Tom D. Stodhill was a polarizing figure, and like most such men, had a knack for bringing out the worst in some people. Stodghill had a habit of making unfounded, controversial, and non-falsifiable statements like:

“…black and tan English Shepherds are recognized as the world’s most beautiful dog … the black and tan are the best working dogs and the most intelligent of all English Shepherds…the black and tan English Shepherd with long, glossy hair is the GENUINE Pre-Historic English Shepherd of long, long ago.”

– Tom D. Stodghill

Presumably, he intended for such statements to help him sell dogs.  But longer term, this behavior fostered a great deal of animus among other English Shepherd community members.  So much so that, after his death, many people wanted not only to forget him but his dogs as well.  Some of Stodghill’s writings, which I won’t quote here, even took a misogynistic slant.  As you might expect, this didn’t go over well with a good portion of the community.  Many reasons were used to mask the underlying animus for Stodghill: the dogs are too gritty; the dogs have bad hips; the dogs are inbred; the dogs are crossbred with other breeds.  The net result has been a dramatic decrease in black and tans over the past 40 years. 

But if we try to put emotions aside, and just deal with the facts, we might be able to see more clearly what remains of our black and tan English shepherd bloodline.  I have several black and tans, I’ve bred dozens and met at least a hundred decides.  They are not gritty or mean.  Their OFA hips results come back mostly Good – I’ve had one Fair.  They are not inbred, the dozen-and-a-half I have tested genetically (Embark) have an average Genetic COI of about 9%.  That’s barely above the breed average.

The creation of regional subisolates in dog breeding refers to the development of distinct subgroups within a breed that emerge due to geographical isolation or selective breeding practices concentrated in a specific region. These subisolates can develop unique characteristics due to the local breeders’ focus on certain traits, whether they are physical (like coat type) or behavioral (such as herding ability). Over time, these focused breeding efforts can lead to genetic, phenotypic, and behavioral differences that distinguish the subisolates from the main population of the breed. This process is influenced by factors like the breeders’ goals, environmental conditions, and the genetic diversity available within the regional breeding pool.

The collaborative breeding efforts started by Tom D. Stodghill, followed by John and Pauline Blankenship, and continued by decades of downstream breeders, especially in the southeastern United States, illustrate a significant case of bloodline development within the English Shepherd breed. The concentration of these breeding efforts in the south and southeastern United States suggests a regional influence on the sub-type’s development. Regional breeding practices such as those that occurred within the English Shepherd population in the mid-twentieth century can lead to variations within a breed, reflecting the specific needs, preferences, and environments of a particular area. This divergence is what creates a distinct sub-type, in a specific region, with unique characteristics that set it apart from other lines or types within the breed.

If you’d like to learn more about our black and tan English shepherd bloodline, you are invited to read my book:

The King of the Cattle Dogs: English Shepherds in Mid-Twentieth Century America

http://esbt.us/king ^[1]

A history of cultural use and appreciation

History

Historically, the old-fashioned Black and Tan English Shepherd dog was essential to the lives of many rural Tennesseans. But the utility of these all-purpose working dogs spread beyond the borders of Tennessee. In the Southeast, Black and Tan English Shepherds were well-known to farmers in Alabama, Mississippi, and Georgia. To the west, these gritty dogs were especially useful to cattle ranchers in Texas.^[1]Guy Gordon, “How the English Shepherd Became a Cattle Herder,” The San Francisco Examiner, December 1, 1957 To the north, Black and Tan English Shepherd dogs earned a living on farmsteads throughout the Ohio river valley. And further east, these dogs were well-known to rural citizens of the Mid-Atlantic region. But the history and use of these farm dogs is particularly well-documented in Tennessee.

• Watch a video about Black and Tan English Shepherd History ^[2]

An All Purpose Farm Dog

On dairy farms, cattle ranches and rural family homes, the Black and Tan English Shepherd provided a variety of essential services. Horace Curtis of Algood, Tennessee writes about his black and tans from the early nineteen-hundreds. He tells of their ability to eradicate vermin, protect children, and drive cattle.A Match for Man or Beast ^[3],” English Shepherd Club of America’s Who’s Who Breeder Manual, pg 22 At about the same time, John Blankenship’s father rode into town on his old gray mare. Charles Blankenship of Wilson County kept a black and tan English Shepherd at his side to drive home the livestock he purchased.The Blankenships’ Best Friend ^[4],” English Shepherd Club of America’s Who’s Who Breeder Manual, pg 19 Further east, Walter Oglivie of Allisona pushed his cattle through the thick woods of the Cumberland Mountains. He followed on horseback while a pair of black and tan English shepherds drove his stock over the narrow mountain paths. Upon reaching the rail head in Pikeville, Oglivie and his two dogs loaded the steers into train cars.Pete The Mountain Dog ^[5],” English Shepherd Club of America’s Who’s Who Breeder Manual, pg 25

