Maternal Care and Early Life in Dogs: How the Mother Shapes the Puppy

1. Why the First Weeks Matter

1.1 A Puppy Born Immature

A newborn puppy is a dependent animal. Its eyes and ear canals are closed, it cannot regulate its body temperature well, and it relies on its mother for food, warmth and, in the first days, even for the reflexes that trigger urination and defecation. Over the following weeks the senses open, the puppy begins to move purposefully, and contact with littermates, the environment and people increases. Researchers who study maternal care in dogs often start from this point: because the nervous system is still immature at birth, the early environment has an unusually good chance of influencing how it develops (Foyer et al., 2016).

For most of these weeks, the mother is the largest and most constant part of that environment. It is plausible that differences between mothers translate into differences between their puppies. Whether they do, how strongly and in which direction is an empirical question, and the answer is more interesting than the expectation that more care simply produces better dogs.

The first weeks also overlap with the beginning of the period in which puppies are especially receptive to social and environmental experience, a period that is covered in more detail elsewhere (how the sensitive period shapes the puppy brain). Maternal care sits at the start of this development. It is not the whole story, and as later sections show, what happens after the puppy leaves the litter also matters.

1.2 What "Maternal Care" Means in Research

In everyday language, a "good mother" is one who is attentive, protective and calm. Research has to be more specific. Studies of maternal care in dogs usually measure observable behaviors: how much time the mother spends in the whelping box, how much time she spends in body contact with the puppies, how long she nurses and in which position, and how often she licks and grooms them (Santos et al., 2020). Some studies add anogenital licking, sniffing and nudging, or the mother's orientation toward or away from the litter.

These measures are combined into a score in which more time with the litter, more nursing, more licking and more contact mean a higher value. This treats more of each behavior as more care. As the guide-dog studies show, the style of care may matter in ways that a simple sum does not capture.

1.3 Where the Evidence Comes From

Controlled research on maternal care in dogs is recent and comes from a small number of settings. The most detailed studies were carried out in breeding programs for working dogs: a Swedish program for military dogs (Foyer et al., 2016) and an American guide-dog school (Bray et al., 2017a, 2017b). A second line of work comes from Italian research groups that observed laboratory-bred Beagle litters and, later, litters reared in family homes (Guardini et al., 2016, 2017). Further evidence comes from large owner questionnaires that ask about a dog's puppyhood years after the fact (Tiira & Lohi, 2015).

Each setting has strengths and limits. Working-dog programs offer standardized records and outcome tests but involve a few breeds selected for specific tasks. Laboratory litters live in conditions unlike a family home. Owner surveys reach thousands of dogs but rely on memory. Newer studies broaden the evidence base. A 2025 study of 27 litters in a detection-dog program confirmed that maternal care declined across the first three weeks while individual differences among dams remained relatively stable; lower maternal care was associated with a greater chance that offspring would later be selected for operational work (Montgomery et al., 2025). A 2026 study following 59 litters found associations among maternal care, rearing environment and selected puppy behavior and cognition measures, with some associations extending to owner reports through 16 months of age (Bray et al., 2026). These findings are still observational and population-specific.

Reviews of early life experience in dogs have brought maternal care, attachment formation and socialization together as interconnected influences on later behavior problems (Dietz et al., 2018). Throughout this article, findings are treated as results from particular populations, and the question of how far they transfer to puppies bred in private homes is raised where it matters.

2. What Mothers Do

2.1 Nursing, Licking and Contact

The core maternal behaviors in dogs resemble those described in laboratory rodents: nursing, licking of the whole body and of the anogenital region, nest building and defense of the litter (Li, 2024). In the first days, licking stimulates elimination, body contact keeps the puppies warm, and nursing provides food.

Nursing itself is not a single behavior. In the guide-dog studies, three nursing positions were distinguished: lateral nursing, with the mother lying on her side; ventral nursing, with the mother lying on her stomach; and vertical nursing, with the mother sitting or standing while the puppies feed (Bray et al., 2017a). These positions place different demands on the puppies. Lying on her side, the mother makes the teats easily accessible. Standing or sitting, she makes the puppies work harder to reach and hold on. This distinction turned out to be relevant in the analysis of later outcomes, as section 4 shows.

