Mini review

Microbial enzymes in the Mediterranean Sea: relationship with climate changes

  • Received: 10 June 2019 Accepted: 10 September 2019 Published: 12 September 2019
  • In most of the aquatic ecosystems, microorganisms are major players in the biogeochemical and nutrients cycles (Carbon Nitrogen, Phosphorus), through their enzymatic activities (leucine aminopeptidase, alkaline phosphatase and beta-glucosidase) on organic polymers such as polypeptides, organophosphate esters and polysaccharides, respectively. The small monomers released by decomposition are metabolised by microbes, supporting their growth. Most of the extracellular enzymes are adaptative and their synthesis and activity is strongly affected by environmental factors, consequently the relative importance of leucine aminopeptidase, alkaline phosphatase and beta-glucosidase reflects differences in the composition of organic matter and assume a different meaning.
    Since more than two decades, at the CNR the influence of climate changes, seasonal variability, depth and coastal input on the patterns of enzymatic activities in the Mediterranean Sea have been studied. Its particular characteristics of a semi-closed basin, high summer evaporation and the occurrence of important water dynamics, make this ecosystem particularly suitable as a model site for climate changes-related observations.
    The present paper reviews the current information of environmental changes on extracellular enzymatic activity obtained in the Mediterranean areas with the aim of evaluating the effects of environmental changes on the microbial activities. The obtained results revealed significant variations in the rates of hydrolytic activities in relation to space and time, with the highest levels generally found in the epipelagic layer (0–100m) and in coastal zones during warm periods. In the Central Mediterranean Sea their relationship with temperature changes was demonstrated.
    Spatial variations in the relative enzyme activities also suggested a modulation in the metabolic profiles of the prokaryotic communities, with biogeochemical implications in nutrient regeneration.
    Long term studies on microbial activity and abundances in relation with rising temperatures can have a predictive value to describe the evolutionary scenario of microbial processes and the response of microbial metabolism to climate changes in the Mediterranean Sea.

    Citation: Renata Zaccone, Gabriella Caruso. Microbial enzymes in the Mediterranean Sea: relationship with climate changes[J]. AIMS Microbiology, 2019, 5(3): 251-271. doi: 10.3934/microbiol.2019.3.251

    Related Papers:

    [1] Sung Woo Choi . Explicit characteristic equations for integral operators arising from well-posed boundary value problems of finite beam deflection on elastic foundation. AIMS Mathematics, 2021, 6(10): 10652-10678. doi: 10.3934/math.2021619
    [2] Moh. Alakhrass . A note on positive partial transpose blocks. AIMS Mathematics, 2023, 8(10): 23747-23755. doi: 10.3934/math.20231208
    [3] Xinfeng Liang, Mengya Zhang . Triangular algebras with nonlinear higher Lie n-derivation by local actions. AIMS Mathematics, 2024, 9(2): 2549-2583. doi: 10.3934/math.2024126
    [4] Cui-Xia Li, Long-Quan Yong . Modified BAS iteration method for absolute value equation. AIMS Mathematics, 2022, 7(1): 606-616. doi: 10.3934/math.2022038
    [5] Sara Smail, Chafika Belabbaci . A characterization of Wolf and Schechter essential pseudospectra. AIMS Mathematics, 2024, 9(7): 17146-17153. doi: 10.3934/math.2024832
    [6] Yuna Zhao . Construction of blocked designs with multi block variables. AIMS Mathematics, 2021, 6(6): 6293-6308. doi: 10.3934/math.2021369
    [7] Wen-Ning Sun, Mei Qin . On maximum residual block Kaczmarz method for solving large consistent linear systems. AIMS Mathematics, 2024, 9(12): 33843-33860. doi: 10.3934/math.20241614
    [8] Shakir Ali, Amal S. Alali, Atif Ahmad Khan, Indah Emilia Wijayanti, Kok Bin Wong . XOR count and block circulant MDS matrices over finite commutative rings. AIMS Mathematics, 2024, 9(11): 30529-30547. doi: 10.3934/math.20241474
    [9] James Daniel, Kayode Ayinde, Adewale F. Lukman, Olayan Albalawi, Jeza Allohibi, Abdulmajeed Atiah Alharbi . Optimised block bootstrap: an efficient variant of circular block bootstrap method with application to South African economic time series data. AIMS Mathematics, 2024, 9(11): 30781-30815. doi: 10.3934/math.20241487
    [10] Ziqiang Wang, Qin Liu, Junying Cao . A higher-order numerical scheme for system of two-dimensional nonlinear fractional Volterra integral equations with uniform accuracy. AIMS Mathematics, 2023, 8(6): 13096-13122. doi: 10.3934/math.2023661
  • In most of the aquatic ecosystems, microorganisms are major players in the biogeochemical and nutrients cycles (Carbon Nitrogen, Phosphorus), through their enzymatic activities (leucine aminopeptidase, alkaline phosphatase and beta-glucosidase) on organic polymers such as polypeptides, organophosphate esters and polysaccharides, respectively. The small monomers released by decomposition are metabolised by microbes, supporting their growth. Most of the extracellular enzymes are adaptative and their synthesis and activity is strongly affected by environmental factors, consequently the relative importance of leucine aminopeptidase, alkaline phosphatase and beta-glucosidase reflects differences in the composition of organic matter and assume a different meaning.
    Since more than two decades, at the CNR the influence of climate changes, seasonal variability, depth and coastal input on the patterns of enzymatic activities in the Mediterranean Sea have been studied. Its particular characteristics of a semi-closed basin, high summer evaporation and the occurrence of important water dynamics, make this ecosystem particularly suitable as a model site for climate changes-related observations.
    The present paper reviews the current information of environmental changes on extracellular enzymatic activity obtained in the Mediterranean areas with the aim of evaluating the effects of environmental changes on the microbial activities. The obtained results revealed significant variations in the rates of hydrolytic activities in relation to space and time, with the highest levels generally found in the epipelagic layer (0–100m) and in coastal zones during warm periods. In the Central Mediterranean Sea their relationship with temperature changes was demonstrated.
    Spatial variations in the relative enzyme activities also suggested a modulation in the metabolic profiles of the prokaryotic communities, with biogeochemical implications in nutrient regeneration.
    Long term studies on microbial activity and abundances in relation with rising temperatures can have a predictive value to describe the evolutionary scenario of microbial processes and the response of microbial metabolism to climate changes in the Mediterranean Sea.


    A problem that occurs frequently in a variety of mathematical contexts, is to find the common invariant subspaces of a single matrix or set of matrices. In the case of a single endomorphism or matrix, it is relatively easy to find all the invariant subspaces by using the Jordan normal form. Also, some theoretical results are given only for the invariant subspaces of two matrices. However, when there are more than two matrices, the problem becomes much harder, and unexpected invariant subspaces may occur. No systematic method is known. In a recent article [1], we have provided a new algorithms to determine common invariant subspaces of a single matrix or of a set of matrices systematically.

    In the present article we consider a more general version of this problem, that is, providing two algorithms for simultaneous block triangularization and block diagonalization of sets of matrices. One of the main steps in the first two proposed algorithms, consists of finding the common invariant subspaces of matrices using the new method proposed in the recent article [1]. It is worth mentioning that an efficient algorithm to explicitly compute a transfer matrix which realizes the simultaneous block diagonalization of unitary matrices whose decomposition in irreducible blocks (common invariant subspaces) is known from elsewhere is given in [2]. An application of simultaneous block-diagonalization of normal matrices in quantum theory is presented in [3].

