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Rapid Method for Computing the Inverse of a

Relationship Matrix

C. R. HENDERSON
Department of Animal Science
Cornell University, Ithaca, NY 14850

ABSTRACT We wish to predict additive genetic merits of


The accuracy of prediction of breeding many animals from a set of records with
values can be improved by utilizing all unknown fixed effects and random variables in
relationships among animals to be evalu- addition to breeding values. Let a general mixed
ated. The inverse of the numerator rela- model for such prediction be
tionship matrix is required for best pre-
diction but is usually much too large to y = X/3 + Z l u + Z 2 a + e [1]
be computed by conventional inversion
routines. A simple and rapid technique where y is the vector of records, X is a known
for finding this inverse directly from a list matrix, (3 is an unknown fixed vector, Z1 is a
of sires and dams, bypassing computation known matrix, u is a nonobservable random
of the matrix whose inverse is wanted, is vector, Z2 is a known matrix, a is a nonobser-
presented. vable random vector representing additive
genetic merits, and e is a nonobservable random
vector of "errors." We assume that u, a, and
INTRODUCTION e all have null means and that Var(u) = Ga 2,
Var(a) = kAo 2 where k is a scalar, and Var(e) =
The use of best linear unbiased prediction
Io 2. G and A are known and nonsingular, and
(BLUP) methods for predicting breeding values
A is the numerator relationship matrix de-
has been described by Henderson (1, 2, 3). The fined below. It is assumed that u, a, and e are
matrix of relationships of animals to be evalu- mutually uncorrelated. The scalar cr2 is possibly
ated could be incorporated in these methods,
unknown.
but as presented in these papers it was neces- Henderson (1) has shown that BLUP of a is
sary either to invert the relationship matrix or ~, any solution to
to transform involving extensive matrix multi-
plications. The former was prohibitive compu-
rationally if the matrix was too large to store in
_1 . . . . . .
the memory of the computer, and the latter t ~ tI
destroys the customary sparseness of the co- tX Iz~zl+~ I Z'~Z2 = 't .[21
efficient matrix and thereby reduces efficiency
of an iterative method of solution. A simple
~X Z~Zl I Z~Z2 + k-1 ^ Lz' ,J
method for inverting the relationship matrix
has the unusual feature that the inverse can The solution to ~ is not necessarily unique, but
be written without having to compute the and ~ are unique regardless of the solution to
matrix to be inverted. In a noninbred popula- because of the addition of G-1 and k-1 A-1 to
tion, the inverse can be computed by using only the usual least squares matrix of coefficients.
a list of parents. An inbred population requires In some important applications, either
in addition that a triangular matrix be gener- [~' ~ ~'] has relatively few elements or
ated, but still no conventional matrix inversion
is required. This paper presents the method
with examples. The derivation has been re- x XZll
ported (4). ix zlz,.

Received October 17, 1974. has a large submatrix that is easy to invert (is

1727
1728 HENDERSON

diagonal, for example), and consequently m o s t 4) If only one parent is k n o w n , say p,


"f A P
or all of the elements of [3 : u ] can be ab-
sorbed easily. F o r example, in the Cornel1 ~ij = 9.pj/2 for j = 1 . . . . . p
m e t h o d o f sire evaluation used in the Northeast
= 0 f o r j = p + l . . . . . i--1, provided
(3), 3 represents a few groups and a large num-
ber of fixed herd-year-season effects. There is p<i-1.
no u in the model. Herd-year-seasons are ab- ~i~ = (1 - ~ ~2).s
sorbed easily, leaving a set o f equations with a j=l ~1
few fixed groups and a large n u m b e r of sires 5) If neither parent is known,
( a p p r o x i m a t e l y 1,500 Holsteins). Presently, the n

relationships among sires are ignored. T h a t is, ~ij = 0 f o r ~ . . . . , i - 1 ;


A is regarded as having the value I. T h e re- j=l
~ii = 1.
sulting equations are well suited to an iterative
solution because o f m a n y zero coefficients and To illustrate, suppose we have the following
in particular because the diagonal coefficients animals, with 1 and 2 being defined as the base
are large relative to the off-diagonal ones. population:
The prohibitive cost o f inverting A has dic-
tated in the past that the r e d u c t i o n in predic-
tion error variance by using A-1 not be utilized. Indi~dual Parents
The ease with which A -1 c a n be written, as
described in this paper, n o w makes feasible in- 1 Unknown
c o r p o r a t i o n of relationships in sire evaluation. 2 Unknown
The relationship m a t r i x could also be used in 3 1 and unknown
4 1 and 2
intraherd evaluation of females.
5 3 and 4
6 1 and 4
C O M P U T A T I O N OF A T R I A N G U L A R M A T R I X 7 5 and 6
NEEDED FOR I N B R E D P O P U L A T I O N S