2.2 The Later Weeks and Weaning

Maternal behavior changes over time. As the puppies become more mobile and begin to take solid food, the mother spends more time away from them, nurses less often and more briefly, and increasingly controls access to the teats. A recent review describes dogs nursing for roughly ten to thirteen weeks when left to themselves and notes that mothers may regurgitate food for their puppies in the period from about the fourth to the ninth week (Li, 2024). In breeding practice, puppies are usually weaned and placed in new homes well before the end of this natural span.

Most controlled studies have focused on the first three weeks, when maternal behavior is most intense and easiest to observe. The weaning period, in which the mother becomes more restrictive, has received much less systematic attention.

2.3 What Shapes a Mother's Style

Maternal behavior is not fixed by the mother alone. A review of maternal behavior in dogs lists litter size, breed, parity, meaning the number of previous litters, and contact with humans among the factors that influence it (Santos et al., 2020). The guide-dog data illustrate several of these. Labrador mothers scored higher on the maternal behavior component than German Shepherd mothers, mothers with smaller litters scored higher, and more experienced mothers scored lower than first-time mothers (Bray et al., 2017a).

The litter-size effect is intuitive: a mother with ten puppies cannot give each one as much contact as a mother with three. In the Swedish military-dog study, litter size was also related both to maternal care and to the offspring's later social engagement (Foyer et al., 2016). This creates a methodological problem that runs through the whole field: if large litters receive less care per puppy and also differ in other ways, for example in birth weight or competition between littermates, it becomes difficult to attribute any later difference to maternal care alone.

The parity finding is less intuitive. Experienced mothers provided less of what was measured (Bray et al., 2017a), which does not necessarily mean that their care was worse. They may simply be more efficient or tolerate more independence. The study cannot distinguish between these interpretations, which is a good example of why a higher score should not automatically be read as better care.

3. Measuring Maternal Style

3.1 From Video to a Single Score

The most detailed description of maternal style in dogs comes from a study of 21 mothers and their 138 puppies at a guide-dog school (Bray et al., 2017a). The mothers were German Shepherds, Labrador Retrievers and Golden Retrievers. Researchers analyzed 6,890 minutes of video from the first three weeks and coded seven variables, including time in the whelping pool, contact and licking per puppy, three types of nursing and the mother's orientation out of the whelping area.

A principal component analysis reduced these variables to one dominant component, which the authors called maternal behavior. It explained 54 percent of the variation in the data. In plain terms, mothers who scored high on one of the behaviors tended to score high on the others: they spent more time in the box, had more contact with each puppy, licked more and nursed more, particularly in the lateral position.

3.2 Is Maternal Style Stable?

For a measure of maternal style to be useful, it has to describe something consistent about a mother, not just a snapshot. In the guide-dog study, rank order was highly stable from week to week. The correlation between scores in the first and second week was 0.82, and between the second and third week 0.89 (Bray et al., 2017a). A mother who was among the most attentive in the first week tended to remain among the most attentive later.

The Swedish military-dog study reached a similar conclusion with a different sample. Video recordings of 22 German Shepherd mothers on days 1, 7, 14 and 21 after birth showed that the females differed consistently in their level of maternal care (Foyer et al., 2016). Two independent studies in different countries, breeds and programs thus agree that maternal style is a stable individual characteristic over the first weeks, at least within one litter.

Whether it is stable across litters is a different question. The guide-dog study observed only one litter per mother, and the authors explicitly noted that it remains unknown whether a mother would show the same style with her next litter (Bray et al., 2017a). Since parity itself was associated with maternal behavior, it is plausible that style changes with experience. For breeders, this matters: a mother's behavior with one litter is not necessarily a reliable prediction of her behavior with the next.

3.3 Validity and Physiology

The researchers also tested the component's validity. In the second week, mothers were given a choice between staying with their puppies and visiting a human experimenter. Mothers with higher maternal behavior scores tended to remain with their puppies, although the association only approached conventional statistical significance (p = 0.053). Mothers with high scores also showed marginally higher baseline cortisol and somewhat larger cortisol responses when separated from their puppies in the first week (Bray et al., 2017a).

