“One of the most bandied about terms among … breeders today seems to be linebreeding. Despite it’s widespread use, however, linebreeding is frequently misunderstood and miscommunicated; in fact, it is not altogether uncommon for an outcrossed pedigree to be mistakenly viewed as linebreeding by the novice. The present discussion defines linebreeding and how we can more accurately define our linebred litters.”
From – “Let’s Talk Linebreeding” written by Claudia Waller Orlandi, Ph.D. published in ‘Tally Ho’ the Basset Club of America Newsletter (July-August ’97). The online article may be found by clicking here.
(While this article was written with the Basset Hound breeder in mind, one can change the name to Gordon Setter, or any breed for that matter, as the material is “one size fits all” when it comes to the topic of breeding.)
From pro-handler Will Alexander a You Tube video chock full of tips about prepping your Setter for the ring on show day. While Will is working on an English Setter in this video, his tips for brushing and grooming are fantastic and will help make your dog look like a million bucks! Will’s posted many more tips and tricks on You Tube for those who seek more, check it out! Thank you Will for this fabulous site.
Jerold s Bell DVM, Clinical Associate Professor of Genetics, Tufts Cummings School of Veterinary Medicine
To some breeders, determining which traits will appear in the offspring of a mating is like rolling the dice – a combination of luck and chance. For others, producing certain traits involves more skill than luck – the result of careful study and planning. As breeders, you must understand how matings manipulate genes within your breeding stock to produce the kinds of offspring you desire.
When evaluating your breeding program, remember that most traits you’re seeking cannot be changed, fixed or created in a single generation. The more information you can obtain on how certain traits have been transmitted by your animal’s ancestors, the better you can prioritize your breeding goals. Tens of thousands of genes interact to produce a single individual. All individuals inherit pairs of chromosomes; one from the mother and one from the father. On the chromosomes are genes; so all genes come in pairs. If both genes in a pair are the same gene (for instance, “aa” or “AA”) the gene pair is called homozygous. If the two genes in a gene pair are unlike (for instance, “Aa”) the gene pair is called heterozygous. Fortunately, the gene pairs that make a cat a cat and not a dog are always homozygous. Similarly, the gene pairs that make a certain breed always breed true are also homozygous. Therefore, a large proportion of homozygous non-variable pairs – those that give a breed its specific standard – exist within each breed. It is the variable gene pairs, like those that control color, size and angulation that produce variations within a breed.
There are ways to measure the genetic diversity of a population. One method is to measure the average inbreeding coefficient (or Wright’s coefficient) for a breed. The inbreeding coefficient is a measurement of the genetic relatedness of the sire and dam. If an ancestor appears on both the sire and dam’s side of the pedigree, it increases the inbreeding coefficient. The inbreeding coefficient gives a measurement of the total percentage of variable gene pairs that are expected to be homozygous due to inheritance from ancestors common to the sire and dam. It also gives the chance that any single gene pair can be homozygous due to inheritance from ancestors common to the sire and dam. It also gives the chance that any single gene pair can be homozygous.
The types of matings that you choose for your breeding animals will manipulate their genes in the offspring, affecting their expression. Linebreeding is breeding individuals more closely related (a higher inbreeding coefficient) than the average of the breed. Outbreeding involves breeding individuals less related than the average of the breed. Linebreeding tends to increase homozygosity. Outbreeding tends to increase heterozygosity. Linebreeding and inbreeding can expose deleterious recessive genes through pairing-up, while outbreeding can hide these recessives, while propagating them in the carrier state.
Most outbreeding tends to produce more variation within a litter. An exception would be if the parents are so dissimilar that they create a uniformity of heterozygosity. This is what usually occurs in a mismating between two breeds, or a hybrid, like a Cockapoo. The resultant litter tends to be uniform, but demonstrates “half-way points” between dissimilar traits of the parents. Such litters may be phenotypically uniform, but will rarely breed true due to a mix of dissimilar genes.
One reason to outbreed would be to bring in new traits that your breeding stock does not possess. While the parents may be genetically dissimilar, you should choose a mate that corrects your breeding animal’s faults but complements its good traits. It is not unusual to produce an excellent quality individual from an outbred litter. The abundance of genetic variability can place all the right pieces in one individual. Many top-winning show animals are outbred. Consequently, however, they may have low inbreeding coefficients and may lack the ability to uniformly pass on their good traits to their offspring. After outbreeding, breeders may want to breed back to individuals related to their original stock, to attempt to solidify newly acquired traits.
Linebreeding attempts to concentrate the genes of specific ancestors through their appearance multiple times in a pedigree. It is better for linebred ancestors to appear on both the sire’s and dam’s sides of the pedigree. That way their genes have a better chance of pairing back up in the resultant offspring. Genes from common ancestors have a greater chance of expression with paired with each other than when paired with genes from other individuals, which may mask or alter their effects.
Linebreeding on an individual may not reproduce a outbred ancestor. If an ancestor is outbred and generally heterozygous (Aa), increasing homozygosity will produce more AA and aa. The way to reproduce ab outbred ancestor is to mate two individuals that mimic the appearance and pedigree of the ancestor’s parents.
Inbreeding significantly increases homozygosity, and increases the expression of both desirable and deleterious recessive genes through pairing up. If a recessive gene (a) is rare in the population, it will almost always be masked by a dominant gene (A). Through inbreeding, a rare recessive gene (a) can be passed from a heterozygous (Aa) common ancestor through both the sire and dam, creating a homozygous recessive (aa) offspring.
