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Abstract
We study Lotka–Volterra coalgebras, a new family of nonassociative coalgebras emerging from population genetics. We focus on their defining algebraic properties and deal with characterizing the existence of counital and character maps. We also provide a classification of their basis elements into (co)algebraically persistent and transient generators resulting in a semidirect sum decomposition of Lotka–Volterra coalgebras.
1. Introduction
Lotka–Volterra algebras were introduced by Itoh [Citation1] as a nonassociative framework for describing ternary interactions in competition systems. Despite being aimed by random models mirroring the kinetic theory of gases, a fact remarked by Itoh in [Citation1], these algebras were later shown to include, as particular examples, normal Bernstein algebras. Bernstein algebras became essential in the study of quadratic stochastic operators in population genetics [Citation2].
Given a field of characteristic not 2, a commutative
-algebra
is Lotka–Volterra if it admits a basis
such that
, for all i, j = 1, …, n, where
is a skew-symmetric matrix. We refer to [Citation3, Citation4] and references therein for the algebraic treatment of Lotka–Volterra algebras. Alternatively, Lotka–Volterra algebras can be defined as follows:
Definition 1.1
A -algebra
is Lotka–Volterra (for short, an LV-algebra) if it is endowed with a basis
and multiplication
, such that for any 1 ≤ i, j, k ≤ n:
,
.
From the genetic viewpoint Lotka–Volterra algebras reproduce the forward evolution of populations where offspring necessarily reproduce the genetic type of one of the progenitors, assuming also panmixia, that is, considering that no progenitor has any specific weight on offspring type. As a result, LV-algebras are, by definition, commutative. Notice however that nonnegativity of the multiplication constants has been later not required [Citation3] as it happens to be the case for (nonassociative) algebras with genetic realization [Citation5]. This was not however the case in [Citation1], where the requirement on the skew-symmetric matrix entries ensured the genetic realization of Itoh's Lotka–Volterra algebras.
Lotka–Volterra algebras are a family of genetic algebras different from the recently studied evolution algebras [Citation6], although both families of algebras, Lotka–Volterra and evolution, are defined by square, not cubic, structure matrices. The simplest example of Lotka–Volterra algebra is the gametic algebra for simple Mendelian inheritance [Citation7, p.135], also called an elementary algebra by Heuch and Holgate, given by the zero matrix A = 0n×n
Our aim here is, retaining those properties characterizing the genetic population under consideration, i.e. offspring reproducing some progenitor type and panmixia, to consider Tian and Li's approach based on the search for new algebraic structures focused on describing the backward inheritance within genetic systems [Citation8]. As a result, Lotka–Voterra coalgebras are introduced, providing a new framework for the treatment of such populations.
Lotka–Volterra coalgebras constitute therefore a new family of noncoassociative coalgebras whose defining conditions reflect the backward dynamics of the transference of genetic inheritance within the considered populations. Even though here we are mainly concerned with the study of their algebraic structure, the connection between Lotka–Volterra coalgebras and the theory of dynamical systems will also become apparent.
After this introduction, the paper is organized as follows. Section two provides the definition of Lotka–Volterra coalgebras, together with examples, and also focuses on their matrix realization. In the third section, we cope with the duality of Lotka–Volterra structures. A well-known result states the correspondence, through dual vector spaces, between algebraic and coalgebraic structures defined on finite-dimensional vector spaces. Here we examine up to what extent the duality between Lotka–Volterra algebras and coalgebras is retained, to continue in Section four tackling with the algebraic description of Lotka–Volterra coalgebras.