These results suggest that a high maternal behavior score may partly reflect a mother who is more aroused or more invested in staying close, not only one who is more competent. Cortisol is an ambiguous measure, however, and the effects were small and marginal. Interpreting them as evidence of maternal stress or anxiety would go beyond the data (why stress measures in dogs need careful interpretation).

3.4 The Limits of Observation

All of these studies observe behavior for limited periods. Short recordings can miss rare behaviors such as anogenital licking (Li, 2024), and sampling schedules differ between studies, so scores are not directly comparable.

There is also a conceptual limit. Most studies measure maternal behavior at the level of the litter. A mother may treat her puppies unequally, and the puppies themselves differ in how actively they seek contact and food. The Swedish study explicitly noted that it could not track variation between individual puppies within a litter (Foyer et al., 2016). Litter-level scores are therefore an approximation of what each individual puppy experienced.

4. Maternal Care and Later Behavior in Working Dogs

4.1 Military Dogs in Sweden

The first study to link measured maternal care in dogs to adult behavior was carried out in the Swedish breeding program for military working dogs (Foyer et al., 2016). Twenty-two German Shepherd mothers were filmed with their litters during the first three weeks, and their offspring, 94 dogs in total, were later assessed in the standardized temperament test used by the Swedish Armed Forces at around 18 months of age.

The level of maternal care was significantly related to the offspring's adult behavior, mainly to behaviors grouped as physical engagement and social engagement, and also to aggression. More maternal care was related to more engagement in social activities with humans. The authors concluded that taking maternal quality into account in breeding programs might improve the selection of working dogs (Foyer et al., 2016).

The study connects observed maternal behavior to a standardized outcome more than a year later. Its limits are equally clear: one breed, one program, few mothers, and, as the authors stated, possible confounding by genetic factors. A mother who provides more care may also pass on genes that influence engagement.

4.2 Guide Dogs in the United States

The second major study followed 98 puppies from birth to adulthood in a guide-dog program (Bray et al., 2017b). Maternal behavior was coded from video during the first three weeks, using the same principal component described above. As young adults, the dogs completed a battery of temperament and cognitive tests. The outcome was whether they succeeded in the program or were released. In this population, 66 of the 98 dogs succeeded.

The results contradicted the intuitive expectation. Higher overall maternal behavior was associated with program failure, not success: dogs whose mothers scored high on the maternal behavior component had higher odds of being released (odds ratio 3.39). A high proportion of ventral nursing, with the mother lying on her stomach, was also associated with failure (odds ratio 4.22). In contrast, more vertical nursing, with the mother sitting or standing so that the puppies had to work harder to feed, was associated with success (odds ratio 0.25 for release) (Bray et al., 2017b).

In the young-adult tests, poor performance on a multistep problem-solving task combined with perseveration, and a short latency to vocalize when confronted with a novel object, were also associated with program failure. The predictive accuracy of the models was moderate. A model based on maternal behavior alone reached an area under the curve of 0.664, a model based on young-adult performance 0.750, and the combined model 0.754 (Bray et al., 2017b). An area under the curve of 0.5 corresponds to chance, and 1.0 to perfect prediction.

4.3 Why More Care Is Not Always Better

The guide-dog authors interpreted their findings within a stress-inoculation framework: moderate challenges early in life may help young animals develop resilience, while very intensive care may reduce such challenges (Bray et al., 2017b). A mother who stands while nursing and does not stay in constant contact may expose her puppies to mild, manageable difficulty, such as having to search, compete and hold on. A mother who lies with them constantly may buffer them from it.

The Swedish and American findings are also not easy to combine. In the military-dog study, more maternal care was related to more social engagement, which is desirable in many working contexts (Foyer et al., 2016). In the guide-dog study, more maternal behavior was associated with a lower chance of success (Bray et al., 2017b). The populations, outcome tests and definitions of care differed, so the studies do not directly contradict each other. Together, they show that the relationship between maternal care and adult behavior is neither simple nor uniform. The idea that "more maternal care is always better" is not supported. A dog released from a guide-dog program may still be a well-adjusted family dog, and it is not known whether the same pattern would appear with other outcomes or in other breeding environments.

