RELATIONSHIPS

AND INBREEDING
Dr N. Chikumba
Importance of studying relationships
and inbreeding
1. To determine the likeness between relatives.
2. To come up with inbreeding computations.
3. To gain knowledge of additive genetic
relationships that can be helpful in:
i. Selecting animals of superior genetic quality on the
basis of relatives records to keep in herds;
ii. Arranging matings to avoid high levels of inbreeding;
and
iii. Establishing lines tracing back to desirable animals.
Relationships
• Relationships refer to genetic connections existing
between definite individuals by virtue of their being
in possession of identical genes.

• There are generally two types of relationships:

i. Direct
ii. Collateral
Types of relationships
1. DIRECT RELATIVES – These are individuals which
have a direct line of descent such as:
i. Parent and progeny
ii. Grandfather and grandson
iii. Great grand mother and great grand daughter
• In this case all individuals are related to each
other on the allele donor-recipient basis i.e.
between ancestor and descendent.
Types of relationships
2. COLLATERAL RELATIVES – These are individuals
with one or more common ancestors but which
are not direct descendents.
Examples of Collateral relatives
1. PATERNAL HALF-SIBS : Calves born by the same
bull with different dams and these have an
average relationship of 25% = ½ x ½ = ¼
2. MATERNAL HALF- SIBS: Calves by the same dam
but different sires and these have an average
relationship of 25% = ½ x ½ = ¼
3. FULL SIBS (brothers and sisters): Calves by the
same dam and sire and these are related to each
other by 50%.
Collateral relatives
• Fraternal twins – they are related by 50% to each
other, 25% through the sire and 25% through the
dam, as are full sibs born at different times.
• Identical twins – are genetically alike, thus they are
100% related to each other, just as an animal to
itself.
Methods of computing relationships
between individuals
1. The Tabular method
2. The Path Coefficient
Characteristics of the methods
• They assume that the relationship between any
random pair of animals at some specified time in
the past is ZERO.
• This specified time period is called the BASE
PERIOD, and animals of this base period are
called the BASE POPULATION.
Computing additive relationships
using Path Coefficients
• axy will be used to denote the additive relationship
between relatives X and Y.
Path coefficients for Direct Relatives
• For example, if C, G, and K are direct descendents of A, the arrow
pedigree is:
•K G C A
I F B

• Individual A transmits ½ of its genes to C, thus the additive

relationship between relatives A and C, aAC, is
aAC= (½)1= ½ .
• The additive relationship, aAC, is equivalent to the probability that
a particular allele of individual A at each locus will be transmitted
to C.
• So the probability of an allele of A at each locus being transmitted
to C is ½.
Path coefficients for Direct
Relatives
• Animal C transmits ½ of its alleles to G; thus aAG =
(½ )(½ )= (½ )2= ¼
• implying that the probability that a particular
allele of A will be transmitted to G is ¼.

• Similarly, animal G transmits a sample ½ of its

alleles to animal K, thus,
aAK = (½)(½)(½)=(½)3=1/8
• Thus the probability that a particular allele of A will be
transmitted to K is 1/8
Rules for calculating the additive
relationship between direct relatives
• More than one path can exist between direct relatives e.g
F C A

G E D
1. Count for each path the number of intervening generations
from the ANIMAL to its DIRECT ANCESTOR.
For example, the number of generations for the 3 paths
between G and A are:
nGEA= 2,
nGEDA= 3
nGFCA= 3
Rules for calculating the additive
relationship between direct relatives
2. Compute
aAG = (½)nGEA + (½)nGEDA + (½)nGFCA
Thus:
aAG = (½)2 + (½)3 + (½)3 = ½
Path coefficients for Collateral Relatives
• Example: A pair of full sibs (C and D) having unrelated
parents (A and B)
C A

D B
• The probability that C will receive an allele from A is ½.
• The probability that D will receive an allele from A is ½.
• Thus C and D are related through A by:
½ x ½ = (½)2= 1/4.
Collateral relationship
• Similarly, C and D are related through B by:
½ x ½ = (½)2= ¼.
• These paths are independent so that the total relationship
of C and D through both common ancestors is ½ x ½ + ½ x ½
= ¼ + ¼ = ½ , thus,
aCD= ½ .
• The rule for calculating the relationship between collateral
relatives is to sum the relationships due to each
independent path.
• The sum of the relationships due to each independent path
listed is known as the RELATIONSHIP COVARIANCE.
Rules for calculating the relationship
between collateral relatives
• Thus, for full sibs the relationships are:
(1/2) nCA + nAD and (1/2) nCB + nBD
where:
• nCA is the number of steps from C to common ancestor A, and
• nAD is the number of steps from D to common ancestor A.