Section five is devoted to a rather classical problem in algebraic genetics [Citation7, p.103], namely here, discussing which (coalgebra) constructions retain the property of being Lotka–Volterra. To finish the algebraic description of Lotka–Volterra coalgebras we achieve in this work, Section six focuses on their algebraic structure. Distinguishing generators of Lotka–Volterra coalgebras (i.e. their natural basis elements) into group-like and nongroup-like elements provides a semidirect sum decomposition (see [Citation6, p.45]) of Lotka–Volterra coalgebras into a subcoalgebra spanned by algebraic persistent elements and a coideal spanned by algebraic transient elements. This classification of the coalgebra natural basis elements brings into a linkage between Lotka–Volterra coalgebras and dynamical systems, to be exploited in a forthcoming work.
2. Lotka Volterrra coalgebras
Throughout the paper, unless otherwise stated, will be a field of characteristic not 2, and all objects will be defined over finite-dimensional
-vector spaces. We refer the reader to [Citation9] for basic notions on coalgebras.
A coalgebra is a
-vector space
endowed with a linear map (comultiplication)
. Given a basis
of the
-vector space
, we write
A subcoalgebra D of a coalgebra
is a subspace D of
such that Δ(D)⊆ D ⊗ D. A vector subspace I of
is a left (resp. right) coideal of
if
(resp.
) and a coideal if
. Subcoalgebras are one-sided coideals and, one-sided coideals are coideals. Intersections and sums of subcoalgebras (resp. left or right coideals) are again subcoalgebras (resp. left or right coideals). Sums of coideals are also still coideals, but this does not apply to intersections.
The comultiplication structure constants can be rearranged into a cubic (n × n × n) matrix , given by
. Cubic matrices are usually unfolded by their frontal slices as n × n2 rectangular matrices P = (P::1|P::2| · · · |P::n), where, for any k = 1, …, n, P::k is the n × n matrix with (i, j)-th entry pijk, i, j = 1, …, n. Thus the k-th frontal slice P::k of P contains the comultiplication constants arising from Δ(ek).
Definition 2.1
A coalgebra is Lotka–Volterra (for short, an LV-coalgebra) if it admits a basis
, satisfying for all i, j, k = 1, …, n:
;
;
;
where is a set of scalars such that
Remark 2.1
Any such a basis will be called a natural basis. All bases considered in the paper will be assumed to be natural bases. Because of LCV-1 and LCV-2 we have that
Moreover, the above relation and LCV-3 imply
or, equivalently,
Definition 2.2
A real (i.e. ) LV-coalgebra
, with natural basis
, has genetic realization if:
for all i, j, k = 1, …, n.
Example 2.3
Let G be a finite group of order n. Then the group coalgebra is an LV-coalgebra with natural basis
and
, for all i, j, k = 1, …, n.
Definition 2.4
An LV-coalgebra will be said to be trivial if
, for all i, j, k = 1, …, n., i.e. if
is a group coalgebra.
Example 2.5
Tian and Li [Citation8, p. 248] considered the case of a randomly mating population of diploid individuals differing in a locus alleles . The n-dimensional
-vector space
with basis
and comultiplication given by
, for all i, j, k = 1, …, n, is an LV-coalgebra. Following [Citation7, Example 1.1, p. 8], we refer to
to be the n-dimensional gametic coalgebra for simple Mendelian inheritance.
The class of LV-coalgebras is a family of (genetic) coalgebras not containing the zygotic coalgebra or the gametic coalgebra of autotetraploids [Citation10]. Moreover, an evolution coalgebra is Lotka–Volterra if and only if
is a trivial LV-coalgebra, that is, a group coalgebra [Citation11].
Displayed by its frontal slices, the cubic matrix associated to an LV-coalgebra has the following form:
which, by LVC-2, is (1,2)-symmetrical with equal accompanying matrices [Citation10, Lemma 2].
Proposition 2.6
Let be an LV-coalgebra. Then, with respect to a natural basis
, the accompanying matrices of
(i.e. those of its associated cubic matrix) are
Proof.
See [Citation10, 3.3].
