Strengthening Relationships between Neural Ideals and Receptive Fields
Angelique Morvant
(March 8, 2018)
Abstract
Neural codes are collections of binary vectors that represent the firing patterns of neurons. The information given by a neural code C can be represented by its neural ideal JCβ. In turn, the polynomials in JCβ can be used to determine the relationships among the receptive fields of the neurons. In a paper by Curto et al., three such relationships, known as the Type 1-3 relations, were linked to the neural ideal by three if-and-only-if statements. Later, Garcia et al. discovered the Type 4-6 relations. These new relations differed from the first three in that they were related to JCβ by one-way implications. In this paper, we first show that the converses of these new implications are false at the level of both the neural ideal JCβ and the larger ideal I(C) of a code. We then present modified statements of these relations that, like the first three, can be related by if-and-only-if statements to both JCβ and I(C). Using the modified relations, we uncover a new relationship involving JCβ, I(C), and the Type 1-6 relations.
1 Introduction
One of the goals of neuroscience is to determine how the firing of neurons helps the brain understand its environment.
Some neurons are observed to fire rapidly in response to particular stimuli; the set of such stimuli is known as the neuronβs
receptive field. One aim of the study of such neurons is to determine how their firing patterns encode the relationships
among their receptive fields. For example, place cells, discovered in 1971 by OβKeefe and Dostrovsky, fire more rapidly when an animal is
in certain regions, allowing the animal to navigate its environment [4]. In this case, the receptive field of a place cell
is the spatial region in which it fires, and the firing patterns of the place cells allow the brain to construct a βmapβ of these regions. We want to
understand how the brain does this.
To this end, Curto, Itskov, Veliz-Cuba, and Youngs introduced the neural ideal [1]. Firing patterns of neurons can be recorded as collections of binary vectors known as neural codes, and the neural ideal JCβ of a code C is a polynomial ideal that contains the same information as the code itself. Curto et al. showed that the presence of certain types of polynomials in the neural ideal gives information about the relationships among receptive fields [1]. For example, consider two neurons that never fire at the same time. The corresponding neural code might be C={[1,0],[0,1],[0,0]}, and this code is associated with the ideal JCβ=β¨x1βx2ββ©. As we will see, the fact that x1βx2ββJCβ implies that the receptive fields of neuron 1 and neuron 2 do not overlap. So if U1β is the receptive field of neuron 1 and U2β is the receptive field of neuron 2, U1ββ©U2β=β
is a receptive field relation that can be read off from JCβ; it is known as a Type 1 relation. Also, it can be shown that the presence of a Type 1 relation implies that x1βx2ββJCβ. Thus, the relationship between JCβ and the Type 1 relation is if-and-only-if.
Curto et al. found three receptive field relationships, known as the Type 1-3 relationships, that can be read off from the neural ideal. The three if-and-only-if statements relating the Type 1-3 relationships to the polynomials in the neural ideal are also loosely called the Type 1-3 relations. Later, Garcia et al. discovered three more such relations, known as the Type 4-6 relations, but proved only one direction of the relations [2]. It is therefore natural to ask whether the converses of any of the Type 4-6 relations might also hold.
In the sections that follow, we will prove that the answer is no for all three relations, but that we can modify the Type 4-6 relations to be if-and-only-if statements. Like the original Type 1-3 relations, the modified versions of the Type 4-6 relations are if-and-only-if at the level of both JCβ and a larger ideal I(C), called the ideal of C. This suggests that, at least for the purposes of receptive field relations, the ideal and neural ideal of a code are interchangeable.
The rest of this paper will be organized as follows. In Section 2, we will introduce the background and definitions needed in the rest of the work. In Section 3, we will show by counterexample that the converses of the Type 4-6 relations do not hold. In Section 4, we will present modified versions of these relations and prove that these modified relations are if-and-only-if. Finally, in Section 5, we will discuss the implications of our results and suggest topics for future research.
2 Background
In this section we introduce neural codes, pseudo-monomials, and neural ideals, as well as the prior results on which our work is based. Our notation matches that in [1, 2]. We begin with neural codes.
Definition 2.1**.**
A neural code (or binary code) C is a set of vectors in F2nβ. A vector cβC is called a codeword.
Each codeword c in a neural code represents a firing pattern of n neurons: the ith component of c is 1 if neuron i is firing and [math] if it is not. For example, the neural code C={[0,1,1],[1,0,1],[0,0,1],[0,0,0]} consists of the codewords c1β=[0,1,1], c2β=[1,0,1], c3β=[0,0,1], and c4β=[0,0,0]. The codeword c1β tells us that neurons 2 and 3 fire together while neuron 1 does not, c2β tells us that neurons 1 and 3 fire together while neuron 2 does not, c3β tells us that neuron 3 fires alone, and c4β tells us that none of the neurons fire.
Alternatively, each codeword cβC can also be represented by the set
[TABLE]
where [n]:={1,2,...,n}. So in the example above, supp(c1β)={2,3}, supp(c2β)={1,3}, supp(c3β)={3}, and supp(c4β)=β
. By dropping the set notation, we can write C in shorthand as C={β
,3,13,23}.