John and Pauline Blankenship

It is difficult to overstate the influence of John and Pauline Blankenship on the Old-fashioned Black and Tan English Shepherd bloodline. Over the span of four decades, the Blankenships bred dogs on their family farm in Rutherford County.Sold on Shepherds ^[6],” The Nashville Tennessean Magazine, May 23, 1953 John and Pauline Blankenship were master farmers,^[6]“Rutherford Countians First Man, Wife Team in Plowing Contest”, Nashville Banner, September 22, 1956^[7]“Farmer Blankenship Will Plow for National Title,”, The Rutherford Courier, September 11, 1955 respected community leaders, and arguably the most prolific breeders^[8]“English Shepherd Training School Set in August,” The Rutherford Courier, July 21, 1953^[9]“Named Board Member of Shepherd Club,” The Daily News-Journal, August 10, 1953^[10]“English Shepherd to Observe Pioneer Days,” The Daily News-Journal, July 17, 1953 of English Shepherds in the history of the breed. The Blankenships’ dogs were selected and bred to bring in cows, herd sheep, catch and hold hogs, and load livestock into loading chutes.^[11]Murphy, op. cit. Advertised as “the world’s best all-purpose dog,” a Blankenship dog was also your faithful companion and an ardent night watchman. John Blankenship writes of a female black and tan English Shepherd named “Queen” whose ancestry could be traced back 100 years through a line of purebreds in Tennessee.^[12]Blankenship, op. cit. With his assertion, Blankenship establishes the existence of Old-fashioned Black and Tan English shepherds in Tennessee around 1850.

• Learn more about John and Pauline Blankenship ^[7]

Tom Stodghill

Mr. and Mrs. Tom Stodghill traveled to middle Tennessee in the early nineteen-fifties.Some Farm Dogs Said True English Shepherds ^[8],” The Rutherford Courier, January 23, 1953 Stodghill’s grandfather was a native of Rutherford County named George Dromgoole. Dromgoole used black and tan English Shepherds to work his cattle. So when Dromgoole moved his family to Texas in the late nineteenth century, he brought his cattle dogs along with him. Sixty years later, his grandson founded the English Shepherd Club of America in Quinlan, Texas. When Stodghill needed new blood for his English Shepherd breeding program, he returned to middle Tennessee. The Stodghills and Blankenships spent several days together, travelling the rural counties of DeKalb, Jackson, Putnam and Wilson. Their search bore fruit, finding middle Tennessee families which had been using these dogs for generations to manage their livestock. The markings, performance and behavior of the unregistered dogs led Stodghill and Blankenship to assert that they had been bred true down through the years. A number of these middle Tennessee dogs were registered and used to form the foundation of the Old-fashioned Black and Tan English Shepherd bloodline.All Fifty States ^[9],” English Shepherd Club of America’s Who’s Who Breeder Manual, pg 15

• Learn more about Tom Stodghill ^[11]

Old-fashioned Black and Tan English Shepherd Association

Today, the bloodline of the Old-fashioned Black and Tan English Shepherd is at risk. As lifestyles in America have shifted, these rare and historic dogs have dwindled in population. But fortunately, a few like-minded advocates have formalized as a Tennessee nonprofit corporation. The mission of the Old-fashioned Black and Tan English Shepherd Association is to assist with breeding, registration and training. OBTESA works to preserve the unique qualities of the bloodline, to preserve the integrity of the bloodline, and to protect the longevity of the bloodline. The OBTESA Breeder Incubator program identifies and supports select breeders who are committed to the revitalization of the Old-fashioned Black and Tan English Shepherd bloodline by providing mentorship, resources and quality breeding stock.

• Visit www.obtesa.com ^[12] for more information.

Next Steps

• Find a Black and Tan English Shepherd Puppy ^[13]
• Browse a photo gallery of Black and Tan English Shepherds ^[14]
• Read more articles about Black and Tan English Shepherds ^[15]

^[16]

This work (The Black and Tan English Shepherd, by Tony Bierman) is free of known copyright restrictions.

PI_HAT stands for “proportion IBD hat.” It is a statistical measure used in genetics to estimate the proportion of alleles shared identical by descent (IBD) between pairs of individuals.  The PI_HAT value ranges from 0 to 1, where 0 indicates no alleles shared IBD and 1 indicates complete allele sharing IBD. It provides an estimate of the genetic relatedness between individuals and is commonly used in studies of population genetics, pedigree analysis, and genetic association studies.  In the context of paternity testing, the PI_HAT value is often used to assess the probability of a father-child relationship by estimating the degree of genetic similarity between them. Higher PI_HAT values indicate a greater likelihood of a biological relationship, although additional factors and analyses are typically considered for conclusive paternity determination.

