Journal of Cell Science & Therapy

Journal of Cell Science & Therapy
Open Access

ISSN: 2157-7013

+44 1300 500008

Research Article - (2019) Volume 10, Issue 2

Arrangement of Human Cell Metaphase Chromosomes on the Equatorial Plate

Tao Jiang1*, Chunxiao Wu2, Ali Wajid3, Donyun Jiang2, Han Huan1, Zhiren Zhan4 and Jiacheng Huang5
1The Second Affiliated Hospital of Nanjing University of Chinese Medicine, Nanjing, P.R. China
2The First Affiliated Hospital of Soochow University, Suzhou, P.R. China
3Jiangsu Key Laboratory for Molecular and Medical Biotechnology, Nanjing Normal University, Wen Yuan Road, Nanjing, P.R. China
4Xishanqiao Community Health Service Centre, Nanjing, P.R. China
5Nanjing University of Chinese Medicine, Nanjing, P.R. China
*Corresponding Author: Tao Jiang, The Second Affiliated Hospital of Nanjing University of Chinese Medicine, Nanjing, 210023, P.R. China Email:

Received Date: Apr 12, 2019 / Accepted Date: May 03, 2019 / Published Date: May 10, 2019

Abstract

Chromosome is an important hereditary material that maintains the genetic stability of an organism. The structure and location of different chromosomes in the cell governs the basis, how it expresses the genetic effects in the cell and thereby in the whole organism. The chromosome territories (CTs) suggest the location of the chromosomes within the cell by analyzing the karyotypes of human cells. We assume the location and arrangement of chromosomes in the cells is by the interrelationship between chromosomes. More than 100 years ago a Cytologist named Theodor, proposed the idea of chromosome territories. It is indicated that chromosomes are not randomly distributed in Metaphase. This hypothesis is still not directly supported by any laboratory work. We have found in our clinical research that there is a high degree of karyotype expression, so we took a supportive position on this hypothesis but, in order to find the existence of such interrelationships between chromosomes, we conducted a statistical analysis of a large number of existing karyotypic data. In analyzed data we found a clear dominance of long arm over short arm of chromosome according to its activity, with exceptions in specific chromosomes. From the chromosomes statistical data that karyotype occurring between the short arms is relatively small then that of long arm. From the present data it was clear that the exchange of genetic material between the long-long arm is > than exchange of genetic material among long-short arm and short-short arm of the chromosome. Based on the frequency of chromosomal material exchange patterns for arrangement can be considered as circular arrangement, or honeycomb arrangement.

Keywords: Chromosome territories; Metaphase; Karyotype expression; Statistical analysis

Introduction

With few exceptions [1], genomes of living organisms are made of several segments of DNA, they together form a highly organized structure called chromosome. Chromosomes have highly packed DNA that interacted with basic proteins i.e., histones and non-histone, which also play a significant role in the regulation of gene expression [2]. Thus, Chromosomes are genetic vehicles, which facilitate the reproduction and maintenance of a specie’s traits [3,4]. Theodor a cytologist in 1909 for the first time introduced the term chromosome territory [5], and currently several reviews have focused on this topic [6].

Two models have been proposed that explained different aspects of nuclear Chromosome territory distribution [7]. One model is based on the radial arrangement of CTs found in various types of mammalian cells suggests that gene-dense chromosomes are located more internally as compared gene-poor ones [8-10]. However, in non-cycling cells no such arrangement was found by Bridger et al. [11]. The second model proposed a specific neighbourhood relationship between distinct chromosome domains [7]. However, none of the above models have succeeded in proving beyond any doubt the actual mechanism of the chromosome arrangement.

It is imperative to note that, most of the diseases due to genetic defects are directly related to the chromosomal abnormalities. These abnormalities can be better understood by cytogenetics, the field that deals with the structure and properties of chromosomes and cell division, which is based on various methods, one of them known as “karyotyping.” It is a photographic representation of chromosomes organized in a standard manner. For the first in 1960, fibroblasts were used for karyotyping [12]. Karyotypes are prepared from mitotic cells that are arrested in the metaphase state of the cell cycle, the stage when chromosomes reassume their condensed from. As a source of these cells a variety of tissue can be used. In order to diagnose cancer, typical specimens include bone marrow samples or tumour biopsies [13]. With the development of karyotyping, it has made possible to visualize previously undetectable chromosomal abnormalities such as small deletions of chromosomes and translocations between minute segments of chromosomes with each other [14].

The aim of the current study is to demonstrate to certain extent that the location and arrangement of chromosomes in metaphase in the cells is established by the interrelationship between chromosomes based on karyotype expression data analysis. Further, this analysis may shed light on the understanding of mechanism involved in chromosomal defects in rearrangements and possibly the derivation of methods to prevent them.

Research Methodology

We selected data from the National Cancer Institute [15], database on April 26, 2017. There were 10,676 karyotypes of the data sets, with 66,919 cases including karyotype analysis data within chromosomes and among chromosomes. Then we segregated karyotypic analysis data in which the exchange of materials takes place between chromosomes. The data of conventional karyotyping was selected, while the data of complex karyotyping was excluded. Complex karyotyping has its special significance, but in the present condition, there is some interference to the analysis of the data which would be analyzed in the future. Finally, a total of 43,368 valid data sets were collected.

The segregated data sets contains the specific information such as the chromosome number, the long arm and short arm of the chromosome and chromosome division and summarized results. Because of chromosome difference in the gender we calculated the sex chromosomes, X and Y respectively (the statistical data of X chromosome shows a slight increase but has little effect). All data were obtained by means of ordinary accumulation.