As mentioned in the previous section, neurons such as place cells fire in specific regions of a stimulus space known as receptive fields. For a (nonempty) stimulus space X, let U={U1β,U2β,...,Unβ}, where each UiββX is the receptive field of neuron i. Then we can define the associated receptive field code as
[TABLE]
In addition, for any point pβX, we will let c(p) denote the codeword such that supp(c(p))={iβ[n]Β β£Β pβUiβ}. Going back to our example, the code C={β
,3,13,23} is the receptive field code for the set U={U1β,U2β,U3β} shown in Figure 1.
The information in a code on n neurons can also be encoded in an ideal generated by pseudo-monomials in F2β[x1β,...xnβ].
Definition 2.2**.**
A pseudo-monomial fβF2β[x1β,...,xnβ] is a polynomial with the form
[TABLE]
where Ο,Οβ[n] and Οβ©Ο=β
.
For any vector vβF2nβ, we define
[TABLE]
Such a pseudo-monomial is called the characteristic function for v because pvβ(v)=1, but pvβ(x)=0 for all xβF2nβ such that xξ =v. The neural ideal of a code is defined in terms of characteristic functions.
Definition 2.3**.**
The neural ideal JCβ of a code C is the ideal generated by all pvββF2β[x1β,...,xnβ] such that vβ/C. That is,
[TABLE]
Notice that this implies that if fβJCβ, then f(c)=0 for any cβC. In [1], Curto et al. define the ideal of a code as follows:
Definition 2.4**.**
Let CβF2nβ be a neural code. Then the ideal of C is
[TABLE]
By this definition, JCββI(C). This fact will be important in later sections.
The polynomials in a neural ideal give information about the relationships among the receptive fields in a stimulus space. The first three relations (known as the Type 1-3 relations) were discovered by Curto, Itskov, Veliz-Cuba, and Youngs [1]. To simplify notation, we
let
[TABLE]
for any Οβ[n]. Note that if Ο=β
, then xΟβ=βiβΟβxiβ=1 and UΟβ=X.
Proposition 2.5** (Curto, Itskov, Veliz-Cuba, and Youngs).**
Let X be a stimulus space, let U={Uiβ}i=1nβ be a collection of sets in X, and consider the receptive field code C=C(U). Then for any pair of subsets Ο,Οβ[n], we have the following receptive field relations:
xΟββJCββUΟβ=β
* (where Οξ =β
).*
xΟββiβΟβ(1+xiβ)βJCββUΟβββͺiβΟβUiβ* (where Ο,Οξ =β
).*
βiβΟβ(1+xiβ)βJCββXββͺiβΟβUiβ* (where Οξ =β
), and thus X=βͺiβΟβUiβ.*
In addition, Garcia et al. found three more receptive field relationships [2, Theorem 5.1]:
Proposition 2.6** (Garcia, GarcΓa Puente, et al.).**
Let U={Uiβ}i=1nβ be a collection of sets in a stimulus space X. Let C=C(U) denote the corresponding receptive field code, and let JCβ denote the neural ideal. Then for any subsets Ο1β, Ο2β, Ο1β, Ο2ββ[n], and m indices 1β€iiβ<i2β<...<imββ€n, with mβ₯2, we have receptive field relationships as follows:
xΟ1βββiβΟ1ββ(1+xiβ)+xΟ2βββjβΟ2ββ(1+xjβ)βJCββUΟ1βββ©(β©iβΟ1ββUicβ)=UΟ2βββ©(β©jβΟ2ββUjcβ).
xi1ββ+...+ximβββJCββUikββββͺjβ[m]β{k}βUijββ* for all k=1,...,m, and if, additionally, m is odd, then β©k=1mβUikββ=β
.*
xi1ββ+...+ximββ+1βJCβββͺk=1mβUikββ=X.
Notice that, unlike the Type 1-3 relations, the Type 4-6 relations are not stated as if-and-only-if statements. That is, knowing the relations among receptive fields in a stimulus space, we cannot use the Type 4-6 relations in their current form to conclude anything about the associated neural ideal. It is therefore natural to wonder whether each of these statements can be reversed, and in Section 3 we show that in fact none of them can. However, in Section 4 we will present modified versions of the Type 4-6 relations that are if-and-only-if statements.
3 Disproving the Converses of the Type 4-6 Relations
In this section, we show by counterexample that none of the converses of the Type 4-6 relations hold (Theorem 3.3). The converses of the relations are stated below:
Let U={Uiβ}i=1nβ be a collection of sets in a stimulus space X. Let C=C(U) denote the corresponding receptive field code, and let JCβ be the neural ideal of C. Then for any subsets Ο1β, Ο2β, Ο1β, Ο2ββ[n] and m indices 1β€i1β<i2β<...<imββ€n, with mβ₯2, we have the following:
UΟ1βββ©(β©iβΟ1ββUicβ)=UΟ2βββ©(β©jβΟ2ββUjcβ)βxΟ1βββiβΟ1ββ(1+xiβ)+xΟ2βββjβΟ2ββ(1+xjβ)βJCβ* for any subsets Ο1β, Ο2β, Ο1β, Ο2ββ[n].*
For all mβ₯2 indices 1β€iiβ<i2β<...<imββ€n,
-
m* is even and Uikββββͺjβ[m]β{k}βUijββ for all k=1,...,mβxi1ββ+...+ximβββJCβ, and*
2. 2.
m* is odd, Uikββββͺjβ[m]β{k}βUijββ for all k=1,...,m, and β©k=1mβUikββ=β
βxi1ββ+...+ximβββJCβ.*
βͺk=1mβUikββ=Xβxi1ββ+...+ximββ+1βJCβ* for all mβ₯2 indices 1β€iiβ<i2β<...<imββ€n.*
Again, each of the converses is false. The receptive fields for all of the counterexample codes are shown in Figure 2. In the case of the Type 5 and 6 relations, we will first use the counterexample codes to prove that the converses of these statements do not hold even when JCβ is replaced by the larger ideal I(C).