Using PLINK

We can determine the genetic relatedness of two dogs by processing their Embark Vet lab results using a free tool called PLINK.  The PLINK commands we’ll use look like this:

plink --dog --noweb --tfile alleged_father --make-bed --out alleged_father_data

plink --dog --noweb --tfile son --make-bed --out son_data

plink --dog --noweb --bfile alleged_father_data --bmerge son_data.bed son_data.bim son_data.fam --make-bed --out merged_data

plink --noweb --bfile merged_data –genome

The Results

The resultant plink.genome file created by the final PLINK command will contain several columns of data.  For our purposes, the PI_HAT, Z0, Z1 and Z2 are of interest.  A PI_HAT value above 0.7 is a likely father-child relationship, whereas a PI_SCORE in the mid-range of 0.5 is unlikely.  In a paternity scenario between a father and a child, you would typically expect to see the following ranges of values for the Z0, Z1, and Z2 columns in the PLINK .genome file:

Z0: The Z0 column represents the probability of sharing zero alleles IBD (identical by descent) between the father and son. In a father-child relationship, you would expect a relatively low value for Z0, indicating a low probability of not sharing any alleles IBD. This value is usually close to 0 or small compared to Z1 and Z2.

Z1: The Z1 column represents the probability of sharing one allele IBD between the father and son. In a father-child relationship, you would typically expect a moderate value for Z1, indicating a moderate probability of sharing one allele IBD. This value could vary but is generally lower than Z2.

Z2: The Z2 column represents the probability of sharing two alleles IBD between the father and son. In a father-child relationship, you would expect a relatively high value for Z2, indicating a high probability of sharing both alleles IBD. This value is typically the highest among Z0, Z1, and Z2.

Setting-up Your Computer

Here are the steps I use to run PLINK against Embark Vet data. 

Start by creating three folders:

c:\usr\bin

c:\usr\var

c:\usr\var\paternity

Next, download the PLINK 1.9 binaries and unzip them to c:\usr\bin.

Now, download the RAW DATA from Embark Vet for the alleged father and the son.  Unzip them in c:\usr\var.  Each .zip file will contain a .tped and a .tfam file.  For the alleged father, rename the files to alleged_father.tped and alleged_father.tfam.  For the son, name them son.tfam and son.tped.  Copy the 4 files to c:\usr\var\paternity.

From a command prompt, change to the c:\usr\var\paternity directory.  You’ll have to prefix each PLINK command with c:\usr\bin\ or add that location to your PATH environment variable.  Here again are the commands.

c:\usr\bin\plink --dog --noweb --tfile alleged_father --make-bed --out alleged_father_data

c:\usr\bin\plink --dog --noweb --tfile son --make-bed --out son_data

c:\usr\bin\plink --dog --noweb --bfile alleged_father_data --bmerge son_data.bed son_data.bim son_data.fam --make-bed --out merged_data

c:\usr\bin\plink --noweb --bfile merged_data –genome

Finally, analyze the plink.genome file with a text editor or import it to Excel.

In any breed population, genetic diversity is a key factor in ensuring the breed’s health and longevity. This diversity is often maintained through the presence of distinct bloodlines within the breed.

A bloodline represents the lineage of a dog, tracing its heritage back through many generations. Each bloodline can carry specific traits, abilities, and sometimes health issues, that make it unique. By maintaining distinct bloodlines, breeders can control certain traits in a breed, improve breed characteristics, and reduce the risk of genetic disorders, thereby improving overall breed health.

When we talk about preserving a healthy balance of distinct bloodlines, this means ensuring that no single bloodline dominates the breed. This is important because over-reliance on a single bloodline can lead to a lack of genetic diversity.

In addition, maintaining distinct bloodlines allows breeders to have options when planning matings. It gives them the flexibility to choose the most appropriate mate for a particular dog, based on factors like physical traits, temperament, and health status. This is beneficial for both the individual dogs and the breed as a whole.

For breed populations to be sustainably maintained, a concerted effort by all breeders is required to ensure diverse bloodlines. It’s not enough for one or two breeders to prioritize genetic diversity – it has to be a community-wide effort. Additionally, incorporating modern scientific tools like genetic testing can aid in preserving bloodlines and improving breed health.

Maintaining a balance of distinct bloodlines is a complex process but is crucial for the sustainable health and diversity of a breed. It’s a continuous process that requires careful planning, monitoring, and cooperation among breeders.