Results

Chromosomes short and long arms arrangement

We have found that there is an exchange of materials between the long - long arms, short - long arms and both short arms of chromosome. The high frequency exchange rate was found for long- long arms and short- short arms. It is suggestive that the arrangement of chromosomes in the cells may not be arranged in an “ordered” way, the long arm and the short arm of the chromosome float freely in the cytoplasm of cell during the metaphase. We divided the chromosomes into region 1 and non 1 region to observe the dominance of karyotype analysis data (Tables 1 and 2).

1P P Summary   20 99 2   25 89 2 5   6 2     2 9 2   10 6     12   29
1
Non 1 region   18 99     18   2 2                           12   15
1
1 region   2   2   7 89   3   6 2     2 9 2   10 6         14
0
Non 1 region 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
Q 1 region             9 2 2 2   3 2 7 5 2 4   4     68     11
0
  56 10 8   3 2 13 18 10 2 37 26 11 42   2 24       2       26
6
Summary 56 10 8   3 2 22 20 12 4 37 29 13 49 5 4 28   4   2 68     37
6
  56 30 107 2 3 27 111 22 17 4 43 31 13 49 7 13 30 0 14 6 2 68 12 0 66
7
Non 1 region                                                  
Summary 14           2   4     9       6   5             40
  12           2   2                               16
P 1 region 2               2     9       6   5             24
2P Q   1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
1 region               2     3     3         4     6   2 20
Non 1 region 2   49 4 93   29 101   2 38     52     2 18             39
0
Summary 2   49 4 93   29 103 2 2 41     55     2 18 4     6   2 41
2
  16 0 49 4 93 0 31 103 6 2 41 9 0 55 0 6 2 23 4 0 0 6 0 2 45
2
Summary 2   15 2 2 7       2 2 4             2           38
Non 1 region 2   15                                           17
P 1 region       2 2 7       2 2 4             2           21
3P Q   1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
1         34     13 2                         2     51
region                                                  
Non 1 region 3 9 2   61 2 2 85   2   2 2 8 4   2       2       18
6
Summary 3 9 2   95 2 2 98 2 2   2 2 8 4   2       2 2     23
7
  5 9 17 2 97 9 2 98 2 4 2 6 2 8 4 0 2 0 2 0 2 2 0 0 27
5
Summary 2   2       2               2   4               12
Non 1 region                                                 0
P 1 region 2   2       2               2   4               12
4P Q   1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
1 region 2   2                           2         3     9
Non 1 region 7   10     2               35                     54
Summary 9   12     2               35     2         3     63
  11 0 14 0 0 2 2 0 0 0 0 0 0 35 2 0 6 0 0 0 0 3 0 0 75
Summary 2   4       3                   14               23
Non 1 region                                                 0
P 1 region 2   4       3                   14               23
5P Q   1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
1 region 3   2       6 10             15   19   4           59
Non 1 region 2       2           2     2                     8
Summary 5   2   2   6 10     2     2 15   19   4           67
  7 0 6 0 2 0 9 10 0 0 2 0 0 2 15 0 33 0 4 0 0 0 0 0 90
Summary 25   7       2   8     2 2       2 2 2           52
Non 1 region 18           2   8                               28
P 1 region 7   7                 2 2       2 2 2           24
6P Q   1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
1 region 7   6         5                 2   8           28
Non 1 region   2 5 2 2 2 6 9 146 15 10 6 3 71 2   2               28
3
Summary 7 2 11 2 2 2 6 14 146 15 10 6 3 71 2   4   8           31
1
  32 2 18 2 2 2 8 14 154 15 10 8 5 71 2 0 6 2 10 0 0 0 0 0 36
3
Summary 83 2   2 3       46   82 19       2 5   2           24
6
Non 1 region   2                                             2
P 1 region 83     2 3       46   82 19       2 5   2           24
7P Q   1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
1 region 25
8
      2       6   4     9     8         8     29
5
Non 1 region 5   4     2 20 2     2 2 2 10     6       4       59
Summary 26
3
  4   2 2 20 2 6   6 2 2 19     14       4 8     35
4
  34
6
2 4 2 5 2 20 2 52 0 88 21 2 19 0 2 19 0 2 0 4 8 0 0 60
0
Summary 2               14   3 5       117 12         2     15
5
Non 1 region 2               14                               16
P 1 region                     3 5       117 12         2     13
9
8P Q   1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
1 region 3   2         6       2 16 2     2   6     15     54
Non 1 region 10             7 10     4   2     2       5       40
Summary 13   2         13 10     6 16 4     4   6   5 15     94
  15 0 2 0 0 0 0 13 24 0 3 11 16 4 0 117 16 0 6 0 5 17 0 0 24
9
Summary 5 4       30 46 16 4   7 68 2   4   6 7   3   4     20
6
Non 1 region 2 2       30   16 4                               54
P 1 region 3 2         46       7 68 2   4   6 7   3   4     15
2
9P Q   1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
1 region 14           16   2     2 2   2     4   120 4 27     19
3
Non 1 region 2 2           36 2   48
2
14 2 39 2           4       58
5
Summary 16 2         16 36 4   48
2
16 4 39 4     4   120 8 27     77
8
  21 6 0 0 0 30 62 52 8 0 48
9
84 6 39 8 0 6 11 0 123 8 31 0 0 98
4
Summary     2                                     6     8
Non 1 region                                                 0
P 1 region     2                                     6     8
10P     1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
Q 1 region 2                   56 2                         60
Non 1 region 2                   13
0
  2 9     9               152
  4 0 2 0 0 0 0 0 0 0 18
6
2 2 9 0 0 9 0 0 0 0 6 0 0 22
0
Summary 6   2       84 3 7     2 2   2   2       4 2     11
6
Non 1 region                                                 0
P 1 region 6   2       84 3 7     2 2   2   2       4 2     11
6
11P     1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
Q 1 region   2 6     2 4 2 2 2 2 10   99         4 12   9     15
6
Non 1 region 2 5 2 14 37   12       2     3     4               81
Summary 2 7 8 14 37 2 16 2 2 2 4 10   102     4   4 12   9     23
7
  8 7 10 14 37 2 10
0
5 9 2 4 12 2 102 2 0 6 0 4 12 4 11 0 0 35
3
Summary 2 11 8     6 19 5 68   4   5 3 2 2 4 2 8   5   2   15
6
Non 1 region                                                 0
