|
Development
of Behavior: Evolutionary Background - Normal Dog Behavior
Karen L.
Overall, MA, VMD, PhD, DACVB, ABS Certified Applied Animal
Behaviorist
To understand
canine behaviors and breeds it is critical to remember that,
fundamentally and ancestrally, the story of canine behavior is
that of working with humans at many tasks. Dogs are not just
altered wolves: as part of the co-evolution of our
co-domestication they became their own, and a very different
species. Listen to them and look at them to know this is true.
Outline of
canine social systems:
1. "alpha" (caution for
uncritical use of this terminology) male and female usually
older or larger
2. familial "pack" (caution for uncritical use of this
terminology) system
a. "alpha" animals are primary (or only) breeders
3. "season" may differ from domestic animals:
a. 1-2 times per year (once with African Wild Dogs)
b. doesn't start until 1-2 years of age
4. deference based system - may be controlled by oldest female
5. family groups in large packs for larger canids - wolves
6. monogamous small packs for small canids - foxes
7. cooperative breeding
8. cooperative nurturing
9. similar evolutionary strategies driving canid and human
social systems: needs for protection and cooperative hunting/foraging
10. truly omnivorous
11. regurgitation of food to young - facilitation of social
ties
12. gestation averages 63 d (49-54 d)
a. litters 2 (Fennec Foxes) - 13 (wolves, coyotes, racoon
dogs)
b. frequency and size of litters may be affected by
density dependent population regulation
13. importance of signaling behavior: facial, ears, tail, ruff
14. sexual dimorphism variable - less pronounced in monogamous
individuals
a. size
b. color
c. secondary characteristics (tail, ruff)
15. importance of vocal signals
16. defecation and urination as olfactory / territorial cues
17. social development
a. born blind
b. eyes open about 10-14 d.
c. concept of risk assessment and reduction of cost of
error
1. neonatal period: d. 0-14
2. transition period: d. 14-20
3. "socialization" period - better thought of as developmental
or sensitive period: begins about week 3
a. weeks 3 - 8: other dogs
b. weeks 5/7 - 12: people
c. weeks 5 - 12/16: novel circumstances
4. juvenile period: through 6-9 months of age
5. fearful potential most apparent: 10-12 weeks of age (as
early as 8;
peaks at 11)
6. substrate preference for elimination: 8.5 weeks
The Role of Canid Genetics / Breed
1. consider the purpose for which breeds were chosen
2. consider the decrease in additive genetic variance
3. consider a normal distribution in a "wild" situation
4. critical to distinguish between contextually appropriate and
inappropriate behaviors
5. probability that inappropriate behavior will reflect purpose
for which they were artificially selected
All behavior has environmental and
genetic components. The variation in the genetic component is
sufficient to produce a wide array of individual behavioral
phenotypes in the absence of any specific breed. Hence, not all
domestic long-haired cats or mixed breed dogs look alike or
demonstrate identical behaviors in response to like situations.
This is true within and between litters. Evolutionary biologists
since and including Darwin have recognized these differences.
The extent to which behavioral plasticity is a function of
genetics is a hotly debated issue in the fields of behavioral
ecology and evolution. Some recent experimental evidence on
desert toads living in highly variable environments indicates
that selection may be operating to maintain developmental and
concomitant behavioral plasticity, rather than selecting one or
a few modes, each of which would persevere under alternate
conditions. These issues are important and relevant when one
discusses the issue of breeds.
One function of
establishing and maintaining a breed is to canalize some of this
overall genetic variation. Although domestic canine breeds have
a relative body size that spans two orders of magnitude, such
variation in size is absent in wild canids. The process of
domestication, alone, relieved many of the pressures for which
wild canid body size was a response, allowing the underlying
genetic variation to respond to artificial selection. In the
process of selecting for certain physical and behavioral traits
within any breed, one has also selected for some variation in
that trait. Accordingly, when one discusses what have been
considered breed specific behaviors, caution is urged. Selection,
natural or artificial, cannot act if there is no underlying
genetic variation. Some variation (termed additive genetic
variance by quantitative geneticists) must be present for a
trait to be developed.
