner heads unlike the broader, squarer heads of the breeds that traditionally have long coat on their faces, and tend just to look weird and close-eyed rather than chunky and woolly when groomed like this.

Indi: only gets groomed every 6 weeks

A lady grooming a SWD, a curly water breed whose coat forms cords. The coat has to be clipped off regularly and left to regrow, which is probably one of the lowest maintenance ways of managing this type of coat. Cords do not need to be groomed in the conventional sense but do require attention to stop them becoming mats.

Breeds that are traditionally bearded usually need to have their beards washed frequently, often daily, to keep the dog clean and comfortable. Any mats that develop in the dog's face are a serious welfare issue, as trying to brush out the mats will hurt the sensitive vibrissae -- vibrissae are the coarse 'whiskers' you can see on a smooth-coated dog. They help dogs sense things like wind direction, but they don't work so well on bearded breeds because the long hair on the face grows around them, and on poodles they just get clipped off. Dogs with hairy feet and bottoms need careful attention to stop mud and poo collecting on the coat there. Most of the water dog breeds when they are not being shown are kept in pet trims that are much shorter and easier to manage than the show coat. So some people who like this look may be being unrealistic about the work needed to achieve the show coat appearance of some of the water dog breeds.

Unfortunately some people seem to be under the impression that a mix-breed dog that has poodle in its ancestry will have a water-dog-like coat that will lend itself to being groomed in this way with very little maintenance. This seems to be one of many myths put about by unscrupulous persons who produce mix-breed dogs as novelties under silly marketing 'portmanteau breed' names. It is of course possible for a mongrel to have a coat that is superficially similar to a water dog's, and if you find such a dog as an adult you should be able to get some idea of its coat maintenance needs. It is also of course quite likely that a dog that is a cross of two water breeds will have a predictable water breed coat -- and mix-breed dogs generally are more stable and more predictable when the parent breeds come from related families. However, the novelty mix-breeds being promoted usually combine poodle with something totally different with a completely incompatible coat, and are promoted as having the coat of a poodle but with 'easier maintenance'. While a mix-breed dog with a poodle ancestor could chance to look a bit like a Barbet, the results are unpredictable and it could also look like a lanky lurcher, a not-quite-right Schnauzer, Curly, or Spinone, or even like an enormous donkey creature with the pelage of a yak. I know this not because I take an interest in such things, but because if I go out in public with my dogs, strangers tend to approach me with such dogs to tell me that their strange animal is half a poodle like mine, as though they think I am writing a thesis on this subject and want to know. All of the aforementioned could inherit various degrees of double, single, straight, or curly coat spanning the whole gamut of maintenance requirements, which will result in coats that shed or grow constantly, or that do both and are a matting nightmare.

The bottom line is, if you like the look of the Barbet or Lagotto or any other water dog breed, these are great breeds and you can help them to survive by supporting a good breeder. But you also need to be realistic and research the work that is needed in caring for their coats before you commit to one. You also need to be patient if you want a dog of any rare breed.

Barbets, image by Pleple2000

Meet the Pups

The pups are four weeks old today, and their personalities are starting to become apparent. The pups' owners-to-be will come to visit them starting this week, and they have moved into a pen in the room adjoining the sitting room where they will have more space and light. The pups who have names just have them as 'placeholders' and their new owners will choose names for them when they go home.

One person on the waiting list for this litter has had to cancel, unfortunately due to being ill, and as the other two people who were waiting have made other commitments and decided to wait until the next litter, there is a space available, probably for a brown girl. If you are interested in offering her a loving home, please get in touch.

Paris

Paris is the only little boy in the litter! Despite being the smallest pup, he is bright and lively and interested in people and toys.

Ebony

Ebony is the only black puppy in the litter. She was the smallest pup at birth, but has caught up with the other ones. Ebony was the first pup to open her eyes and shows a lot of interest in people and things, but she is also noisy and is likely to be the sort of pup who will make her own amusement and get up to mischief if she's not kept entertained.

Miss Piggy

Miss Piggy was the biggest pup at birth, but is now a similar size to the other three girls. She is a calm and chilled-out pup.

Purple collar pup

Purple collar pup and Green collar pup were very similar from birth and it's only recently as their personalities have developed that their differences are becoming apparent. Purple is slightly more laid back.

Green collar pup

Green collar pup is slightly darker than Purple pup and a bit more outgoing. She and Ebony in the litter are the two pups who are so far showing the most enthusiasm about food.

Hobsey and Saffi Jump in a Snowdrift

Poodles in Snow

Otto isn't in these pictures as Saffi has decided to come in heat for the first time. Unfortunately Saffi's puppy coat got the better of me under these circumstances along with everything else that's going on, and I ended up 7-blading her body and legs.

It's obvious without her fluff how narrow and spidery Saffi still is. While poodles generally don't get much taller after a year old, they still have a lot of growing to do in terms of muscle and bone development. This is why breeders who care about their dogs won't breed from a bitch under two years old.

Snowdrift

I can't say I have much appreciation for this weather, but it's part of the seasonal progression, and I don't suppose it will last very long.