Maternal style also does not predict outcomes on its own. In the guide-dog study, young-adult temperament and cognition improved prediction more than maternal behavior did (Bray et al., 2017b). Early care is one influence among several, and its predictive value for an individual dog is limited (how temperament and coping styles develop).

5. Laboratory Litters and Family Litters

5.1 Beagle Puppies in a Laboratory Setting

An Italian research group observed maternal care in eight litters of laboratory-bred Beagles during the first three weeks after birth, recording licking, anogenital licking, nursing and mother-puppy contact during morning observations (Guardini et al., 2016). At 58 to 60 days of age, the puppies were tested in an arena test and an isolation test.

Higher maternal care was associated with more exploration and a longer latency to the first yelp in the tests, while lower maternal care was associated with more vocalization and destructive behavior during isolation (Guardini et al., 2016). In this setting, more care appeared to go with a more confident response to novelty and separation at eight weeks.

5.2 Puppies Reared in Family Homes

The same group repeated the approach with 12 litters reared in family homes, 72 puppies of different breeds and mixed breeds (Guardini et al., 2017). The arena test included an unfamiliar person, and the isolation test lasted three minutes. The pattern was different. The amount of maternal care was positively associated with some stress behaviors, such as whining, yelping and yawning in the arena, and standing, paw lifting and howling in isolation. It was also positively associated with interest in the unfamiliar person and negatively associated with exploration and individual play.

The authors suggested that these responses resemble those of young mammals separated from an attachment figure with whom they have a strong bond. They also noted that the difference from the laboratory study suggests that puppies' development is influenced by a combination of maternal behavior and the social and physical environment (Guardini et al., 2017). In a barren laboratory environment, maternal care may be one of the few sources of stimulation. In a family home, puppies receive many other inputs, and the role of maternal care may change.

5.3 What the Two Studies Show Together

Taken together, the two Italian studies show the same measure of care associated with different behavioral profiles depending on the rearing environment, consistent with early influences interacting rather than adding up.

Both studies were small, both measured outcomes at around two months of age, and both were correlational. Behavior in an arena or isolation test at eight weeks is a snapshot of a developing puppy. Whether these early differences persist into adulthood was not tested. More vocalization during brief isolation at eight weeks is not the same as separation-related problems in an adult dog (what is known about separation-related problems).

6. Owner-Reported Maternal Care

A different kind of evidence comes from large questionnaire studies. In a Finnish survey of 3,264 family dogs, fearful dogs had fewer socialization experiences (p = 0.002) and a lower owner-reported quality of maternal care (p < 0.0001) during puppyhood (Tiira & Lohi, 2015). The sample size is large, and the association is strong in statistical terms.

The limits are those of all retrospective owner reports. Most owners saw the mother only briefly, and their judgment may be colored by the dog's later behavior: an owner of a fearful dog may be more inclined to remember the mother as nervous. The study shows that owner-reported maternal care and fearfulness go together, not that one caused the other.

7. Mechanisms: What Rodent Research Suggests

7.1 The Rat Model

The idea that maternal care can program offspring stress responses comes mainly from research in laboratory rats. In rats, mothers differ naturally in how much they lick and groom their pups. Offspring of mothers who lick and groom more show differences in DNA methylation of the glucocorticoid receptor gene in the hippocampus and differences in adult stress responses (Weaver et al., 2004). A review of this line of research describes it as environmental programming of stress responses through DNA methylation (Meaney & Szyf, 2005).

These studies show a plausible pathway from maternal behavior to lasting changes in the stress system, and the rodent literature uses experimental tools that dog research lacks, such as cross-fostering, which separates the effects of care from inherited traits.

7.2 Why Rodent Findings Cannot Be Transferred Directly

Rats and dogs differ in development, social structure and life history. A recent review that proposes a research agenda for maternal behavior in dogs based on rodent work emphasizes that, in dogs, hormonal mechanisms have been described but not tested experimentally, and that the neural basis of maternal behavior is largely unknown (Li, 2024).

The rodent work therefore provides a hypothesis for dogs, not a finding. When dog studies discuss "stress programming" or "epigenetic effects of maternal care", they borrow a mechanism from rats. Whether the same mechanism operates in dogs has not been shown (how the stress system responds to long-term strain).