P 1 region 2 11 8     6 19 5 68   4   5 3 2 2 4 2 8   5   2   15
6
12P Q   1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
1 region 4 4   27 4 5 6 16 2   2 4 37 21 14   52 6 2 2   74     28
Non 1 region 10 5 31   42 6 29 4 6 3       2 6 2 2       39       54
Summary 14 9 31 27 46 11 35 20 8 3 2 4 37 23 20 2 54 6 2 2 39 74     82
  16 20 39 27 46 17 54 25 76 3 6 4 42 26 22 4 58 8 10 2 403 74 2 0 98
4
Summary           2     2     5 2 4 5           4 8     32
Non 1 region                                                 0
P 1 region           2     2     5 2 4 5           4 8     32
13P     1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
Q 1 region 4 2                         2       2     2     12
  Non 1 region                                                 0
  Summary 4 2                         2       2     2     12
    4 2 0 0 0 2 0 0 2 0 0 5 2 4 7 0 0 0 2 0 4 10 0 0 44
  Summary                       3 4 4 5   2       7 7 2   34
  Non 1 region                                                 0
P 1 region                       3 4 4 5   2       7 7 2   34
14P     1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
Q 1 region 6                   2 2     3   2               15
  Non 1 region                                   2             2
  Summary 6                   2 2     3   2 2             17
    6 0 0 0 0 0 0 0 0 0 2 5 4 4 8 0 4 2 0 0 7 7 2 0 51
  Summary 5     2 2       4   2 2 5 5   2 4       9 8     50
  Non 1 region                                                 0
P   5     2 2       4   2 2 5 5   2 4       9 8     50
15P Q   1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
1 region 23                                   2 2   6     33
Non 1 region                                                 0
Summary 23                                   2 2   6     33
  28 0 0 2 2 0 0 0 4 0 2 2 5 5 0 2 4 0 2 2 9 14 0 0 83
Summary 11 6         2 11
7
  4   2     2                   14
4
Non 1 region                                                 0
P 1 region 11 6         2 11
7
  4   2     2                   14
4
16P     1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
Q 1 region 13
5
          2 5     7 96     2   5     2 2 3     25
9
Non 1 region 9   3       11   3   63 3                 66       15
Summary 14
4
  3       13 5 3   70 99     2   5     2 68 3     41
7
  15
5
6 3 0 0 0 15 12
2
3 4 70 10
1
0 0 4 0 5 0 0 2 68 3 0 0 2
Summary 6     4 12   5 10 6   2 6   2   2   8   8         71
Non 1 region                                                 0
p 1 region 6     4 12   5 10 6   2 6   2   2   8   8         71
17P     1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
Q 1 region 11   3 2 30     4 8   13   4   2   2 3   7 5 6     10
Non 1 region 54                 20 2         6                 82
Summary 65   3 2 30     4 8 20 15   4   2 6 2 3   7 5 6     18
2
  71 0 3 6 42 0 5 14 14 20 17 6 4 2 2 8 2 11 0 15 5 6 0 0 25
3
Summary   5       2     7     2         8               24
Non 1 region                                                 0
P 1 region   5       2     7     2         8               24
18P Q   1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
1 region 5 2                     2     2 2     2   6     21
Non 1 region 2                                               2
Summary 7 2                     2     2 2     2   6     23
  7 7 0 0 0 2 0 0 7 0 0 2 2 0 0 2 10 0 0 2 0 6 0 0 47
Summary 6   2     2 2         5                   2     19
Non 1 region                                                 0
P 1 region 6   2     2 2         5                   2     19
19P Q   1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
1 region 45   2   2     2     9 2 2 2 2   2     2 2 5     79
Non 1 region 47
5
        5 2       33
4
    11     28               85
0
Summary 52
0
  2   2   2 2     34
3
2 2 13 2   30     2 2 5     92
9
  52
6
0 4 0 2 2 4 2 0 0 34
3
7 2 13 2 0 30 0 0 2 2 7 0 0 94
8
Summary                 3               2     34 2 2     43
Non 1 region                                                 0
P 1 region                 3               2     34 2 2     43
20 P Q   1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y 25
3
1 region 2     2       2     2   2 5     2 4   16   2     39
Non 1 region 23             2                                 25
Summary 25     2       4     2   2 5     2 4   16   2     64
  25 0 0 2 0 0 0 4 3 0 2 0 2 5 0 0 4 4 0 50 2 4 0 0 10
7
Summary                     4 2 11   9         2 18 2     48
Non 1 region                                                 0
P 1 region                     4 2 11   9         2 18 2     48
21P Q   1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
1 region 4         2 2             2       2     2 4     18
Non 1 region 4                                               4
Summary 8         2 2             2       2     2 4     22
  8 0 0 0 0 2 2 0 0 0 4 2 11 2 9 0 0 2 0 2 20 6 0 0 70
Summary               3 2   2   8 7 8         2 10 6     48
Non 1 region                                                 0
P 1 region               3 2   2   8 7 8         2 10 6     48
22P Q   1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
1 region 14             2         2 4   2       3         27
Non 1 region 4           2   2                               8
Summary 18           2 2 2       2 4   2       3         35
  18 0 0 0 0 0 2 5 4 0 2 0 10 11 8 2 0 0 0 5 10 6 0 0 83
Summary 12     2           4   2   2                     22
Non 1 region 12                                               12
P 1 region       2           4   2   2                     10
XP Q   1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
1 region                             2     16   2   2     16
9
Non 1 region 32                     2   42   3 24               10
3
Summary 32                     2   42 2 3 24 16
3
  2   2     27
2
  44 0 0 2 0 0 0 0 0 4 0 4 0 44 2 3 24 16
3
0 2 0 2 0 0 29
4
Summary                                                 0
Non 1 region                                                 0
P 1 region                                                 0
YP     1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
Q 1 region                                                 0
Non 1 region                           14                     14
Summary                           14                     14
  0 0 0 0 0 0 0 0 0 0 0 0 0 14 0 0 0 0 0 0 0 0 0 0 14