• Similarly, nCB and nBD are the number of steps from C to B

and B to D.
• Each pair of ‘n’s reflects the total number of steps from
C through the common ancestor to its relative D.
Rules for calculating the relationship
between collateral relatives
• Thus aCD = (1/2)nCA + nAD + (1/2) nCB + nBD = (1/2)1+1 +
(1/2) 1+1
=½
Combined Direct and Collateral
relatives

• Relatives can be both direct and collateral

simultaneously e.g the relationship between B and D
in the pedigree below:

B A

D
C
 Combined Direct and Collateral
relatives
• There is a direct relationship between B and D, which
is ½
• There is also collateral relationship between B and D
via the collateral path’
B A C D with a collateral relationship (1/2)3=
1/8.
• Thus, the total relationship, aBD = ½ + 1/8
= 5/8
INBREEDING
• Inbreeding refers to matings between related
individuals.
• Inbreeding coefficient: it is the probability that
both alleles at a locus are identical by descent.
• IDENTICAL BY DESCENT (IBD): alleles are said to be
identical by descent when the alleles are exact
replicas from a common ancestor.
INBREEDING
• The symbol for inbreeding coefficient of animal X is
Fx.
• The inbreeding coefficient of an animal is one- half
the additive relationship between its parents i.e.
Fx = (½)aSxDx
• Where aS D , is the additive relationship between the Sire
x x

(Sx) and the Dam (Dx) of individual X.

• The additive relationship of individual X to itself is:
axx = 1 + Fx
Computing relationships for inbred common
ancestors or direct ancestors using Path
Coefficients
• If a common ancestor is inbred, the method is
modified to account for the increased chance of
both alleles at a locus of the ancestor being
identical by descent (IBD).
• The modification requires the inclusion of the term:
1 + Fz
• where Z is the direct ancestor if the relationship is
between direct relatives,
OR
• the common ancestor in the path between relatives.
INBREEDING COEFFICIENTS
• If XS and XD are the parents of X, then

Fx = ∑[(1/2)n(1 + Fz)]
Where: Fx = inbreeding coefficient of X
n = number of arrows connecting the sire and dam
through a common ancestor.
Fz= the inbreeding coefficient of the common
ancestor.
INBREEDING COEFFICIENTS
• NOTE: If the common ancestor is not inbred, the
formula for calculating the inbreeding coefficient is:
Fx = ∑[(1/2)n]
Steps in calculating the inbreeding coefficient
using the Path coefficient method
1. Compute the additive relationship between the
parents of the individual by tracing the paths.

2. Multiply the additive relationship of the parents

by ½.
Rules for calculating additive
relationships for collateral relatives
• Example: Consider the following pedigree of full sib
matings and calculate the inbreeding coefficient for E,
i.e FE
E C A
G
F D B
SOLUTION
1. Identify the parents ( i.e the sire and the Dam) for
E, i.e the sire is C and the dam is D.
2. Compute the additive relationship between the
parents of E i.e
aCD= (1/2)n + n (1 + FA) + (1/2)n + n (1+ FB)
CA AD CB BD

If FA= 0 and FB= 0, then

aCD = (1/2)2 + (1/2)2 = 1/2

SOLUTION
• Compute FE :
FE = ½aCD
= (½)(1/2)
=¼
• Compute FG???
Computing relationships for inbred common ancestors or direct
ancestors using Path Coefficients

Consider the following pedigree for direct relatives:

F C A

G E D
• To calculate the relationship between A and G, aAG,
do the following;
Computing relationships for inbred common
ancestors or direct ancestors using Path
Coefficients
1. Count for each path the number of intervening
generations between individual G and its direct
ancestor A for each separate path:
• The no. of generations for the three paths between G
and A are:
nGEA= 2,
nGEDA= 3
nGFCA= 3
Computing relationships for inbred common
ancestors or direct ancestors using Path
Coefficients
2. Calculate the additive relationship between A and
G as:
aAG = (1/2)nGFCA(1 + FA) + (1/2)nGEA(1 + FA) + (1/2)nGEDA (1 + FA)

