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A. (Sorted by C.PC1)
B. (Sorted by C.PC2)
Trait
C37
Trait description
C.PC1
-86
C.PC2
Trait
Trait description
C.PC1 C.PC2
*
Aggressive sounds
8
14
12
8
4
12
20
-14
-24
-33
-41
-39
1
-26
29
24
4
10
19
13
14
20
14
20
-18
-15
0
-12
7
-10
C7
C38
C4
C35
C3
C13
C14
C55
C16
C15
C8
C50
C6
C36
C12
C37
C32
C204
C33
C31
C25
C18
C29
Observer can first touch fox in zones 5-6
-2
-41
**
C34
C31
C32
C36
C33
C30
C55
C3
C4
C7
C38
C50
C35
C39
C2
C6
C204
C19
C25
C18
C17
C29
C24
C13
C14
C8
C16
C12
C15
Follows the hand (aggr.)
Attack alert
Pinned ears (aggr.)
Triangle ears directed back (aggr.)
Trying to bite
Attack
Leans on back or side walls in zones 5-6
Fox is in zones 3-4-5-6 at the beginning of step C
Spends more than 30 seconds in zones 3-4-5-6
Observer can first touch fox in zones 5-6
Fox remains only in zones 3-5-6
Tail is up for at least for 3 seconds
Narrow ears directed back
Moved forward at least one zone during the step
Fox is in zones 1-2-3-4 at the beginning of step C
Observer can first touch fox in zones 3-4
Tame sounds (combined)
Comes into zone 2 at the end of step C
Wagging tail
Fox holds observer's hand with its mouth
Fox rolls onto its side, inviting observer to touch its belly
Comes to and sniffs observer's hand at the end of step C
Loud breathing
Fox allows observer to touch the rear part of its back
Fox allows observer to touch its back
Lies down during a contact for at least 5 seconds
Fox allows observer to touch its head
Tame ears
Fox allows observer to touch its nose
-84
-83
-56
-42
-33
-33
-5
-4
-3
-2
-1
-1
3
3
6
18
18
21
25
26
27
32
36
49
73
88
105
107
115
Fox remains only in zones 3-5-6
-1
-39
Spends more than 30 seconds in zones 3-4-5-
-3
-33
Narrow ears directed back
3 6
Fox is in zones 3-4-5-6 at the beginning of step C
-4
-24
Fox allows observer to touch the rear part of its back
49
-18
Fox allows observer to touch its back
73
-15
Leans on back or side walls in zones 5-6
-5
-14
Fox allows observer to touch its head
105
-12
Fox allows observer to touch its nose
115
-10
****
Lies down during a contact for at least 5 seconds
88
0
**
Tail is up for at least for 3 seconds
-1
1
Observer can first touch fox in zones 3-4
18
4
Triangle ears directed back (aggr.)
-42
4
Tame ears
107
7
***
Aggressive sounds
-86
8
*
Pinned ears (aggr.)
-56
8
Tame sounds (combined)
18
10
Trying to bite
-33
12
Attack alert
-83
12
Wagging tail
25
13
Fox holds observer's hand with its mouth
26
14
Comes to and sniffs observer's hand at the end of step C
32
14
C34
C19
C30
C17
C24
C2
C39
Follows the hand (aggr.)
-84
14
Comes into zone 2 at the end of step C
21
19
Attack
-33
20
Fox rolls onto its side, inviting observer to touch its belly
27
20
Loud breathing
36
20
Fox is in zones 1-2-3-4 at the beginning of step C
6
24
***
****
Moved forward at least one zone during the step
3
29
FIGURE 10.3
Comparison of behavioral traits contributing to the first two principal compo-
nents of silver fox behavior at test step C (C.PC1 and C.PC2). All observations from one step of
the test (Step C “Observer attempts to touch the fox”) are shown. Behavioral observations (traits)
contributing to C.PC1 and C.PC2 are ranked according to their loadings (eigenvalues) from prin-
cipal component analysis. The significances for each trait loading for C.PC1 and C.PC2 are
shown as the number of standard errors from zero (negative or positive) as established by boot-
strap trials. Note in particular the difference in ranking for the traits marked with asterisks.
C.PC1 and C.PC2 are very different behavioral gestalts. PC1 is comprised of traits that contrib-
ute to overall aggressiveness (*) or tameness (****), whereas PC2 represents traits that are
marked by either passivity (**) or activity (***).
definition, the phenotypes measured by PC1 (tameness vs aggressiveness)
and PC2 (bold vs shy) are not entirely unrelated, as PC2 can enhance the
observed expression of PC1. That is, if an animal is aggressive, passive
behavior will reduce the perceived expression of that trait whereas active
behavior will enhance the expression, and the same effect applies if an ani-
mal is tame. In backcross populations the distribution of behavior is skewed
toward the extreme of the recurrent parent, reducing the range of tame vs
aggressive behaviors. Under these circumstances, PC2 acts to increase that
range. We would therefore expect that whereas PC1 and PC2 are distinct
principal components in a matrix composed of all populations, they could be
correlated in particular backcross populations. This is in fact the case for the
backcross-to-tame populations (r
0.8). In contrast, in F2 populations
where the behaviors are more normally distributed,
0.75
5
this is not
the case
(r
0.06). It could well be, therefore, that the PC2 QTL on VVU12 in the
backcross-to-tame population reflects enhancement of the expression of PC1.
52
TRANSCRIPTOME ANALYSIS
A good example of an approach that can be applied for identification of
genes and regulatory elements involved in fox behavioral phenotypes is the
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