Summary 56 8 9 9 5 7 26
3
13 20 4 2 14 4 6 53 142 20 7 52 8 8 18 32   12
34
Non 1 region 56 8 9       5 10 6                           32   12
6
P 1 region       7 5 7 25
8
3 14 4 2 14 4 6 53 14
2
20 7 52 8 8 8     11
06
1Q Q   1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
1 region   2 4   5 10   12 12   2 2 11 5   53 9 2 21 7 13 20   4 19
4
Non 1 region   21 10 21 10 18 9 4 13 12 43 18 20 39 7 67 8 11     20   2   35
3
Summary   23 14 21 15 28 9 16 25 12 45 20 31 44 7 12
0
17 13 21 7 33 20 2 4 54
7
  56 31 23 30 20 35 27
2
29 45 16 47 34 35 50 60 26
2
37 20 54 15 41 38 34 4 17
81
Summary 8   9     2     4   7 14 2     6   13             65
1 region 6   9     2     2                               19
P   2               2   7 14 2     6   13             46
2Q Q 1 region 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
Non 1 region               2     5   3           4     10 2 2 28
Summary 23   15 4 4       5   3 8 56 10 3               4   13
5
  23   15 4 4     2 5   8 8 56 13 3       4     10 6 2 16
3
  31 0 24 4 4 2   2 9 0 15 22 58 13 3 6 0 13 4 0 0 10 6 2 22
8
Summary 44 59 76 12 2 11 5 2     6 60       30 3   2           31
2
Non 1 region 44 59 76     5 5                                   18
9
P 1 region       12 2 6   2     6 60       30 3   2           12
3
3Q Q   1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
1 region 4         5 2 7     2 2 2       2 2 2 2   52   2 86
Non 1 region 10 15 37
6
4 61 8 31 31     2 43 8 14
1
    14       14
6
      89
0
Summary 14 15 37
6
4 61 13 33 38     4 45 10 14
1
    16 2 2 2 14 52   2 97
6
  58 74 45
2
16 63 24 38 40 0 0 10 105 10 14
1
0 30 19 2 4 2 14
6
52 0 2 12
88
Summary   4       2         14 23         2     2         47
Non 1 region   4       2                                     6
P 1 region                     14 23         2     2         41
4Q Q   1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
1 region               2           2         8     13     25
Non 1 region 21 4 4   2 2     2   67
0
2 2       4               71
3
Summary 21 4 4   2 2   2 2   67
0
2 2 2     4   8     13     73
8
  21 8 4 0 2 4   2 2 0 68
4
25 2 2 0 0 6 0 8 2 0 13 0 0 4
Summary 3 93 118   2 2 2       37 50         30   2           33
9
Non 1 region 3 93 95     2                                     19
3
P 1 region     23   2   2       37 50         30   2           14
6
5Q Q   1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
1 region 5   9               2 2 2 2     14   5   2 4     47
Non 1 region 10 4 37 2 5   4     4 14 6   25     3 4     2       12
0
Summary 15 4 46 2 5   4     4 16 8 2 27     17 4 5   4 4     16
7
  18 97 164 2 7 2 6   0 4 53 58 2 27 0 0 47 4 7 0 4 4 0 0 50
6
Summary 2   2 2   2         2 11                 2   3   26
Non 1 region 2   2     2                                 3   9
P 1 region       2             2 11                 2       17
6Q Q   1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
1 region 8   5               2   2 2 2                   21
Non 1 region 15   6       4 3   2 11
9
4   13                 2   16
8
Summary 23   11       4 3   2 12
1
4 2 15 2               2   18
9
  25 0 13 2   2 4 3 0 2 12 15 2 15 2 0 0 0 0 0 2 0 5 0 21
5
Summary 34 19 2     6 16   16   17 34       13     2   2 2     16
3
Non 1 region 15 19 2     6 16                                   58
P 1 region 19               16   17 34       13     2   2 2     10
5
7Q     1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
Q 1 region     2           9   4 3 2 2     2   4 2   6     36
Non 1 region 7   11   4 4   7 5 23 4 5 2 15 2 2                 91
Summary 7   13   4 4   7 14 23 8 8 4 17 2 2 2   4 2   6     12
7
  41 19 15 0 4 10 16 7 30 23 25 42 4 17 2 15 2 0 6 2 2 8 0 0 29
0
Summary 20 10
3
96   10 17 2 32 38   2 18       5 4   2 6   2     35
7
Non 1 region 18 10 85     12 2 26 38                               28
2
P 1 region 2 2 11   10 5   6     2 18       5 4   2 6   2     75
8Q Q   1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
1 region 12   7 2         6     6 2 10 6   9       8 203     27
1
Non 1 region 4   31       7   6 4 4 4 5 10
61
4           17 30       28 60
Summary 16   38 2     7   12 4 4 10 7 10
71
10   9       17 38 203     31
31
  36 10
3
134 2 10 17 9 32 50 4 6 28 7 10
71
10 5 13 0 2 6 17 38 20 0 0 34 88
Non 1 region 12   2     13
6
6 12 4   6 10       3 4               19
5
1 region 10         13
6
2 10 2                               16
0
P   2   2       4 2 2   6 10       3 4               35
9Q Q 1 region 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
Non 1 region 12           9 6     2 3 2 2 6   4     4   37 25     37 75
Summary 13 5   2     39 10 4 2 10 2 2   5   6       2       10
2
  25 5   2     48 16 4 2 12 5 4 2 11   10     4 2 37 25     38
77
  37 5 2 2   13
6
54 28 4 2 18 15 4 2 11 3 14 0 0 4 2 37 25 0 0 40 68
Summary 8 2 2     15         2 3         20               52
Non 1 region 6 2 2     15                                     25
P 1 region 2                   2 3         20               27
10 Q Q   1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
1 region               2       2 2 2 2   2         12     24
Non 1 region 12       6 2 23 4 2   5 9 6 75 2   2 2     4       15
4
Summary 12       6 2 23 6 2   5 11 8 77 4   4 2     4 12     17
8
  20 2 2 0 6 17 23 6 2 0 7 14 8 77 4 0 24 2 0 0 4 12 0 0 23
0
Summary 43 43     2 10 10   50 2 23
6
2 2   2   23 15   37
9
2         12 71
Non 1 region 41 38       10 6   50 2                               59
7
P 1 region 2 5     2   4     23 6 2 2   2   23 15   37 9 2         67
4
11Q Q   1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
1 region 2       2 2 4   2     6 2 2 2   2   13     33 5     37
4
Non 1 region 55 2 2 67
0
14 11 9 4 4 11 5   2   96
3
7 4 93 29     2   4   19 90
Summary 57 2 2 67
0
16 12
1
8 4 13 5   8 2 96
5
9 4 95 29 13   2 33 5 4   23 64
  10 45 2 67
0
18 13
1
18 4 51 5 24
1
2 10 2 96
7
9 27 11 0 29 39
2
2 2 335 4 0 36 35
Summary 41   2     2 2 6 18     2   2   99     4       2   18
0
Non 1 region 28   2     2 2 4 16                           2   56
P 1 region 13             2 2     2   2   99     4           12
4
12Q Q   1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
1 region 2   5   2   3 6 3 2 6 2 2   5   2 2       21     63
Non 1 region 20 8 49 2 17 4 4 4 4 7 2     54     2               17
7
Summary 22 8 54 2 19 4 7 10 7 9 8 2 2 54 5   4 2       21     24
0