Lemma 3.1**.**
For each of the following relations, there exists a collection of sets U={Uiβ}i=1nβ in a stimulus space X (with corresponding neural code C=C(U)) and mβ₯2 indices 1β€iiβ<i2β<...<imββ€n such that the relation does not hold:
-
m* is even and Uikββββͺjβ[m]β{k}βUijββ for all k=1,...,mβxi1ββ+...+ximβββI(C), and*
2. 2.
m* is odd, Uikββββͺjβ[m]β{k}βUijββ for all k=1,...,m, and β©k=1mβUikββ=β
βxi1ββ+...+ximβββI(C).*
βͺk=1mβUikββ=Xβxi1ββ+...+ximββ+1βI(C)**
Proof.
Converse of Type 5: To disprove the converse of the Type 5 relation, it is enough to prove that the first implication is false. To this end, let C={β
,12,13,14,123}. The corresponding stimulus space is shown in Figure 2(b). From the figure, we can see that Uikββββͺjβ[4]β{k}βUijββ, where i1β=1,...,i4β=4. However, if f:=x1β+x2β+x3β+x4β, then for any point pβ(U1ββ©U2ββ©U3β)βU4β, f(c(p))=(x1β+x2β+x3β+x4β)(c(p))=1+1+1+0β‘1mod2, so f=x1β+x2β+x3β+x4ββ/I(C).
There are also counterexamples for the second implication. For instance, the code Cβ²={β
,12,13,14,123,145} (shown in Figure 3) with i1β=1,...,i5β=5 satisfies Uikββββͺjβ[5]β{k}βUijββ for all k=1,...,5, and β©k=15βUikββ=β
. But, again, if g:=x1β+x2β+x3β+x4β+x5β and pβ(U1ββ©U2ββ©U3β)β(U4ββͺU5β), then g(c(p))=(x1β+x2β+x3β+x4β+x5β)(c(p))=1+1+1+0+0β‘1mod2. So gβ/I(Cβ²).
Converse of Type 6: Now consider the code C={1,2,12} on 2 neurons pictured in Figure 2(c). We compute that
[TABLE]
and we can see that βͺi=12βUiβ=X. If the converse of the Type 6 relation held, then it would be true that
[TABLE]
However,
[TABLE]
Adding expressions (1) and (2) would then imply that f:=x1βx2ββI(C). However, for a point pβU1ββ©U2β, f(c(p))=1β
1=1 and thus fβ/I(C), a contradiction. So x1β+x2β+1β/I(C).
β
Now we are ready to show that none of converses of the Type 4-6 relations hold.
Theorem 3.2**.**
For each of the following relations, there exists a collection of sets U={Uiβ}i=1nβ in a stimulus space X (with corresponding neural code C=C(U)) such that the relation does not hold:
UΟ1βββ©(β©iβΟ1ββUicβ)=UΟ2βββ©(β©jβΟ2ββUjcβ)βxΟ1βββiβΟ1ββ(1+xiβ)+xΟ2βββjβΟ2ββ(1+xjβ)βJCβ* for any subsets Ο1β, Ο2β, Ο1β, Ο2ββ[n].*
For all mβ₯2 indices 1β€iiβ<i2β<...<imββ€n,
-
m* is even and Uikββββͺjβ[m]β{k}βUijββ for all k=1,...,mβxi1ββ+...+ximβββJCβ, and*
2. 2.
m* is odd, Uikββββͺjβ[m]β{k}βUijββ for all k=1,...,m, and β©k=1mβUikββ=β
βxi1ββ+...+ximβββJCβ.*
βͺk=1mβUikββ=Xβxi1ββ+...+ximββ+1βJCβ* for all mβ₯2 indices 1β€iiβ<i2β<...<imββ€n.*
Proof.
We present a counterexample of each of the three statements, beginning with the converse of the Type 4 relation. The counterexamples for the Type 5 and 6 relations are the same as in the proof of Lemma 3.1. As we will see, we can use the same counterexamples for these two relations because JCββI(C).
Converse of Type 4: Consider the code C={β
,1,2,12} on 2 neurons pictured in Figure 2(a). Also let Ο1β=Ο1β=1 and Ο2β=Ο2β=2. Then U1ββ©U1cβ=β
=U2ββ©U2cβ, but JCβ=β¨0β©, so x1β(1+x1β)+x2β(1+x2β)β/JCβ.
Converse of Type 5: Again let C={β
,12,13,14,123}. The realization of this code is shown in Figure 2(b). From the proof of Lemma 3.1, we know that x1β+x2β+x3β+x4ββ/I(C), so x1β+x2β+x3β+x4ββ/JCββI(C). This shows that the first implication is false.