    In this article we shall be concerned with finite dimensions only. Of course the fact that a single complex matrix can always be put into triangular form follows readily from the Jordan normal form theorem [4]. For a set of matrices, Jacobson in [5] introduced the notion of a composition series for a collection of matrices. The idea of a composition series for a group is quite familiar. The Jordan-Hölder Theorem [4] states that any two composition series of the same group have the same length and the same composition factors (up to permutation). Jacobson in [5] characterized the simultaneous block triangularization of a set of matrices by the existence of a chain $ \{0\} = V_0 \subset V_1 \subset... \subset V_t = \mathbb{C}^n $ of invariant subspaces with dimension $ dim(V_i/V_{i-1}) = n_i $. Therefore, in the context of a collection of matrices $ \Omega = \{A_i \}_{ i = 1}^N $, the idea is to locate a common invariant subspace $ V $ of minimal dimension $ d $ of a set of matrices $ \Omega $. Assume $ V $ is generated by the (linearly independent) set $ \mathcal{B}_1 = \{u_1, u_2, ..., u_d \} $, and let $ \mathcal{B} = \{u_1, u_2, ..., u_d, u_{d+1}, u_{d+2}, ..., u_n\} $ be a basis of $ \mathbb{C}^n $ containing $ \mathcal{B}_1 $. Upon setting $ S = (u_1, u_2, ..., u_d, u_{d+1}, u_{d+2}, ..., u_n) $, $ S^{-1}A_i S $ has the block triangular form

    $ S1AiS=(Bi1,1Bi1,20Bi2,2),
    $

    for $ i = 1, ..., n $. Thereafter, one may define a quotient of the ambient vector space, and each of the matrices in the given collection will pass to this quotient. As such, one defines

    $ Ti=Bi2,2=(0(nd)×dInd)S1AiS(0d×(nd)Ind).
    $

    Then one may begin again the process of looking for a common invariant subspace of minimal dimension of a set of matrices $ \{T_i \}_{ i = 1}^N $ and iterate the procedure. Since all spaces and matrices are of finite dimension, the procedure must terminate at some point. Again, any two such composition series will be isomorphic. When the various quotients and submatrices are lifted back to the original vector space, one obtains precisely the block-triangular form for the original set of matrices. It is important to find a composition series in the construction in order to make the set of matrices as "block-triangular as possible."

    Dubi [6] gave an algorithmic approach to simultaneous triangularization of a set of matrices based on the idea of Jacobson in [5]. In the case of simultaneous triangularization, it can be understood as the existence of a chain $ \{0\} = V_0 \subset V_1 \subset... \subset V_t = \mathbb{C}^n $ of invariant subspaces with dimension $ dim(V_i) = i $. We generalize his study to cover simultaneous block triangularization of a set of matrices. The generalized algorithm depends on the novel algorithm for constructing invariant subspaces of a set of matrices given in the recent article [1].

    Specht [7] (see also [8]) proved that if the associative algebra $ \mathcal{L} $ generated by a set of matrices $ \Omega $ over $ \mathbb{C} $ satisfies $ \mathcal{L} = \mathcal{L}^{*} $, then $ \Omega $ admits simultaneous block triangularization if and only if it admits simultaneous block diagonalization, in both cases via a unitary matrix. Following a result of Specht, we prove that a set of matrices $ \Omega $ admits simultaneous block diagonalization if and only if the set $ \Gamma = \Omega \cup \Omega^{*} $ admits simultaneous block triangularization. Finally, an algorithmic approach to simultaneous block diagonalization of a set of matrices based on this fact is proposed.

    The latter part of this paper presents an alternate approach for simultaneous block diagonalization of a set of $ n \times n $ matrices $ \{A_s\}_{ s = 1}^N $ by an invertible matrix that does not require finding the common invariant subspaces. Maehara et al. [9] introduced an algorithm for simultaneous block diagonalization of a set of matrices by a unitary matrix based on the existence of a Hermitian commuting matrix. Here, we extend their algorithm to simultaneous block diagonalization of a set of matrices by an invertible matrix based on the existence of a commuting matrix which is not necessarily Hermitian. For example, consider the set of matrices $ \Omega = \{A_i \}_{i = 1}^2 $ where

    $ A1=(100220111),A2=(000210010).
    $
    (1.1)

    The only Hermitian matrix commuting with the set $ \Omega $ is the identity matrix. Therefore, we cannot apply the proposed algorithm given in [9]. However, one can verify that the following non Hermitian matrix $ C $ commutes with all the matrices $ \{A_i \}_{ i = 1}^2 $

    $ C=(000210010).
    $
    (1.2)

    The matrix $ C $ has distinct eigenvalues $ \lambda_1 = 0, \lambda_2 = 1 $ with algebraic multiplicities $ n_1 = 2, n_2 = 1 $, respectively. Moreover, the matrix $ C $ is not diagonalizable. Therefore, we cannot construct the eigenvalue decomposition for the matrix $ C $. However, one can decompose the matrix $ C $ by its generalized eigen vectors as follows:

    $ S1CS=(010000001)=(0100)(1),
    $
    (1.3)

    where

    $ S=(0120011101).
    $
    (1.4)

    Initially, it is noted that the matrices $ \{A_i \}_{ i = 1}^2 $ can be decomposed into two diagonal blocks by the constructed invertible matrix $ S $ where

    $ S1A1S=(11201)(2),S1A2S=(0100)(1).
    $
    (1.5)

    Then, a new algorithm is developed for simultaneous block diagonalization by an invertible matrix based on the generalized eigenvectors of a commuting matrix. Moreover, a new characterization is presented by proving that the existence of a commuting matrix that possesses at least two distinct eigenvalues is the necessary and sufficient condition to guarantee the simultaneous block diagonalization by an invertible matrix.

    An outline of the paper is as follows. In Section 2 we review several definitions pertaining to block-triangular and block-diagonal matrices and state several elementary consequences that follow from them. In Section 3, following a result of Specht [7] (see also [8]), we provide conditions for putting a set of matrices into block-diagonal form simultaneously. Furthermore, we apply the theoretical results to provide two algorithms that enable a collection of matrices to be put into block-triangular form or block-diagonal form simultaneously by a unitary matrix based on the existence of invariant subspaces. In Section 4, a new characterization is presented by proving that the existence of a commuting matrix that possesses at least two distinct eigenvalues is the necessary and sufficient condition to guarantee the simultaneous block diagonalization by an invertible matrix. Furthermore, we apply the theoretical results to provide an algorithm that enables a collection of matrices to be put into block-diagonal form simultaneously by an invertible matrix based on the existence of a commuting matrix. Sections 3 and 4 also provide concrete examples using the symbolic manipulation system Maple.

    Let $ \Omega $ be a set of $ n \times n $ matrices over an algebraically closed field $ \mathcal{F} $, and let $ \mathcal{L} $ denote the algebra generated by $ \Omega $ over $ \mathcal{F} $. Similarly, let $ \Omega^{*} $ be the set of the conjugate transpose of each matrix in $ \Omega $ and $ \mathcal{L}^{*} $ denote the algebra generated by $ \Omega^{*} $ over $ \mathcal{F} $.

    Definition 2.1. An $ n \times n $ matrix $ A $ is given the notation $ BT(n_1, ..., n_t) $ provided $ A $ is block upper triangular with $ t $ square blocks on the diagonal, of sizes $ n_1, ..., n_t $, where $ t \geq 2 $ and $ n_1+... +n_t = n $. That is, a block upper triangular matrix $ A $ has the form

    $ A=(A1,1A1,2A1,t0A2,2A2,t00At,t)
    $
    (2.1)

    where $ {\bf{A}}_{i, j} $ is a square matrix for all $ i = 1, ..., t $ and $ j = i, ..., t $.

    Definition 2.2. A set of $ n \times n $ matrices $ \Omega $ is $ BT(n_1, ..., n_t) $ if all of the matrices in $ \Omega $ are $ BT(n_1, ..., n_t) $.

    Remark 2.3. A set of $ n \times n $ matrices $ \Omega $ admits a simultaneous triangularization if it is $ BT(n_1, ..., n_t) $ with $ n_i = 1 $ for $ i = 1, ..., t $.

    Remark 2.4. A set of $ n \times n $ matrices $ \Omega $ is $ BT(n_1, ..., n_t) $ if and only if the algebra $ \mathcal{L} $ generated by $ \Omega $ is $ BT(n_1, ..., n_t) $.