The total inbreeding coefficient is the sum of the inbreeding from the close relatives (first cousin mating), and the background inbreeding from common ancestors deep in the pedigree. Such founding ancestors established the pedigree base for the breed.
The total inbreeding coefficient is the sum of the inbreeding
from the close relatives (first cousin mating), and the
background inbreeding from common ancestors deep in the
pedigree. Such founding ancestors established the pedigree
base for the breed.
Knowledge of the degree of inbreeding in a pedigree does not necessarily help you unless you know whose genes are being concentrated. The relationship coefficient, which can also be approximated by what is called the percent blood coefficient, represents the probable genetic likeness between the individual whose pedigree is being studied, and a particular ancestor.
We know that a parent passes on an average of 50% of its genes, while a grandparent passes on 25%, a great-grandparent 12.5%, and so on. For every time the ancestor appears in the pedigree, its percentage of passed on genes can be added up and its “percentage of blood” estimated. In many breeds, an influential individual may not appear until later generations, but then will appear so many times that it necessarily contributes a large proportion of genes to the pedigree.
The average inbreeding coefficient of a breed is a measurement of its genetic diversity. When computing inbreeding coefficients, you have to look at a deep pedigree to get accurate numbers. An inbreeding coefficient based on 10 generation pedigrees is standardly used, but requires a computerized pedigree database to compute.
The average inbreeding coefficient for a breed will be based on the age and genetic background of the breed. A mating with an inbreeding coefficient of 14 percent based on a ten generation pedigree, would be considered moderate inbreeding for a Labrador Retriever (a popular breed with a low average inbreeding coefficient), but would be considered outbred for an Irish Water Spaniel (a rare breed with a higher average inbreeding coefficient).
Most breeds start from a small founding population, and consequently have a high average inbreeding coefficient. If a breed is healthy and prolific, the breadth of the gene pool increases, and the average inbreeding coefficient can go down over time. Some dog breeds were established on a working phenotype, and not on appearance. These breeds usually start with low inbreeding coefficients due to the dissimilar backgrounds of the founders. As certain individuals are linebred on to create a uniform physical phenotype, the average inbreeding coefficient can increase.
There is no specific level or percentage of inbreeding that causes impaired health or vigor. If there is no diversity (non-variable gene pairs for a breed) but the homozygote is not detrimental, there is no effect on breed health. The characteristics that make a breed reproduce true to its standard are base on non-variable gene pairs. There are pure-bred populations where smaller litter sizes, shorter life expectancies, increased immune-mediated disease, and breed-related genetic disease are plaguing the population. In these instances, prolific ancestors have passed on detrimental recessive genes that have increased in frequency and homozygosity. With this type of documented inbreeding depression, it is possible that an outbreeding scheme could stabilize the population. However, it is also probable that the breed will not thrive without an influx of new genes; either from a distantly related (imported) population, or crossbreeding.
Fortunately, most breeds do not find themselves in the position of this amount of limited diversity and inbreeding depression. However, the perceived problem of a limited gene pool has caused some breeders to advocate outbreeding of all individuals. Studies in genetic conservation and rear breeds have shown that his practice contributes to the loss of genetic diversity. By uniformly crossing all “lines” in a breed, you eliminate the differences between them, and therefore the diversity between individuals. Eventually, there will not be any “unrelated line” to be found. Everyone will have a mixture of everyone else’s genes. The practice in livestock breeding has significantly reduced diversity, and caused the reduced diversity, loss of unique rare breeds.
A basic tenet of population genetics is that gene frequencies do not change from generation to generation. This will occur regardless of the homozygosity or heterozygosity of the parents, or whether the mating is an outbreeding, linebreeding, or inbreeding. This is the nature of genetic recombination. Selection, and not the types of matings used affect gene frequencies and breed genetic diversity.
If two parents are both heterozygous (both Aa) for a gene pair, on the average, they would produce 25% AA, 50% Aa, and 25% aa. (These are the averages when many litters are combined. In reality, any variety of pairing up can occur in a single litter.) If a prolific male comes out of this litter, and he is homozygous aa, then the frequency of the “a” gene will increase in the population, and the frequency of the “A” gene will decrease. This is known as the popular sire syndrome. Of course, each individual has thousands of genes that vary in the breed, and everyone carries some deleterious recessive genes. The overuse of individual breeding animals contributes the most to decreased diversity (population bottlenecks), and the increased spread of deleterious recessive genes (the founders effect). Again, it is selection (use of this stud to the exception of others), and not the types of matings he is involved in that alters gene frequencies. Breeders should select the best individuals from all lines, so as to not create new genetic bottlenecks.
Decisions to linebreed, inbreed or outbreed should be made based on the knowledge of an individuals traits and those of its ancestors. Inbreeding will quickly identify the good and bad recessive genes the parents share, based on their expression in the offspring. Unless you have prior knowledge of what the offspring of milder linebreedings on the common ancestors were like, you may be exposing your litters (and buyers) to extraordinary risk of genetic defects. In your matings, the inbreeding coefficient should only increase because you are specifically linebreeding (increasing the percentage of blood) to selected ancestors.
Don’t set too many goals in each generation, or your selective pressure for each goal will necessarily become weaker. Genetically complex or dominant traits should be addressed early in a long-range breeding plan, as they may take several generations to fix. Traits with major dominant genes become fixed more slowly, as the heterozygous (Aa) individuals in a breed will not be readily differentiated from the homozygous-dominate (AA) individuals. Desirable recessive traits can be fixed in one generation because individuals that show such characteristics are homozygous for the recessive genes. Individuals that pass on desirable traits for numerous matings and generations should be preferentially selected for breeding stock. This prepotency is due to homozygosity of dominate (AA) and recessive (aa) genes. However, these individuals should not be overused, to avoid the popular sire syndrome.