Mimicking the algebra setting, LV-coalgebras are proposed to model backward dynamics in genetic systems when offspring necessarily reproduce the genetic type of one progenitor (LVC-1). Panmixia, i.e. considering that paternal or maternal type have no specific weight on offspring type, is also assumed, resulting in assumption LVC-2. Finally, for real LV-coalgebras, having genetic realization amounts to:
3. Duality
As remarked in the introduction, when posing the problem of introducing coalgebras as an alternative mathematical framework for modelling backward genetic inheritance, Tian and Li [Citation8] stressed on the fact that looking for new coalgebraic structures should imply more than just dualizing already existing genetic-like algebras. In this section, it is shown that, as previously defined, LV-coalgebras give rise to a new family of algebraic structures playing their own role in the mathematical study of population genetics.
The dual vector space of any coalgebra
is endowed with an algebra structure with multiplication induced by Δ*. Conversely, for any finite dimensional algebra
the isomorphism
allows us to define a coalgebra structure on
[Citation9].
Let be a n-dimensional
-vector space with basis
. We denote by
the dual basis of
in
, i.e. fi(ej) = δij, for all i, j = 1, …, n.
Theorem 3.1
Let be a coalgebra satisfying LVC-1 and LVC-2 (with respect to
). Then the induced multiplication in
is given by:
Proof.
Use Δ*(fr ⊗ fs)(ek) = (fr ⊗ fs)Δ(ek) to obtain:
for all r, s, k = 1, …, n.
Remark 3.1
Note the multiplication in the dual algebra is indeed defined by
, for all r, s = 1, …, n.
Corollary 3.2
Let be an LV-coalgebra with genetic realization. Then
is not an LV-algebra (with respect to
).
Proof.
Assume with the induced multiplication (see Theorem 3.1) is an LV-algebra. Then we have
for all k = 1, …, n, and LVC-3 and LVC-4 together imply that
. Hence frfs = fsfr = 0 if r ≠ s, contradicting then that
is LV-algebra.
We remark that, given an LV-coalgebra with genetic realization whose dual algebra
is also an LV-algebra Corollary 3.2 shows that
is then a group coalgebra, i.e. a trivial LV-coalgebra, and
a (nonzero) trivial evolution algebra.
Next given a coalgebra we provide necessary and sufficient conditions for its dual algebra
to be an LV-algebra.
Theorem 3.3
Let be a coalgebra with basis
. Then, there exists a skew-symmetric matrix
such that Δ(ek) = ek ⊗ Tk + Tk ⊗ ek, where
if and only if
is an LV-algebra with respect to the dual basis
.
Proof.
Let be a coalgebra with comultiplication Δ(ek) = ek ⊗ Tk + Tk ⊗ ek, with
defined by a skew-symmetric matrix
. We note this results into:
Then, by Remark 3.1,
, for all r, s = 1, …, n. Moreover,
is commutative and, since A is skew-symmetric,
is an LV-algebra.
Assume conversely is a coalgebra, whose dual algebra
with the inherited multiplication m = Δ* is a (commutative) LV-algebra, with respect to
, and write, for all r, s = 1, …, n:
for a skew-symmetric matrix
. We consider the obvious identifications
and (Δ*)* ≅ Δ resulting from the finite-dimensionality of
[Citation9, p.10]. Then Δ(ek)(fr ⊗ fs) = <ek, frfs > implies
, with
, for all k, r, s = 1, …, n and, as a result:
If
, then
,
If k = r ≠ s, then
,
If k = r = s, then
.
Moreover for all k, r, s = 1, …, n. Thus, taking into account that akk = 0,
Hence the comultiplication in
is therefore given by the skew-symmetric matrix A defining the elements Tk's.
We note that, as defined in Theorem 3.3, satisfies LVC-1 and LVC-2, but not LVC-3. Indeed we have
for all k = 1, …, n, that is, comultiplication of the basis elements is not normalized.