In the realm of canine studies, research into behavior—particularly non-pathological behaviors—proves intriguing for both dog owners and scientists alike (Serpell, 1995). These behaviors span a wide range from herding and pointing to more abstract qualities like temperament and trainability (Svartberg, 2006). A deeper understanding of the genetics behind these behaviors not only expands our knowledge about dogs but could also shed light on behavioral genetics in other animals, including humans (Overall, 2000).

Dog behaviors are complex traits, determined by a combination of potentially hundreds or even thousands of genes, coupled with environmental factors (vonHoldt et al., 2010). While the specific genes and mechanisms involved are not yet fully understood, studies have highlighted certain behaviors as largely influenced by genetics (Jones et al., 2008). Breed-specific behaviors such as herding in English Shepherds, fetching in Retrievers, and scent tracking in Hounds are thought to be primarily genetically determined. These traits were selected over generations of breeding to meet specific needs (Farrell et al., 2015). However, the genetic influence extends beyond these tasks and impacts a dog’s temperament, physical abilities, and even its trainability and intelligence (Hare et al., 2002).

A pivotal study by Jones et al., (2008) offers significant insights into the genetics of these behaviors. The research analyzed the genetic makeup of 2801 dogs across 147 breeds, using breed averages for specific behaviors and traits. They identified quantitative trait loci (QTLs) significantly associated with behaviors such as pointing, herding, boldness, trainability, and excitability. QTLs are regions of DNA associated with variation in a specific phenotype (observable trait), suggesting that these behaviors are not solely the result of environmental factors but are at least partially influenced by genetics. However, it’s important to emphasize that the identification of QTLs related to behaviors does not imply that these behaviors are entirely determined by genetics. Behaviors are complex traits, and while genetics set the stage, environmental factors and training play significant roles in shaping an individual dog’s behaviors (Scott and Fuller, 1965).

Looking ahead, the identification of intermediate phenotypes or endophenotypes could bring significant advancements (Gottesman and Gould, 2003). Endophenotypes are heritable traits or characteristics that act as an intermediary link between a gene and a complex phenotype. They often manifest as measurable biological markers seen in individuals with a particular genetic predisposition, even before the full-blown phenotype manifests (Gould and Gottesman, 2006). In the context of dog behavior, endophenotypes might manifest as variations in brain structure or activity patterns, hormone levels, or subtler behavioral indicators. As endophenotypes for behaviors are identified and understood, researchers may be able to better pinpoint the genes and genetic variants influencing different behaviors. This type of research may enable more targeted breeding programs if desired, and also could guide potential interventions, such as tailored training strategies or even medications, to manage or modify behaviors (Houpt, 2007).

The exploration of the genetic basis for non-pathological behaviors in dogs is an exciting field of study. As our understanding improves, it may well revolutionize the ways we train, breed, and care for our dogs.

References:

1. Farrell, L. L., Schoenebeck, J. J., Wiener, P., Clements, D. N., & Summers, K. M. (2015). The challenges of pedigree dog health: approaches to combating inherited disease. Canine Genetics and Epidemiology, 2(1), 3.
2. Gottesman, I. I., & Gould, T. D. (2003). The endophenotype concept in psychiatry: etymology and strategic intentions. American Journal of Psychiatry, 160(4), 636-645.
3. Gould, T. D., & Gottesman, II. (2006). Psychiatric endophenotypes and the development of valid animal models. Genes, Brain and Behavior, 5(2), 113-119.
4. Hare, B., Brown, M., Williamson, C., & Tomasello, M. (2002). The domestication of social cognition in dogs. Science, 298(5598), 1634-1636.
5. Houpt, K. A. (2007). Genetics of canine behavior. Acta Veterinaria Brno, 76, S11-S16.
6. Jones, P., Chase, K., Martin, A., Davern, P., Ostrander, E. A., & Lark, K. G. (2008). Single-nucleotide-polymorphism-based association mapping of dog stereotypes. Genetics, 179(2), 1033-1044.
7. Overall, K. L. (2000). Natural animal models of human psychiatric conditions: assessment of mechanism and validity. Progress in Neuro-Psychopharmacology and Biological Psychiatry, 24(5), 727-776.
8. Scott, J. P., & Fuller, J. L. (1965). Genetics and the Social Behavior of the Dog. The University of Chicago Press.
9. Serpell, J. A. (1995). The domestic dog: its evolution, behavior, and interactions with people. Cambridge University Press.
10. Svartberg, K. (2006). Breed-typical behaviour in dogs—Historical remnants or recent constructs?. Applied Animal Behaviour Science, 96(3-4), 293-313.
11. vonHoldt, B. M., Pollinger, J. P., Lohmueller, K. E., Han, E., Parker, H. G., Quignon, P., … & Wayne, R. K. (2010). Genome-wide SNP and haplotype analyses reveal a rich history underlying dog domestication. Nature, 464(7290), 898-902.