Breeders should plan their matings based on selecting toward a breed standard, based on the ideal temperament, performance, and conformation, and should select against the significant breed related health issues. Using progeny and sib-based information to select for desirable traits and against detrimental traits will allow greater control.
This article can be reproduced with the permission of the author. Jerold.Bell@tufts.edu
Sometimes tube feeding is the only way to save newborn puppies, however there are other options that can be tried first, and this article offers advice on that topic. By clicking on the title below”To Tube or Not to Tube” you will be taken to Mary Wakeman’s website where many other useful articles scan be found. Enjoy!
March 16th, 117 The Best of Breed of Online Show Dog Magazines
The answer to this depends entirely upon whether you want your puppies to live or not. What! You say, tubing is the ONLY way to save puppies. And besides, it’s fast. Fast, yes, and deadly. It’s one of those things that sounds too good (easy) to be true; and if it sounds too good to be true it is; we know that it is in our most private thoughts.
Fast and deadly isn’t doing your part by the bitch or the puppies. You may be certain that you are getting the tube in the esophagus (which leads to the stomach) and not the trachea (which leads to the lungs). But, this isn’t the problem I’m referring to. Consider this: when we eat, the process of eating stimulates waves of contraction throughout our entire GI tract. You know very well that as puppies nurse they defecate. That reaction is due to these waves of contraction, which are called peristalsis.
OK. So, we have a sluggish or weak puppy. We put it on the bitch and it won’t nurse. What to do! TUBE. NO! If the puppy does not have a good sucking reflex, it will not have any peristalsis. This means the milk we force in through the tube will just sit there. When the tube is removed, it forces itself back up the esophagus, into the trachea, and ends up in the lungs. It does not travel down through the stomach into the intestine.
Now, how big is the stomach of a newborn puppy in your breed? 1/2 cc? Less? As much as 1cc? Probably not much more. That stomach is just a slightly wide spot on a narrow tube.
So; let’s stick 2 1/2 cc into it . Fast and Deadly. The stomach and esophagus will stretch a bit, then return to it’s original shape and size after the milk runs into the lungs. Not going to raise many puppies that way.
Well then, what do we do? Easy. We give them sub-cutaneous dextrose and saline. Sugar in salt water. The solution which is used for IV therapy. All puppies need 3 things. Warmth. Water. Sugar. That’ all they need right away and for an additional few days if necessary. So, we take the weak puppy out of the whelping box. We drop a few drops of colostrum onto its tongue several times in the first few hours. Got that immunity taken care of. We keep it in a confined box with a heat source – a heating pad or light bulb, and we give subQ dextrose in saline to supply the sugar and water. We gently stimulate it to urinate and defecate. We’ve met all the puppies needs.
How much fluid do we give? We give enough to satisfy any current dehydration debt and to provide a cushion for an hour or two in the future. How much is that? It is enough so that when we refill the syringe with dextrose and saline, the last 10 cc injection we gave hasn’t already disappeared. And it will disappear, just that fast, if the puppy is already dehydrated.
So first, we need to satisfy the back log, and then we put in some more. We want to raise a good sized lump – say the size of a golf ball on a 12-16 oz puppy. We want that golf ball to stay there a while. If it does, we can safely leave the puppy for a couple of hours. As time goes by, the fluids in this reservoir will be absorbed and the lump will disappear. Also, gravity will take a hand in removing the lump, shifting any spare fluids down around the neck. We can keep this puppy going in this way for 2 to 4 days easily. There no danger here, if the area is clean when and where we inject, and as long as the needle is parallel to the body – not pointed down at the body. We don’t want to pith the puppy (look it up). With the needle parallel to the body, the worst we can do is squirt the wall. The wall can take it.
Fluids given intravenously, by contrast, would run the risk of drowning the puppy – excess fluids in the veins will force their way out through the lungs. This result is essentially the same as that of tubing. Not good. SubQ fluids are essentially outside the circulatory system – just in a repository under the skin. If a fluid defecit exists, they can be instantly drawn into the blood stream. Until then, they have no other effect on the body.
While we are satisfying the puppy’s needs in this way, we will also repeatedly present a nipple to the puppy, several minutes after we have placed a drop of Karo syrup on its tongue. The Karo give the puppy an energy boost, so that when we place it on the bitch, it will make as strong an attempt to nurse as it can muster. We will also present the puppy with a bottle, as it will be easier for it to get milk from the bottle’s nipple than from the bitch, most of the time, during the first couple of days.
One of the greatest deterrents to getting puppies started, after tubing, is the ‘Pet Nurser’ which is widely available. Few if any breeds will nurse off of this thing – maybe a couple of toy breeds I’ve never encountered. Rather, puppies from 4.5 oz to 2# and up will readily take a Playtex preemie, or Playtex 0-3 months nipple (slow flow), one which has a flat, button-like shape. ANY puppy which does want to suck, but is unable to get enough from the bitch, should be asked to take the Playtex nurser. And if they don’t learn to nurse from it within the first few minutes, as soon as an hour or two after birth, it’s your fault, because they like this nipple just fine.
