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Research article

Long lasting effect of acute restraint stress on behavior and brain anti-oxidative status

  • Received: 08 November 2021 Revised: 18 January 2022 Accepted: 18 January 2022 Published: 27 January 2022
  • Exposure to certain acute stressors results in an immediate behavioral and physiological response to these situations during a significant period of days. The goal of the current study is to evaluate the long-lasting effect of single exposure of restraint stress among mice after 0 h, 24 h, 48 h and 72 h. Five groups of mice are under experiment: a control group and four groups exposed to one session of restraint stress. All these groups have been studied for behavioral tests in order to evaluate their memories. This is done through a Y-labyrinth and an object recognition test, and anxiety by using open field device. In the second part of the study, enzymatic assays (concerning catalase, glutathione s transferase, glutathione peroxidase and superoxide dismutase) are used to evaluate oxidative stress. The enzymatic activity of the antioxidant system is assessed in five brain structures, including the cerebellum, olfactory bulb, spinal bulb, hypothalamus, and hippocampus.

    The obtained results show that acute restraint stress leads to a decrease in memory function and to the development of an anxious state; concomitant to an increase of locomotor activity afterword. It causes disturbance of antioxidant balance in the brain by developing a state of oxidative stress. Indeed, restraint stress causes a change in anti-oxidant stress enzymatic activity in the brain, notably in post-stress period. In conclusion, acute restraint stress is responsible for altering cognitive functions, especially memory, and the development of anxious behavior, which could be a result of the generation of oxidative stress; effects that are persistent over an important period after the cessation of stress.

    Citation: Nouhaila Chaoui, Hammou Anarghou, Meriem Laaroussi, Oumaima Essaidi, Mohamed Najimi, Fatiha Chigr. Long lasting effect of acute restraint stress on behavior and brain anti-oxidative status[J]. AIMS Neuroscience, 2022, 9(1): 57-75. doi: 10.3934/Neuroscience.2022005

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  • Exposure to certain acute stressors results in an immediate behavioral and physiological response to these situations during a significant period of days. The goal of the current study is to evaluate the long-lasting effect of single exposure of restraint stress among mice after 0 h, 24 h, 48 h and 72 h. Five groups of mice are under experiment: a control group and four groups exposed to one session of restraint stress. All these groups have been studied for behavioral tests in order to evaluate their memories. This is done through a Y-labyrinth and an object recognition test, and anxiety by using open field device. In the second part of the study, enzymatic assays (concerning catalase, glutathione s transferase, glutathione peroxidase and superoxide dismutase) are used to evaluate oxidative stress. The enzymatic activity of the antioxidant system is assessed in five brain structures, including the cerebellum, olfactory bulb, spinal bulb, hypothalamus, and hippocampus.

    The obtained results show that acute restraint stress leads to a decrease in memory function and to the development of an anxious state; concomitant to an increase of locomotor activity afterword. It causes disturbance of antioxidant balance in the brain by developing a state of oxidative stress. Indeed, restraint stress causes a change in anti-oxidant stress enzymatic activity in the brain, notably in post-stress period. In conclusion, acute restraint stress is responsible for altering cognitive functions, especially memory, and the development of anxious behavior, which could be a result of the generation of oxidative stress; effects that are persistent over an important period after the cessation of stress.



    Let n and k be two positive integers. Denote by p(n,k) the number of partitions of the positive number n on exactly k parts. Then the partition class k is the sequence p(1,k),p(2,k),,p(n,k), We already know, see [1], all these values can be divided into the highest d0=LCM(1,2,,k) sub sequences, each of which is calculated by the same polynomial.

    Choose a sequence of k natural numbers such that: the first member is arbitrary, and the rest form an arithmetic progression with a difference d=md0,mN, starting from the chosen first member. For example:

    x1=j,x2=j+d,,xk=j+(k1)d,jN. (1.1)

    The corresponding number of partitions of the class k for the elements of the previous arithmetic progression's values is:

    p(x1,k),p(x2,k),,p(xk,k). (1.2)

    If the values, which are calculated using the same polynomial, multiplied by the corresponding binomial coefficients, form the alternate sum, we notice that the sum always has a value which is independent of x1, no matter how we form the sequence (1.1).

