The Number of Canalyzing Functions over Any Finite Set
*
Yuan Li1#, David Murrugarra2, John O. Adeyeye1, Reinhard Laubenbacher3
1Department of Mathematics, Winston-Salem State University, Winston-Salem, USA 2School of Mathematics, Georgia Tech, Atlanta, USA
3Virginia Bioinformatics Institute, Virginia Tech, Blacksburg, USA Email: #[email protected]
Received March 1, 2013; revised May 20, 2013; accepted June 16, 2013
Copyright © 2013 Yuan Li et al. This is an open access article distributed under the Creative Commons Attribution License, which
permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.
ABSTRACT
In this paper, we extend the definition of Boolean canalyzing functions to the canalyzing functions of multi-state case. Namely, f Q: nQ, where
. We obtain its cardinality and the cardinalities of its various subsets (They may not be disjoint). When , we obtain a combinatorial identity by equating our result to the formula in [1]. For a better understanding to the magnitude, we obtain the asymptotes for all the cardinalities as either or.
1, , ,2 qQ a a a
2 q
n
q
Keywords: Canalyzing Function; Inclusion and Exclusion Principle
1. Introduction
The idea of canalization was initiated from Waddington, C. H. [2]. When comparing the class of canalyzing functions to other classes of functions with respect to their evolutionary plausibility as emergent control rules in genetic regulatory systems, it is informative to know the number of canalyzing functions with a given number of input variables [1]. However, the Boolean network modeling paradigm is rather restrictive, with its limit to two possible functional levels, ON and OFF, for genes, proteins, etc. Many discrete models of biological net- works therefore allow variables to take on multiple states. Common used discrete multi-state model types are so- called logical models [3], Petri nets [4], and agent- based models [5].
In this paper, we generalize the concept of Boolean canalyzing rules to the multi-state case. By generalizing the results in [1], we provide formulas for the cardi- nalities of various subsets of canalyzing functions. We also obtain the asymptotes of these cardinalities as either or q . We obtain a combinatorial identity by equating our result to the formula in [1].
n
2. Preliminaries
In this section we introduce the definition of a canalyzing
function.
Let
n
1, 2, , n Q
,
a a1, , ,2 aq
and : nf Q Q.
A function is canalyzing if there is a variable xi and an element aQ so that the value of the function is fixed once variable xi is fixed at . More precisely, we have the following definitions.
a
Definition 2.1
1) The function f x x
1, , ,2 xn
is i a b: : canaly- zing if f x
1, ,xi 1, , i 1, , n
1 1, 1,
a x x
, n
b, for all , , i i
x x x x 2) The function
.
1, , ,2 n
f x x x is i a: : canaly- zing if there exists bQ such that
1, , i 1, , i 1, , n
f x x a x x b
1, , 1, 1,
,
, for all
i i n
x x x x 3) The function
.
1, , ,2 n
f x x x is : :a b ca- nalyzing if there exists i
n such that
1, , i 1, , i 1, , n
f x x a x x b
1, , i1, i1, , n
, for all
x x x x .
4) The function f x x
1, ,2 ,xn
is i: : b canaly- zing if there exists aQ such that
1, , i 1, , i 1, , n
f x x a x x b
1, , 1, 1,
,
, for all
i i n
x x x x 5) The function
.
1, , ,2 n
f x x x is : :a ca- nalyzing if there exist i
n b, Q such that
1, , i 1, , i 1, , n
f x x a x x b
1, , i1, i1, ,
n
, for all x x x x .
*Supported by an award from the USA DoD # W911NF-11-10166.
#Corresponding author. 6) The function
, , ,
1 2 n
canalyzing if there exist a, bQ such that
1, , i1, , i 1, , n
f x x a x
1, , i1, i1, , n
x
b, for all
x x x x .
7) The function f x x
1, , ,2 xn
is : :b ca- nalyzing if there exist i
n ,a
1, , i1, , i 1, , n
Qsuch that f x x a x
1, , i1, i1, , n
x
b, for all
x x x x .