Of course, you have to put the right stuff in it. The concept of using a formulated synthetic milk replacer seems a bit bizarre. Cow’s milk is good, it’s complete, it contains the same things as dog milk. It’s not quite as good as dog’s milk, however, because it’s too dilute. Cow’s milk is 1/2 as concentrated as dog milk. So, all we have to do is go to the store and buy evaporated milk. Nothing could be simpler; comes in a can, easy to store and have on hand, useful for other purposes. We use the evaporated cow’s milk, in the slow flow nipple (no modifications to the nipple, we want it to go in slowly, and to require some exercise from the puppy to make it work). We add a dollop of Karo syrup for energy and palatability, warm slightly, and that’s it; it’s perfect.
Some of us seem to have a need to make life more complicated than it has to be. If you think your puppies suffer from the rare human problem where the size of the cow butterfat globule is too large for comfort, you can search out a source for evaporated, canned goat’smilk. And you might wish to do that because it will make it seem as though your puppies have a special problem, not a routine, ordinary problem. However, goat’s milk has no special benefit for dogs. It also must be fed undiluted from the can, with some Karo.
Note: The only puppies I have ever seen which were nutritionally stunted – and didn’t recoup their early deficits when put on solid food – were 2 giant breed siblings which were fed fresh goat’s milk. To this day these two are ‘minis’. Fresh ruminant milk has 50% too much water in it. Evaporated ruminant milk is just fine as long as you don’t screw it up by adding water. If you are faced with total milk replacement due to the death of a bitch, you will eventually have to add an egg yolk (without the white) to a can of evaporated milk with Karo, in order to raise the protein level even more. But, there is no need for this when we’re simply supplementing.
These puppies which are eager to nurse, but just can’t get anything from the bitch’s nipples, will have good peristalsis. They will work at the nipple and develop their lungs and their body muscles, though only a fraction as well as they would if they were working on the bitch’s nipples. One caution when supplementing the large litter to lessen the stress on the bitch. You must be careful not to OVER feed. The idea is to take some load off her, so you should keep her out of the box for some time every day. We don’t want to supplement and then let them drink their fill from their mother as well, then we’ll only have fat and colicy puppies, not a mother in better shape.
The next question is, will their mother lick them and stimulate the urination and defecation reflexes? If she’s not yet into that, we also have to wash their tummies with a warm wet tissue. This will stimulate the elimination reflexes. We can’t skip this part either. If we do, they’ll all colic. Some bitches, even though they have milk and the puppies nurse with no problem, just don’t like to clean their puppies. If so, then it’s our job. We caused these puppies to be born, the buck stops with us; if they need to be cleaned we have to do the job. We have to be gentle, but we have to be just as certain that we’re successful in stimulating defecation and urination as we are that the puppies are getting enough to eat. What goes in must come out!
One good way to help you be certain you’re getting each one fed and cleaned is to place colorful yarn collars around their necks. This way we can identify each puppy at a glance, no waking them or dislodging them from a nipple in order to check markings. And later, when one puppy is repeatedly striking a pose we can see from a distance which one it is. Helps us identify that BIS Puppy.
One of the things we almost never talk about is the personal safety of our exhibitors. We are constantly On The Road Again, flying into strange cities, driving to new locations. Some of these journeys, and even show sites, are in sketchy areas, at best. A significant number of us are traveling alone through all of it.
Our sport also features a unique mix of ages, genders and positions of power that have been known to be abused.
With that in mind, we offer our “Five Best Tips for Safety” while traveling. And, as a bonus, “Best Practices for Handling Inappropriate Advances.” Obviously, much of this advice applies anywhere, anytime.
Ray Helmken, retired from the Honolulu Police Department and Akita fancier; along with GWP lover Guy Miner, owner of GMM Defense, who offers self-defense courses for small groups, provide these insights.
1. Stay alert and aware. Pay attention to your surroundings. If you want to check Facebook or send a text, do it from the safety of your locked vehicle. Wandering aimlessly into a rest area bathroom while staring at your phone sets you up for someone to take advantage of your distraction. Body posture has a lot to do with how the bad guys choose victims. Head up and watchful, shoulders back, strong core and purposeful movement is our first line of defense.
2. Stay in well-lighted, populated areas. Dim parking lots, deserted (or seemingly so) rest areas with no other people around leaves the predators amongst us with too many places to lurk. “Situational awareness,” Miner says, “is critical. Know what’s going on around you. Avoiding conflict is vastly better than fighting.”
3. Keep keys and cell phones with you. Calling for help in a bad situation isn’t possible if you left your phone in the vehicle while you ran in to pee or exercise a dog or grab a bite to eat. Your car’s key fob also may have an alarm option that you can push which will set up enough racket for people to look up and see a problem. In an absolute worst case, keys wedged between your fingers, with your hand in a fist, make an adequate weapon. Aim for eyes or throat and make it count. Male attackers will always expect and be prepared for a kick to the genitals. “If you must fight, cheat!” says Miner. “Win. Defend yourself.”
4. Stay in touch with family/friends. Someone should always know your route, intended destination and ETA. Always have a travel buddy you check in with when you stop for the night. This sets up a built in alarm system — if you don’t check in, your buddy should check up on you.
5. Make use of available non-lethal self-defense and deterrent options. Pepper spray, whistles, self-defense alarms and barking dogs all work. Keep in mind that most attackers are looking for targets of opportunity. They don’t want to get caught. “The more noise you make,” Helmken says, “the more the individual will divert, go a different direction.”
*Don’t* Touch This….
As disappointing and upsetting as it is, inappropriate touching or advances are not confined to billionaires and Hollywood starlets. From copping a cheap feel to offering hotel room keys, and worse, it does happen, even in our sport.