    Proposition 2.5. [7] (see also [8]) Let $ \Omega $ be a nonempty set of complex $ n \times n $ matrices. Then, there is a nonsingular matrix $ S $ such that $ S \Omega S^{-1} $ is $ BT(n_1, ..., n_t) $ if and only if there is a unitary matrix $ U $ such that $ U \Omega U^{*} $ is $ BT(n_1, ..., n_t) $.

    Theorem 2.6. [5,Chapter Ⅳ] Let $ \Omega $ be a nonempty set of complex $ n \times n $ matrices. Then, there is a unitary matrix $ U $ such that $ U \Omega U^{*} $ is $ BT(n_1, ..., n_t) $ if and only if the set $ \Omega $ has a chain $ \{0\} = V_0 \subset V_1 \subset... \subset V_t = \mathbb{C}^n $ of invariant subspaces with dimension $ dim(V_i/V_{i-1}) = n_i $.

    Definition 2.7. An $ n \times n $ matrix $ A $ is given the notation $ BD(n_1, ..., n_t) $ provided $ A $ is block diagonal with $ t $ square blocks on the diagonal, of sizes $ n_1, ..., n_t $, where $ t \geq 2 $, $ n_1+... +n_t = n $, and the blocks off the diagonal are the zero matrices. That is, a block diagonal matrix $ A $ has the form

    $ A=(A1000A2000At)
    $
    (2.2)

    where $ {\bf{A}}_k $ is a square matrix for all $ k = 1, ..., t $. In other words, matrix $ {\bf{A}} $ is the direct sum of $ {\bf{A}}_1, ..., {\bf{A}}_t $. It can also be indicated as $ {\bf{A}}_{\text{1}} \oplus {\bf{A}}_{\text{2}} \oplus... \oplus {\bf{A}}_{\text{t}} $.

    Definition 2.8. A set of $ n \times n $ matrices $ \Omega $ is $ BD(n_1, ..., n_t) $ if all of the matrices in $ \Omega $ are $ BD(n_1, ..., n_t) $.

    Remark 2.9. A set of $ n \times n $ matrices $ \Omega $ admits a simultaneous diagonalization if it is $ BD(n_1, ..., n_t) $ with $ n_i = 1 $ for $ i = 1, ..., t $.

    Remark 2.10. A set of $ n \times n $ matrices $ \Omega $ is $ BD(n_1, ..., n_t) $ if and only if the algebra $ \mathcal{L} $ generated by $ \Omega $ is $ BD(n_1, ..., n_t) $.

    Proposition 2.11. [7] (see also [8]) Let $ \Omega $ be a nonempty set of complex $ n \times n $ matrices and let $ \mathcal{L} $ be the algebra generated by $ \Omega $ over $ \mathbb{C} $. Suppose $ \mathcal{L} = \mathcal{L}^{*} $. Then, there is a nonsingular matrix $ S $ such that $ S \mathcal{L} S^{-1} $ is $ BT(n_1, ..., n_t) $ if and only if there is a unitary matrix $ U $ such that $ U \mathcal{L} U^{*} $ is $ BD(n_1, ..., n_t) $.

    Dubi [6] gave an algorithmic approach to simultaneous triangularization of a set of $ n \times n $ matrices. In this section, we will generalize his study to cover simultaneous block triangularization and simultaneous block diagonalization of a set of $ n \times n $ matrices. The generalized algorithms depend on the novel algorithm for constructing invariant subspaces of a set of matrices given in the recent article [1] and Theorem 3.3.

    Lemma 3.1. Let $ \Omega $ be a nonempty set of complex $ n \times n $ matrices, $ \Omega^{*} $ be the set of the conjugate transpose of each matrix in $ \Omega $ and $ \mathcal{L} $ be the algebra generated by $ \Gamma = \Omega \cup \Omega^{*} $. Then, $ \mathcal{L} = \mathcal{L}^{*} $.

    Proof. Let $ A $ be a matrix in $ \mathcal{L} $. Then, $ A = P(B_1, ..., B_m) $ for some multivariate noncommutative polynomial $ P(x_1, ..., x_m) $ and matrices $ \{B_i\}_{i = 1}^m\in \Gamma $. Therefore, $ A^{*} = P^*(B_1, ..., B_m) = Q(B_1^*, ..., B_m^*) $ for some multivariate noncommutative polynomial $ Q(x_1, ..., x_m) $ where the matrices $ \{B_i^*\}_{i = 1}^m\in \Gamma^* = \Gamma $. Hence, the matrix $ A^* \in \mathcal{L} $

    Lemma 3.2. Let $ \Omega $ be a nonempty set of complex $ n \times n $ matrices and $ \Omega^{*} $ be the set of the conjugate transpose of each matrix in $ \Omega $, and $ \Gamma = \Omega \cup \Omega^{*} $. Then, there is a unitary matrix $ U $ such that $ U \Gamma U^{*} $ is $ BD(n_1, ..., n_t) $ if and only if there is a unitary matrix $ U $ such that $ U \Omega U^{*} $ is $ BD(n_1, ..., n_t) $.

    Proof. Assume that there exists a unitary matrix $ U $ such that $ U \Omega U^{*} $ is $ BD(n_1, ..., n_t) $. Then, $ (U \Omega U^{*})^{*} = U \Omega^{*} U^{*} $ is $ BD(n_1, ..., n_t) $. Hence, $ U \Gamma U^{*} $ is $ BD(n_1, ..., n_t) $.

    Theorem 3.3. Let $ \Omega $ be a nonempty set of complex $ n \times n $ matrices and $ \Omega^{*} $ be the set of the conjugate transpose of each matrix in $ \Omega $, and $ \Gamma = \Omega \cup \Omega^{*} $. Then, there is a unitary matrix $ U $ such that $ U \Omega U^{*} $ is $ BD(n_1, ..., n_t) $ if and only if there is a unitary matrix $ U $ such that $ U \Gamma U^{*} $ is $ BT(n_1, ..., n_t) $.

    Proof. Let $ \mathcal{L} $ be the algebra generated by $ \Gamma $. Then, $ \mathcal{L} = \mathcal{L}^{*} $ using Lemma 3.1. Now, by applying Proposition 2.11 and Lemma 3.2, the following statements are equivalent :

    There is a unitary matrix $ U $ such that $ U \Gamma U^{*} $ is $ BT(n_1, ..., n_t) $.

    $ \iff $ There is a unitary matrix $ U $ such that $ U \mathcal{L} U^{*} $ is $ BT(n_1, ..., n_t) $.

    $ \iff $ There is a unitary matrix $ U $ such that $ U \mathcal{L} U^{*} $ is $ BD(n_1, ..., n_t) $.

    $ \iff $ There is a unitary matrix $ U $ such that $ U \Gamma U^{*} $ is $ BD(n_1, ..., n_t) $.

    $ \iff $ There is a unitary matrix $ U $ such that $ U \Omega U^{*} $ is $ BD(n_1, ..., n_t) $.

    (1) Input: the set $ \Omega = \{A_i \}_{ i = 1}^N $.

    (2) Set $ k = 0, \mathcal{B} = \phi, s = n, T_i = A_i, S_2 = I $.

    (3) Search for a $ d $-dimensional invariant subspace $ V = \langle v_1, v_2, ..., v_d \rangle $ of a set of matrices $ \{T_i \}_{ i = 1}^N $ starting from $ d = 1 $ up to $ d = s-1 $. If one does not exist and $ k = 0 $, abort and print "no simultaneous block triangularization". Else, if one does not exist and $ k\ne 0 $, go to step (8). Else, go to next step.

    (4) Set $ V_{k+1} = (S_2 v_1\; S_2 v_2\; ...\; S_2 v_d), \mathcal{B} = \mathcal{B} \cup \{S_2 v_1, S_2 v_2, ..., S_2 v_d\}, S_1 = (V_1\; V_2\; ...\; V_{k+1}) $.

    (5) Find a basis $ \{u_1, u_2, ..., u_l \} $ for the orthogonal complement of $ \mathcal{B} $.