As before, however, we can also show that the second implication is false. The code Cβ²={β
,12,13,14,123,145} again serves as a counterexample. By the proof of Lemma 3.1, x1β+x2β+x3β+x4β+x5ββ/I(Cβ²), so x1β+x2β+x3β+x4β+x5ββ/JCβ²ββI(Cβ²).
Converse of Type 6: Finally, the proof of Lemma 3.1 tells us that for the code C={1,2,12} pictured in Figure 2(c), x1β+x2β+1β/I(C). Therefore, x1β+x2β+1β/JCββI(C).
β
Notice that if we replace JCβ with I(C) in the Type 4 relation, the counterexample we used in the proof of Theorem 3.2 no longer works since x1β(1+x1β)+x2β(1+x2β)βI(C). In fact, we will prove in the next section that the converse of the Type 4 relation is actually true when JCβ replaced with I(C). This suggests that the Type 4-6 relations may be modified so that their converses hold. Possible modifications will be discussed in Section 4.
We finish this section by showing that the counterexample codes presented all have the smallest possible numbers of neurons.
Theorem 3.3**.**
Let U={Uiβ}i=1nβ be a collection of sets in a stimulus space X. Let C=C(U) denote the corresponding receptive field code, and let JCβ denote the neural ideal. Then for any subsets Ο1β, Ο2β, Ο1β, Ο2ββ[n], and m indices 1β€i1β<i2β<...<imββ€n, with mβ₯2, the following hold:
When n=1, UΟ1βββ©(β©iβΟ1ββUicβ)=UΟ2βββ©(β©jβΟ2ββUjcβ)βxΟ1βββiβΟ1ββ(1+xiβ)+xΟ2βββjβΟ2ββ(1+xjβ)βJCβ.
When nβ€3,
-
m* is even and Uikββββͺjβ[m]β{k}βUijββ for all k=1,...,mβxi1ββ+...+ximβββJCβ, and*
2. 2.
m* is odd, Uikββββͺjβ[m]β{k}βUijββ for all k=1,...,m, and β©k=1mβUikββ=β
βxi1ββ+...+ximβββJCβ.*
When n=1, βͺk=1mβUikββ=Xβxi1ββ+...+ximββ+1βJCβ.
Proof.
Converse of Type 4: For n=1, the possible neural codes are C1β={β
,1} and C2β={1}.
In the case of C1β, JCβ=β¨0β©, and the possible values for UΟ1βββ©(β©iβΟ1ββUicβ) are U1ββ©U1cβ=β
(when Ο1β=Ο1β=1), U1ββ©X=U1β (when Ο1β=1 and Ο1β=β
), Xβ©U1cβ=U1cβ (when Ο1β=β
and Ο1β=1), and Xβ©X=X (when Ο1β=Ο1β=β
). Since the possible values of UΟ2βββ©(β©jβΟ2ββUjcβ) are also β
, U1β, U1cβ, and X, and since none of these four sets are equal, UΟ1βββ©(β©iβΟ1ββUicβ)=UΟ2βββ©(β©jβΟ2ββUjcβ) only when Ο1β=Ο2β and Ο1β=Ο2β. This implies that xΟ1βββiβΟ1ββ(1+xiβ)+xΟ2βββjβΟ2ββ(1+xjβ)=xΟ1βββiβΟ1ββ(1+xiβ)+xΟ1βββjβΟ1ββ(1+xjβ)=0βJCβ.
In the case of C2β, JCβ=β¨1+x1ββ©, U1β=X, and U1cβ=β
. This means that there are two choices of Ο1β and Ο1β for which UΟ1βββ©(β©iβΟ1ββUicβ)=β
: Ο1β=Ο1β=1, or Ο1β=β
and Ο1β=1. Similarly, UΟ1βββ©(β©iβΟ1ββUicβ)=X when either Ο1β=1 and Ο1β=β
, or Ο1β=Ο1β=β
. The same is true for UΟ2βββ©(β©iβΟ2ββUicβ), so UΟ1βββ©(β©iβΟ1ββUicβ)=UΟ2βββ©(β©jβΟ2ββUjcβ) implies that we have one of the following cases:
Ο1β=Ο2β and Ο1β=Ο2β
Ο1βξ =Ο2β and Ο1β=Ο2β={1}
Ο1βξ =Ο2β and Ο1β=Ο2β=β
In case 1, we can use the same reasoning we used when dealing with C1β, so it is enough to show that the converse of the Type 4 relation holds in cases 2 and 3. For these cases, we can assume without loss of generality that Ο1β=1 and Ο2β=β
.
In case 2,
[TABLE]
because 1+x1ββJCβ.
Similarly, in case 3,
[TABLE]
So the implication holds.
Converse of Type 5: We consider the cases where n=1, n=2, and n=3 separately.
In the case where n=1, we cannot have mβ₯2 indices , so the implication is vacuously true.