  63 8 56 2 19 6 9 16 25 9 8 4 2 56 5 99 4 2 4 0 0 21 2 0 42
0
Summary 13   2   2 3 2 16 4 2   17         4 2 2 2   2     73
Non 1 region 11   2     3 2   2                               20
P 1 region 2       2     16 2 2   17         4 2 2 2   2     53
13Q Q   1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
1 region 11   2   2 2 2 2 2 2 2 8 14 33 17   9       12 8     12
8
Non 1 region 20 56 8 2     2 5 3 6     6 6 2 2   2         2   12
2
Summary 31 56 10 2 2 2 4 7 5 8 2 8 20 39 19 2 9 2     12 8 2   25
0
  44 56 12 2 4 5 6 23 9 10 2 25 20 39 19 2 13 4 2 2 12 10 2 0 32
3
Summary 51 55 11 35 2 76 19 4 39 9 10 2 24             13 5 2 2 49 21 51
9
Non 1 region 44 52 11     76 10 2 39                           49   28
3
P 1 region 7 3   35 2   9 2   9 10 2 24             13 5 2 2   21 23
6
14 Q Q   1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
1 region 5     2 2 2 2 9 2 2 54           6   18 9 18 13 27     33 3
Non 29 10 13 8   25 8 15 10
93
  7 94 8 48 11 63 2 13 2 17 27     2   4   41 45
Summary 34 10 13 8 2 27 10 17 11
2
2 9 94 8 48 65 63 2 13 8 17 27 18 9 18 15 27 4   44 78
  85 65 14 9 37 29 86 36 11
6
41 18 10 50 72 65 63 2 13 8 17 27 20 2 23 17 29 53 21 49 97
Summary 5   4   9 2     4     13 2 3   2 2   2       2   50
Non 1 region     4     2     2                               8
P 1 region 5       9       2     13 2 3   2 2   2       2   42
15 Q Q   1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
1 region 27         2   6 6 2 2 5 17 21 4   2   2   12 4     11
2
Non 1 region 7 3         3 4 7 2 7   2 2     11
64
      4       12
5
Summary 34 3       2 3 10 13 4 9 5 19 23 4   11
66
  2   16 4     13 17
  39 3 4 0 9 4 3 10 17 4 9 18 21 26 4 2 11
68
0 4 0 16 4 2 0 13
67
Summary 4                 6   2       46 6 2       2     68
Non 1 region 2                                               2
P 1 region 2                 6   2       46 6 2       2     66
16 Q Q   1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
1 region 53                               2         5     60
Non 1 region 68           2       4   2 13     2       25       11
6
Summary 12           2       4   2 13     4       25 5     17
6
  12 5 0 0 0     2   0 6 4 2 2 13 0 46 10 2 0 0 25 7 0 0 24
4
Summary 26 2 2 2 19 4 14 6   11 2 39   2 2 5 2   25 2     24   18
9
Non 1 region 22 2 2     2 6 2                             24   60
P 1 region 4     2 19 2 8 4   11 2 39   2 2 5 2   25 2         12
9
17Q Q   1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
1 region 9   2   14   2 9 4 2 2 6 15   4 2   32 2   10 16     13
1
Non 1 region 8   14 5 3       11 2 93 2   7 12 22 2 9       2   2   13 82
Summary 17   16 5 17   2 9 15 4 95 8 15 7 12 26 4 9 32 2   12 16 2   15 13
  43 2 16 7 36 4 16 15 15 15 97 47 15 9 12 28 9 11 32 27 2 12 16 26 0 17 00
Summary 2 18 3           4     4         3     4 2   163   20
3
Non 1 region 2 18                                             20
P 1 region     3           4     4         3     4 2   16
3
  18
3
18Q Q   1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
1 region 2   2                 2 2       30   2   6 33     79
Non 1 region 11       4     2   2 29     17
8
            4       17 60
Summary 13   2   4     2   2 29 2 2 18
8
    30   2   10 33     18 39
  15 18 5 0 4     2 4 2 29 6 2 17
8
0 0 33 0 2 4 12 33 16 3 0 20 42
Summary 4 4 2   4 8   6     4 2 2   2                   38
Non 1 region 4   2     6   2                                 14
P 1 region   4     4 2   4     4 2 2   2                   24
19Q Q   1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
1 region 41   2 8 5   4       15 2   18
9
2   6 2 7     5     28
8
Non 1 region                                                 0
Summary 41   2 8 5   4       15 2   18
9
2   6 2 7     5     28
8
  45 4 4 8 9 8 4 6 0 0 19 4 2 18
9
4 0 6 2 7 0 0 5 0 0 32
6
Summary                 12
0
  12 2     2 2 7 2 2     3 2   15
4
Non 1 region                 4                               4
P 1 region                 11 6   12 2     2 2 7 2 2     3 2   15
0
20 Q Q   1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
1 region 7   2       2   2   2     18           32   2 2   69
Non 1 region                                                 0
Summary 7   2       2   2   2     18           32   2 2   69
  7 0 2 0     2   12
2
0 14 2 0 18 2 2 7 2 2 32 0 5 4 0 22
3
Summary 4   2       4 2 8     38 8       68 5   2   4       48
7
Non 1 region 4   2       4 2 4                               16
P 1 region                 4     38
8
      68 5   2   4       47
1
21 Q Q   1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
1 region 13       2     8         12 12 12   10 6     4 17     96
Non 1 region 18   14 6   2     17 22 2 4 2     2 4 25 2 4             19 33
Summary 31   14 6   4     17 30 2 4 2   12 14 16 25 12 10     4 17     20 29
  34 0 24 8 0 4   4 17 32 10 4 2 38 8 12 14 16 93 17 10 2 0 8 17 0 0 26 15
Summary 68 6 4 3     11 19     9 56     6 3 6 2 4 2 4   2   20
5
Non 1 region 10 2 4       11                               2   29
P 1 region 58 4   3       19     9 56     6 3 6 2 4 2 4       17
6
22 Q Q   1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
1 region 20 6 52 13 4   6 22
1
36 74 12 33 5 8 8 27 4 5 16 45 5 2 19 14     44   96
Non 1 region                                                 0
Summary 20 6 52 13 4   6 22
1
36 74 12 33 5 8 8 27 4 5 16 45 5 2 19 14     44 96
  88 12 56 16 4   17 24
0
36 74 12 34 4 64 8 27 10 8 22 47 9 4 23 14 2 0 47
01
Summary 2 4       2         4   2 2     2               18
Non 1 region 2 4       2                                     8
P 1 region                     4   2 2     2               10
X Q Q   1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
1 region                                     2           2
Non 1 region                                                 0
Summary                                     2           2
  2 4 0 0   2     0 0 4   2 2     2   2 0 0 0 0 0 20
Summary   2                                             2
Non 1 region                                                 0
P 1 region   2                                             2
YQ Q   1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 x y  
1 region 4   2                                           6
Non 1 region                                                 0
Summary 4   2                                           6
P   4 2 2 0         0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 8