Helmken says, “If somebody touches or grabs at you without your approval, step back. Get space between you. Get loud and verbal. This is not a time to avoid making a scene.”
From the “School of Hard Knocks” files:
Your momma was right when she told you to make good decisions and use your common sense.
Gracefully extricating oneself from an awkward or even ugly situation is much more difficult than avoiding it in the first place.
There is safety in numbers. Don’t allow yourself to be singled out of a group in social settings.
Everyone has a slightly different comfort level of what is “harmless” and what is not. Be true to yourself.
While it’s pretty to think that in today’s society people know the boundaries of what is and is not acceptable, each and every one of us need to be able and willing to say, “Back off” if a line is crossed.
In the Year of Living Well, stay safe, stay aware and stand strong.
Our family always had dogs. Mutt dogs, purebred dogs, but always dogs. I grew up with dogs everywhere. My mother eventually enrolled me in dog care 4-H because I was “shy and retiring and lacked people skills”….. I am the living testimonial to the success of the 4-H program! I continued into AKC shows as my family transitioned from “dogs” to the wonderful world of Purebred Dogs. I showed all of our family dogs in conformation and participated in Junior Showmanship competition. I went to college, earned a degree and worked as a newspaper reporter and freelance writer. Today, Today, I am an AKC Breeder of Merit, a member of the Professional Handlers Association and the host of pure dog talk http://puredogtalk.com/, THE podcast about purebred dogs.
“Furnishings” doesn’t refer just to furniture. The word itself is quite old and can be traced to back the 16th century and the Middle French word, “fournir,” which morphed into “fourniture” to mean “a supply,” or the act of furnishing. In the dog world, “furnishings” refers to long hair on the extremities of certain breeds. In some wire-haired breeds, it can refer to a longer mustache, beard and eyebrows, while in setters, furnishings refers to the flowing hair coming off the dog’s body.
Interestingly, variants in only three genes govern coat length, curl and furnishings. It was something discovered in 2009 by Edouard Cadieu and Elaine A. Ostrander of the National Human Genome Research Institute who looked at some 900 dogs representing 80 breeds. They were able to identify mutations at specific points, or loci, on three genes linked to fur length, curliness and growth pattern (what we call “furnishings”). When they looked at the three loci on the genes of another 662 dogs representing 108 breeds — from Old English Sheepdogs to Pugs – they found that the presence of the mutations or not, in various combinations, accounted for the variation in coat in 95 percent of the breeds. Only a few breeds, including Afghan hounds, have coats that can’t be explained by these genes.
Coat color and type are essential characteristics of domestic dog breeds. Although the genetic basis of coat color has been well characterized, relatively little is known about the genes influencing coat growth pattern, length, and curl. We performed genome-wide association studies of more than 1000 dogs from 80 domestic breeds to identify genes associated with canine fur phenotypes. Taking advantage of both inter- and intrabreed variability, we identified distinct mutations in three genes, RSPO2, FGF5, and KRT71 (encoding R-spondin–2, fibroblast growth factor–5, and keratin-71, respectively), that together account for most coat phenotypes in purebred dogs in the United States. Thus, an array of varied and seemingly complex phenotypes can be reduced to the combinatorial effects of only a few genes.
The tremendous phenotypic diversity of modern dog breeds represents the end point of a >15,000-year experiment in artificial and natural selection (1, 2). As has been demonstrated for traits such as body size (3) and coat color (4), marker-based associations with phenotypic traits can be explored within single breeds to initially identify regions of genetic association, and then expanded to multiple breeds for fine-mapping and mutation scanning (5, 6). Coat (pelage) phenotypes are particularly amenable to this strategy as they show a huge amount of variation across breeds but still allow for simple variation within single breeds (7). This offers a unique strategy for advancing the genetic understanding of a complex phenotype.
We used the structured pattern of fur variation in dogs to localize the genetic basis of three characteristics of the canine coat: (i) the presence or absence of “furnishings,” the growth pattern marked by a moustache and eyebrows typically observed in wire-haired dogs; (ii) hair length; and (iii) the presence or absence of curl. To accomplish this, we generated three genome-wide single-nucleotide polymorphism (SNP) data sets using the Affymetrix version 2.0 canine SNP chip (8, 9). The first data set consisted of 96 dachshunds segregating three coat varieties: wire-haired with furnishings, smooth, and long-haired without furnishings. The second data set comprised 76 Portuguese water dogs (PWDs), segregating the curl phenotype. The final data set, termed CanMap, included 903 dogs from 80 breeds representing a wide variety of phenotypes. An additional data set used to map furnishings included a panel of microsatellite markers (10), genotyped on a 96-dachshund pedigree segregating all three coat varieties.
The same strategy was used to map all three traits. First, a genome-wide association study (GWAS) within a breed segregating the phenotype was conducted to determine the most strongly associated locus. To rule out false-positives caused by population structure within the breeds (11), we did a second GWAS that used the CanMap data set divided into cases and controls based on the presence or absence of the phenotype in question. Fine-mapping of significant, concordant peaks was used to define the smallest shared haplotype, followed by sequencing to identify the putative causative mutations. Each mutation was validated in a large panel of at least 661 dogs from 108 breeds, including cases and controls for all phenotypes (table S1).