    For the partition function of classes we already know the following results, see [1,2] for some details:

    ⅰ) The values of the partition function of classes is calculated with one quasi polynomial.

    ⅱ) For each class k the quasi polynomial consists of at most LCM(1,2,,k) different polynomials, each of them consists of a strictly positive and an alternating part.

    ⅲ) All polynomials within one quasi polynomial p(n,k) are of degree k1.

    ⅳ) All the coefficients with the highest degrees down to [k2] are equal for all polynomials (all of strictly positive) and all polynomials differ only in lower coefficients (alternating part).

    ⅴ) The form of any polynomial p(n,k) is:

    p(n,k)=a1nk1+a2nk2++ak, (1.3)

    where the coefficients a1,a2,,ak are calculated in the general form.

    Let us forget for a moment that the coefficients a1,a2, are known in general form. Knowing that all values for partitions class of the sequence (1.1) are obtained by one polynomial p(n,k), it is possible to determine all unknown coefficients in a completely different way from that given in papers [1,2]. To determine k unknowns, a k equation is required. For this purpose, it is sufficient to know all the values of the sequence (1.2). To this end, we must form the system (1.4) and solve it. (For k=10, see [3]).

    a1xk11+a2xk21++ak=p(x1,k)a1xk12+a2xk22++ak=p(x2,k)a1xk1k+a2xk2k++ak=p(xk,k) (1.4)

    The system (1.4) can be solved by Cramer's Rule. For further analysis, we need to find the following determinants. We will start with the known Vandermonde determinant, see [4].

    Δm=|xm11xm211xm12xm221xm1mxm2m1|=1i<jm(xixj),m>1. (1.5)

    When we remove the first column and an arbitrary row from the previous determinant we obtain the Vandermonde determinant of one order less. The following results are known, see [4] and are needed for further exposure. If we remove the second column and an arbitrary a-th row from the determinant (1.5) we get

    |xm11xm31x11xm1a1xm3a1xa11xm1a+1xm3a+1xa+11xm1mxm3mxm1|=(ia0imxi)i,ja0i<jm(xixj) (1.6)

    If we remove the third column and an arbitrary a-th row from the determinant (1.5) we get

    |xm11xm21xm41x11xm1a1xm2a1xm4a1xa11xm1a+1xm2a+1xm4a+1xa+11xm1mxm2mxm4mxm1|=i,ja1i<jmxixji,ja0i<jm(xixj) (1.7)

    Generaly, if we remove the b-th column and an arbitrary a-th row from the determinant (1.5) we get

    Δ(a,b)m=|xm11xm21xb+11xb11x11xm1a1xm2a1xb+1a1xb1a1xa11xm1a+1xm2a+1xb+1a+1xb1a+1xa+11xm1mxm2mxb+1mxb1mxm1|=(t1,,tb1a1t1<t2<tb1mxt1xt2xtb1)i,ja0i<jm(xixj).

    The label Δ(a,b)m means that from Δm remove the a-th row and b-th column from the set of variables xa.

    Theorem 1. Let m,j and k be three positive integers and

    I1(k,j,d)=k1i=0(1)i(k1i)p(j+id,k),

    where d=mLCM(1,2,3,,k). Then I1(k,j,d)=(1)k1dk1k! and is independent of j. (I1(k,j,d) is the first partition invariant which exists in all classes.)