When n=2, in order for the left-hand side of the implication to be true, we must have m=2, U1ββU2β, and U2ββU1β, which implies that U1β=U2β. This means that neither 1 nor 2 is in the associated code C, so x1β(1+x2β),x2β(1+x1β)βJCβ. Therefore, x1β(1+x2β)+x2β(1+x1β)=x1β+x2ββJCβ
Finally, when n=3, we can have m=2 or m=3. If m=2, then Ui1ββ=Ui2ββ as in the n=2 case. If we let i3β be the remaining element of {1,2,3}, then i1β, i2β, i1βi3β, i2βi3ββ/C, which implies that the following pseudo-monomials are in JCβ:
[TABLE]
Adding these four terms together gives us xi1ββ+xi2βββJCβ.
If m=3, similar reasoning applies, but we also must assume that β©k=13βUikββ=β
. We know that the codewords 1, 2, 3β/C since Uikββββͺjβ[3]β{k}βUijββ for all k=1,2,3, and we know that 123β/C because β©k=13βUikββ=β
by assumption. Therefore, the following are in JCβ:
[TABLE]
Adding the four terms again gives x1β+x2β+x3ββJCβ.
Converse of Type 6: Again, when n=1, it is impossible to have mβ₯2 different indices, so the implication is vacuously true.
β
4 Modifying the Type 4-6 Relations
In this section, we discuss modified versions of the Type 4-6 relations and prove that their converses hold. The modifications are marked in bold in the following result.
Theorem 4.1**.**
Let U={Uiβ}i=1nβ be a collection of sets in a stimulus space X. Let C=C(U) denote the corresponding receptive field code, and let JCβ be the neural ideal of C. Then for any subsets Ο1β, Ο2β, Ο1β, Ο2ββ[n] such that Ο1ββͺΟ2β and Ο1ββͺΟ2β are disjoint, and m indices 1β€i1β<i2β<...<imββ€n, with mβ₯2, we have the following equivalences:
xΟ1βββiβΟ1ββ(1+xiβ)+xΟ2βββjβΟ2ββ(1+xjβ)βJCββUΟ1βββ©(β©iβΟ1ββUicβ)=UΟ2βββ©(β©jβΟ2ββUjcβ)**
xi1ββ+...+ximβββJCββ* for any Οβ[m] such that β£Οβ£ is odd, β©kβΟβUikββββͺjβ[m]βΟβUijββ*
xi1ββ+...+ximββ+1βJCβββͺk=1mβUikββ=X* whenever Ui1ββ,...,Uimββ are pairwise disjoint.*
Proof.
By Proposition 2.6, we know that the forward direction of each of the three implications is true, so it only remains to prove the reverse implications.
Modified Type 4: Given a neural code C that satisfies a relation of the form UΟ1βββ©(β©iβΟ1ββUicβ)=UΟ2βββ©(β©jβΟ2ββUjcβ), we will construct the desired sum by adding pseudo-monomials known to be in JCβ. There are two cases: one where Ο1ββͺΟ2β is empty, and the remaining case where Ο1ββͺΟ2β is nonempty. We deal with each of these cases in turn.
Case 1 (Ο1ββͺΟ2β=β
): In this case, UΟ1ββ=UΟ2ββ (we ignore the trivial cases where Ο1β or Ο2β is empty or where Ο1β=Ο2β). Then any codeword c for which Ο1ββc but Ο2βξ βc, or Ο2ββc but Ο1βξ βc, cannot be in C (this would contradict the fact that UΟ1ββ=UΟ2ββ), and so pcβ=βiβcβxiββjβ/cβ(1+xjβ)βJCβ for every such c. Letting C1β={cβ[n]β£Ο1ββc but Ο2βξ βc}, C2β={cβ[n]β£Ο2ββc but Ο1βξ βc}, and f=βcβC1ββͺC2ββpcβ, we claim that f=xΟ1ββ+xΟ2ββ and that fβJCβ as desired.
To see this, first note that fβJCβ because each pcβ is in JCβ by the remarks above. To prove that f=xΟ1ββ+xΟ2ββ, we will consider xΟβ for all Οβ[n] and determine whether xΟβ appears as a summand in f.
First consider xΟβ, where Ο1βξ βΟ and Ο2βξ βΟ. Then xΟβ is not a term in the pseudo-monomial pcβ for any cβC1ββͺC2β since all terms in such a pseudo-monomial must be of the form xΟ1ββxΟβ²β or xΟ2ββxΟβ²β for some Οβ²β[n]. So xΟβ is not a term in f.
If Ο1ββΟ but Ο2βξ βΟ, then xΟβ is not a term in pcβ for any cβC2β or for any cβC1β such that cξ βΟ. But xΟβ appears once as a term in pcβ for all cβC1β such that cβΟ. There are 2m such c, where m=β£ΟβΟ1ββ£. Therefore, if mβ₯1, xΟβ will be added an even number of times in the expansion of f and will therefore cancel out since we are working in F2β[x]. We get m=0 only when Ο=Ο1β, which tells us that xΟ1ββ is a monomial term in f. Similar reasoning applies when Ο2ββΟ but Ο1βξ βΟ.