We initially mapped furnishings in the dachshund using smooth-coated and long-haired dogs as controls and wire-haired dogs as cases (Fig. 1A). Single-marker analysis of the dachshund GWAS data set and concurrent linkage analysis of the dachshund pedigree identified the same locus on canine chromosome 13 (CFA13) surrounding nucleotide 11,095,120 [P = 3.4 × 10−27, lod score (logarithm of the odds ratio for linkage) = 5.6; Fig. 1B]. We confirmed the association on CFA13 in the CanMap data set at nucleotide 11,659,792 (P = 10−241; Fig. 1C and table S2). A 718-kb homozygous haplotype in all dogs fixed with furnishings was located within both the original 3.4-Mb haplotype observed in the dachshund-only GWAS, and a 2.8-Mb haplotype identified in crossover analysis within the dachshund pedigree (Fig. 1D).
GWAS and fine-mapping identify RSPO2 as the associated gene for moustache and eyebrow growth pattern (furnishings). (A) Three types of coat segregate in dachshunds: (from left to right) smooth-coated, long-haired, and wire-haired with furnishings. (B…
Fine-mapping allowed us to reduce the homozygous region to 238 kb spanning only the R-spondin–2 (RSPO2) gene, excluding the 5′ untranslated region (5′UTR) and the first exon (Fig. 1D, fig. S1, and table S3). RSPO2 is an excellent candidate for a hair-growth phenotype as it synergizes with Wnt to activate β-catenin (12), and Wnt signaling is required for the establishment of the hair follicles (13, 14). Moreover, the Wnt/β-catenin pathway is involved in the development of hair-follicle tumors, or pilo-matricomas (15), which occur most frequently in breeds that have furnishings (16). Recent studies have shown that a mutation in the EDAR gene, also involved in the Wnt pathway, is responsible for a coarse East-Asian hair type found in humans (17), with some similarity to canine wirehair.
All exons and conserved regions of RSPO2 were sequenced in dogs from seven breeds (table S4). Only an insertion of 167 base pairs (bp) within the 3′UTR at position 11,634,766 was perfectly associated with the furnishings trait in dogs from both the case/control study and the extended pedigree (table S5). The result was further confirmed in a set of 704 dogs of varying phenotypes. In total, 297 of 298 dogs with furnishings were either homozygous (268) or heterozygous (29) for the insertion, and all 406 dogs lacking the trait were homozygous for the ancestral state, as is consistent with a dominant mode of inheritance (table S1).
This mutation does not affect the protein-coding region of the RSPO2 gene. However, because the 3′UTR frequently encodes elements that influence mRNA stability [reviewed in (18)], we examined whether the insertion was associated with a change in the expression level of the RSPO2 gene. We found a threefold increase in RSPO2 transcripts in muzzle skin biopsies of dogs with furnishings, consistent with a transcript effect (fig. S2).
We applied the same mapping strategy to hair length. Previously, mutations in the FGF5 gene were identified in Welsh corgis segregating an atypical “fluffy” or long-haired phenotype (19) and associated with excess hair growth in mice and cats (20–22). Our study replicates these findings in an extended breed set. Indeed, association analyses in both the dachshund and CanMap data sets highlight the region on CFA32 containing FGF5 with P values of 3 × 10−27 and 9 × 10−44, respectively. After fine-mapping, a 67-kb homozygous region highlighted the FGF5 gene (Fig. 2A, fig. S3, and table S6). The strongest association was observed at position 7,473,337 (P = 1 × 10−157), in which a highly conserved Cys is changed to Phe (Cys95→Phe) in exon 1 of FGF5, consistent with the previous study (19). Sequencing within the homozygous haplotype revealed no SNPs with stronger association (table S7).
Regions of homozygosity identify genes for pelage length and curl. (A) Homozygous region found on CFA32 defining the length locus. The red bar indicates the 520-kb associated haplotype from 29 long-haired dachshunds; the blue bar spans the 125-kb homozygous …
This diagnostic SNP was typed in several hundred additional dogs of varying hair length. Within the dachshunds, all long-haired dogs had the TT genotype, whereas all short or wire-haired dogs had either the GT or GG genotypes, suggesting a recessive mode of inheritance, as predicted previously (23). Across all breeds, the T allele was found in 91% of the long-haired dogs, in only 3.9% of the short-haired dogs, and accounts for ~30% of genotypes found in medium-haired dogs. Three breeds with very long hair, including the Afghan hound, neither carry the Cys95→Phe variant nor show an association with CFA32, suggesting that additional loci exist that contribute to hair length in dogs (table S1).
To identify the gene that causes curly coat, we conducted a GWAS using PWDs (Fig. 2B) and identified a single associated SNP at position 5,444,030 on CFA27 (P = 4.5 × 10−7). A SNP in close proximity (5,466,995; P = 6.9 × 10−28) was associated with curly coat in the CanMap data set. Fine-mapping revealed a shared homozygous haplotype that included two keratin genes (Fig. 2C, fig. S4, and table S8). Sequence data covering 87% of the homozygous region identified one SNP at position 5,542,806 that segregated with the trait. Non–curly haired dogs carried the CC genotype; curly coated dogs had the TT genotype. In breeds where the trait segregates, such as PWDs, all three genotypes were observed. The relevant SNP is located in the KRT71 gene (previously called K6irs1, Kb34, and K71) and causes a nonsynonymous Arg151→Trp alteration (table S9). Genotyping an additional 661 samples at this SNP validated the association (P = 3 × 10−92) (table S1).