7.3 What Has Been Measured in Dogs

Canine epigenetic research is still at an early stage. One small study reported that early life history in dogs was associated with methylation of the glucocorticoid receptor and oxytocin receptor genes, cortisol levels and attachment styles (Awalt et al., 2024). Another found an association between peripheral methylation of the oxytocin receptor gene and social behavior in pet dogs (Cimarelli et al., 2017). Both studies measured methylation in peripheral tissue, not in the brain, and neither measured maternal care directly.

These studies show that methylation differences can be detected in dogs and that they vary with early life history and behavior. They do not show that maternal care in dogs produces specific epigenetic marks that then change behavior. The chain from maternal licking to methylation to adult behavior has not been demonstrated in dogs (what epigenetic research in dogs can and cannot show). The oxytocin system is often named as a link between early social contact and later bonding, but in dogs this remains a hypothesis (what oxytocin does in dogs).

8. Genes, Care and Confounding

8.1 The Central Problem

Every observational study of maternal care in dogs faces the same difficulty. Puppies inherit half of their genes from their mother. If a mother's caring style is partly influenced by her own temperament, and if temperament is partly heritable, then an association between maternal care and puppy behavior may reflect shared genes rather than the effect of care. Both the Swedish and American authors acknowledged this explicitly (Foyer et al., 2016; Bray et al., 2017b).

8.2 Heritability and Prediction

Research on heritability in dogs shows why the genetic explanation must be taken seriously. In a large Swedish dataset of 1,235 German Shepherd puppies tested at eight weeks, several tested behavior traits were heritable, yet the correspondence between puppy test results and adult performance was negligible (Wilsson & Sundgren, 1998). Heritability and predictability are different things. A trait can be partly inherited and still be difficult to predict from early behavior (what puppy tests can predict).

The same caution applies to maternal care. Even if maternal care contributes to later behavior, the contribution is one of many, and it interacts with genes and later experience. A study of 4,497 dogs illustrates this interaction: adverse experiences in the first six months, including separation from the primary caregiver, were associated with more aggression and fear in adulthood, and the strength of this association varied between breeds (Espinosa et al., 2025). Early environment and genetic background do not act separately (how breed and individual behavior relate).

8.3 What Would Settle the Question

The classic method for separating genetic and maternal effects is cross-fostering: puppies from one mother are raised by another. If puppies resemble their foster mother's style more than their biological mother's, care has an effect beyond genes. The guide-dog authors suggested cross-fostering studies as the next step (Bray et al., 2017b). Cross-fostering is difficult in dogs, because litters must be born at almost the same time and not every mother accepts foreign puppies.

Until such studies exist, the most accurate description of the current evidence is that maternal care in dogs is associated with later behavior in specific populations, that a causal effect is plausible given rodent research, and that genetic confounding has not been excluded.

9. Beyond the Mother: Early Handling and Structured Programs

9.1 Early Gentling

Maternal care is not the only early experience that has been studied. Experimental studies have tested the effects of brief, gentle human handling during the first weeks. In one study, 43 puppies from seven litters were divided so that half of each litter received daily gentle handling from day 3 to day 21 after birth, while the other half did not. Some litters were raised in a professional breeding kennel and the others in family homes. At eight weeks, the puppies were tested in an isolation test and an arena test (Gazzano et al., 2008).

Handled puppies explored more than non-handled puppies: 119.00 seconds and 97.12 seconds in the handled kennel and family groups, compared with 64.90 and 57.00 seconds in the non-handled groups. Handled puppies from the kennel showed a longer latency to the first yelp in isolation, 89.46 seconds compared with 13.10 to 45.90 seconds in the other groups (Gazzano et al., 2008). The authors concluded that early gentling had a positive influence on emotional development in both rearing environments.

Because each litter was split, handled and non-handled puppies shared mother and genetic background, so their differences cannot be explained by maternal care or genetics. The limitations are the small sample, the follow-up to eight weeks only, and the possibility that handling also changed how the mother treated the returned puppies.