Social learning, the process by which animals acquire information or behaviors by observing others, is an important aspect of the development and behavior of many species, including domestic dogs. For domestic dogs, social learning can happen in a variety of contexts and through different mechanisms.

1. Observational Learning: Dogs can learn new behaviors by watching others, including other dogs and humans. For example, they can learn how to perform tricks or navigate obstacles by watching another dog or human demonstrate. In one study, dogs were shown to be capable of using humans as a social reference to solve an otherwise unsolvable task.
2. Imitation: Closely related to observational learning, imitation involves copying the behaviors of others. There is evidence that dogs can imitate human actions, even after a delay. In a process known as “Do as I Do” training, dogs are taught to imitate human actions on command, demonstrating a form of imitative learning.
3. Social Facilitation: The presence of conspecifics (members of the same species) can influence a dog’s behavior, even if the dog isn’t directly imitating the other’s actions. For example, dogs may be more likely to explore a new environment, play with toys, or eat certain foods when they see other dogs doing so.
4. Local Enhancement and Stimulus Enhancement: Local enhancement occurs when the presence of another individual at a particular location draws an observer’s attention, while stimulus enhancement is the increased attention to an object due to another individual’s interaction with it. Both these processes can lead dogs to explore new locations or objects, thereby facilitating learning.
5. Social Transmission of Avoidance: Dogs can learn what to avoid from the reactions of other dogs. For instance, if a dog sees another dog react fearfully to a certain object or situation, they may also learn to avoid it.
6. Mother-Pup Interaction: Puppies learn much of their behavior from their mothers, including social skills, foraging behaviors, and fear responses. This is particularly important in the early stages of life.
7. Human-Dog Interaction: Dogs can learn from humans through direct training, but they also pick up on subtle cues and behaviors through daily interaction. Dogs have been shown to understand human pointing gestures and even the direction of human gaze, both of which can guide their learning.

Social learning mechanisms in domestic dogs are varied and complex, and are likely influenced by a combination of genetic predispositions, early life experiences, and ongoing social interactions. While social learning provides a less costly way of acquiring new behaviors and information, it is important to note that the efficacy of social learning may depend on the specific context and the individual dog’s characteristics.

Introduction

Play, a cornerstone of canine behavior, is a multifaceted activity providing multiple benefits to dogs and their human companions. The high levels of play in dogs, a product of domestication and selection for playfulness, are integral to their physical, cognitive, and social development, and continue to be an enduring trait into adulthood. The core focus of this entry is to delve into the nature and nuances of play in dogs, examining its forms, its communicative aspects, and its implications for canine welfare.

Forms of Play

Play in dogs manifests in three primary forms: locomotor-rotational, object, and social play. Locomotor-rotational play involves movements like running, spinning, and jumping, stimulating a dog’s physical agility. Object play encompasses interactions with toys or other items, often reflecting predatory instincts. Social play, the most complex, involves interaction with other dogs or humans, facilitating social skill development. While the structure and motivations behind dog play with humans and conspecifics (dogs of the same species) differ, both serve to enhance their social intelligence and relationship-building capabilities.

Play Behavior and Behavioral Repertoires

The actions dogs perform during play often mirror those used in other contexts such as predation, fighting, and mating. These mimicry behaviors are crucial, providing dogs with a safe platform to practice and hone these skills, preparing them for real-life scenarios. The significant difference, however, lies in the consequences of these actions. In a real fight, for instance, a bite can lead to serious injury. In play, the bite is inhibited and harmless, emphasizing the importance of distinguishing between play and aggression, primarily for the safety and welfare of the dogs involved.

Communication in Play

To differentiate play actions from potentially harmful ones, dogs rely on ‘play signals’. These are explicit behaviors primarily used during play to communicate the dog’s playful intent. Two widely documented visual play signals in dogs are the play face – a relaxed, open-mouthed expression, and the play bow – a specific posture where the front end is lowered, and the rear end stays up. In addition to visual signals, acoustic play signals also hold relevance. These play sounds differ from their counterparts used during aggressive situations, further distinguishing play from potentially harmful interactions.

Metacommunication in Play

Play signals don’t just communicate the intent to play; they are a part of a metacommunication system where the signals define the context for interpreting other actions and behaviors. This concept, initially proposed by Bateson (1955), allows dogs to display actions that might seem aggressive in other contexts without escalating into actual aggression.

Strategies to Instigate Play

Dogs often resort to strategies such as self-handicapping and simple contagion to instigate and facilitate play. Self-handicapping involves a dog intentionally putting itself at a disadvantage to keep the play fair, such as a larger dog letting a smaller one ‘win’. This strategy helps maintain the positive, reciprocal nature of play, building social bonds, and learning social skills.