• If FA is zero, then
aAG = (1/2)3(1 + 0) + (1/2)2(1 + 0) + (1/2)3 (1 + 0)
=(1/2)3+ (1/2)2 + (1/2)3
= 1/2
Computing relationships for inbred common
ancestors or direct ancestors using Path
Coefficients
• What is the relationship between A and G, if the
inbreeding coefficient of A is 0.5?
Computing relationships for inbred common
ancestors or direct ancestors using Path
Coefficients
For Collateral relatives:
• For example, calculate the relationship between E and F,
aFE
• Simply count the number of generations from F to E
through the common ancestor A for the 2 different
paths;
nFCA + nAE = 3
nFCA + nADE = 4
Computing relationships for inbred common
ancestors or direct ancestors using Path
Coefficients
• So, aFE = (1/2)3 (1 + FA) + (1/2)4(1 + FA)
• If FA is zero, then;
aFE = (1/2)3 + (1/2)4)= 3/16
And if FA = 0.5
aFE = (1/2)3 (1 + 0.5) + (1/2)4(1 + 0.5)
= 9/32
Rules for calculating additive
relationships for collateral relatives
• If two or more ancestors are included in the
pedigree, you should include all the different paths
through all common ancestors making sure that:
1. An animal can occur only once in a single path
2. A path follows direct descendents only, i.e, it does
not zig-zag but goes directly to the common
ancestor, from offspring to parents etc.
3. Inbreeding of intermediate animals is ignored,
with only the inbreeding coefficient of the
common ancestor considered in each path.
Rules for calculating additive relationships for
collateral relatives

• Example: What is the additive relationship between

animal G and animal E in the pedigree below
E C A
G
F D B
To calculate aGE, there is;
i. a direct path G E
ii. Six collateral paths through common ancestors C, D, A and B
 Rules for calculating additive
relationships for collateral relatives

1. Count the number of generations between E and G.

i. For the direct path E G, nEG = 1
ii. For the indirect paths
E C F G n1= 1 + 2 = 3
E D F G n2= 1 + 2 = 3
E C A D F G n3= 2 + 3 = 5
E C B D F G n4= 2 + 3 = 5
E D A C F G n5= 2 + 3 = 5
E D B C F G n6= 2 + 3 = 5
Rules for calculating additive relationships
for collateral relatives
• Thus
aGE=(½)1 (1 + FE) + (½)3(1 + FC) + (½)3(1 + FD) + (½)5(1 +FA) + (½)5(1+FB) +
(½)5(1+FA) + (½)5 (1 + FB)
• If the common ancestors are not inbred, and FE =1/4
• Remember Fx = ½ aSxDx, the Sire is C and the Dam is D
C A D; n1 = 2
D B C; n2 = 2
aCD= (½)2 + (½)2 = ½
FE= ½ aCD= ½ x ½ = 1/4
Then:
aGE=(½) (1 + 1/4) + (½)3 + (½)3 + (½)5 + (½)5 + (½)5 + (½)5 = 1
Rules for calculating additive
relationships for collateral relatives
• If A and B are inbred, aGE, will depend on FA and FB.
• For the calculation of aGE, the paths are:
nGECA = 3
nGEDA = 3
nGFCA = 3
nGFDA = 3
• So that,
aGE = (½)3 (1 + FA) + (½)3(1 + FA) + (½)3(1 + FA) + (½)3(1 +FA)
COEFFICIENT OF
RELATIONSHIP
• Definition: an expression of the probability that
two individuals possess duplicate alleles, because
of their common line of descent, over and above
those found in the base population.
Computing relationship coefficients
between collateral relatives
• Methods are similar to those used for calculating Fx
and arrow diagrams are of value in this respect.
• The formula for calculating relationship coefficients
between collateral relatives X and Y is:
• RXY = ∑[(1/2)n(1 + FA)
(1 + FX)(1 + FY)
Computing relationship coefficients
between collateral relatives
• Where:
• RXY = relationship coefficient between animals X and Y
• ∑ = Greek symbol meaning to sum/add.
• N = number of arrows connecting individual X with Y through the
common ancestor for each path.
• FX = inbreeding coefficient of animal X.
• FY = inbreeding coefficient of animal Y.
• FA = inbreeding coefficient of the common ancestor.
• Note: If individuals X and Y and their common ancestor are not
inbred, the formula becomes;
• RXY = ∑[(1/2)n]
Computing relationship coefficients
between collateral relatives
• Example 1: Relationship coefficient between half
brothers and half sisters:
• Let X be the male and Y be the female.
• Pedigree
X 1
2
Y 1
3
Computing relationship coefficients
between collateral relatives
• Arrow diagram:
X
1
Y
• Pathway:
X 1 Y n=2
Thus RXY = (1/2)2 = ¼
• Interpretation: The two individuals X and Y are related by 25%,
or they probably have an increase in the percentage of
duplicate alleles over that found in the base or non-inbred
population.
Computing relationship coefficients
between collateral relatives
• Example 2: Compute the relationship coefficient for
full sibs i.e brothers and sisters
Pedigree: X 1
2
Y 1
2
Arrow diagram: X 1

Y 2
Computing relationship coefficients
between collateral relatives
• Pathways:
X 1 Y n= 2
X 2 Y n= 2

RXY = (1/2)2 + (1/2)2

=¼+¼
=½