8) The function is f x x
1, , ,2 xn
is : : ca- nalyzing if there exist i
n , ,a b
1, , i1, , i 1, , n
Q such thatf x x a x
1, , i1, i1, , n
x
b, for all
x x x x .
By abuse of notation, we also use i: : b to stand for the set of all the i: : b canalyzing functions,
: :
i a b will stand for the set of all the i a b: : canalyzing functions and etc. We use to stand for the empty set.
By the definitions, we immediately have the following propositions.
Proposition 2.2 If b1b2, then
1 2
: : : :
i a b i a b b b
.
Proposition 2.3 If 1 2 and i1i2, then 1: : 1 2: :b2
i b i .
By the definitions, we have
: : : : : : : : ,
: : : : : : ,
: : : : : : ,
: : : : : : ,
: : : : ,
: : : : ,
: : : : .
b Q a Q i n
i n a Q
b Q i n
a Q b Q
a Q
b Q
i n
b a i
b i b a b
a a b i a
i i a i b
i b i a b
i a i a b
a b i a b
For any set S , we use S to stand for its cardinality.
We use
!
,
! !
n C n k
k n k
to stand for the binomial
coefficients. As usual, should be explained as
zero once .
, C n k
knObviously, for the above notations, the cardinality are same for different values of i a, and . In other words, b we have i a b1: :1 1 i2:a2:b2 ,
: : : :
i b j c , : :a b : :c d and etc.
3. Enumeration
Theorem 3.1 Given i
n a b, , Q, the number of: :
i a b canalyzing functions is qqnqn1. In other words, we have i a b: : qqnqn1
.
Proof: A function in the set i a b: : is uniquely determined by its value on inputs
x1, , xn
withi
x a . There are
q1
q q1
n n n
q q
1 such inputs,
and the function can take different values. Thus
1
: : qn qn i a b q . □
Because : :
i a b: : b Q i a
, by Proposition 2.2,
we get
Theorem 3.2 The number of all the i a: : canaly- zing function is
1
11.
n n n n
q q q q
q q q
Lemma 3.3 We have k1 : : qn kqn1 j
j i a b q
forany
a a1, , ,2 ak
Q.Proof: A function in the set
j1 is uniquely determined by it values on inputs : : k
i aj b
x1, , xn
with
1, ,
i k
x a a . There are such
inputs. □
qk qn1qnkqn1
Theorem 3.4 Given i
n and bQ, the numberof i: : b canalyzing functions is qq n
other
1
1n qn q
q
. I
words, we have : : n
n1 1
q
q q
i b q q .
and Exclusion Principle, we ve
Proof: By Inclusion ha
1 2
1 2
1 1
1
1
1
1 2
,
1 1
, , , 1 2
1 1
1 1
1
: : : :
: : : : : :
1 : :
,1 , 2
1 , 1
1 ,
,
1
k
n n n n
n n
n n
n n
n
a Q
a a Q
a Q
k
k q
j
a a a Q j
q q q q
k q kq q
q
k q kq
k
q k
q q
k
q
i b i a b
i a b i a b i a b
i a b
C q q C q q
C q k q
C q k q
q C q k q
q
1
1
1
1 qn q qn qn 1q
q q q
.
Similar to Lemma 3.3, we have
Lemma 3.5 If
i i1 2, , , ik
n , then
k
q1 q
1
: : k n k
j j
i a b q
.Based on this lemma, we can get the following result.
Theorem 3.6 We have
1
11
: : 1k , q kqn k.
k n
a b C n k q
12
1 2
1 1
1
1
1 1 1
1 1
1 1 1
1 1
1
: : : :
: : : : : :
1
: : 1 : :
,1 , 2
1 , 1
1 , .
k
n n
k n k
k n k
i n
i n i j n
k
k
n j
i i n j j
q q q q
k q q q
k q q
k n
a b i a b
i a b i a b j a b
i a b j a b
C n q C n q
C n k q q
C n k q
1n
n
n
From the above theorem, we can get the following re
m 3.7 We have
sult.