Table 1: Karyotype analysis by chromosome long arm short arm, division summary.

1 P 56 30 107 2 3 27 11
1
22 17 4 43 31 13 49 7 13 30 0 14 6 2 68 12 0 66
7
Q 56 31 23 30 20 35 27
2
29 45 16 47 34 35 50 60 26
2
37 20 54
7
15 41 38 34 4 17
81
11
2
61 13
0
32 23 62 38
3
51 62 20 90 65 48 99 67 27
5
67 20 56
1
21 43 10
6
46 4 24
48
2 P 16 0 49 4 93 0 31 10
3
6 2 41 9 0 55 0 6 2 23 4 0 0 6 0 2 45
2
Q 31 0 24 4 4 2   2 9 0 15 22 58 13 3 6 0 13 4 0 0 10 6 2 22
8
  47 0 73 8 97 2 31 105 15 2 56 31 58 68 3 12 2 36 8 0 0 16 6 4 68
0
3 P 5 9 17 2 97 9 2 98 2 4 2 6 2 8 4 0 2 0 2 0 2 2 0 0 27
5
Q 58 74 45
2
16 63 24 38 40 0 0 10 10
5
10 14
1
0 30 19 2 4 2 14
6
52 0 2 12
88
  63 83 46
9
18 16
0
33 40 13
8
2 4 12 11
1
12 14
9
4 30 21 2 6 2 14
8
54 0 2 15
63
4 P 11 0 14 0 0 2 2 0 0 0 0 0 0 35 2 0 6 0 0 0 0 3 0 0 75
Q 21 8 4 0 2 4   2 2 0 68
4
25 2 2 0 0 6 0 8 2 0 13 0 0 78
5
  32 8 18 0 2 6 2 2 2 0 68
4
25 2 37 2 0 12 0 8 2 0 16 0 0 86
0
5 P 7 0 6 0 2 0 9 10 0 0 2 0 0 2 15 0 33 0 4 0 0 0 0 0 90
Q 18 97 16
4
2 7 2 6   0 4 53 58 2 27 0 0 47 4 7 0 4 4 0 0 50
6
  25 97 17
0
2 9 2 15 10 0 4 55 58 2 29 15 0 80 4 11 0 4 4 0 0 59
6
6 P 32 2 18 2 2 2 8 14 15
4
15 10 8 5 71 2 0 6 2 10 0 0 0 0 0 36
3
Q 25 0 13 2 0 2 4 3 0 2 12
3
15 2 15 2 0 0 0 0 0 2 0 5 0 21
5
  57 2 31 4 2 4 12 17 15
4
17 13
3
23 7 86 4 0 6 2 10 0 2 0 5 0 57
8
7 P 34
6
2 4 2 5 2 20 2 52 0 88 21 2 19 0 2 19 0 2 0 4 8 0 0 60
0
Q 41 19 15 0 4 10 16 7 30 23 25 42 4 17 2 15 2 0 6 2 2 8 0 0 29
0
  387 21 19 2 9 12 36 9 82 23 113 63 6 36 2 17 21 0 8 2 6 16 0 0 89
0
8 P 15 0 2 0 0 0 0 13 24 0 3 11 16 4 0 11
7
16 0 6 0 5 17 0 0 24
9
Q 36 10
3
13
4
2 10 17 9 32 50 4 6 28 7 10
71
10 5 13 0 2 6 17
38
20
5
0 0 34
88
  53 10
3
13
6
2 10 17 9 45 74 4 9 39 23 10
75
10 12
2
29 0 8 6 1743 222 0 0 3739
9 P 21 6 0 0 0 30 62 52 8 0 48
9
84 6 39 8 0 6 11 0 12
3
8 31 0 0 98
4
Q 37 5 2 2 0 13
6
54 28 4 2 18 15 4 2 11 3 14 0 0 4 2 37
25
0 0 40
68
  58 11 2 2 0 16
6
11
6
80 12 2 50
7
99 10 41 19 3 20 11 0 12
7
10 37
56
0 0 50
52
10 P 4 0 2 0 0 0 0 0 0 0 186 2 2 9 0 0 9 0 0 0 0 6 0 0 22
0
Q 20 2 2 0 6 17 23 6 2 0 7 14 8 77 4 0 24 2 0 0 4 12 0 0 23
0
  24 2 4 0 6 17 23 6 2 0 19
3
16 10 86 4 0 33 2 0 0 4 18 0 0 45
0
11 P 8 7 10 14 37 2 10
0
5 9 2 4 12 2 10
2
2 0 6 0 4 12 4 11 0 0 35
3
Q 10
0
45 2 67
0
18 131 18 4 51
5
24
1
2 10 2 96
7
9 27 110 29 39
2
2 2 33
5
4 0 36
35
  10
8
52 12 68
4
55 133 118 9 52
4
24
3
6 22 4 10
69
11 27 116 29 39
6
14 6 34
6
4 0 39
88
12 P 16 20 39 27 46 17 54 25 76 3 6 4 42 26 22 4 58 8 10 2 40
3
74 2 0 98
4
Q 63 8 56 2 19 6 9 16 25 9 8 4 2 56 5 99 4 2 4 0 0 21 2 0 42