Keratins are obvious candidates for hair growth [reviewed in (24)], and mutations in KRT71 have been described in curly coated mice (25). The mutation described in our study is within the second exon of the gene and may affect either or both of two protein domains: a coiled-coil and a prefoldin domain (www.ensembl.org/Canis_familiaris/). Conceivably, sequence alterations in these domains could affect cellular targeting, receptor binding, or proper folding of the protein after translation [reviewed in (26)].
Notably, these three mutations in various combinations explain the observed pelage phenotype of 95% of dogs sampled, which include 108 of the ~160 American Kennel Club (AKC)–recognized breeds. A total of 622 dogs representing all identifiable coat phenotypes were genotyped at all three loci (table S10). By analyzing each of the three major traits both within and across multiple breeds, we show that combinations of these genotypes give rise to at least seven different coat types, encompassing most coat variation in modern domestic dogs (Fig. 3). Specifically, short-haired breeds display the ancestral state in all three genes. Wire-haired breeds, all of which have furnishings, carry the RSPO2 insertion. Dogs that carry both the RSPO2 and KRT71 mutations display “curly-wire” hair that is similar in texture to wire-hair but longer and curled or kinked rather than straight. Long-haired breeds carry the variant form of FGF5. Dogs carrying the FGF5 mutation, along with the RSPO2 insertion, have furnishings and long soft coats, rather than wiry ones. When dogs carry variants in both FGF5 and KRT71, the pelage is long and curly. Not surprisingly, coats must be of sufficient length to curl, and all curly haired dogs in our study were homozygous for the FGF5 mutation. Finally, if all three mutations are present, the phenotype is long and curly with furnishings.
Combinations of alleles at three genes create seven different coat phenotypes. Plus (+) and minus signs (−) indicate the presence or absence of variant (nonancestral) genotype. A characteristic breed is represented for each of the seven combinations …
None of the mutations we observed were found in three gray wolves or the short-haired dogs, indicating that short-haired dogs carry the ancestral alleles (table S1). Our finding of identical haplotypes surrounding the variants in all dogs displaying the same coat type suggests that a single mutation occurred for each trait and was transferred multiple times to different breeds through hybridization. Because most breeds likely originated within the past 200 years (27), our results demonstrate how a remarkable diversity of phenotypes can quickly be generated from simple genetic underpinnings. Consequently, in domesticated species, the appearance of phenotypic complexity can be created through combinations of genes of major effect, providing a pathway for rapid evolution that is unparalleled in natural systems. We propose that in the wake of artificial selection, other complex phenotypes in the domestic dog will have similar tractable architectures that will provide a window through which we can view the evolution of mammalian form and function.
We gratefully acknowledge support from NSF grants 0733033 (R.K.W.) and 516310 (C.D.B.), NIH grants 1RO1GM83606 (C.D.B.) and GM063056 (K.G.L. and K.C.), the Nestlé Purina company, the AKC Canine Health Foundation, the University of California–Davis Veterinary Genetics Laboratory, and the Intramural Program of the National Human Genome Research Institute. We thank L. Warren and S. Stafford for providing pictures. Finally, we thank the many dog owners who generously provided us with samples from their pets.
I sure hope I’m still on good terms with my guardian angel because I’m about to walk barefoot on hot coals. Now folks, before I move on, you need to know, I love my fellow breeder/exhibitors and am not, in any way, shape, or form finding fault with anyone’s breeding or dogs. What I do intend is to help newbies learn what more experienced breeders and judges see as they wade through a class of Gordon Setters or sort through a litter of puppies. So bear with me, and know that I’ve randomly chosen from a huge group of photos. I did the best I could to crop those photos to prevent identification, so if you spot your own dog and don’t like the way it looks…KEEP QUIET…you can pretend it’s not your dog and no one will be any wiser! Also, everyone needs to remember that this…
Thanks to Barbara Manson for sharing this article from the AKC website. Gordon Setters are not normally an anxiety ridden breed, however they can have their anxious moments just like every other dog, cat, pig, sheep or goat – not that we’re talking about cats, pigs, sheep or goats here, I just get to rambling sometimes.
As owners and breeders though, we do need to know how to read a dog’s body language, and we especially need to understand when we are seeing signs of anxiety so we can help to stabilize our dogs emotionally. We don’t want to be adding fuel to this fire!
We thought this article was a good starting place, but we also need you to share your advice, comments and suggestions to round it out, especially where your insight pertains to our Gordon Setters. Let’s give new and inexperienced Gordon owners a resource to guide them in building a calm, well behaved and socially adjusted dog. Thanks so much for contributing advice or stories in the comment section below.
Meanwhile you can read this article about anxiety below or follow the link embedded in the title to the AKC website for access to this and many other excellent articles.
Sally Gift, Mesa AZ
Feature Photo by Susan Roy Nelson, Casper, WY (This boy looks cool as a cucumber, obviously anything but anxious.)
If you or a loved one suffers from anxiety, then you know how difficult it can be to get through the day. What you might not know is that some dogs also suffer from anxiety.
Dog anxiety affects all breeds of dogs and can lead to serious behavioral problems if left untreated. Luckily, there are steps owners can take to help their dogs live with canine anxiety. Here are the symptoms, treatment options, and prevention techniques owners need to know about.
Fear-related anxiety can be caused by loud noises, strange people or animals, visual stimuli like hats or umbrellas, new or strange environments, specific situations like the vet’s office or car rides, or surfaces like grass or wood floors. These fears may seem inconsequential to us, but they create a lot of anxiety for dogs.