Simple contagion refers to the tendency of dogs to mimic the behavior of their peers. Observing other dogs play might entice a dog to join the fun, spreading the playful behavior like a ripple effect in a social setting. This ‘contagious’ nature of play underlines its role in strengthening social cohesion.

The Play Bow

One classic example of metacommunication in dog play is the “play bow.” This is a specific posture where a dog lowers its front legs and chest while keeping its rear end in the air. It’s a signal that’s almost universally recognized among dogs and is used almost exclusively in the context of play.

The play bow serves a dual purpose:

1. Solicitation: It is used to invite another dog to play. When a dog performs a play bow, it’s essentially saying, “I want to play with you.”
2. Metacommunication: It helps set or maintain the playful context for the subsequent actions. If a dog play bows and then performs an action that could otherwise be interpreted as aggressive (like a mock bite or a chase), the other dog understands that this is a part of play, not a real threat. The play bow is communicating about the nature of the interaction—”This is play, not a fight.”

So, in this way, the play bow serves as a metacommunicative signal, defining the context of the interaction and helping to prevent misunderstandings that could escalate into actual aggression.

Ontogenetic Ritualization

Ontogenetic ritualization is a process that explains how certain communicative behaviors can develop between two individual animals over time. Imagine two puppies from the same litter who spend a lot of time together. Over time, they begin to develop a unique set of play behaviors and signals that they consistently use when interacting with each other. For instance, one puppy might develop a unique wag of its tail that it only uses when it wants to play with its sibling, and the other puppy learns to recognize and respond to this signal. The signal is not a hardwired, instinctual behavior (see phylogenetic ritualization) — rather, it’s a product of their repeated interactions and mutual feedback. The consistent use of these specific signals in their interactions can be seen as an example of ontogenetic ritualization.

Conclusion

Play in dogs can take various forms, including locomotor-rotational, object, and social play. These types of play enhance physical agility, fulfill predatory instincts, and facilitate social skill development, respectively.

We discussed how play actions often mimic behaviors used in other contexts such as predation, fighting, and mating, but in a safe and harmless way. This is achieved through the use of play signals, which help differentiate playful actions from potentially harmful ones. Play signals include both visual cues, such as the play face and play bow, and acoustic signals.

We touched on the concept of metacommunication, where play signals also help define the context of interpreting other behaviors during play. This system allows dogs to display actions that might appear aggressive without escalating into actual aggression.

Strategies such as self-handicapping and simple contagion were discussed as methods dogs may use to instigate and maintain play. We also elaborated on the topic of ontogenetic ritualization, which explains how unique communicative behaviors can develop between individual animals over time.

Further Reading

Canine ontogeny, or the development of dogs from conception to adulthood, is a complex and intriguing process. It is characterized by a series of six “natural” stages: (1) the prenatal period; (2) the neonatal period; (3) the transition period; (4) the socialization period; (5) the juvenile period; and (6) the pubertal period. Understanding these stages is crucial in guiding strategies for training, socialization, and overall care for puppies, ensuring they grow into well-adjusted adult dogs. In this entry, I’m going to briefly describe each of these stages (aka “critical” or “sensitive” periods). At the end of the entry, I will provide you with recommendations for further reading.

1. Prenatal Period

The journey begins with the prenatal period, which stretches from conception to birth. During this time, the developing puppy relies entirely on its mother for nutrition and protection. The mother’s health and well-being can significantly impact the puppy’s development. Nutritional deficiencies, stress, or exposure to harmful substances during this time can have long-lasting implications for the puppy’s health and behavior.

2. Neonatal Period

Immediately following birth, puppies enter the neonatal period, which lasts until about two weeks of age. This period is characterized by limited sensory and motor capabilities. Newborn puppies are both deaf and blind, spending most of their time sleeping and nursing. Despite their limited interaction with the world, early tactile experiences and their mother’s care are still vital for their development during this stage.

3. Transition Period

The transition period is a short but significant stage, beginning around day 13 (plus or minus 3 days) and lasting until about day 18-20. During this phase, puppies undergo rapid sensory development. Their eyes and ears, closed at birth, begin to open, introducing them to the world of sights and sounds for the first time. This marks the beginning of a crucial period of sensory learning and development.

4. Socialization Period

From about four weeks to twelve weeks of age, puppies enter the socialization period. This stage is arguably the most critical period in a dog’s life when it comes to behavioral development. During this time, puppies become more aware of their surroundings and start to interact with other dogs and humans. They are highly receptive to learning about their environment, and positive experiences during this window can have a profound impact on their future behavior. Conversely, lack of exposure or negative experiences during this stage can lead to behavior problems later in life.