0
  79 28 95 29 65 23 63 41 10
1
12 14 8 44 82 27 10
3
62 10 14 2 40
3
95 4 0 14
04
13 P 4 2 0 0 0 2 0 0 2 0 0 5 2 4 7 0 0 0 2 0 4 10 0 0 44
Q 44 56 12 2 4 5 6 23 9 10 2 25 20 39 19 2 13 4 2 2 12 10 2 0 32
3
  48 58 12 2 4 7 6 23 11 10 2 30 22 43 26 2 13 4 4 2 16 20 2 0 36
7
14 P 6 0 0 0 0 0 0 0 0 0 2 5 4 4 8 0 4 2 0 0 7 7 2 0 51
Q 85 65 149 37 29 86 36 11
06
41 18 10
50
72 65 63 2 13 8 17
27
20
2
23 17 29 53 21 49
97
  91 65 14
9
37 29 86 36 11
06
41 18 10
52
77 69 67 10 13 12 17
29
202 23 24 36 55 21 50
48
15 P 28 0 0 2 2 0 0 0 4 0 2 2 5 5 0 2 4 0 2 2 9 14 0 0 83
Q 39 3 4 0 9 4 3 10 17 4 9 18 21 26 4 2 11
68
0 4 0 16 4 2 0 13
67
  67 3 4 2 11 4 3 10 21 4 11 20 26 31 4 4 11
72
0 6 2 25 18 2 0 14
50
16 P 15
5
6 3 0 0 0 15 122 3 4 70 101 0 0 4 0 5 0 0 2 68 3 0 0 56
1
Q 12
5
0 0 0     2   0 6 4 2 2 13 0 46 10 2 0 0 25 7 0 0 24
4
  28
0
6 3 0 0 0 17 12
2
3 10 74 10
3
2 13 4 46 15 2 0 2 93 10 0 0 80
5
17 P 71 0 3 6 42 0 5 14 14 20 17 6 4 2 2 8 2 11 0 15 5 6 0 0 25
3
Q 43 2 16 7 36 4 16 15 15 15 97 47 15 9 12
28
9 11 32 27 2 12 16 26 0 17
00
  114 2 19 13 78 4 21 29 29 35 114 53 19 11 12
30
17 13 43 27 17 17 22 26 0 19
53
18 P 7 7 0 0 0 2 0 0 7 0 0 2 2 0 0 2 10 0 0 2 0 6 0 0 47
Q 15 18 5 0 4     2 4 2 29 6 2 17
08
0 0 33 0 2 4 12 33 16
3
0 20
42
  22 25 5 0 4 2 0 2 11 2 29 8 4 17
08
0 2 43 0 2 6 12 39 163 0 20
89
19 P 52
6
0 4 0 2 2 4 2 0 0 34
3
7 2 13 2 0 30 0 0 2 2 7 0 0 94
8
Q 45 4 4 8 9 8 4 6 0 0 19 4 2 18
9
4 0 6 2 7 0 0 5 0 0 32
6
  57
1
4 8 8 11 10 8 8 0 0 36
2
11 4 20
2
6 0 36 2 7 2 2 12 0 0 12
74
20 P 25 0 0 2 0 0 0 4 3 0 2 0 2 5 0 0 4 4 0 50 2 4 0 0 10
7
Q 7 0 2 0     2   12
2
0 14 2 0 18 2 2 7 2 2 32 0 5 4 0 22
3
  32 0 2 2 0 0 2 4 12
5
0 16 2 2 23 2 2 11 6 2 82 2 9 4 0 33
0
21 P 8 0 0 0 0 2 2 0 0 0 4 2 11 2 9 0 0 2 0 2 20 6 0 0 70
Q 34 0 24
8
0 4   4 17
32
10 4 2 38
8
12 14 16 93 17 10 2 0 8 17 0 0 26
15
  42 0 24
8
0 4 2 6 17
32
10 4 6 39
0
23 16 25 93 17 12 2 2 28 23 0 0 26
85
22 P 18 0 0 0 0 0 2 5 4 0 2 0 10 11 8 2 0 0 0 5 10 6 0 0 83
Q 88 12 56 16 4   17 24
0
36
74
12 34
4
64 8 27 10 8 22 47 9 4 23 14 2 0 47
01
  10
6
12 56 16 4 0 19 24
5
36
78
12 34
6
64 18 38 18 10 22 47 9 9 33 20 2 0 47
84
x P 44 0 0 2 0 0 0 0 0 4 0 4 0 44 2 3 24 16
3
0 2 0 2 0 0 29
4
Q 2 4 0 0   2     0 0 4   2 2     2   2 0 0 0 0 0 20
  46 4 0 2 0 2 0 0 0 4 4 4 2 46 2 3 26 16
3
2 2 0 2 0 0 31
4
y P 0 0 0 0 0 0 0 0 0 0 0 0 0 14 0 0 0 0 0 0 0 0 0 0 14
Q 4 2 2 0         0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 8
  4 2 2 0 0 0 0 0 0 0 0 0 0 14 0 0 0 0 0 0 0 0 0 0 22