Finally, if Ο1ββͺΟ2ββΟ, then xΟβ is a term in pcβ for any cβC1ββͺC2β such that cβΟ. As before, we need to determine how many cβC1ββͺC2β are contained in Ο. We know there are 2m codewords c such that Ο1ββcβΟ and 2k codewords cβ² such that Ο2ββcβ²βΟ, where m=β£ΟβΟ1ββ£β₯1 and k=β£ΟβΟ2ββ£β₯1 111Note that if m=0, then Ο1ββͺΟ2ββΟ=Ο1β, which would imply that Ο1β=Ο2β. We can ignore this case because in this case xΟ1ββ+xΟ2ββ=0, and 0βJCβ for all C.. However, if we count all such c and cβ², we will also count twice the 2l codewords cβ²β² such that Ο1ββͺΟ2ββcβ²β²βΟ. (Here l=β£Οβ(Ο1ββͺΟ2β)β£β₯0.) These codewords cβ²β² are not in C1ββͺC2β and therefore must be excluded twice from our count of codewords in C1ββͺC2β contained in Ο. So there are 2m+2kβ2(2l) codewords contained in Ο that are in C1ββͺC2β. This implies that xΟβ will be added 2m+2kβ2(2l) times in f and therefore will cancel out since 2m+2kβ2(2l) is even.
Thus, the only terms that appear in f are xΟ1ββ and xΟ2ββ, so f=xΟ1ββ+xΟ2βββJCβ as desired.
Case 2 (Ο1ββͺΟ2βξ =β
): In the case where Ο1ββͺΟ2βξ =β
and UΟ1βββ©(β©iβΟ1ββUicβ)=UΟ2βββ©(β©jβΟ2ββUjcβ), we can apply the same reasoning used in the previous case if we first define a new stimulus space {Uiβ²β} (with receptive field code Cβ²) as follows:
[TABLE]
Because Ο1ββͺΟ2β and Ο1ββͺΟ2β are disjoint and UΟ1βββ©(β©iβΟ1ββUicβ)=UΟ2βββ©(β©jβΟ2ββUjcβ), we know that UΟ1ββ²ββ©(β©iβΟ1ββUiβ²β)=UΟ2ββ²ββ©(β©jβΟ2ββUjβ²β) as well. Therefore, we can say from the first case that xΟ1ββͺΟ1ββ+xΟ2ββͺΟ2βββJCβ²β.
Now we need to show that the corresponding pseudo-monomial xΟ1βββiβΟ1ββ(1+xiβ)+xΟ2βββjβΟ2ββ(1+xjβ) is in JCβ. Consider the i-th bit flip map Ξ΄iβ : F2β[x1β,...,xnβ]βF2β[x1β,...,xnβ] given by
[TABLE]
as defined in Jeffs, Omar, and Youngs [3], and let Ξ΄Ο1ββͺΟ2ββ be the composition of all Ξ΄iβ such that iβΟ1ββͺΟ2β. Also as in [3], for a neural code C we define the following code:
[TABLE]
where β denotes the symmetric difference. Then by Theorem 2.13 in [3], we know that Ξ΄iβ(JCβ²β)=JΞ΄iβ(Cβ²)β for each iβΟ1ββͺΟ2β, so Ξ΄Ο1ββͺΟ2ββ(JCβ²β)=Ξ΄i1βββΞ΄i2βββ...βΞ΄imββ(JCβ²β)=JΞ΄i1βββΒ Ξ΄i2βββ...βΒ Ξ΄imββ(Cβ²)β=JΞ΄Ο1ββͺΟ2ββ(Cβ²)β for i1β,i2β,...imββΟ1ββͺΟ2β. But from (3), we can see that Ξ΄Ο1ββͺΟ2ββ maps Cβ² into C, and because Ξ΄Ο1ββͺΟ2ββ(Ξ΄Ο1ββͺΟ2ββ(c))=c for any cβCβ², Ξ΄Ο1ββͺΟ2ββ is a bijection, which implies that JΞ΄Ο1ββͺΟ2ββ(Cβ²)β=JCβ. Since each Ξ΄iβ is a homomorphism [3], Ξ΄Ο1ββͺΟ2ββ is also a homomorphism from JCβ²β to JCβ, so Ξ΄Ο1ββͺΟ2ββ(xΟ1ββͺΟ1ββ+xΟβͺβΟ2ββ)=xΟ1βββiβΟ1ββ(1+xiβ)+xΟ2βββjβΟ2ββ(1+xjβ)βJCβ.
Modified Type 5: Suppose that for any Οβ[m] such that β£Οβ£ is odd, β©kβΟβUikββββͺjβ[m]βΟβUijββ. Then by the Type 2 relation, βkβΟβxikβββjβ[m]βΟβ(1+xijββ)βJCβ for any such Ο. So if S is the set of all Οβ[m] such that β£Οβ£ is odd, then
[TABLE]
We claim that f=xi1ββ+...+ximββ. To see this, consider the monomial βkβΟβxikββ, where Οβ[m]. (Note that such terms are the only possible monomials that can appear in the expansion of f.) We know that βkβΟβxikββ will appear exactly once in the expansion of every summand βkβΟβxikβββjβ[m]βΟβ(1+xijββ) for which ΟβΟ and ΟβS, and never in the expansions of the other summands. Call the number of such summands NΟβ. If β£Οβ£=1, then the only odd-sized subset of Ο is Ο itself, so the term βkβΟβxikββ must appear exactly once in the expansion of f. That is, NΟβ=1. If β£Οβ£β₯2, then NΟβ=β1β€iβ€β£Οβ£,Β iΒ oddβ(iβ£Οβ£β)=2β£Οβ£β1, and since β£Οβ£β₯2, NΟβ must be even. This means that βkβΟβxikββ cancels out of f when β£Οβ£β₯2, so the only terms that appear in the expansion of f are xi1ββ,...,ximββ, which implies that f=xi1ββ+...+ximββ.