    (6) Set $ S_2 = (u_1\; u_2\; ...\; u_l), S = (S_1\; S_2) $, and

    $ T_i = \left({0(sd)×dIsd

    } \right)S^{-1}A_i S \left({0d×(sd)Isd
    } \right) $.

    (7) Set $ k = k+1, s = s-d $, and return to step (3).

    (8) Compute the QR decomposition of the invertible matrix $ S $, by means of the Gram–Schmidt process, to convert it to a unitary matrix $ Q $.

    (9) Output: a unitary matrix $ U $ as the conjugate transpose of the resulting matrix $ Q $.

    Remark 3.4. If one uses any non-orthogonal complement in step 5 of Algorithm $ A $, then the matrix $ S $ is invertible such that $ S^{-1} \Omega S $ is $ BT(n_1, ..., n_t) $. However, in such a case, one cannot guarantee that $ U \Omega U^{*} $ is $ BT(n_1, ..., n_t) $.

    Example 3.5. The set of matrices $ \Omega = \{A_i \}_{i = 1}^2 $ admits simultaneous block triangularization where

    $ A1=(321011050000014012131113020025010006),A2=(441244840360001012320444168524404102880400040).
    $
    (3.1)

    Applying Algorithm $ A $ to the set $ \Omega $ can be summarized as follows:

    Input: $ \Omega $.

    Initiation step:

    We have $ k = 0, \mathcal{B} = \phi, s = 6, T_1 = A_1, T_2 = A_2, S_2 = I $.

    In the first iteration:

    We found two-dimensional invariant subspace $ V = \langle e_1, e_4 \rangle $ of a set of matrices $ \{T_i \}_{ i = 1}^2 $. Therefore, $ \mathcal{B} = \{e_1, e_4\}, S_1 = (e_1, e_4), S_2 = (e_2, e_3, e_5, e_6) $,

    $ T1=(5000141220251006),T2=(360011232444128840040),
    $
    (3.2)

    $ k = 1 $, and $ s = 4 $.

    In the second iteration: We found two-dimensional invariant subspace $ V = \langle e_2, e_3 \rangle $ of a set of matrices $ \{T_i \}_{ i = 1}^2 $. Therefore, $ \mathcal{B} = \{e_1, e_4, e_3, e_5\}, S_1 = (e_1, e_4, e_3, e_5), S_2 = (e_2, e_6) $,

    $ T1=(5016),T2=(361440),
    $
    (3.3)

    $ k = 2 $, and $ s = 2 $.

    In the third iteration: There is no one-dimensional invariant subspace of a set of matrices $ \{T_i \}_{ i = 1}^2 $. Therefore, $ S = (e_1\; e_4\; e_3\; e_5\; e_2\; e_6) $, and the corresponding unitary matrix is

    $ U = \left(100000000100001000000010010000000001
    \right) $

    such that the set $ U \Omega U^{*} = \{U A_i U^{*}\}_{i = 1}^2 $ is $ BT(2, 2, 2) $ where

    $ UA1U=(301121111133004112000225000050000016),UA2U=(444481244528416400324124001284800003610000440).
    $
    (3.4)

    (1) Input: the set $ \Omega = \{A_i \}_{ i = 1}^N $.

    (2) Construct the set $ \Gamma = \Omega \cup \Omega^{*} $.

    (3) Find a unitary matrix $ U $ such that $ U \Gamma U^{*} $ is $ BT(n_1, ..., n_t) $ using Algorithm $ A $.

    (4) Output: a unitary matrix $ U $.

    Remark 3.6. Algorithm $ B $ provides the finest block-diagonalization. Moreover, the number of the blocks equals the number the of the invariant subspaces, and the size of each block is $ n_i \times n_i $, where $ n_i $ is the dimension of the invariant subspace.

    Example 3.7. The set of matrices $ \Omega = \{A_i \}_{i = 1}^2 $ admits simultaneous block diagonalization where

    $ A1=(3000000020000000200000001000000010000000100000003),A2=(0000000000000001000000000000000000000010001000000).
    $
    (3.5)

    Applying Algorithm $ B $ to the set $ \Omega $ can be summarized as follows:

    Input: $ \Gamma = \Omega \cup \Omega^{*} $.

    Initiation step:

    We have $ k = 0, \mathcal{B} = \phi, s = 7, T_1 = A_1, T_2 = A_2, T_3 = A_2^T, S_2 = I $.

    In the first iteration:

    We found one-dimensional invariant subspace $ V = \langle e_5 \rangle $ of a set of matrices $ \{T_i \}_{ i = 1}^3 $. Therefore, $ \mathcal{B} = \{e_5\}, S_1 = (e_5), S_2 = (e_1, e_2, e_3, e_4, e_6, e_7) $,

    $ T1=(300000020000002000000100000010000003),T2=(000000000000010000000000000100100000),T3=TT2,
    $
    (3.6)

    $ k = 1 $, and $ s = 6 $.

    In the second iteration: We found two-dimensional invariant subspace $ V = \langle e_4, e_5 \rangle $ of a set of matrices $ \{T_i \}_{ i = 1}^3 $. Therefore, $ \mathcal{B} = \{e_5, e_4, e_6\}, S_1 = (e_5\; e_4\; e_6), S_2 = (e_1, e_2, e_3, e_7) $,

    $ T1=(3000020000200003),T2=(0000000001001000),T3=TT2,
    $
    (3.7)

    $ k = 2 $, and $ s = 4 $.

    In the third iteration: We found two-dimensional invariant subspace $ V = \langle e_2, e_3 \rangle $ of a set of matrices $ \{T_i \}_{ i = 1}^3 $. Therefore, $ \mathcal{B} = \{e_5, e_4, e_6, e_2, e_3\}, S_1 = (e_5\; e_4\; e_6\; e_2\; e_3), S_2 = (e_1, e_7) $,

    $ T1=(3003),T2=(0010),T3=(0100),
    $
    (3.8)

    $ k = 3 $, and $ s = 2 $.

    In the fourth iteration: There is no one-dimensional invariant subspace of a set of matrices $ \{T_i \}_{ i = 1}^3 $. Therefore, $ S = (e_5\; e_4\; e_6\; e_2\; e_3\; e_1\; e_7) $, and the corresponding unitary matrix is

    $ U = \left( 0000100000100000000100100000001000010000000000001
    \right) $

    such that the set $ U \Omega U^{*} = \{U A_i U^{*}\}_{i = 1}^2 $ is $ BD(1, 2, 2, 2) $ where

    $ UA1U=(1)(1001)(2002)(3003),UA2U=(0)(0010)(0010)(0010).
    $
    (3.9)

    Example 3.8. The set of matrices $ \Omega = \{A_i \}_{i = 1}^2 $ admits simultaneous block diagonalization where

    $ A1=(3000000020000000200000001000000010000000100000003),A2=(0000000000100001000000000000000010000001001000000).
    $
    (3.10)

    Similarly, applying Algorithm $ B $ to the set $ \Omega $ provides the matrix $ S = (e_6\; e_5\; e_7\; e_1\; e_3\; e_2\; e_4) $. Therefore, the corresponding unitary matrix is

    $ U = \left( 0000010000010000000011000000001000001000000001000
    \right) $

    such that the set $ U \Omega U^{*} = \{U A_i U^{*}\}_{i = 1}^2 $ is $ BD(2, 2, 3) $ where

    $ UA1U=(1001)(3003)(200020001),UA2U=(0101)(0100)(010001000).
    $
    (3.11)

    Example 3.9. The set of matrices $ \Omega = \{A_i \}_{i = 1}^3 $ admits simultaneous block diagonalization where

    $ A1=(000000000020000000001000000000200000000000000000001000000000100000000010000000000),A2=(000100000100010000000001000000000000000100000000000000000000000000000100000000000),A3=(010000000000000000000000000100010000010000000001000000000000010000000000000000000).
    $
    (3.12)