Example 3.4
Let be a 3-dimensional
-vector space with basis
. We define the following multiplication in
:
and extend it by linearity to
. Let us denote
the matrix associated to the multiplication m (as a linear map) in bases
of
and
of
. Writing
, we have
, for all i = 1, …, 9, and
Then
defines a coalgebra structure on
, with associated matrix (as a linear map) the transpose matrix MT of M. Moreover
satisfies LVC-1 and LVC-2 but not LVC-3, since columns of MT are not normalized. Hence
is not an LV-coalgebra. If, assuming then
, we column-normalize MT, rows of the resulting matrix
i.e. columns of
, define a new comultiplication
given by:
also satisfying LVC-3. Hence
is an LV-coalgebra.
4. Algebraic properties of Lotka–Volterra coalgebras
Cocommutativity of LV-coalgebras follows from LVC-2. Recall a coalgebra is cocommutative if τΔ = Δ, where
is the twist map given by τ(a ⊗ b) = b ⊗ a for all
. LV-coalgebras are however not necessarily coassociative.
Example 4.1
Let be a 2-dimensional coalgebra with comultiplication (with respect to a basis
):
Clearly
is an LV-coalgebra and (id ⊗ Δ)Δ(e2) ≠ (Δ ⊗ id)Δ(e2). Hence
is not coassociative.
Definition 4.2
A coalgebra is counital if it exists a linear map (counit)
such that
.
Proposition 4.3
Let be an LV-coalgebra. Then
is counital if and only if there exists a nonzero
such that, for all k = 1, …, n,
(1)
(1)
(2)
(2)
Proof.
Assume is a (nonzero) counit for
and write
, i = 1, …, n. Then, by LVC-2, we have:
Thus, ε is a counit of
if and only if, for all k = 1, …, n it holds that
or equivalently, by LVC-3,
Conversely, any such nonzero
defines a (nonzero) counit for
.
Example 4.4
Let
be a trivial LV-coalgebra, i.e. a group coalgebra. Since then we have
for all i, j, k = 1, …, n,
holds straightforwardly for all k, and
becomes
. Thus, by Proposition 4.3,
has a unique counit defined by
, for all k = 1, …, n.
The 2-dimensional LV-coalgebra defined in Example 4.1 admits no nonzero counit.
It follows from Example 4.4(ii) above that having natural basis elements that are group-like is not enough for ensuring LV-coalgebras to be counital. However, having a nonzero counit is a sufficient condition for the existence of group-like elements (i.e. elements such that Δ(x) = x ⊗ x) in the case of LV-coalgebras with genetic realization.
Theorem 4.5
Let be an LV-coalgebra with genetic realization and natural basis
. If
has a nontrivial counit ε, then
, where
is a (group) subcoalgebra of
. Moreover γk = 1 for all k ∈ Λ.
Proof.
Let ε be a nonzero counit for . Then
. Take k0 ∈ Λ. Then, by Proposition 4.3(2), we have
, which implies
and, therefore, by LVC-4,
, for all r, s = 1, …, n. Thus
and
is a group-like element, and, as a result,
is a subcoalgebra of
. The last assertion, γk = 1 for all k ∈ Λ, follows from Proposition 4.3(1).
Definition 4.6
A coalgebra is baric if there exists a nonzero linear map
such that (ϕ ⊗ ϕ)Δ = ϕ. The map ϕ is called character or weight function.
Any character ϕ is an element of and therefore can be uniquely written
, for some
.
Proposition 4.7
Let be an LV-coalgebra. A linear map
is a character of
if and only if
(3)
(3)
Proof.
Let . Then ϕ is a character of
if and only if, for all
Hence, ϕ is a character if and only if for all k = 1, …, n,
Corollary 4.8
The linear map is a character of any LV-coalgebra
.
Proof.
Let a = (α1, …, αn)T = 1. By Proposition 4.7, ϕ is a character if and only if
which follows straightforwardly for any LV-coalgebra
. (See also [Citation8, Theorem 4.3].)