    Proof. Among the values of the class k we choose the ones corresponding to the sequence (1.1), and they are given with the sequence (1.2). According to [2], all the elements in (1.2) can be calculated using the same polynomial p(n,k) with degree k1. Elements of the following sequence:

    q,q+d,,q+(k1)d,qj,

    are calculated with not necessarily the same polynomial as the previous one. Let the polynomial p(n,k) have the form as in (1.3). To determine the coefficients a1,a2,,ak it suffices to know the k values: p(x1,k),p(x2,k),,p(xk,k) where x1=j,x2=j+d,,xk=j+(k1)d are different numbers. Since Δk0, system (1.4) always has a unique solution, because all the elements of the set {x1,x2,,xk} are different from one another. According to Cramer's Rule, to determine the coefficient of the highest degree of the polynomial (1.3), which calculates the value of the number of partitions of class k, we have the following formula:

    a1=p(x1,k)Δ(1,1)kp(x2,k)Δ(2,1)k++(1)k1p(xk,k)Δ(k,1)kΔk (2.1)

    Determinants Δ(a,1)k,(1ak) are also Vandermonde and their values are equal to Δk1. Let {xi}1ik satisfy (1.1) then for 1ak it holds that

    Δ(a,1)k=Δk1=Δka1i=1(xixa)ki=a+1(xaxi)=Δk(1)a1(a1)!da1(1)ka(ka)!dka=(1)k1Δk(a1)!(ka)!dk1.

    Replacing in (2.1), after shortening with Δk we have

    a1=(1)k1(p(x1,k)0!(k1)!dk1p(x2,k)1!(k2)!dk1++(1)k1p(xk,k)(k1)!0!dk1).

    The coefficient a1 is already defined in [2] where it is shown that a1=1k!(k1)!. Substituting into the previous equality and multiplying by (1)k1, we obtain

    (1)k1k!(k1)!=p(x1,k)0!(k1)!dk1p(x2,k)1!(k2)!dk1++(1)k1p(xk,k)(k1)!0!dk1.

    Multiplying the last equality with (k1)!dk1 we obtain

    (1)k1dk1k!=k1i=0(1)i(k1i)p(j+id,k),

    which was to be proved. As these values are equal to each observed number of objects (1.2) within a class, the sum is invariant for any observed class.

    All classes of the partition do not contain all the invariants we will list. This primarily refers to the classes from the beginning. Only the first invariant appears in all classes. The second invariant holds starting from the third class. The third invariant holds starting from the fifth class. Fourth, from the seventh class, etc. This coincides with the appearance of the common coefficients {ak} in quasi polynomials p(n,k), kN.

    Theorem 2. Let m, j and k be three positive integers, k3 and

    I2(k,j,d)=k1i=0(1)i+1(j(k1)+((k2)i)d)(k1i)p(j+id,k)

    where d=mLCM(2,3,,k). Then I2(k,j,d)=(1)k(k3)dk14(k2)! and is independent of j.

    Remark. In the previous expression, we should not simplify as then the value for k=3 cannot be obtained. However, the value for k=3 exists and is equal to zero.

    Proof. Analogously to Theorem 1, the fact that the sum does not depend on the parameter j is a consequence of the periodicity per modulo LCM(2,3,,k) using the same polynomial to calculate the partition class values.

    In [2] it is shown how the system of linear equations can determine the other unknown coefficient of the polynomials which are calculated values of the partition classes. This coefficient is obtained from Cramer's Rule on system (1.4) and a2 is given by

    a2=p(x1,k)Δ(1,2)k+p(x2,k)Δ(2,2)k(1)kp(xk,k)Δ(k,2)kΔk. (2.2)

    Considering (1.6), knowing that {xi}i=1,2,,k is an arithmetic progression, determinants Δ(a,2)k can be written for 1ak with

    Δ(a,2)k=(ia1ikxi)Δk1=((k1)j+((k2)a+1)d)Δka1i=1(xixa)ki=a+1(xaxi)=((k1)j+((k2)a+1)d)Δk(1)a1(a1)!da1(1)ka(ka)!dka=(1)k1((k1)j+((k2)a+1)d)Δk(a1)!(ka)!dk1.

    Knowing the value of the coefficient a2=k34(k1)!(k2)! [2] and substituting in (2.2), and after multiplication with (1)k(k1)!dk1 we obtain

    (1)kk34(k2)!dk1=((k1)j+(k2)d)(k10)p(j,k)((k1)j+((k2)+1)d)(k11)p(j+d,k)+=k1i=0(1)i+1(j(k1)+((k2)i)d)(k1i)p(j+id,k).