9.2 Structured Early Socialization

A study in guide-dog puppies tested a standardized early socialization program from birth to six weeks of age. Puppies in the program scored better in an assessment at six weeks and, at eight months, showed less separation-related behavior, less distraction, less general anxiety and less body sensitivity in handler questionnaires (Vaterlaws-Whiteside & Hartmann, 2017). This is one of the few studies in dogs that followed an early intervention beyond the litter period.

Together with the gentling study, it suggests that what breeders do in the first weeks can make a measurable difference. Both studies were small and carried out in specific populations, and neither identifies a single optimal program.

9.3 Learning from the Mother

Mothers may also influence puppies through example. In a study of working dogs, observational learning of a behavior pattern that the mother had acquired through training was investigated in her puppies (Slabbert & Rasa, 1997). The study belongs to a broader body of research showing that dogs can learn by observing others (how dogs learn from observing others). For maternal care, it suggests that the mother's influence may not be limited to nursing, licking and contact.

10. Leaving the Litter: Age at Separation

10.1 Early and Later Separation

The age at which puppies leave their mother and littermates is one of the few early factors that buyers can easily find out. An Italian study compared 70 adult dogs that had been separated from the litter at 30 to 40 days with 70 dogs separated at two months (Pierantoni et al., 2011). According to owner reports, the odds of destructiveness, excessive barking, fearfulness on walks, reactivity to noises, toy possessiveness, food possessiveness and attention-seeking were significantly greater in the dogs separated early. Early-separated dogs from pet shops showed particularly high frequencies of problem behaviors.

A more recent study of 107 dogs adopted at ages ranging from one month or younger to four months or older found that puppies adopted at one or two months showed higher fear, anxiety and attachment or attention-seeking scores on a standardized owner questionnaire than those adopted later (Cocco et al., 2025). This study was small, and the adoption age groups overlap with differences in source and early environment.

10.2 Interpreting Separation Age

The Italian findings are consistent with the view that very early separation, at around one month, deprives puppies of important experiences with their mother and littermates. They do not establish a specific optimal age. The early and later groups may also differ in other ways: puppies that leave very early may come from sources in which other aspects of early care are also poorer. A systematic assessment of studies on early experience and human-directed aggression likewise identified the source of the dog and the age at which it was rehomed among the relevant factors, but stressed the methodological limits of the underlying studies (Baslington-Davies et al., 2023).

11. Commercial Breeding and Early Environment

11.1 Pet-Store Puppies

A large questionnaire study compared 413 dogs obtained as puppies from pet stores with 5,657 dogs obtained from noncommercial breeders. Pet-store dogs scored significantly less favorably on 12 of 14 behavioral measures, including separation-related problems and house soiling (McMillan et al., 2013). The study is cross-sectional and cannot determine which aspect of the commercial background is responsible: genetics of dogs bred for the market, maternal care under commercial conditions, early separation, transport, the pet-store environment or the type of owner who buys from pet stores (why house soiling has several causes).

11.2 The Mothers in Commercial Breeding

A related study examined the mothers rather than the puppies. In 1,169 dogs formerly used as breeding stock in commercial breeding establishments, owners reported higher rates of social and non-social fear, house soiling and compulsive staring, and lower trainability, than in pet dogs in the general population (McMillan et al., 2011). These are the dogs that raise the puppies sold through commercial channels.

The study does not measure maternal care, and whether chronically fearful mothers under restricted conditions care for their litters differently has not been studied directly. The question links the welfare of breeding females to the early environment of their puppies.

11.3 Early Adversity More Broadly

The study of 4,497 dogs mentioned above also measured adversity such as intense fright or being chained outdoors (Espinosa et al., 2025). Its data come from owner reports and extend well beyond the litter period. For maternal care, its main relevance is that early adversity and breed interact, which cautions against simple, universal predictions.

12. After the Litter: What Comes Next Still Counts

A focus on maternal care can create the impression that a puppy's future is decided before it leaves the breeder. The evidence does not support this. In a guide-dog program, aspects of the environment between puppyhood and twelve months, such as the experience of the puppy raiser, the presence of another dog in the household and frightening experiences with people or dogs, were associated with aggression and fear at twelve months (Serpell & Duffy, 2016).