Group-like elements of LV-coalgebras behave, with respect to weight maps (or characters), as do idempotent elements in genetic algebras [Citation7, Lemma 4.1, p.65]. Indeed from the biological viewpoint, the existence of group-like elements can be understood as equilibrium states for the underlying biological system.
Proposition 4.9
Let ϕ be a character of an LV-coalgebra . For any group-like element
, either ϕ(x) = 0 or ϕ(x) = 1.
Proof.
Let be group-like. Then ϕ(x) = (ϕ ⊗ ϕ)Δ(x) = (ϕ ⊗ ϕ)(x ⊗ x) = ϕ(x)2. Hence either ϕ(x) = 0 or ϕ(x) = 1.
Corollary 4.10
Characters of LV-coalgebras, if there exist, are not necessarily unique.
Proof.
Let be the LV-coalgebra introduced in Example 3.4. The cubic matrix P associated to
unfolded by its frontal slices is
and Proposition 4.7(3) gives that
is a (nonzero) character of
if and only if a = (α1, α2, α3)T is of the form:
Thus the LV-coalgebra
defined in Example 3.4 has two 1-parameter families of characters together to one additional nontrivial character.
5. Constructions of LV-coalgebras
Achieving the construction of general objects enclosing simultaneously different (genetic) features has been a recurrent topic in the literature of genetic algebraic structures. See, for instance, [Citation7, p.103] or [Citation8, Section 5]. Here we consider different constructions of coalgebras preserving the property of being Lotka–Volterra.
Following [Citation8, Subsection 5.1], given a n-dimensional -vector space V we denote by
the set of all coalgebra structures defined on V or, equivalently, the set of all linear maps Δ: V → V ⊗ V. The set
was shown to be a vector space in [Citation8, Proposition 5.1].
Let and
be an (ordered) basis of V. Given elements
, with
and
, we write
. Then
where for all i, j, k = 1, …, n, we have
.
Theorem 5.1
Convex linear combinations of real LV-coalgebras are Lotka–Volterra.
Proof.
Let be real LV-coalgebras and
be nonnegative and such that
. We claim that
, with
, is also Lotka–Volterra. Indeed LVC-1 and LVC-2 follow straightforwardly from being each
LV-coalgebra, and LVC-3 follows from the Cauchy–Schwarz inequality.
Corollary 5.2
Any convex linear combination of real LV-coalgebras with genetic realization has genetic realization.
Proof.
LVC-4 follows from the nonnegativity of .
Remark 5.1
The direct sum of any finite family of coalgebras
has a coalgebra structure resulting from
being isomorphic to
and, therefore, containing a copy of
[Citation9, p.50].
Let , with Vt being a nt-dimensional
-vector space with basis
. If
is a coalgebra, then Δ:V → V ⊗ V, given by Δ(xt) = Δt(xt), for all xt ∈ Vt, defines a coalgebra structure on V. We write
Theorem 5.3
Let be LV-coalgebras. Then, the direct sum
is an LV-coalgebra. If, moreover, each
is a real LV-coalgebra with genetic realization, so is
.
Proof.
Write, for k = 1, …, nt and t = 1, …, p,
On the other hand,
. Thus we have
Assume now
is an LV-coalgebra, for all t = 1, …, p. Then
clearly satisfies LVC-1 and LVC-2, whereas LVC-3 follows from the fact that Δ(xt) = Δt(xt), for all xt ∈ Vt. The last statement about the genetic realization (i.e.
satisfying LVC-4) is clear.
The family of LV-coalgebras is therefore closed under convex linear combinations (when ) and direct sums, preserving also both constructions the genetic realization (i.e. LVC-4). LV-coalgebras are not however closed under tensor products or the cocommutative duplication.
Example 5.4
Let be a coalgebra. It is a well-known result that the (
-vector space)
admits a comultiplication
, where τ denotes the twist map, endowing
with a coalgebra structure [Citation9, p.49].