Let A be the n u m e r a t o r relationship m a t r i x


for n animals. The first t < n of these are de- T h e n L has the value in [3]. Starting with the
fined as the " b a s e " population. T h a t is, these third row, the first two elements are (1 + 0)/2
animals are regarded as noninbred, unrelated, and 0 which is [.5 0]. The third e l e m e n t is
and with unspecified parents. A lower triangu- (1 - . 5 2 ) .5 = .866025. The first three elements
lar matrix, say L, exists such t h a t LL ~ = A. If o f the f o u r t h row are (1 + 0)/2, (0 + 1)/2,
we can c o m p u t e L, we can use this to c o m p u t e and 0 which is [.5 .5 0]. The f o u r t h e l e m e n t
A. In fact, L is easy to c o m p u t e recursively, and of this ,row is 11 + [(1)(0) + ( 0 ) ( 1 ) ] / 2 -- (.5) 2
its c o m p u t a t i o n is well adapted to computeriza- _ (.5)2t .s = (.5).5. The first f o u r elements
tion in contrast to the path coefficient m e t h o d . of row five are (.5 + .5)/2, (0 + .5)/2, (.866025
A simple c o m p u t i n g algorithm follows: + 0)2, and .707107/2 which is [.5 .25 .43301c3
. 3 5 3 5 5 3 ] . The diagonal e l e m e n t o f row 5 is 41
1) N u m b e r the animals 1, 2 , . . . , n, ordered + [(.5)(.5) + (0)(.5) + ( . 8 6 6 0 2 5 ) ( 0 ) 1 / 2 ~ (.5) 2
such that parents precede their progeny. _ (.25)2 _ (.433013)~ _ (.353553)2I .s =
2) The upper t 2 submatrix of L is I. (.5) "5. The last two rows are generated in a
3) Given that p < q are the parents of i, similar manner.
N o w if we m u l t i p l y matrix [3] b y its trans-
£ij = (l~pj + ~qj)/2 f o r j = 1 . . . . . p pose, we obtain the A m a t r i x in [4] which is
inverted to d e m o n s t r a t e t h a t the same result
= ~qj/2 f o r j = p + 1 . . . . . q can be obtained by the simple m e t h o d of this
paper. Since this is a matrix of small order, it is
= 0 for] = q + 1..... i- 1, provided easy to invert by conventional methods. A-1 is
tabulated in [5] so we can check this result
q<i-1;
with our simpler m e t h o d . A m u c h easier
P q
E E ~.2vs m e t h o d for obtaining this inverse is in the n e x t
~ii = (1 + .5:1j= ~pj~qj q, section.

Journal of Dairy Science Vol. 58, No. 11


TECHNICAL NOTE 1 729

(1) (2) (3) (4) (5) (6) (7)


(1) ~000000 .000000 .000000 .000000 .000000 .000000 .000000
(2) .000000 1.000000 .000000 .000000 000000 .000000 .000000
(3) .500000 .000000 .866025 .000000 .000000 .000000 .000000
(4) .500000 .500000 .000000 .707107 .000000 .000000 .000000 [31
(5) .500000 .250000 .433013 .353553 .707107 .000000 .000000
(6) .750000 .250000 .000000 .353553 .000000 .707107 .000000
(7) .625000 .250000 .216506 .353553 .353553 .353553 .637378

(1) (2) (3) (4) (5) (6) (7)


(1) ~-.00000 .00000 .50000 .50000 .50000 .75000 .62500-
(2) .00000 1.00000 .00000 .50000 .25000 .25000 .25000
(3) .50000 .00000 1.00000 .25000 .62500 .37500 .50000
(4) .50000 .50000 .25000 1.00000 .62500 .75000 .68750 [41
(5) .50000 .25000 .62500 .62500 1.12500 .56250 .84375
(6) .75000 .25000 .37500 .75000 .56250 1.25000 .90625
(7) .62500 .25000 .50000 .68750 .84375 .90625 1.28125

(1) (2) (3) (4) (5) (6) (7)


(1) 2.333333 .500000 -.666667 -.500000 .000000 -1.000000 .000000
(2) .500000 1.500000 .000000 -1.000000 .000000 .000000 .000000
(3) -.666667 .000000 1.833333 .500000 -1.000000 -1.000000 .000000
(4) -.500000 -1.000000 .500000 3.000000 -1.000000 -1.000000 .000000 151
(5) .000000 .000000 -1.000000 -1.000000 2.615384 .615384 -1.230767
(6) -1.000000 .000000 .000000 -1.000000 .615384 2.615384 -1 230767
(7) .000000 .000000 .000000 .000000 -1.230767 -1.230767 2.461534

INVERSE OF R E L A T I O N S H I P M A T R I X --.5 where the first t e r m arises as a con-