Separation anxiety is estimated to affect around 14 percent of dogs. Dogs with separation anxiety are unable to find comfort when they are left alone or separated from their family members. This anxiety often manifests itself in undesirable behaviors, such as urinating and defecating in the house, destroying furniture and furnishings, and barking.
Age-related anxiety affects older dogs and can be associated with cognitive dysfunction syndrome (CDS). In dogs with CDS, memory, learning, perception, and awareness start to decline, similar to the early stages of Alzheimer’s disease in humans. This understandably leads to anxiety in senior dogs.
SYMPTOMS OF ANXIETY
So how can you tell if your dog has anxiety? There are several important symptoms to look out for:
By far the most dangerous symptom of dog anxiety is aggression. This aggression can be targeted directly or indirectly, depending on the situation. Direct aggression occurs when a dog acts aggressively toward people or other animals. Indirect aggression can be equally dangerous, and often happens when a person comes between the dog and the source of the dog’s aggression, such as another dog. Even if a dog is prevented from harming others, aggressive behaviors such as growling or barking can lead to dangerous situations for humans and dogs, alike.
Urinating and defecating in the house is a common symptom of separation anxiety. Anxious dogs often work themselves up to the point that they pee or poop in the house, even if they are housebroken. This is frustrating for owners and can cause damage to property, not to mention the unpleasantness of the cleanup.
Destructive behavior is also common with separation anxiety. The damage is usually located around entry and exit points, like doorways and windows, but dogs in a state of heightened anxiety are also at risk of harming themselves. Attempts to break out of dog crates, windows, and even doors can result in painful injuries and expensive veterinary treatments.
Treating Dog Anxiety
The best way to treat anxiety is to talk with a veterinarian. She can help you identify the type of anxiety your dog suffers from and the possible causes and triggers. Veterinarians can also rule out any other medical conditions that could be causing your dog’s symptoms.
Your vet will help you come up with a treatment plan. Since anxiety is often caused by a variety of factors, the best way to treat it is usually through a combination of training, preventative strategies, and in some cases, medications.
TRAINING AND COUNTERCONDITIONING:
There are several training strategies dog owners can use to treat anxiety. One way is counterconditioning. The purpose of counterconditioning is to change your dog’s response to the stimuli responsible for anxiety, usually by replacing the anxious or aggressive behavior with a more desirable behavior, like sitting or focusing on the owner.
Another training strategy is desensitization. The owner slowly introduces the dog to the source of anxiety, preferably in small doses and at a decreased intensity. Repeated exposure and rewarding positive behavior can go a long way toward managing anxiety.
Some cases of anxiety are so severe that your veterinarian may recommend medications or natural therapies. SSRIs and antidepressants are occasionally prescribed for dogs with anxiety, including fluoxetine and clomipramine. For predictable anxiety-producing events like thunderstorms, fireworks, or car rides, your vet might prescribe a medication such as benzodiazepine in conjunction with an antidepressant to help your dog cope with the stress.
Senior dogs with cognitive dysfunction syndrome may benefit from the drug selegiline, which can help reduce some of the symptoms of CDS. Selegiline is also used for treating chronic anxiety in Europe.
The Merck Veterinary Manual also states that natural therapies and products can help dogs with anxiety. Some products work best in conjunction with other medications, while others can be used alone, depending on your dog’s case. Natural products use pheromones and aromatherapy to reduce anxiety. Talk to your vet about the natural products best suited for your dog.
Preventing Dog Anxiety
It is hard to predict if a pet will develop anxiety, but there are ways to help a new dog or puppy avoid anxiety-related problems.
One of the best things you can do is learn to read dog body language. Knowing when your dog is uncomfortable or scared can help you avoid negative experiences or use them as a positive training moment. Body language can also tell you when a dog is getting anxious, which is especially useful if your dog has a history of aggression-related anxiety.
Proper socialization can prevent the development of anxiety. Introducing your dog to new people, dogs, animals, places, and experiences can help them avoid an exaggerated response down the road, and also helps them become well-adjusted canine citizens.
Obedience training is an essential tool for preventing and managing anxiety. It lays the foundation of a healthy relationship and establishes trust. A well-trained dog is easier to socialize than a dog without training, and obedience classes are a great place for dogs to meet other dogs in a controlled environment.
Exercise and Nutrition
Regular exercise and stimulation are crucial for a dog’s development, physical, and mental well-being. A stimulated dog is less likely to pick up destructive behaviors, and good nutrition is equally important for your dog’s health. Making sure you take care of your dog’s physical and mental needs can help you prevent any behavior problems that don’t stem from anxiety, letting you know the areas where your dog needs the most help.
If your dog has been diagnosed with anxiety, you can also try to avoid or prevent situations that trigger your dog’s anxiety. For example, if you know that your dog grows anxious around large groups of dogs, you should avoid dog parks. Avoidance does not mean that you need to put your life on hold, but it can reduce some of the stress on you and your dog.
If the source of the anxiety cannot be avoided, preventative measures like leashes, body harnesses, and, in some cases, basket muzzles can prevent dangerous situations. Once you know your dog’s triggers, you can prepare for these situations ahead of time.
Take Action Now
Don’t let your dog’s anxiety take control of your life. With the right treatment strategy, you can help your dog overcome his anxiety and prevent dangerous and destructive situations from happening in the first place. If you think your dog might have anxiety, talk to your veterinarian today about a treatment plan that best fits your dog and your lifestyle.
We are dedicated to building a knowledge base and a sharing site for those who are involved in all of the various aspects of competition with Gordon Setters, competitions that showcase the Gordon Setter’s Beauty, Brains and Bird-Sense.