Modified Type 6: Finally, suppose that βͺk=1mβUikββ=X and that Ui1ββ,...,Uimββ are pairwise disjoint. Also let M={i1β,...,imβ}. Then by the Type 3 relation we know that βiβMβ(1+xiβ)βJCβ. But βiβMβ(1+xiβ)=βΟβMβxΟβ, and since Ui1ββ,...,Uimββ are pairwise disjoint, xΟββJCβ for all ΟβM such that β£Οβ£β₯2 by the Type 1 relation. Therefore,
[TABLE]
β
Recall that in the previous section we showed that the converses of the original Type 5 and 6 relations are false when JCβ is replaced by I(C). However, it turns out that the converses of the modified Type 4-6 relations all hold when JCβ is replaced by I(C).
Theorem 4.2**.**
Let U={Uiβ}i=1nβ be a collection of sets in a stimulus space X. Let C=C(U) denote the corresponding receptive field code, and let I(C) be the ideal of C. Then for any subsets Ο1β, Ο2β, Ο1β, Ο2ββ[n] such that Ο1ββͺΟ2β and Ο1ββͺΟ2β are disjoint, and m indices 1β€i1β<i2β<...<imββ€n, with mβ₯2, we have the following equivalences:
xΟ1βββiβΟ1ββ(1+xiβ)+xΟ2βββjβΟ2ββ(1+xjβ)βI(C)βUΟ1βββ©(β©iβΟ1ββUicβ)=UΟ2βββ©(β©jβΟ2ββUjcβ)**
xi1ββ+...+ximβββI(C)β* for any Οβ[m] such that β£Οβ£ is odd, β©kβΟβUikββββͺjβ[m]βΟβUijββ*
xi1ββ+...+ximββ+1βI(C)ββͺk=1mβUikββ=X* whenever Ui1ββ,...,Uimββ are pairwise disjoint.*
Proof.
The backward directions of all the statements are true by Theorem 4.1 and the fact that JCββI(C). Therefore, it remains to prove the forward implications. We do this by essentially repeating the proofs in [2].
Modified Type 4: Let f1β:=xΟ1βββiβΟ1ββ(1+xiβ), let f2β=xΟ2βββjβΟ2ββ(1+xjβ), and suppose that f1β+f2ββI(C). Also suppose that pβUΟ1βββ©(β©iβΟ1ββUicβ). Then because f1β+f2ββI(C), f1β(c(p))+f2β(c(p))=0, which implies that f1β(c(p))=f2β(c(p)). But pβUΟkβββ©(β©jβΟkββUjcβ) if and only if fkβ(c(p))=1, so pβUΟ2βββ©(β©jβΟ2ββUjcβ). The same argument can be used to show that if pβUΟ2βββ©(β©jβΟ2ββUjcβ), then pβUΟ1βββ©(β©iβΟ1ββUicβ).
Modified Type 5: Suppose g:=xi1ββ+...+ximβββI(C). Let Οβ[m] such that β£Οβ£ is odd, and let pββ©kβΟβUikββ. (We can assume that β©kβΟβUikββξ =β
because the desired containment would be automatic otherwise.) Then because gβI(C), g(c(p))=0=c(p)i1ββ+...+c(p)imββ=βkβΟβc(p)ikββ+βjβ[m]βΟβc(p)ijββ=1+βjβ[m]βΟβc(p)ijββ in F2β since β£Οβ£ is odd. Then βjβ[m]βΟβc(p)ijββ=1, which implies that c(p)ijββ=1 for some jβ[m]βΟ, so pβUijββ for this j. Therefore, β©kβΟβUikββββͺjβ[m]βΟβUijββ.
Modified Type 6: Finally, suppose h:=xi1ββ+...+ximββ+1, and let pβX. Since hβI(C), 0=c(p)i1ββ+...+c(p)imββ+1. Since we are working in F2β, this implies that for some kβ[m], c(p)ikββ=1, and thus pβUikββββͺk=1mβUikββ.
β
Directly from Theorems 4.1 and 4.2, we get the following corollary, which we view as the βcorrectedβ Type 4-6 relations.
Corollary 4.3**.**
Let U={Uiβ}i=1nβ be a collection of sets in a stimulus space X, and let C=C(U) denote the corresponding receptive field code. Then for any subsets Ο1β, Ο2β, Ο1β, Ο2ββ[n] such that Ο1ββͺΟ2β and Ο1ββͺΟ2β are disjoint, and m indices 1β€i1β<i2β<...<imββ€n, with mβ₯2, we have the following:
xΟ1βββiβΟ1ββ(1+xiβ)+xΟ2βββjβΟ2ββ(1+xjβ)βJCββxΟ1βββiβΟ1ββ(1+xiβ)+xΟ2βββjβΟ2ββ(1+xjβ)βI(C)βUΟ1βββ©(β©iβΟ1ββUicβ)=UΟ2βββ©(β©jβΟ2ββUjcβ).* ***
xi1ββ+...+ximβββJCββxi1ββ+...+ximβββI(C)β* for any Οβ[m] such that β£Οβ£ is odd, β©kβΟβUikββββͺjβ[m]βΟβUijββ.*
xi1ββ+...+ximββ+1βJCββxi1ββ+...+ximββ+1βI(C)ββͺk=1mβUikββ=X* whenever Ui1ββ,...,Uimββ are pairwise disjoint.*
Essentially, Corollary 4.3 tells us that we get the same information from the modified relations regardless of whether we consider JCβ or I(C). In fact, this is true for the Type 1-3 relations as well, as shown in [1, Lemma 4.2].