5. Juvenile Period

Following the socialization period is the juvenile period, which lasts until the onset of puberty (varying by breed and individual dog). Dogs start exploring their environment more during this stage, and their personalities continue to develop. Continued training and socialization are essential during this period to reinforce good behavior and prevent the development of unwanted behaviors.

6. Pubertal Period

The final stage of early development is the pubertal period, marking the onset of sexual maturity. This usually occurs between six months to a year, depending on the breed. Dogs will begin to exhibit adult sexual behaviors during this time, and females will experience their first heat cycle.

Conclusion

Understanding canine ontogeny is essential for dog owners, breeders, and trainers. It allows them to provide the necessary care, socialization, and training that’s appropriate to each stage of development, promoting the growth of healthy, well-adjusted dogs. Moreover, this knowledge contributes to our understanding of the profound and lasting effects of early life experiences on an individual’s development and behavior.

Further Reading

To understand why maintaining a balanced ratio of males to females in a breeding population is desirable, in this entry we’ll explore Mendelian inheritance and its implications for genetic diversity.

Mendelian inheritance refers to the principles of genetics discovered by Gregor Mendel. Mendel’s laws of inheritance explain how traits are passed from parents to offspring. The principles include:

1. Principle of Segregation: Every individual has two alleles for each trait, and these alleles segregate (separate) during the formation of gametes (egg and sperm cells), each gamete carrying only one allele for each trait.
2. Principle of Independent Assortment: The alleles of two (or more) different traits get sorted into gametes independently of one another.

The principles of segregation and independent assortment show that each individual has a unique combination of genes to contribute to the next generation. In order for this potential diversity to be fully realized in the offspring, there must be a wide array of mating possibilities, which is facilitated by a balanced sex ratio. By maintaining a 1:1 sex ratio in a breeding population, we maximize the potential combinations of alleles that can be produced, thus supporting genetic diversity and overall population health.

Given this common understanding of Mendel’s principles, here are reasons why a balanced 1:1 ratio of males to females is important for breeders:

Genetic Diversity: This is one of the most fundamental principles for the health of any breed. In dogs, it helps prevent the occurrence of inherited diseases. When there’s a balanced ratio of males to females, there are more potential combinations of genetic material. This means that there’s a better chance of producing healthy offspring and a lower risk of expressing harmful recessive traits.

Avoiding Inbreeding: In a small breeding population, or where males significantly outnumber females (or vice versa), there’s a higher risk of inbreeding, which can magnify the risk of harmful genetic conditions. A balanced ratio helps ensure a greater variety of potential mates, thus reducing this risk.

Population Management: A balanced 1:1 ratio helps breeders manage their population better. It’s easier to ensure that every dog gets a chance to breed and contribute to the next generation. Overproduction of one gender can lead to individuals not being used for breeding, thus their genes not contributing to the genetic diversity of the breed.

Enhancing Breed Traits: A balanced male-to-female ratio provides breeders with a broader genetic pool to select from when aiming to enhance specific breed traits, whether they’re physical traits (e.g. size) or behavioral traits (e.g. herding ability or temperament). This allows for a more nuanced selection process that can benefit the breed overall.

These points focus on a healthy breeding program that benefits both the individual dogs and the breed as a whole. Unbalanced ratios may lead to genetic bottlenecks, inbreeding depression, and an overall reduction in the health and resilience of the breed. Hence, for the good of their own breeding program and the breed as a whole, dog breeders strive to maintain as close to a 1:1 ratio of males to females in their breeding population as possible.

Proof

Let’s consider a sexually reproducing population with diploid organisms (i.e., organisms with two sets of chromosomes, like dogs) and two sexes. In accordance with Mendel’s principles, each parent contributes one allele of each gene to their offspring. If we have ‘n’ unique alleles in the population, each parent can contribute ‘n’ different alleles.

Suppose we have a population with a total of ‘P’ individuals, ‘M’ males and ‘F’ females, such that:

M + F = P

1. If the sex ratio is not balanced (say, skewed towards males), then even though males could contribute a variety of alleles, not all of these combinations would be utilized because there are fewer females to mate with. Each female may mate with multiple males, but this does not fully utilize the variety of alleles in the male population.
2. On the other hand, with a balanced 1:1 sex ratio (M = F = P/2), each male and each female has the potential to contribute their unique set of alleles to the next generation.