    These invariants are in all classes starting from the fifth. For simplicity we denote them by

    R(i,j,k,d)=12(((k1)j+((k2)i)d)2(k1)j2(16k(k1)(2k1)i2)d22dj((k2)i)).

    Theorem 3. Let m,j and k be three positive integers, k5 and

    I3(k,j,d)=ki=0(1)iR(i,j,k,d)(k1i)p(j+id,k),

    where d=mLCM(2,3,,k). Then I3(k,j,d)=(1)k19k358k2+75k2288(k3)!dk1 and is independent of j.

    Proof. For the third invariant we need the value of the third polynomial coefficient of p(n,k), and it is shown [2] that this is

    a3=9k358k2+75k2288(k1)!(k3)!,k5.

    On the other hand, we have

    a3=p(x1,k)Δ(1,3)kp(x2,k)Δ(2,3)k++(1)k1p(xk,k)Δ(k,3)kΔk (2.3)

    From formula (1.7) we find Δ(a,3)k. The required sum 1i<jkxixj is convenient to calculate from the equality

    i,ja1i<jkxixj=12((ia1ikxi)2ia1ikx2i),

    where the sequence {xi} satisfies (1.1). Then, we should determine the quotient which can be simplified by reducing the following:

    Δ(a,3)kΔk=R(a1,j,k,d)a1i=1(xixa)ki=a+1(xaxi).

    By multiplying (2.3) with (1)k1(k1)!dk1 and after shortening we obtain:

    I3(k,j,d)=k1i=0(1)iR(i,j,k,d)(k1i)p(j+id,k).

    In every subsequent invariant, the proceedings become more complex. But, it is quite clear how further invariants can be calculated.

    For each partitions class k, kN we determine d0=LCM(1,2,3,,k), and then form d=md0, mN. In addition arbitrarily choose the natural number j and than form sequences (1.1) and (1.2). Finally, we form an appropriate sum which is for the first invariant:

    k1i=0(1)i(k1i)p(j+id,k)=k1i=0(1)i(k1i)p(xi+1,k),jN. (3.1)

    Sum (3.1) has a constant value in each partitions class and can be nominated as the first partitions class invariant.

    For k=1, sum (3.1) has a constant value of 1.

    For k=2, d0=2. If we choose some mN and set d=2m, the sum (3.1) has the form: p(j,2)p(j+d,2),jN. According to [1], it is known that p(n,2)=[n2]. Distinguishing between even and odd numbers of j (j and j+d have the same parity) and substituting into the sum, we obtain that the result, in both cases, is equal to d2=m.

    For k=3, d0=6. If we choose some mN and set d=6m the sum (3.1) has the form:

    p(j,3)2p(j+d,3)+p(j+2d,3),jN. (3.2)

    According to [1], it is known that:

    p(n,3)=n2+ωi12,i=nmod6,ωi{0,1,4,3,4,1}. (3.3)

    By replacing (3.3) in relation (3.2) we get

    j2+wi1122(j+d)2+wi212+(j+2d)2+wi312.

    Note that: i1=jmod6, i2=(j+d)mod6, i3=(j+2d)mod6 and wi1=wi2=wi3. Finally, we get the unique sum 6m2.

    For k=4, d0=12. If we choose some mN and set d=12m the sum (3.1) has the form:

    p(j,4)3p(j+d,4)+3p(j+2d,4)p(j+3d,4),jN. (3.4)

    According to [1], it is known that:

    p(n,4)=1144n3+148n2+{wi144,n even,116n+wi144,n odd,inmod12, (3.5)
    wi{0,5,20,27,32,11,36,5,16,27,4,11}.

    Similar to case k=3, by distinguishing the even and odd j and replacing (3.5) in relation (3.4) we obtain that the corresponding sums in both cases are equal to: 72m3. (Note that: i1=jmod12, i2=(j+d)mod12, i3=(j+2d)mod12, i4=(j+3d)mod12 and wi1=wi2=wi3=wi4.)