As a final note, we can use Theorem 4.1 and part of the proof of Theorem 4.2 to show that, as stated in the previous section, the original Type 4 relation is if-and-only-if when JCβ is replaced by I(C).
Corollary 4.4**.**
Let U={Uiβ}i=1nβ be a collection of sets in a stimulus space X, and let C=C(U) denote the corresponding receptive field code. Then for any subsets Ο1β, Ο2β, Ο1β, Ο2ββ[n], we have the following:
xΟ1βββiβΟ1ββ(1+xiβ)+xΟ2βββjβΟ2ββ(1+xjβ)βI(C)βUΟ1βββ©(β©iβΟ1ββUicβ)=UΟ2βββ©(β©jβΟ2ββUjcβ).
Proof.
We prove the forward direction using the same reasoning used to prove the forward direction of the modified Type 4 relation in Theorem 4.2. Thus, it remains to prove the backward direction.
To this end, assume that UΟ1βββ©(β©iβΟ1ββUicβ)=UΟ2βββ©(β©jβΟ2ββUjcβ), and consider the following three cases.
Case 1 ((Ο1ββͺΟ2β)β©(Ο1ββͺΟ2β)=β
): If Ο1ββͺΟ2β and Ο1ββͺΟ2β are disjoint, then by Theorem 4.1, we know that xΟ1βββiβΟ1ββ(1+xiβ)+xΟ2βββjβΟ2ββ(1+xjβ)βJCββI(C).
Case 2 (Ο1ββ©Ο1βξ =β
or Ο2ββ©Ο2βξ =β
): Assume without loss of generality that Ο1ββ©Ο1βξ =β
. This implies that UΟ1βββ©(β©iβΟ1ββUicβ)=β
, which implies by hypothesis that UΟ2βββ©(β©iβΟ2ββUicβ)=β
as well. Now let f1β=xΟ1βββiβΟ1ββ(1+xiβ) and f2β=xΟ2βββjβΟ2ββ(1+xjβ). If pβX, then f1β(c(p))=1 would imply that pβUΟ1βββ©(β©iβΟ1ββUicβ)=β
, a contradiction. Since any codeword c is associated with some pβX, f1β(c)=0 for all cβC. Thus, f1ββI(C). By the same reasoning, f2ββI(C), and so f1β+f2ββI(C) as well.
Case 3 (Ο1ββ©Ο2βξ =β
or Ο2ββ©Ο1βξ =β
): Assume without loss of generality that Ο1ββ©Ο2βξ =β
. This means that there is some kβΟ1ββ©Ο2β. Then UkββUΟ1βββ©(β©iβΟ1ββUicβ)=UΟ2βββ©(β©jβΟ2ββUjcβ)βUkcβ, which implies that UΟ1βββ©(β©iβΟ1ββUicβ)=UΟ2βββ©(β©jβΟ2ββUjcβ)=β
. So by the same reasoning used in Case 2, xΟ1βββiβΟ1ββ(1+xiβ)+xΟ2βββjβΟ2ββ(1+xjβ)βI(C).
β
5 Discussion
In this work we proved that not only are the converses of the Type 4-6 relations in [2] false as stated, but the converses of the Type 5 and 6 relations are also false even when the neural ideal JCβ is replaced by the larger ideal I(C). However, our modified versions of the Type 4-6 relations are if-and-only-if statements at the level of both JCβ and I(C). From this we concluded that, in the case of these modified relations, JCβ and I(C) give the same information about the stimulus space. In fact, this is true for the Type 1-3 relations as well [1]. These observations suggest that future receptive field relationships should only involve polynomials that are in JCβ if and only if they are in I(C). Identifying such receptive field relationships is an interesting direction for future work.
In addition, there may be other modifications of the Type 4-6 relations that are also if-and-only-if statements. For instance, replacing JCβ with I(C) in the original Type 4 relation also gives an if-and-only-if statement. If each of the relations could be modified in more than one way, it would be natural to ask which modifications were the βrightβ modifications; that is, which would reveal the most useful information about the corresponding receptive fields.
Finally, for some codes the original Type 4-6 relations are already if-and-only-if. Therefore, we can ask if all such codes have some property in common, and, conversely, we can ask if all codes with a certain property (convexity, for instance β see [1]) also have the property that the reverse implications of the Type 4-6 relations hold.
Acknowledgments
This research was conducted under the mentorship of Dr. Anne Shiu in the Department of Mathematics at Texas A&M University. The author would like to thank Dr. Shiu for all of her guidance and support, as well as Dr. Kaitlyn Phillipson for her comments on an earlier draft of this paper.