BY THE SIMPLE M E T H O D sequence of 4 being a progeny of 1 and
the second t e r m results f r o m the mating
Let d be a v e c t o r of the inverse of the
squares o f each diagonal e l e m e n t of L. T h a t is, of 1 and 4 to produce 6.
d i = 1 / ~ i as c o m p u t e d in the previous section.
In our example, matrix [3], d' = [1, 1, 1.333, F U R T H E R S I M P L I F I C A T I O N IN N O N I N B R E D
2, 2, 2, 2 . 4 6 1 5 3 4 ] . POPULATIONS
T h e n the elements of A-1 can be c o m p u t e d
using d and a list of sires and dams. We denote The m o s t laborious aspect of the preceding
the i j t h e l e m e n t of A-1 by aij. section is the c o m p u t a t i o n of L to obtain d.
In a n o n i n b r e d population, however, this com-
1) a ii = d i + .25 ff d k where k refers to the p u t a t i o n is not needed since d can be written
progeny of the ith sire. In our example, immediately. Only three values are possible,
a 11 = 1 + (.25) (1.33) . . . + 2 + 2) = n a m e l y d i = 1 if neither parent o f i is known,
2.33 . . . and a 44 = 2 + (.25)(2 + 2) = 3. d i = 4/3 if only one parent o f i is known, or
d i = 2 if b o t h parents of i are known. Thus, in
2) aiJ (i < j) = - . 5 d ] + .25 ~ dk i f j i s a a n o n i n b r e d population, all o f the elements of
progeny of i, or = .25 ~ d k f r o m the A-1 can be c o m p u t e d directly b y inspection of
mating of i with j. Of course, aJi = alJ the list of parents. Also in such a p o p u l a t i o n
since A-1 is symmetric. For example, the c o n t r i b u t i o n to aij (i < j) can be either
a 12 = (.25)(2) = .5 as a c o n s e q u e n c e of parent-progeny relationship or c o m m o n prog-
their c o m m o n progeny, n u m b e r 4; eny, b u t n o t both.
a 67 = (--.5)(2.461534) = -1.230767 These c o m p u t a t i o n s for a n o n i n b r e d popula-
as a consequence of 7 being a progeny tion are illustrated with the following list of
of 6; and a 14 --- ( - . 5 ) ( 2 ) + (.25)(2) = parents.

J o u r n a l o f D a i r y S c i e n c e V o l . 58, N o . 1 1
1730 HENDERSON

PROGRAM FOR C O M P U T I N G A - 1
Individual Parents
A F O R T R A N p r o g r a m has b e e n w r i t t e n
t o c o m p u t e A-1. It is e x t r e m e l y fast a n d uses
1 Unknown
2 Unknown little m e m o r y . T h e p r o g r a m requires t h a t a list
3 1 and 2 o f individuals w i t h t h e i r p a r e n t s b e s t o r e d in
4 1 and 2 m e m o r y . If t h e p o p u l a t i o n is specified n o n -
5 1 and 2 i n b r e d or if o n e c h o o s e s t o use d o f a n o n i n b r e d
6 1 and unknown
7 1 and unknown population regardless o f i n b r e e d i n g , t h e pro-
8 2 and unknown gram computes d and then proceeds to gen-
9 7and8 e r a t e e l e m e n t s o f A -1 . If i n b r e e d i n g is speci-
fied, d m u s t b e s t o r e d in m e m o r y . A n o t h e r
p r o g r a m has b e e n w r i t t e n to c o m p u t e this
T h e n d' = [1 1 2 2 2 4/3 4/3 4/3 2]. vector.

19 9 -6 -6 -6 4 -4 0 0
REFERENCES
17 -6 -6 -6 0 0 -4 0
-6 12 0 0 0 0 0 0
1. Henderson, C. R. 1963. Selection index and
- -6 0 12 0 0 0 0 0 expected genetic advance. Page 141 in: Statistical
K I = I/6 -6 o 0 ~2 o o 0 0 genetics and plant breeding. Hanson, W. D. and H.
0 o 0 0 8 0 o o
F. Robinson, eds. NAS--NRC, Pub. 982.
o 0 0 o o I1 3 -6
-~ o 0 o 0 3 11 -6
2. Henderson, C. R. 1973. Sire evaluation and
0 0 0 0 0 -6 -6 12 genetic trends. Page 10 in Proc. of the Anim.
Breeding and Genet. Symp. in Honor of Dr. Jay
L. Lush. ASAS and ADSA, Champaign, IL.
T o illustrate c o m p u t a t i o n of s o m e of the 3. Henderson, C. R. 1974. General flexibility of
e l e m e n t s of this matrix, a 11 = 1 + ( . 2 5 ) ( 2 + linear model techniques for sire evaluation. J.
2 + 2 + 4/3 + 4 / 3 ) = 19/6;a 12 = (.5)(2 + 2 + Dairy Sci. 57:963.
4. Henderson, C.R. 1976. A simple method for
2) = 9/6 since 1 a n d 2 have c o m m o n p r o g e n y computing the inverse of a numerator relationship
3, 4, a n d 5; a n d a ~9 = ( - . 5 ) ( 2 ) = - 1 since 9 is matrix used in prediction of breeding values.
a p r o g e n y o f 7. Biometrics (In press.)

Journal of Dairy Science Vol. 58, No. 11

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