Theore
1
11
: : 1k , q kqn k
k n
a q C n k q
Proof: Because : : : :
b Q
a a
b , by Theorem3.6, we just need to show : :a b1 : :a b2 if 1 2
b b . Suppose f :a: 1 : : 2 ere
1
i and
b a b , then th
exist i2 n
such that1: : 1 2: : 2
f i a b i a b since
: : : :
i n
a b i a b
.If i1i2, we get a contradiction by Proposition 2.2. If
1i2, w
i e get a contradiction by Proposition 2.3 since 1: :b1 i1: : b1 and
i a i2: :a b2 i2: : b2 . □
to find umb
Now, we are going the formula for the n er
of all the canalyzing functions with given canalyzed value b. In other words, the formula of : :b .
Let Sb
i: :a b i
n a, Q
for any bQ. By Inclusion and Exclusion Principle, we have
1
1
: : : : nq 1 k .
k k
i n a Q
b i a b N
where
,
.
b
k
s S s k T s
N T
In order to evaluate Nk, we write all the members in
b as the following n q matrix. S
1 2
1 2
1 2
1: : 1: : 1: :
2 : : 2 : : 2 : :
: : : : : :
q
q
q
a b a b a b
a b a b a b
A
n a b n a b n a b
For any , we will choose ele-
m
k
ents from the above matrix to form s.
Suppose k1 of its elements are fro the first row m
(th
b sS
ere are
q k, 1
ways to do so). Let these k1elements be C
1
12 1
, 1: : , , 1: k :
b a b a b .
Suppose 11 ond row
1:a :
2
k of its elements are from the sec (there are
q k, 2
ways to do so). Let these k2elements be C
2
21
2 :a :b , 2 :a22: , , 2 :b a2k :b .
Su ose of its elements are from the last row (th
pp kn
ere are
q k, n
ways to do so). Let these kn elements beC
1 b , n a: n2: , ,b n a: nkn:b .
1 2
k k
: n an :
,0 , 1, 2, ,
n i
k k k q i n
.
Similar to Lemma 3.3, we have
Lemma 3.8 Let s be the subset of Sb as mentioned
above, then T qq k1q k2 q kn.
T s
Hence,
.
We get
3.9 For any
1 11
, , n
n
q k q k
k n
k k k
C q k C q k q
N
Theorem bQ, we have
1 1 2
1
1 1
: :b
1 , .
n j j n
q k
nq n
k
j
k k k k k j
C q k q
In order to evaluate : :i , we need two more
lemmas. Their proofs are similar to that of Lemma 3.3 and we omit them.
Lemma 3.10 If
a a1, , ,2 ak
Q and
b1, , ,b2 bk
Q, then 1
1
: : n
k
q k q
j j
j
i a b q
.Lemma 3.11 If rk
ar 1 2
11, , 1k; 21, , 2k ; ; r1, , r
a a a a a a
e k1 kr distinct elements of Q,
b1,,br
Q. Then,
1 2
1 1 2
1 1 2 2
1 1 1
: : : : r : :
n r
k k k
j j
j j j
q k k k q
i a b i a b i a b
q
rj r
Now, we are ready to find the cardinality of
: : i .
Theorem 3.12 We have
1
1
1
1 ,0 1 2
: :
i
1 1
! .
! ! ! !
n
q i
k q k q
q
k k k k k q q
q q
q k k k k
Proof: First, we have : : : :
a Q b Q
i i
a b .Let Si
i a b a b: : , Q
, we get
2
1 1
: :
i 1 N ere
q k
k k
. Wh,
.
i
k
s S s k T s
N T
In order to evaluate , we write all the elements in as the following k matrix.
N q q i
S
1 1 2 1
: : :
i a b a b
1
1 2 2 2 2
1 2
: : :
: : : : : :
: : : : : :
q
q
q q q q
i i a b
i a b i a b i a b
B
i a b i a b i a b
For any
i
sS with s k , we will choose
elements fro e matrix to form
k
m the abov s.
Suppose k1 it elem re from the first ro
(There are C q k
, 1
ways to do so). Let theseof ents a w
elements be 1
k
11 1 12 1 11 1
: : , : : , , : k :
i a b i a b i a b .