    Similarly, applying Algorithm $ B $ to the set $ \Omega $ provides the matrix $ S = (e_1+e_5\; e_9\; e_3\; e_6\; e_8\; -e_7\; e_1-e_5, e_2\; e_4) $. Therefore, the corresponding unitary matrix is

    $ U = \left( 12200012200000000000010010000000000010000000000100000001001220001220000010000000000100000
    \right) $

    such that the set $ U \Omega U^{*} = \{U A_i U^{*}\}_{i = 1}^3 $ is $ BD(1, 1, 2, 2, 3) $ where

    $ UA1U=(0)(0)(1001)(1001)(000020002),UA2U=(0)(0)(0100)(0100)(002200000),UA3U=(0)(0)(0010)(0010)(020000200).
    $
    (3.13)

    This section focuses on an alternate approach for simultaneous block diagonalization of a set of $ n \times n $ matrices $ \{A_s\}_{ s = 1}^N $ by an invertible matrix that does not require finding the common invariant subspaces as Algorithm $ B $ given in the previous section. Maehara et al. [9] introduced an algorithm for simultaneous block diagonalization of a set of matrices by a unitary matrix based on the eigenvalue decomposition of a Hermitian commuting matrix. Here, we extend their algorithm to be applicable for a non-Hermitian commuting matrix by considering its generalized eigen vectors. Moreover, a new characterization is presented by proving that the existence of a commuting matrix that possesses at least two distinct eigenvalues is the necessary and sufficient condition to guarantee the simultaneous block diagonalization by an invertible matrix.

    Proposition 4.1. Let $ V $ be a vector space, and let $ T:V \rightarrow V $ be a linear operator. Let $ \lambda_1, ..., \lambda_k $ be distinct eigenvalues of $ T $. Then, each generalized eigenspace $ G_{\lambda_i}(T) $ is $ T $-invariant, and we have the direct sum decomposition

    $ V = G_{\lambda_1}(T)\oplus G_{\lambda_2}(T) \oplus ...\oplus G_{\lambda_k}(T). $

    Lemma 4.2. Let $ V $ be a vector space, and let $ T:V \rightarrow V $, $ L:V \rightarrow V $ be linear commuting operators. Let $ \lambda_1, ..., \lambda_k $ be distinct eigenvalues of $ T $. Then, each generalized eigenspace $ G_{\lambda_i}(T) $ is $ L $-invariant.

    Proof. Let $ V $ be a vector space and $ \lambda_1, ..., \lambda_k $ be distinct eigenvalues of $ T $ with the minimal polynomial $ \mu(x) = (x-\lambda_1)^{n_1}(x-\lambda_2)^{n_2}...(x-\lambda_k)^{n_k} $. Then, we have the direct sum decomposition $ V = G_{\lambda_1}(T)\oplus G_{\lambda_2}(T) \oplus...\oplus G_{\lambda_k}(T) $.

    For each $ i = 1, .., k $, let $ x \in G_{\lambda_i}(T) $, and then $ (T-\lambda_i I)^{n_i}x = 0 $. Then, $ (T-\lambda_i I)^{n_i}Lx = L(T-\lambda_i I)^{n_i}x = 0 $. Hence, $ L x \in G_{\lambda_i}(T) $.

    Theorem 4.3. Let $ \{A_s\}_{ s = 1}^N $ be a set of $ n \times n $ matrices. Then, the set $ \{A_s\}_{ s = 1}^N $ admits simultaneous block diagonalization by an invertible matrix $ S $ if and only if the set $ \{A_s\}_{ s = 1}^N $ commutes with a matrix $ C $ that possesses two distinct eigenvalues.

    Proof. $\Rightarrow $ Assume that the set $ \{A_s\}_{ s = 1}^N $ admits simultaneous block diagonalization by the an invertible matrix $ S $ such that

    $ S^{-1} A_s S = B_{s,1} \oplus B_{s,2} \oplus ... \oplus B_{s,k}, $

    where the number of blocks $ k\geq 2 $, and the matrices $ B_{s, 1}, B_{s, 2}, ..., B_{s, k} $ have sizes $ n_1 \times n_1, n_2 \times n_2, ..., n_k \times n_k $, respectively, for all $ s = 1, .., N $.

    Now, define the matrix $ C $ as

    $ C = S (\lambda_1 I_{n_1 \times n_1} \oplus \lambda_2 I_{n_2 \times n_2} \oplus ... \oplus \lambda_k I_{n_k \times n_k}) S^{-1}, $

    where $ \lambda_1, \lambda_2, ..., \lambda_k $ are any distinct numbers.

    Clearly, the matrix $ C $ commutes with the set $ \{A_s\}_{ s = 1}^N $. Moreover, it has the distinct eigenvalues $ \lambda_1, \lambda_2, ..., \lambda_k $.

    $\Leftarrow$ Assume that the set $ \{A_s\}_{ s = 1}^N $ commutes with a matrix $ C $ that posseses distinct eigenvalues $ \lambda_1, \lambda_2, ..., \lambda_k $.

    Using Proposition 4.1, one can use the generalized eigenspace $ G_{\lambda_i}(C) $ of the matrix $ C $ associated to these distinct eigenvalues to decompose the matrix $ C $ as a direct sum of $ k $ matrices. This can be achieved by restricting the matrix $ C $ on the invariant subspaces $ G_{\lambda_i}(C) $ as follows:

    $ S^{-1}{C}S = {\big[ C \big]}_{G_{\lambda_1}(C)} \oplus {\big[ C \big]}_{G_{\lambda_2}(C)} \oplus ... \oplus {\big[ C \big]}_{G_{\lambda_k}(C)} $

    where

    $ S = \big( G_{\lambda_1}(C), G_{\lambda_2}(C) ,...,G_{\lambda_k}(C) \big). $

    Using Lemma 4.2, one can restrict each matrix $ A_s $ on the invariant subspaces $ G_{\lambda_i}(C) $ to decompose the matrix $ A_s $ as a direct sum of $ k $ matrices as follows:

    $ S^{-1}{A_s}S = {\big[ A_s \big]}_{G_{\lambda_1}(C)} \oplus {\big[ A_s \big]}_{G_{\lambda_2}(C)} \oplus ... \oplus {\big[ A_s \big]}_{G_{\lambda_k}(C)}. $

    Remark 4.4. For a given set of $ n \times n $ matrices $ \{A_s\}_{ s = 1}^N $, if the set $ \{A_s\}_{ s = 1}^N $ commutes only with the matrices having only one eigenvalue, then it does not admit a simultaneous block diagonalization by an invertible matrix.

    Algorithm $ C $:

    (1) Input: the set $ \Omega = \{A_s \}_{ s = 1}^N $.

    (2) Construct the the following matrix:

    $ X=(IA1AT1IIA2AT2I...IANATNI).
    $

    (3) Compute the null space of the matrix $ X $ and reshape the obtained vectors as $ n \times n $ matrices. These matrices commute with all the matrices $ \{A_s \}_{ s = 1}^N $.

    (4) Choose a matrix $ C $ from the obtained matrices that possesses two distinct eigenvalues.

    (5) Find the distinct eigenvalues $ \lambda_1, ..., \lambda_k $ of the matrix $ C $ and the corresponding algebraic multiplicity $ n_1, n_2, ..., n_k $.

    (6) Find each generalized eigenspace $ G_{\lambda_i}(C) $ of the matrix $ C $ associated to the eigenvalue $ \lambda_i $ by computing the null space of $ (C-\lambda_i I)^{n_i} $.

    (7) Construct the invertible matrix $ S $ as

    $ S = \big( G_{\lambda_1}(C), G_{\lambda_2}(C) ,...,G_{\lambda_k}(C) \big). $

    (8) Verify that

    $ S^{-1} A_s S = B_{s,1} \oplus B_{s,2} \oplus ... \oplus B_{s,k}, $

    where the matrices $ B_{s, 1}, B_{s, 2}, ..., B_{s, k} $ have sizes $ n_1 \times n_1, n_2 \times n_2, ..., n_k \times n_k $, respectively, for all $ s = 1, .., N $.