Consider now the 2-dimensional coalgebra with basis
and comultiplication:
Clearly
is Lotka–Volterra with respect to
, but it is not difficult to check that
is not an LV-coalgebra (with respect to the basis
).
Finally let be the cocommutative duplication of the (cocommutative) coalgebra
[Citation8, Definition 3.1], that is,
, where
is the symmetric tensor product of the (cocommutative) coalgebra
. Recall
is a cocommutative coalgebra with comultiplication (using Sweedler notation [Citation9, p.32])
Then
is a 3-dimensional coalgebra with basis
and, it holds that
Hence
is not Lotka–Volterra (with respect to
).
6. Structure of LV-coalgebras
The different algebraic properties of the generators (i.e. natural basis elements) of algebraic objects with genetic significance usually reproduce their different behaviour within the underlying genetic system, when considered from the dynamical viewpoint. The classification of a natural basis elements into algebraically persistent and transient generators considered in [Citation6, Subsection 3.4.2] led Tian to consider semi-direct sum decompositions of evolution algebras. Here we recover the notion of semi-direct sum decomposition, and apply it to Lotka–Volterra coalgebras.
Let be an LV-coalgebra with natural basis
. We write [n] = {1, …, n} and
, for all k ∈ [n].
Proposition 6.1
Let k ∈ [n]. If Ik is a subcoalgebra of , then ek is a group-like element. Otherwise Ik is a coideal.
Proof.
Take k ∈ [n] and assume Ik is a subcoalgebra of . This implies
and, therefore, by LVC-3,
. Hence ek is group-like. Assume otherwise, ek is not group-like. Then
for some i ≠ k, and
Therefore
. Hence Ik is a coideal.
Let us write (disjoint union), where
and,
. Obviously
or
remains possible. Sufficient conditions for
are given in Theorem 4.5.
Lemma 6.2
D = ⊕k∈Λ Ik is a subcoalgebra of
.
is a subcoideal of
.
Proof.
It follows from [Citation9, p.18].
Theorem 6.3
Let be an LV-coalgebra. Then
, where
denotes the direct sum of vector subspaces.
Proof.
Clear.
Remark 6.1
Following [Citation6, Theorem 11] we will call such an LV-coalgebra decomposition to be a semidirect sum decomposition of . In such a decomposition (co)algebraic persistent elements can be identified to group-like elements, being (co)algebraic transient otherwise.
The dynamical behaviour of LV-coalgebras basis elements becomes apparent when considering their accompanying matrices. Write r = |Λ| and t = n−r. Then, by Theorem 6.3, the accompanying matrices of P have the following form (see Proposition 2.6):
Corollary 6.4
Reordering the natural basis elements, if necessary, so that, , being ek group-like, for k = 1, …, r:
Proof.
It suffices to note that for group-like elements, i.e. for all k = 1, …, r, it holds that , i, j = 1, …, n and, therefore,
. On the other hand, by Theorem 6.3, for all j = 1, …, t, there exists some i = 1, …, r, such that
.
6.1. Further comments
Under the additional assumption of being endowed with genetic realization (i.e. LVC-4), accompanying matrices of LV-coalgebras evidence the connection between these new family of coalgebras and dynamical systems. Indeed, then P(i) turns out to be not only nonnegative, by LVC-4, but also column stochastic as a result of LVC-3. This settles a new connection between LV-coalgebras and Markov processes, where the topology of P(i) classifying the matrix indices into essential and inessential classes translates into the (co)algebraic persistency and transiency of the natural basis elements. It is a forthcoming work to provide a detailed approach to the ergodic behaviour of LV-coalgebras from the viewpoint of Markov chains.
Acknowledgments
This paper was written while the third author was visiting the Department of Mathematics of the University of Chile, supported by FONDECYT 1170547, and wants to thank the members of the department there for their hospitality. The authors want to thank the referee for his/her comments.
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No potential conflict of interest was reported by the author(s).
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