This is
extremely easy to visualize if one is considering a physical
trait such as coat or feather color. It is less easy to see that
selecting for a behavioral trait, such as protectiveness, which
is really a constellation of behaviors, is going to produce a
continuum of protective behaviors, some of which will not be
what the selector desired. In fact, some of these behaviors are
going to be inappropriate, because they are not complete or
forceful enough, and some will be unacceptable because they are
too forceful and out of context. Under natural, instead of
artificial, selection these behaviors would have been selected
against at their extremes; however, it would be an error to
regard the wild environment as producing absolute phenotypes.
The demographic
and local climatic environments act in concert to determine what
scope of the continuum of variation will survive. In a very good
year even the most inept hunter might live to reproduce and
contribute genes to the next generation. This is the source of
the additive genetic variance. That such genetic variance exists
is demonstrated by the extent to which artificial selection has
developed so many and such varied breeds in a few hundred years,
while thousands of years of natural selection has not developed
that degree of canine variability, although the initial stock
should have been similar.
Hence, if one
has developed a breed for certain specific behaviors (rather
than overall survivability, as in the wild situation), one
should expect that there will be variation around that behavior
and that some of this variation will result in inappropriate,
out of context behavior. It is in this light that charges made
about breed predilections should be viewed. This means that if
one has selected a breed for protectiveness or guarding, some of
the individuals in that breed may inappropriately protect or
guard against objects which pose no threat. Some believe that
herding behavior was developed from the first phases of
predatory behavior. It is conceivable that unless selection were
extremely discrete, sufficient variation should exist so that
the occasional herding animal exhibits inappropriate predatory
behavior. This is commonly known by sheep ranchers. Such
concepts of genetic variability in the development of behaviors
are difficult, but given the amount of misinformation regarding
some breeds and prejudice towards others, the issue should be
addressed.
Attempts have
been made to group breeds of dogs according to certain
constellations of behavioral attributes. Although their
classifications are based on subjective opinions (which can
invoke prejudice and folklore) rather than objective
classifications of individual behaviors, and their categories
confound discrete behaviors and behavioral diagnoses, some
overall patterns of behaviors correlated with breeds are
apparent. Cluster analysis grouped animals according to
reactivity, "trainability", and aggression. It is no surprise
that high trainability characterized most working and guard dogs;
prior to selecting for any specific other behavior the ability
to work with and be trained by, people would have to be
elaborated. Less reliable are the characterizations of
aggression and reactivity, since these are both diagnoses and
descriptions of amalgam behaviors.
More recent
studies have attempted to focus on specific behaviors (growling
when disturbed while sleeping; stalking small animals, barking
at approaching strange people, et cetera). This is important
because there is scant documentation of the frequency, duration,
intensity, and pattern of occurrence of the actual behaviors
that are involved in behavioral problems. It is only in this
context that fair evaluations of breed related behaviors should
be made. Furthermore, because so little is known about normal
behavior and behavioral precursors of serious problems, early
signs are not recognized. A survey of the faculty and senior
students at VHUP, and local practitioners revealed that
virtually all individuals thought that there were more and less
aggressive breeds and could rank these, the ranks of the three
survey groups were different within and between groups with no
rank being statistically significant, and, with the exception of
the majority of the students, few individuals in the other two
survey groups could, when provided with a list of discrete
behaviors, accurately identify those that were outright
aggressive or precursors of future aggressive behavior.
In summary,
caution is urged regarding any generalizations about breed based
behaviors. It is best to view selection for specific behaviors
as a risk assessment analysis: breeds that have been selected
for one or a few particular and specific behaviors may be more
at risk for developing unsavory variation for those behaviors.
This does not mean that dogs selected for protective behaviors
are more aggressive than dogs for which this selective pressure
was absent. It does mean that that breed may be more at risk for
developing a disproportionate number of dogs who exhibit
inappropriate, out of context protective aggression. Inherent in
this concept is that any dog, regardless of breed, can also
exhibit the inappropriate behavior. A further corollary is that
dogs who are selected for tenacity and jaw strength in their
in-context work (Bull Terriers, Rottweilers, Rhodesian
Ridgebacks), will, when they respond inappropriately or
out-of-context in another behavioral setting, exhibit this same
tenacity. Coupled with the physical traits attendant with such
selection (large jaws, heavy musculature), they can and will do
large amounts of damage on a first strike. These factors, rather
than increased breed-specific aggression, are the cause of the
severity of wounds, when inflicted. |