    (9) Output: an invertible matrix $ S $.

    Remark 4.5. Algorithm $ C $ provides the finest block-diagonalization if one chooses a matrix $ C $ with maximum number of distinct eigenvalues. Moreover, the number of the blocks equals the number the of the distinct eigenvalues, and the size of each block is $ n_i \times n_i $, where $ n_i $ is the algebraic multiplicity of the eigenvalue $ \lambda_i $.

    Example 4.6. Consider the set of matrices $ \Omega = \{A_i \}_{i = 1}^6 $ where

    $ A1=(000000000100000010010000001000000000),A2=(000100000000000001100000000000001000),A3=(000010000001000000000000100000010000),A4=(010000100000000000000000000001000010),A5=(001000000000100000000001000000000100),A6=(000000001000010000000010000100000000).
    $
    (4.1)

    The set $ \Omega $ admits simultaneous block diagonalization by an invertible matrix. An invertible matrix can be obtained by applying algorithm $ C $ to the set $ \Omega $ as summarized below:

    A matrix $ C $ that commutes with all the matrices $ \{A_i \}_{ i = 1}^6 $ can be obtained as

    $ C=(000001000010000100001000010000100000).
    $
    (4.2)

    .

    The distinct eigenvalues of the matrix $ C $ are $ \lambda_1 = -1, \lambda_2 = 1 $ with algebraic multiplicities $ n_1 = 3, n_2 = 3 $, respectively..

    The generalized eigenspaces of the matrix $ C $ associated to the distinct eigenvalues are

    $ Gλ1(C)=N(Cλ1I)3=e6e1,e2+e5,e4e3,Gλ2(C)=N(Cλ2I)3=e1+e6,e5e2,e3+e4.
    $
    (4.3)

    The invertible matrix $ S = \big(G_{\lambda_1}(C), G_{\lambda_2}(C) \big) $ is

    $ S=(100100010010001001001001010010100100).
    $
    (4.4)

    The set $ S^{-1} \Omega S = \{S^{-1} A_i S\}_{i = 1}^6 $ contains block diagonal matrices where

    $ S1A1S=(000001010)(000001010),S1A2S=(001000100)(001000100),S1A3S=(010100000)(010100000),S1A4S=(010100000)(010100000),S1A5S=(001000100)(001000100),S1A6S=(000001010)(000001010).
    $
    (4.5)

    It is well known that a set of non-defective matrices can be simultaneously diagonalized if and only if the matrices commute. In the case of non-commuting matrices, the best that can be achieved is simultaneous block diagonalization. Both Algorithm B and the Maehara et al. [9] algorithm are applicable for simultaneous block diagonalization of a set of matrices by a unitary matrix. Algorithm C can be applied for block diagonalization by an invertible matrix when finding a unitary matrix is not possible. In case block diagonalization of a set of matrices is not possible by a unitary or an invertible matrix, then one may utilize block triangularization by Algorithm A. Algorithms A and B are based on the existence of invariant subspaces; however, Algorithm C is based on the existence of a commuting matrix which is not necessarily Hermitian, unlike the Maehara et al. algorithm.

    The authors declare they have not used Artificial Intelligence (AI) tools in the creation of this article.

    Ahmad Y. Al-Dweik and M. T. Mustafa would like to thank Qatar University for its support and excellent research facilities. R. Ghanam and G. Thompson are grateful to VCU Qatar and Qatar Foundation for their support.

    The authors declare that they have no conflicts of interest.

    Figure Listing 1.  Step 5 in Algorithm $ A $.
    Figure Listing 2.  Step 6 in Algorithm $ A $.
    Figure Listing 3.  Steps 8 & 9 in Algorithm $ A $.
    Figure Listing 4.  Steps 2 & 3 in Algorithm $ C $.
    Figure Listing 5.  Steps 6 & 7 in Algorithm $ C $.