Suppose k elements are from the second row,
we must cho2 lu
of its os e i
e these elements from different co mns, otherwise th ntersection will be by Proposition 2.2 (There are C q
k k1, 2
ways to do so). Let these k2elements be i a: 21:b2 , :i a22:b2 , :i a2k2:b2
Suppose
,
of its elements are from the last row
(The ways to do so).
Le
q
k C q
e
re are
k1 k2 kq1,kq
t these kq lements be1 2
: q : q , : q : q , ,
i a b i a b : : .
q
qk q
i a b
where . We
have 1 2
,0 , 1, 2, ,
q i
k k k k k q i q
1
1 2
,
1 1 2
,0
1 1
, ,
,
i
q i
q
s S s k T sk k k k q
q q k k k
T
C q k C q k k
C q k k k I
where k N
1 2 q
k k k
I
1 2
1 1 2 2
1 1 1
: : : : q : :
k
k k
j j qj q
j j j
i a b i a b i a b
By Lemma 3.11, we know
This number is zero if
A straightforward computing shows that
1 2
1 2 q
q k k k k k
I q
1 1
.
n n
q
k q q k q
q
kq.
1 1 2 1
!
C k
q
1
1 2
,
.
! ! ! !
q q
q
k k
k k k q k
Hence, we get
, ,
C q k C qk k q
2
q q
1 1
1
1 1
1 1
1
1 ,0 1 2
1 1
1 ,0 1 2
: :
1 1
! 1
! ! ! !
1 1
! .
! ! ! !
n
q i
q i
k k
k k
k k
q
k q k q
k k k k k q q
n
k q k q
q
k k k k k q q
i
N N
q
q
k k k q k
q q
q k k k k
□ Now we begin to evaluate : : .
Theorem 3.13 We have
1
11 1
: :
1 1 .
nq
k k
k k
k k
U V
where
q
1 1 2
1 2
1 1 2
,0
,0 1 2
!
! ! ! !
! 1
! !
n
q q i
n
q i
k
q k q
t t t k t q
q k q
t t t k t q q
U
q
n q
t t t q k
nq q
q k t t ! t !
and
Proof: Let
1 1 ,0 1,0 1
,
n j j n i i
q k n
k j
k k k k k k q j
V q C q k q
: : , ,
.S i a b a bQ i n
We have
: : : :
i n a Q b Q i a b
2
1 1
: : nq 1k
k k
N
, whereand
,
.
k
s S s k T s
N T
We write all the elements of as the following matrices.
2
nq S
1 1 1 2 1
2 1 2 2 2
1
1 2
1 1 1 2 1
2 1 2 2 2
2
1 2
1 1 1 2
1: :a b 1: :a b 1: :a bq
1: : 1: : 1: :
1: : 1: : 1: :
2 : : 2 : : 2 : :
2 : : 2 : : 2 : :
2 : : 2 : : 2 : :
: : : : :
q
q q q q
q
q
q q q q
n
a b a b a b
M
a b a b a b
a b a b a b
a b a b a b
M
a b a b a b
n a b n a b n a
M
1
2 1 2 2 2
1 2
:
: : : : : :
: : : : : :
q
q
q q q q
b
n a b n a b n a b
n a b n a b n a b
We combine all the above Mi to form a nq q matrix M whose first q rows are M1, the sec ro
ond q ws are M2, , the last q rows are Mn. In words, we have
other
1 1 1:a b1 2 1 1:bq
2 1 2 2 2
1 2
1 1 1 2 1
2 1 2 2 2
1 2
1 1 1 2 1
2 1 2 2 2
1: : : :
1: : 1: : 1: :
1: : 1: : 1: :
2 : : 2 : : 2 : :
2 : : 2 : : 2 : :
2 : : 2 : : 2 : :
: : : : : :
: : : : : :
: :
q
q q q q
q
q
q q q q
q
q
q
a b a
a b a b a b
a b a b a b
a b a b a b
a b a b a b
M
a b a b a b
n a b n a b n a b
n a b n a b n a b
n a b
1 n a: q:b2 n a: q:bq
We are going to choose elements from
k M to
form the intersection. In or o get a possi non
em
der t ble
s m pty intersection, we know all these k element ust
come from either the same Mi (for some fixed i) or all of them from the same column of M by Proposition 2.3.