Therefore, the number of potential genetic combinations in the offspring generation is maximized when:

M = F

To illustrate this with a simple example, suppose we consider a single gene with two possible alleles (A and a). If we have two males and two females (a balanced sex ratio), with the following genotypes:

• Male 1: AA
• Male 2: Aa
• Female 1: aa
• Female 2: Aa

Then we can get the following combinations in the offspring:

• From Male 1 and Female 1: Aa
• From Male 1 and Female 2: AA, Aa
• From Male 2 and Female 1: Aa, aa
• From Male 2 and Female 2: AA, Aa, aa

Thus, all possible combinations (AA, Aa, aa) are represented. But if we remove one female (unbalanced sex ratio), we cannot achieve all possible combinations.

While this is a simplification, it helps demonstrate the principle at play. A balanced sex ratio allows for a greater diversity of genetic combinations in the offspring, which is beneficial for the overall health and adaptation of the population.

A point of clarification

Sometimes, the terms “breeding population” and the “number of dogs a breeder owns” are confused with one another. But, these terms refer to different concepts in the context of dog breeding.

Breeding Population: The breeding population refers to the subset of dogs a breeder owns or uses that are used for breeding purposes. These are the dogs that are physically mature, healthy, and have traits the breeder wants to pass on to the next generation. This population may consist of both males and females. The size of the breeding population can vary significantly depending on the breeder’s goals, resources, and ethical considerations. Outsourced stud dogs are considered part of a breeder’s breeding population, even though the breeder does not own these dogs.

Number of Dogs a Breeder Owns: This is simply the total number of dogs a breeder has in their care, regardless of their involvement in the breeding process. This includes not only the breeding population, but also dogs that are too young or too old to breed, retired breeding dogs, dogs that have been neutered or spayed, dogs kept as pets, and dogs that are not suitable for breeding due to health, temperament, or other reasons.

So, while there will likely be overlap between the two groups, they are not identical. The number of dogs a breeder owns is typically larger than the breeding population. Ethical breeders strive to maintain a number of dogs they can adequately care for and provide a high quality of life, and they responsibly manage their breeding population to ensure the health and well-being of the dogs and the quality of the puppies produced.

Conclusion

This entry discussed the significance of maintaining a balanced 1:1 male to female ratio in a dog breeder’s breeding population, based on principles of Mendelian inheritance. A balanced ratio helps maintain genetic diversity, avoid inbreeding, manage population, and enhance breed traits. The distinction between a breeder’s total number of dogs and the breeding population was also explained, with the breeding population referring to dogs actively involved in breeding and meeting certain standards. The number of dogs owned by the breeder includes all dogs in their care, regardless of their involvement in breeding.

Further Reading

As an interesting note, in Fisher’s Principle (aka Fisherian Sex Ratio named after the British statistician and geneticist Ronald Fisher), he hypothesized that a 1:1 sex ratio is evolutionarily stable. According to Fisher’s Principle, any deviation from a 1:1 sex ratio should be self-correcting over time due to the pressures of natural selection. Obviously, dog breeds are under the pressure of artificial selection. Nonetheless, Fisher’s Principle provides a foundational baseline from which to explore and understand sex ratio dynamics.

Play is a critical aspect of the behavior of domestic dogs. It’s not just an outlet for excess energy or a way to pass the time, but rather, a complex behavior with numerous implications for a dog’s physical health, mental well-being, and social relationships.

Play and Its Forms

Dogs engage in various forms of play, namely locomotor-rotational, object, and social play (Rooney et al., 2003). Locomotor-rotational play involves self-directed physical activities such as running, jumping, twisting, and rolling. Object play focuses on interactions with inanimate objects, such as toys or balls, while social play involves interactions with other dogs or humans.

Play Throughout the Lifespan

Interestingly, dogs, compared to many other species, maintain high levels of play throughout their lives. Although play peaks at around 8-9 weeks of age, it continues into adulthood at relatively high levels (Pal, 2010). This extended playfulness may be a result of selective breeding for playful traits or a product of neoteny, the retention of juvenile traits into adulthood. Both factors are thought to be byproducts of the domestication process.

Intrinsic and External Benefits of Play

Research suggests that play is not only intrinsically rewarding for dogs (Burghardt, 2005) but also brings about numerous immediate and long-term benefits. Play contributes to a dog’s positive welfare (Held & Spinka, 2011), reduces social tension (Arelis, 2006), and can even indicate a dog’s well-being (Boissy et al., 2007). Play also fosters a successful dog-human relationship, promoting bonding and understanding between the two (Rooney & Bradshaw, 2003).

Implications for Canids in General

While dogs are a focus due to their close relationship with humans, play behavior also extends to wild canids. Even captive adult wolves have been observed to engage in play (Cordoni, 2009). This underscores the possibility that play has intrinsic benefits for canids in general, beyond just being a “luxury” activity.

Understanding and appreciating the importance of play in dogs not only enhances the quality of life for our dogs but also further deepens our bond with them.

References