In a survey of around six thousand pet dogs, less socialization between 7 and 16 weeks had the strongest association with fear of unfamiliar people and dogs (Puurunen et al., 2020). Most of this period falls after the puppy has moved to its new home. What the new owner does in these weeks therefore matters as well, not only what happened in the whelping box.

Fear-related development also continues after the early weeks, and there are periods in which young dogs react more sensitively to frightening experiences (what is known about fear periods). Maternal care may influence the starting point, but later experience shapes the path, including the bond with new caregivers (how attachment between dogs and people develops).

13. What This Means for Breeders and Puppy Buyers

13.1 For Breeders

The research suggests that the mother's welfare and behavior are part of the puppies' early environment, and that mothers differ consistently in how they care for their litters (Foyer et al., 2016; Bray et al., 2017a). It does not provide a target level of maternal care, and the guide-dog findings warn against assuming that a mother who never leaves the box is ideal (Bray et al., 2017b).

In practice: In our experience, breeders benefit from observing the mother calmly over the first weeks rather than judging her from a single moment. A mother who leaves the box for short periods, returns reliably and nurses in varying positions is, from a practical point of view, not a cause for concern. Persistent restlessness, signs of stress or rough handling of the puppies are a reason to involve a veterinarian.

The studies on gentle handling and structured socialization indicate that the breeder's own contribution in the first weeks can add to maternal care (Gazzano et al., 2008; Vaterlaws-Whiteside & Hartmann, 2017).

In practice: From our work with breeders, we see that a calm environment for the pregnant and nursing mother, a whelping area where she can withdraw, and short, calm daily handling of the puppies by familiar people are easy to implement and fit the direction of the research, even though no study has tested this exact combination.

13.2 For Puppy Buyers

For buyers, the most important practical point is that maternal care cannot be assessed reliably in a short visit, and that no single observation predicts how an individual puppy will develop. What buyers can find out is whether the puppies were raised with their mother, at what age they will leave, and under what conditions the mother lives. The findings on early separation and commercial sources point in the same direction (Pierantoni et al., 2011; McMillan et al., 2013).

In practice: In our experience, it is worth asking to see the mother with her puppies and to observe how she behaves toward the litter and toward visitors. A relaxed, approachable mother is a good sign, but a mother who is cautious with strangers while she has young puppies is not necessarily a problem. What matters more is whether the overall picture is consistent: a stable home environment, puppies that are used to people, and a breeder who can describe the first weeks in concrete terms.

In practice: From our training work, we know that owners sometimes attribute every later difficulty to the breeder or to the mother. The research does not justify this. Early care is one influence, and the weeks and months after the puppy moves in also shape its further development, which is something owners can actively influence.

13.3 For Assessing an Individual Dog

Maternal history is rarely known for adult dogs and cannot explain an individual dog's behavior on its own. In behavioral assessments, the litter and the age at separation are background information to be weighed alongside genetics, later experience, health and current environment (how behavioral assessments are structured).

14. Summary at a Glance

Mothers differ consistently in maternal behavior over the first three weeks. In one guide-dog population, maternal style was summarized by a single component explaining 54 percent of the variation, was highly stable from week to week, and was associated with litter size, breed and parity (Bray et al., 2017a; Santos et al., 2020).

In military German Shepherds, maternal care was related to adult physical engagement, social engagement and aggression (Foyer et al., 2016). In guide dogs, higher overall maternal behavior and more ventral nursing were associated with program failure, and more vertical nursing with success (Bray et al., 2017b). In laboratory Beagles, more care went with more exploration at eight weeks, while in home-reared litters it went with more stress behaviors, more interest in a stranger and less exploration (Guardini et al., 2016, 2017).

Epigenetic programming by maternal care is established in rats but not demonstrated in dogs (Weaver et al., 2004; Awalt et al., 2024). Very early separation and commercial origin are associated with more owner-reported behavior problems (Pierantoni et al., 2011; McMillan et al., 2013), and experience after the litter remains strongly associated with adult behavior (Serpell & Duffy, 2016; Puurunen et al., 2020).

15. Research Gaps and Critical Appraisal

15.1 Small Samples and Few Populations

The core observational studies are based on 21 or 22 mothers each (Foyer et al., 2016; Bray et al., 2017a, 2017b), and the Italian studies on 8 and 12 litters (Guardini et al., 2016, 2017). Since maternal care varies between mothers, the effective sample size is the number of mothers, so effect estimates are uncertain and may not replicate elsewhere.