    [1] Azam F, Smith DC, Steward GF, et al. (1993) Bacteria-organic matter coupling and its significance for oceanic carbon cycling. Microb Ecol 28: 167–179.
    [2] Hoppe HG, Arnosti C, Herndl GJ (2002a) Ecological significance of bacterial enzymes in the marine environment. In: Burns RG, Dick RP, Enzyme in the environment: activity ecology and application, New York: Marcel Dekker, 73–108.
    [3] Azam F, Fenchel T, Field JG, et al. (1983) The ecological role of water column microbes in the sea. Mar Ecol Prog Ser 10: 257–263. doi: 10.3354/meps010257
    [4] Jiao N, Robinson C, Azam F, et al. (2014) Mechanisms of microbial carbon sequestration in the ocean -future research directions. Biogeosciences 11: 5285–5306. doi: 10.5194/bg-11-5285-2014
    [5] Wang L (2018) Microbial control of the carbon cycle in the ocean. Natl Sci Rev 5: 287–291. doi: 10.1093/nsr/nwy023
    [6] Chróst RJ (1991) Environmental control of the synthesis and activity of aquatic microbial ectoenzymes. In: Chróst RJ, Microbial enzymes in aquatic environments, Germany: Springer-Verlag, 29–59.
    [7] Hoppe HG (1993) Use of fluorogenic model substrates for extracellular enzyme activity (EEA) measurement of bacteria, In: Kemp PF, Sherr BF, Sherr EB, et al., Handbook of methods in aquatic microbial ecology. Boca Raton: Lewis Publishers, 423–431.
    [8] Arnosti C (2011) Microbial extracellular enzymes and marine carbon cycle. Ann Rev Mar Sci 3: 401–425. doi: 10.1146/annurev-marine-120709-142731
    [9] Caruso G, Azzaro M, Caroppo C, et al. (2016) Microbial community and its potential as descriptor of environmental status. ICES J Mar Sci 73: 2174–2177. doi: 10.1093/icesjms/fsw101
    [10] Turley CM (1999) The changing Mediterranean Sea - a sensitive ecosystem? Progr Oceanogr 44: 387–400. doi: 10.1016/S0079-6611(99)00033-6
    [11] Malanotte-Rizzoli P, Manca BB, Ribera d'Alcalà M, et al. (1997) A synthesis of the Ionian Sea hydrography, Circulation and water mass pathways during POEM phase I. Progr Oceanogr 39: 153–204. doi: 10.1016/S0079-6611(97)00013-X
    [12] Powley HR, Krom M.D, Van Cappellen P (2017) Understanding the unique biogeochemistry of the Mediterranean Sea: Insights from a coupled phosphorus and nitrogen model. Global Biogeochem Cy 31: 1010–1031. doi: 10.1002/2017GB005648
    [13] Siokou-Frangou I, Christaki U, Mazzocchi MG, et al. (2010) Plankton in the open Mediterranean Sea: a review. Biogeosciences 7: 1543–1586. doi: 10.5194/bg-7-1543-2010
    [14] Zaccone R, Azzaro F, Azzaro M, et al. (2018) Trophic structure and microbial activity in a spawning area of Engraulis encrasicolus. Estuar Coast Shelf Sci 207: 215–222. doi: 10.1016/j.ecss.2018.04.008
    [15] Briand F (2008) Climate warming and related changes in Mediterranean marine biota, In CIESM Workshop Monographs, Monaco, 35: 152.
    [16] Civitarese G, Gacic M, Lipizer M, et al. (2010) On the impact of the Bimodal Oscillating System (BiOS) on the biogeochemistry and biology of the Adriatic and Ionian Seas (Eastern Mediterranean). Biogeosciences 7: 3987–3997. doi: 10.5194/bg-7-3987-2010
    [17] Cunha A, Almeida A, Coelho FJRC, et al. (2010) Bacterial Extracellular Enzymatic Activity in Globally Changing Aquatic Ecosystems. In: Mendez-Vilas A, Current Research, Technology and Education, Topics in Applied Microbiology and Microbial Biotechnology, 124–135. FORMATEX Research Center, Badajoz, Spain.
    [18] Sarmento H, Montoya JM, Vázquez-Domínguez E, et al. (2010) Warming effects on marine microbial food web processes: how far can we go when it comes to predictions? Philos Trans R Soc Lond B Biol Sci 365: 2137–2149. doi: 10.1098/rstb.2010.0045
    [19] IPCC (2018) Summary for Policymakers. In: Global warming of 1.5 ℃ . An IPCC Special Report on the impacts of global warming of 1.5 ℃ above pre-industrial levels and related global greenhouse gas emission pathways, in the context of strengthening the global response to the threat of climate change, sustainable development, and efforts to eradicate poverty, Geneva, Switzerland: World Meteorological Organization, 32.
    [20] Cramer W, Guiot J, Fader M, et al. (2018) Climate changes and interconnected risks to sustainable development in the Mediterranean. Nat clim change. V8: 972–980.
    [21] Shaltout M, Omstedt A (2014) Recent sea surface temperature trends and future scenarios for the Mediterranean Sea. Oceanologia 56: 411–443. doi: 10.5697/oc.56-3.411
    [22] Wohlers J, Engel A, Zollner E, et al. (2009) Changes in biogenic carbon flow in response to sea surface warming. PNAS 106: 7067–7072. doi: 10.1073/pnas.0812743106
    [23] Hoppe HG, Giesenhagen HC, Koppe R, et al. (2013) Impact of change in climate and policy from 1988 to 2007 on environmental and microbial variables at the time series station Boknis Eck, Baltic Sea. Biogeosciences 10: 4529–4546. doi: 10.5194/bg-10-4529-2013
    [24] Walsh DA (2014) Consequences of climate changes on microbial life in the ocean. Microbiology Today.
    [25] Šolić M, Krstulović N, Šantić D, et al. (2017) Impact of the 3 ℃ temperature rise on bacterial growth and carbon transfer towards higher trophic levels: Empirical models for the Adriatic Sea. J Marine Syst173: 81–89.
    [26] Zaccone R., Azzaro M, Caruso G, et al. (2019) Effects of climate changes on the microbial activities and prokaryotic abundances in the euphotic layer of the Central Mediterranean Sea. Hydrobiologia 1–26.
    [27] Hoppe HG, Gocke K, Koppe R, et al. (2002b) Bacterial growth and primary production along a north -south transect of the Atlantic Ocean. Nature 416: 168–172.
    [28] Hoppe HG (1983) Significance of exoenzymatic activities in the ecology of brackish water: measurements by means of methylumbelliferyl-substrates. Mar Ecol Progr Ser 11: 299–308. doi: 10.3354/meps011299
    [29] Sala F, Balagué V, Boras JA, et al. (2016) Contrasting effects of ocean acidification on the microbial food web under different trophic conditions ICES J Mar Sci 73: 670–679.
    [30] Celussi M, Malfatti F, Franzo A, et al. (2017) Ocean acidification effect on prokaryotic metabolism tested in two diverse trophic regimes in the Mediterranean Sea Estuar Coast Shelf Sci 186: 125–138.
    [31] Catalano G, Azzaro M, Bastianini M, et al. (2014) The carbon budget in the northern Adriatic Sea, a winter case study. J Geophys Res Biogeosci 119: 1339–1417.
    [32] Zaccone R, Caruso G, Calì C (2002) Heterotrophic bacteria in the Northern Adriatic Sea: seasonal changes and enzyme profile. Mar Env Res 54: 1–19. doi: 10.1016/S0141-1136(02)00089-2
    [33] Alonzo Saez L, Vàzquez -Domìnguez E, Cardelùs C, et al. (2008) Factors controlling the year-round variability in carbon flux trough bacteria in a coastal marine system. Ecosystem 11: 397–409. doi: 10.1007/s10021-008-9129-0
    [34] Baltar F, Arístegui J, Gasol JM, et al. (2010) High dissolved extracellular enzymatic activity in the deep central Atlantic Ocean. Aquat Microb Ecol 58: 287–302. doi: 10.3354/ame01377
    [35] Baltar F, Legrand C, Pinhassi J (2016) Cell-free extracellular enzymatic activity is linked to seasonal temperature changes: a case study in the Baltic Sea. Biogeosciences 13: 2815–2821. doi: 10.5194/bg-13-2815-2016
    [36] Baltar F, Anxelu GX, Lonborg C (2017) Warming and organic matter sources impact the proportion of dissolved to total activities in marine extracellular enzymatic rates. Biogeochemistry 133: 307–316. doi: 10.1007/s10533-017-0334-9
    [37] Ivančić I, Fuks D, Radic T, et al. (2010) Phytoplankton and bacterial alkaline phosphatase activity in the northern Adriatic Sea. Mar Environ Res 69: 85–94. doi: 10.1016/j.marenvres.2009.08.004
    [38] Ivančić I, Radic T, Lyons DM, et al. (2009) Alkaline phosphatase activity in relation to nutrient status in the northern Adriatic Sea. Mar Ecol Prog Ser 378: 27–35. doi: 10.3354/meps07851
    [39] Bochdansky AB, Puskaric S, Herndl GJ (1995) Influence of zooplankton grazing on free dissolved enzymes in the sea. Mar Ecol Prog Ser 121: 53–63. doi: 10.3354/meps121053
    [40] Zaccone R, Caruso G, Leonardi M, et al. (2015) Seasonal changes on microbial metabolism and biomass in the euphotic layer of Sicilian Channel. Mar Environ Res 112: 20–32. doi: 10.1016/j.marenvres.2015.07.007
    [41] Ivančić I, Pagliaca P, Pfannkuchen M, et al. (2018) Seasonal variations in extracellular enzymatic activity in marine snow-associated microbial communities and their impact on the surrounding water. FEMS Microbiol Ecol 94: 12: 198.
    [42] Hoppe HG, Giesenhagen HC, Gocke K (1998) Changing patterns of bacterial substrate decomposition in a eutrophication gradient. Aquat Microb Ecol 15: 1–13. doi: 10.3354/ame015001