Each Mi is in fact the transpose f B and each column of
o
M is all the elements of A (As sets, they are equal). Hence, a typical intersection is e her the one eorem .9 or the one in Theorem 3.1 t these
two cases ar ot disjoint.
Suppose we choose ki elements from
1 2
, 1, 2, , , ,
i n
it
in Th 3 2. Bu
e n
0 i , 1,2, ,
M i n k k k k
k k i n
.
If there exist such that i kik, then kj 0, j i.
e one in Lemma
This implies the intersection h
3.11 and
looks like t kq.
If 0 ki k 1, i
n , th e intersection lookslike the one in Lemma 3.8 and knq. The above two cases are disjoint now. By Le
en th
mma 3.11 and Lemma 3.8, we get
1
,
k
,0 , 1, 1, ,
n i
i i
s S s k
N T
k k k k k
T s
k k
i k k k k i n
U V
where (Note: there are n matrices M M1, 2, , Mn and columns of
q M )
1 1 2 ,0
!
! ! !
1 1 2
1 1
1 2
,0 1 2
,0 1,0 1
!
! 1
! ! ! !
,
n
q i n
q i n
j j n i i
q k q
t q q
q k q
t t t k t q q
q k n
k j
k k k k k k q j
q k
nq q
q k t t t
V q C q k q
Hence, k
t t k t
q
U n
t t t q
2
2
1 1
1 1
1 1 1
: : 1
1 1
nq k
k k
nq q nq
k k
k k k k
k k k
N
U V U V
11 k , □
In the following, we will reduce the formula : :
when q2 and compare it with the one in [1]. We have
1
1: : 1k 1 k .
k k
U V
2
2 1 1
n
k k
where
1 1 2
1 1
2 2 ,0 2 1 2
2 ,0 1,0 2 1
2! 2
! ! 2 !
2 2,
n
i
n
2j j
n i i
k k
t t k t
k n
k j
k k k k k k j
U n
t t k
V C
k
A simple calculation shows that
2
1 1
2 2
1 4 2 ,1 2 2
n n
U n C n
2 4 .
U n
1
V 0 since the condition of the sum is not satisfied.
2
1
2
2
3 2
2 2,
, 2 2 2
n
1
2
2,0 1 1
j j
n
k n
V C k
C n
n i
j k k k j
Note, is the number of solutions of the
equation When
, 2C n
1 k k
3 k 2nn, 2,0 ki 1.
1
1 2
k
k
2
2 2
n j j
k n
V
,0 1
1 2 2 1
1 2
2,
, 2 2 , , 2 2
n i n k
j
k k k j
k k
k t
C k
C n k C n t C n t k t
Note, 2 is the number of solu
tions of , with
exactly
he en ,
t
, ,
C n t C n t k t the equation 1
t components equal to
-2
n
k k
2.
0 ki 2
nce, wh q2
1 1 2
1
1 2 1
3 2 1
2
: :
k
4n 1 , 2 2
1 , , 2 2
n k
k
k n
k k t
k
k n
t
C n k
C n t C n t k t
When , one can obtain (without calculator)
the sequ 120, 3514. These results are con-
sistent with t By [1], the cardinality of
1, 2,3, 4
n
ence 4, 14, hose in [1]. : :
should be
1: :
1n k n k
n
1 21
2 1 , 2 2k .
k n
C n k
So, we obtain the following combinatorial identity(for any positive integer n).
1 2 1
3 2 1
1 , , 2 2
k
k n
t
C n t C n t k t
2
2 1
k k t
n
n
The left sum should be explained as 0 if
1
n . As
usual, is 0 if
Fro
,
C n k
m Theorem
kn.
3.1, we know : : qn qn1
q
i a b ,
since
: : : :
i n a
i a b
, we obtain b
1 1
2
: : q qn
nq q
.
In order to get an intuitive idea about the magnitude of all the cardinality numbers, We will find their asymptote
as or
We have the followin n
n q .
g notatio
Definition 3.14 We call
x
f x g x if
lim 1
x
f x
g x .