15.2 Narrow Breeds and Specialized Outcomes

The working-dog studies involved German Shepherds, Labrador Retrievers and Golden Retrievers in programs that select for specific traits. Their outcomes, a military temperament test and guide-dog success, are meaningful within those programs but not identical to what matters in a family home. The home-reared Italian study included various breeds but measured outcomes only at two months (Guardini et al., 2017). There is no study following home-reared puppies from observed maternal care to adult behavior in family life.

15.3 Confounding by Genetics

No study in dogs has used cross-fostering to separate maternal care from inherited traits. The authors of the key studies acknowledge genetic confounding explicitly (Foyer et al., 2016; Bray et al., 2017b). The within-litter design of the gentling study shows how genetic effects can be controlled for human handling (Gazzano et al., 2008), but no comparable design exists for maternal care itself.

15.4 Measurement

Maternal care is measured at the litter level, usually over three weeks, with sampling schedules that differ between studies. Rare behaviors may be missed, and care received by individual puppies is not captured (Li, 2024; Foyer et al., 2016). Composite scores assume that more of each behavior means more care, while the guide-dog results show that the type of nursing may matter independently (Bray et al., 2017b). The weaning period has been studied far less than the first three weeks.

15.5 Retrospective Owner Reports

Large studies on maternal care quality, separation age and commercial origin rely on owner questionnaires completed years later (Tiira & Lohi, 2015; Pierantoni et al., 2011; McMillan et al., 2013; Cocco et al., 2025). These designs can show associations in large samples but are vulnerable to recall bias and to confounding by source, owner characteristics and later environment.

15.6 Missing Topics

No primary canine study on prenatal maternal stress was included here, so the question of whether a mother's stress during pregnancy affects her puppies remains open for dogs. Equally, whether maternal style is consistent across successive litters, whether it can be changed by management, and whether the stress-inoculation interpretation holds outside guide-dog programs are all unanswered questions (Bray et al., 2017a, 2017b). Mechanistic work in dogs is limited to small studies of peripheral methylation (Awalt et al., 2024; Cimarelli et al., 2017).

16. Conclusion

The research on maternal care in dogs supports a clear general statement and resists a simple practical rule. Mothers differ consistently in how they care for their puppies, and these differences are associated with later behavior in military dogs, guide dogs and young family puppies. At the same time, more care is not uniformly better. In one guide-dog population, the most intensive mothering was associated with a lower chance of success, and in family litters more care went together with more distress in brief tests at eight weeks.

The evidence base is small, concentrated in a few breeds and programs, and unable so far to separate the effects of care from the effects of shared genes. Rodent research provides a plausible mechanism but not a canine finding. The most useful conclusion for breeders and puppy buyers is therefore not a target level of care, but a broader view: the mother's welfare, the conditions in which the litter is raised, the age at which puppies leave and the experiences they have afterward all contribute to how a dog develops, and none of them alone decides it.

Key Insights (Takeaways)

• Maternal style in dogs is measurable and was stable over the first three weeks within a litter (Bray et al., 2017a; Foyer et al., 2016).

• Maternal care has been associated with later behavior in working dogs, but the direction depends on the population and outcome studied.

• In guide dogs, more intensive mothering was associated with program failure, and nursing that required more effort from the puppies with success (Bray et al., 2017b).

• The same measure of care was associated with different behavior in laboratory and home-reared litters, which suggests that care interacts with the rearing environment (Guardini et al., 2016, 2017).

• Genetic confounding has not been excluded in any canine maternal-care study, and cross-fostering studies are missing.

• Epigenetic programming by maternal care is a rat finding; in dogs, only small studies of peripheral methylation exist (Weaver et al., 2004; Awalt et al., 2024).

• Very early separation and commercial origin are associated with more owner-reported problems, but these factors are intertwined with others (Pierantoni et al., 2011; McMillan et al., 2013).

• What happens after the puppy leaves the litter remains strongly associated with adult behavior (Puurunen et al., 2020; Serpell & Duffy, 2016).

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