    [43] Caruso G, Zaccone R (2000) Estimates of leucine aminopeptidase activity in different marine and brackish environments. J Appl Microbiol 89: 951–959. doi: 10.1046/j.1365-2672.2000.01198.x
    [44] Caruso G (2010) Leucine aminopeptidase, β-glucosidase and alkaline phosphatase activity rates and their significance in nutrient cycles in some coastal Mediterranean sites. Mar Drugs 8: 916–940. doi: 10.3390/md8040916
    [45] Arnosti C (2014) Patterns of microbially driven carbon cycling in the ocean: link between extracellular enzymes and microbial communities. Adv Oceanogr 12.
    [46] Zaccone R, Caruso G, Azzaro M, et al. (2010) Prokaryotic activities and abundance in pelagic areas of the Ionian Sea. Chem Ecol 26: 169–197. doi: 10.1080/02757541003772914
    [47] Hoppe HG (2003) Phosphatase activity in the sea. Hydrobiologia 493: 187–200. doi: 10.1023/A:1025453918247
    [48] Sala M, Karner M, Arin L, et al. (2001) Measurement of ectoenzyme activities as an indication of inorganic nutrient imbalance in microbial communities. Aquatic Microbial Ecology 23: 301–311. doi: 10.3354/ame023301
    [49] Zaccone R, Boldrin A, Caruso G, et al. (2012) Enzymatic activities and prokaryotic abundance in relation to organic matter along a West-East Mediterranean transect (TRANSMED cruise). Microb Ecol 64: 54–66. doi: 10.1007/s00248-012-0011-4
    [50] Thingstad TF, Krom MD, Mantoura RF, et al. (2005) Nature of phosphorus limitation in the ultraoligotrophic eastern Mediterranean. Science 309: 1068–1071. doi: 10.1126/science.1112632
    [51] Van Wambeke F, Christaki U, Giannakourou A, et al. (2002) Longitudinal and vertical trends of bacterial limitation by phosphorus and carbon in the Mediterranean Sea. Microb Ecol 43: 119–133. doi: 10.1007/s00248-001-0038-4
    [52] Van Wambeke F, Christaki U, Bianchi, et al. (2000) Heterotrophic bacterial production in the Cretan Sea. Prog Oceanogr 46: 215–216.
    [53] Van Wambeke F, Ghiglione JF, Nedoma J, et al. (2009) Bottom-up effects on bacterioplankton growth and composition during summer-autumn transition in the open NW Mediterranean Sea. Biogeosciences 6: 705–720. doi: 10.5194/bg-6-705-2009
    [54] Christian JR, Karl DM (1995) Bacterial ectoenzymes in marine waters: activity ratios and temperature responses in three oceanographic provinces. Limnol Oceanogr 40: 1042–1049. doi: 10.4319/lo.1995.40.6.1042
    [55] Zaccone R, Monticelli LS, Seritti A, et al. (2003) Bacterial processes in the intermediate and deep layers of the Ionian Sea in winter 1999: vertical profiles and their relationship to the different water masses. J Geophys Res 108: 8117.
    [56] La Ferla R, Azzaro M, Caruso G, et al. (2010) Prokaryotic abundance and heterotrophic metabolism in the deep Mediterranean Sea. Adv Oceanog Limnol 1: 143–166. doi: 10.4081/aiol.2010.5298
    [57] Azzaro M, La Ferla R, Maimone G, et al. (2012) Prokaryotic dynamics and heterotrophic metabolism in a deep convection site of Eastern Mediterranean Sea (the Southern Adriatic Pit). Continental Shelf Res 44: 106–118. doi: 10.1016/j.csr.2011.07.011
    [58] Monticelli LS, Caruso G, Decembrini F (2014) Role of prokaryotic biomasses and activities in carbon and phosphorus cycles at a coastal, thermohaline front and in offshore waters (Gulf of Manfredonia, Southern Adriatic Sea). Microb Ecol 67: 501–519. doi: 10.1007/s00248-013-0350-9
    [59] Luna GM, Bianchelli S, Decembrini F, et al. (2012) The dark portion of the Mediterranean Sea is a bioreactor of organic matter cycling. Global Biogeochem Cycles 26: GB2017.
    [60] Caruso G, Monticelli L, Azzaro F, et al. (2005) Dynamics of extracellular enzymatic activities in a shallow Mediterranean ecosystem (Tindari ponds, Sicily). Mar Freshwater Res 56: 173–188. doi: 10.1071/MF04049
    [61] Tamburini C, Garcin J, Ragot M, et al. (2002) Biopolymer hydrolysis and bacterial production under ambient hydrostatic pressure though a 2000 m water column in the NW Mediterranean. Deep Sea Res II 49: 2109–2123. doi: 10.1016/S0967-0645(02)00030-9
    [62] Tamburini C, Garel M, Ali BA, et al. (2009) Distribution and activity of Bacteria and Archea in the different water masses of the Tyrrhenian Sea. Deep Sea Res II 56: 700–712. doi: 10.1016/j.dsr2.2008.07.021
    [63] Caruso G, Monticelli L, La Ferla T, et al. (2013) Patterns of prokaryotic activities and abundance among epi- meso- and bathypelagic zones of the Southern Central Tyrrhenian Sea. Oceanography 1: 105.
    [64] Placenti F, Azzaro M, Artale V, et al. (2018) Biogeochemical patterns and microbial processes in the Eastern Mediterranean Deep Water of Ionian Sea. Hydrobiologia 815: 97–112. doi: 10.1007/s10750-018-3554-7
    [65] Misic C, Covazzi Harriague A (2008) Organic matter recycling in a shallow coastal zone [NW Mediterranean]: the influence of local and global climatic forcing and organic matter lability on hydrolytic enzyme activity. Continental Shelf Res 28: 2725–2735. doi: 10.1016/j.csr.2008.09.006
    [66] Misic C, Castellano M, Covazzi Harriague A (2011) Organic matter features, degradation and remineralisation at two coastal sites in the Ligurian Sea [NW Mediterranean] differently influenced by anthropogenic forcing. Mar Environ Res 72: 67–74. doi: 10.1016/j.marenvres.2011.05.006
    [67] Baltar F, Arístegui J, Sintes E, et al. (2009) Prokaryotic extracellular enzyme activity in relation to biomass production and respiration in the meso- and bathypelagic waters of the [sub]tropical Atlantic. Environ Microbiol 11: 1998–2014. doi: 10.1111/j.1462-2920.2009.01922.x
    [68] Misic C, Fabiano M (2006) Ectoenzymatic activity and its relationship to chlorophyll-a and bacteria in the Gulf of Genoa (Ligurian Sea, NW Mediterranean) J Mar Sys 60: 193–206.
    [69] Celussi M, Quero GM, Zoccarato L, et al. (2018) Planktonic prokaryote and protist communities in a submarine canyonsystem in the Ligurian Sea (NW Mediterranean). Progress Oceanography168: 210–221.
    [70] Celussi M., Del Negro P (2012) Microbial degradation at a shallow coastal site: long term spectra and rates of exoenzymatic activities in the NE Adriatic Sea. Estuarine coastal shelf Sci v115: 75–86.
    [71] Céa B, Lefèvre D, Chirurgien L, et al. (2014p) An annual survey of bacterial production, respiration and ectoenzyme activity in coastal NW Mediterranean waters: temperature and resource controls. Environ Sci Pollut Res 22: 13654–13668.
    [72] La Ferla R, Zaccone R, Caruso G, et al. (2001) Enzymatic activities and carbon flux through the microbial compartment in the Adriatic Sea. In: Faranda FM, Guglielmo L, Spezie G, Mediterranean Ecosystems: structures and processes, Springer, Milano, 485–493.
    [73] Zaccone R, Caruso G (2002) Microbial hydrolysis of polysaccharides and organic phosphates in the northern Adriatic Sea. Chem Ecol 18: 85–94. doi: 10.1080/02757540212691
    [74] La Ferla R, Azzaro F, Azzaro M, et al. (2005) Microbial contribution to carbon biogeochemistry in the Mediterranean Sea: variability of activities and biomass. J Mar Syst 57: 146–166. doi: 10.1016/j.jmarsys.2005.05.001
    [75] Zaccone R, Azzaro M, Azzaro F, et al. (2014) Seasonal dynamics of prokaryotic abundance and activities in relation to environmental parameters in a transitional aquatic ecosystem (Cape Peloro, Italy). Microb Ecol 67: 55–66.
    [76] Sinsabough RL and Follstad Shah JJ (2012) Ecoenzymatic Stoichiometry and Ecological Theory. Ann Rev Ecol Syst 43: 313–343. doi: 10.1146/annurev-ecolsys-071112-124414
    [77] Ivančić I, Paliaga P, Pfannkuchen M, et al. (2018) Seasonal variation of extracellular enzymatic activity in marine snow-associated microbial communities and their impact on the surrounding water. FEMS Microbial Ecology 94.
    [78] Azam F, Malfatti F (2007) Microbial structuring of marine ecosystems. Nat Rev Microbiol 5: 782–791. doi: 10.1038/nrmicro1747
    [79] Zaccone R, Azzaro M, Caroppo C, et al. (2004) Deep-Chlorophyll Maximum time series in the Augusta Gulf (Ionian Sea): microbial community structures and functions. Chem Ecol 20: S276–S284.
  • Reader Comments
  • © 2019 the Author(s), licensee AIMS Press. This is an open access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/4.0)
通讯作者: 陈斌, bchen63@163.com
  • 1. 

    沈阳化工大学材料科学与工程学院 沈阳 110142

  1. 本站搜索
  2. 百度学术搜索
  3. 万方数据库搜索
  4. CNKI搜索

Metrics

Article views(5178) PDF downloads(526) Cited by(21)

Figures and Tables

Figures(1)  /  Tables(1)

Other Articles By Authors

/

DownLoad:  Full-Size Img  PowerPoint
Return
Return

Catalog