Now, we can list all the cardinalities asymptotically.
t
Theorem 3.15 If n4 and q2, hen
1 1
1 1
1 1
1 1
1
1 1
1 1
1 1
1 1
1
2 2
: : ; : : ;
: : , : : ;
: : , : : ;
: : , : : ;
: : , : :
n n
n n
n n
n n
n
q q q q
q
q n
q q q q
q n
q q q q
q n
q q q
i a b q i a qq
a b nq a b nq
a nqq a nqq
b nqq b nqq
i q q i q q
1 1
1 1
: : n q qn , : : q q qn ;
i b qq i b qq
n
q q q q
1
1 1
1
1 1
2 2
;
: : , : : .
n
n n
q
q n
q q q q
nq q nq q
Proof:
The first two rows are Theorem 3.1 and Theorem 3.2. We will give a proof of the last row, the others are similar and easier.
1 2 1
1 1
1,0 1 2
1 2
! ! !
.
q i n
q nq
k k
t t t t q q
q q
t t t
nq q
When
1
1 1
! 1
1 !
n
q q
nq
U q
q
1 1
: 1 k 1 k.
k k
U V
:
2 k q, we have
1 1 2
1 1
,0 1 2
2 2
0 , 1,2, ,
! 1
i
t q i q
nq
! 1
! ! ! !
! 1 .
n
q i
n n
q k q k
t t t k t q q
q
q q q q
nq
U q
q k t t t
q nq q q
Hence,
1
1 1
1 1
1
2 1
2
1 2 1
2
1 3
1
! 1 1
.
q
n n
n n
q k
q q q q
k q
k
q q q
q
q q
q q
U
qnq q q q q
U nq q q
q q q
q q
So,
1 21
1
lim 0
q k
k k
n
U
U
.
1 0
V When
1 1
2 2
2
,0 1,0 1 0 , 1,2,
,
1 ! .
n
! 1 2, ( 1)
j j n i i
n
i
q k n
k k k k k k j
n q q
k q i n
V q C q k q
q q q q
n
n
n q q
k
q
j
Note, . Hence,
q
q q
2 21
1 n
n j
j
q k q q
1
12 21
1 1 ! n
nq
k n n q q
k k
V nqq q q q
.We obtain
2 2 1
2 2 2
1
1 1
1 2 1
2
1 1
1
1 !
1 ! 1
.
n
n
n n n
k
n n q q
k k
q q
n n n qn q n
q q q q q
V
nqq q q q
U nq q
q q q q
q
q q
Hence, nq
q
1 11
1
0. lim
nq k
k k
n
V
U
In summary, we obtain
1 1
2 : :
1.
lim q qn
nnq q
In other words,
1 1
2 : :
n q qn . nq q
From the above proof, it is also clear that we have
1 1
2 : :
1.
lim q qn
qnq q
In other words,
1 1
2
: : q q qn .
nq q
□ When , the first equation of the last row in the
ab been obtained in [1].
4. Conclusion
multi-state case. nd Exclusion Principle, we get for- inality all such functions and the car-
cs Institute at Virginia ank Alan Veliz-Cuba and any useful discussions. The
na, “The Number
Physica D, Vol.
197, No. 3-4, 20 doi:10.1016/j.
In this paper, we generalized the definition of Boolean canalyzing functions to the functions of
Using Inclusion a mulas for the card
dinalities of its various subsets. When q2, we derive an interesting combinatorial identity by equating our for- mula to the one in [1]. Finally, for a better understanding to the magnitudes, we provide all the asymptotes of these cardinalities as either n or q
5. Acknowledgements
This work was initiated when the first and the third au- thors visited Virginia Bioinformati
.
Tech in June 2010. We th Franzeska Hinkelmann for m
first and the third authors thank Professor Reinhard Laubenbacher for his hospitality and for introducing them to Discrete Dynamical Systems. The third author was supported in part by a Research Initiation Program (RIP) award at Winston-Salem State University.
We greatly appreciate an anonymous reader. Because of his insightful comments, in this paper, the proofs for many lemmas are simplified, the results are more general (On any finite set instead of finite field).
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