    The number of invariants increases, when the class number increases. Starting with class three, another invariant can be observed.

    Form in the same way as in the previous section: d0, d and the sequences (1.1) and (1.2) as well as the sum:

    k1i=0(1)i(j(k1)+((k2)i)d)(k1i)p(j+id,k).

    Previous sum has a constant value in each partitions class (starting from third class) and can be nominated as the second partitions class invariant.

    For k=3, d0=6. If we choose some mN and set d=6m the general form of the second invariant in the third class can be written as

    (2j+3d)p(j,3)2(2j+2d)p(j+d,3)+(2j+d)p(j+2d,3),jN

    The values p(j,3),p(j+d,3) and p(j+2d,3) are calculated using the same polynomial (3.3). Using (3.3) in the last equality we have

    (2j+3d)j2+wi162(2j+2d)(j+d)2+wi26+(2j+d)(j+2d)2+wi36

    Note that: i1=jmod6, i2=(j+d)mod6, i3=(j+2d)mod6 and wi1=wi2=wi3. The last equality is identical to zero.

    For k=4, d0=12. If we choose some mN and set d=12m the general form of the second invariant in the fourth class can be written as

    (3j+6d)p(j,4)3(3j+5d)p(j+d,4)+3(3j+4d)p(j+2d,4)(3j+3d)p(j+3d,4). (3.6)

    The last equations can be verified in an analogous manner, by using the same form of the known polynomial for the fourth class given in (3.5). Note that: i1=jmod12, i2=(j+d)mod12, i3=(j+2d)mod12, i4=(j+3d)mod12 and wi1=wi2=wi3=wi4. By distinguishing the even and odd j and replacing (3.5) in relation (3.6) we obtain that the corresponding sums in both cases are equal to: 216m3.

    Form in the same way as in the previous two section: d0, d and the sequences (1.1) and (1.2) as well as the sum I3(k,j,d) (Theorem 3). For each class (starting from the fifth) I3(k,j,d) has constant values and can be nominated as the third partitions class invariant. It is known [1] that

    p(n,5)=12880n4+1288n3+1288n2+{124n+wi2880,n even,196n+wi2880,n odd,inmod60, (3.7)

    wi are following numeric respectively:

    0,9,104,351,576,905,216,351,256,9,360,31,576,9,104,225,576,329,216,351,320,9,216,31,576,585,104,351,576,329,360,351,256,9,216,545,576,9,104,351,0,329,216,351,256,585,216,31,576,9,680,351,576,329,216,225,256,9,216,31.

    For k=5, d0=60. If we choose some mN and set d=60m the invariant I3(k,j,d) can be written as:

    12((4j+10d)24j220dj30d2)p(j,5)12((4j+9d)24j218dj29d2)p(j+d,5)+12((4j+8d)24j216dj26d2)p(j+2d,5)12((4j+7d)24j214dj21d2)p(j+3d,5)+12((4j+6d)24j212dj14d2)p(j+4d,5).

    Substituting (3.7) into the previous formula by distinguishing between even and odd j, we obtain a unique value of 1080000m4.

    Remark 1. From the Table 1, see [5], given at the end of the paper it is possible to check all of these explicitly with numerical values. For example:

    Table 1.  Partition classes values.
    d0 1 2 6 12 60 60 420 840 2520 2520
    n/k 1 2 3 4 5 6 7 8 9 10 11 p(n)
    1 1 0 0 0 0 0 0 0 0 0 0 1
    2 1 1 0 0 0 0 0 0 0 0 0 2
    3 1 1 1 0 0 0 0 0 0 0 0 3
    4 1 2 1 1 0 0 0 0 0 0 0 5
    5 1 2 2 1 1 0 0 0 0 0 0 7
    6 1 3 3 2 1 1 0 0 0 0 0 11
    7 1 3 4 3 2 1 1 0 0 0 0 15
    8 1 4 5 5 3 2 1 1 0 0 0 22
    9 1 4 7 6 5 3 2 1 1 0 0 30
    10 1 5 8 9 7 5 3 2 1 1 0 42
    11 1 5 10 11 10 7 5 3 2 1 1 56
    12 1 6 12 15 13 11 7 5 3 2 1 77
    13 1 6 14 18 18 14 11 7 5 3 2 101
    14 1 7 16 23 23 20 15 11 7 5 3 135
    15 1 7 19 27 30 26 21 15 11 7 5 176
    16 1 8 21 34 37 35 28 22 15 11 7 231
    17 1 8 24 39 47 44 38 29 22 15 11 297
    18 1 9 27 47 57 58 49 40 30 22 15 385
    19 1 9 30 54 70 71 65 52 41 30 22 490
    20 1 10 33 64 84 90 82 70 54 42 30 627
    21 1 10 37 72 101 110 105 89 73 55 43 792
    22 1 11 40 84 119 136 131 116 94 75 56 1002
    23 1 11 44 94 141 163 164 146 123 97 77 1255
    24 1 12 48 108 164 199 201 186 157 128 100 1575
    25 1 12 52 120 192 235 248 230 201 164 133 1958
    26 1 13 56 136 221 282 300 288 252 212 171 2436
    27 1 13 61 150 255 331 364 352 318 267 223 3010
    28 1 14 65 169 291 391 436 434 393 340 282 3718
    29 1 14 70 185 333 454 522 525 488 423 362 4565
    30 1 15 75 206 377 532 618 638 598 530 453 5604
    31 1 15 80 225 427 612 733 764 732 653 573 6842
    32 1 16 85 249 480 709 860 919 887 807 709 8349
    33 1 16 91 270 540 811 1009 1090 1076 984 884 10143
    34 1 17 96 297 603 931 1175 1297 1291 1204 1084 12310
    35 1 17 102 321 674 1057 1369 1527 1549 1455 1337 14883
    36 1 18 108 351 748 1206 1579 1801 1845 1761 1626 17977
    37 1 18 114 378 831 1360 1824 2104 2194 2112 1984 21637

     | Show Table
    DownLoad: CSV

    1. Check the first invariant in the third class. Take m=2, j=5. The first invariant formula is

    p(5,3)2p(17,3)+p(27,3).

    From the Table we find: p(5,3)=2, p(17,3)=24, p(29,3)=70. By substitution we find 2224+70=24(=6m2).

    2. Check the second invariant in the forth class. Take m=1, j=3. The second invariant formula is

    81p(3,4)369p(15,4)+357p(27,4)45p(39,4).

    From the Table we find: p(3,4)=0, p(15,4)=27, p(27,4)=150, p(39,4)=441. By substitution we find:

    81036927+35715045441=216(=216m3).

    3. Check the third invariant in the fifth class. Take m=1, j=1. The third invariant formula is

    127806p(1,5)380904p(61,5)+419076p(121,5)206664p(181,5)+40686p(241,5)=1080000.

    Using formulas from (3.7), we find that: p(1,5)=0, p(61,5)=5608, p(121,5)=80631, p(181,5)=393369 and p(241,5)=1220122, and so by checking we are assured of the accuracy.

    Remark 2. Obviously, p(n,k) define values only for nk. The invariants determine very precisely that values for n<k should be taken as zero.

    In this paper, authors have demonstrated a new approach to partitions class invariants, as a way of proving the relevance and accuracy of all formulas given in [1,2]. Also, it I can be considered to be another way to obtain some of the formulas in [2]. The quasi polynomials p(n,k) needed to calculate the number of partitions of a number n to exactly k parts consists of at most LCM(1,2,,k) different polynomials. The invariants claim that the more different polynomials in one quasi polynomial, the more invariable sizes connect them.

    The author thank to The Academy of Applied Technical Studies Belgrade for partial funding of this paper.

    Authors declare no conflicts of interest in this paper.



    Conflict of interest



    The authors declare no conflict of interest.

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