Table 2: Chromosome long arm, short arm summary.

Let’s look at the impact of partitioning on karyotype analysis. Different chromosomes are also divided into different regions. We divided chromosomes into 1 region and non-1 region to observe the dominance of karyotype analysis data. Non-1 region contain regions contains zones 2, 3 and 4, of the chromosome. Since most chromosomes in the short arm have only one region and no other region, the long arm has more than two regions, but there are eight zone in the third region and only four zone in the long arm of chromosome No,1. So we look at the long arm data. The data of the short arm was 11:13, which was relatively close, while the data of long-long arm was 3:21, which showed an obvious difference. It should be suggested that the karyotype analysis data of the region far from the centromere in the chromosome was higher than that of the region close to the centromere, whether the region far from the centromere was suggested that it had a large degree of freedom for activity.

In addition to the quantitative differences, the range of activity in the non-1 region is also higher than the corresponding chromosome region 1. The range of activity in the non-1 region is higher than that in the 1 region, but this is not true for all chromosomes. In general, the non-1 region prevails. This indicates that the activity of the non-region 1 chromosome is greater than that of chromosome region 1. The non 1 and 1 region still have overlapping activity, which is inconsistent on each chromosome, more or less. There may be convergence over a larger sample.

Chromosome long arm and short arm activity (comparisons)

From the numbers of long arm and short arm of the chromosomes, we observed that, there is clear dominancy of long arm over short arm of the chromosomes with exceptions in specific chromosomes as shown in Table 3. The ratio of difference is 9:15, long arm being dominant. It has been seen from the data of Table 4, that the activity in long arm of chromosomes is higher than short arm, with the exceptions in specific chromosomes (chromosomes No, 2, 6, 7, 10, 12, 16, 19, X and Y ) that show higher activity in short arm than long arm.

Comparison between short and long arms
S. No P Q
1 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 19 20 21 X 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 X Y
2 1 3 4  5 7 8 9 10 11 12 14 16 17 18 19 22 Y 1  3  4  5  6  8  9  11  12  13  14  15  16 18  19  22  X  Y
3 1  2  3  4  5  6  7  8  9  10  11  12  13  14  15  17  19  21  22 1  2 3 4  5  6  7  8 11  12  14  16  17  18  19  20  21  22  Y
4 1 3 6 7 14 15 17 22 1 2 3 5 6 8 9 11 12 13 14 17 19 20 22
5 1 3 5 7 8 11 14 15 17 19 1 2 3 4 5 6 7 10 11 12 13 14 17 18 19 21 22
6 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 17 18 19 1 3 4 6 7 8 10 11 12 13 14 15 21 X
7 1 2 3 4 5 6 7 8 9 11 12 13 14 16 17 19 21 22 1 2 3 5 6 7 8 9 11 12 13 14 15 16 17 19 20 21 22
8 1 3 8 9 11 12 13 14 16 17 19 21 22 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 1719 20 21 22
9 1 2 6 7 8 9 11 12 13 14 15 17 18 20 21 22 1 2 3 4 6 7 8 9 10 11 12 13 14 15 16 17 20 21 22
10 1 3 11 12 13 14 17 22 1 2 3 5 6 7 8 9 11 12 13 14 15 17 18 21 22
11 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 17 1920 21 22 1  2  3  4  5  6  7  8  9  10  11  12  13  14  15  16  17  18  19  20  21  22  X
12 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 X 1  2  3  4  5  6  7  8  9  10  11  12  13  14  15  16  17  18  19  22  X
13 1 2  6 9 12 13 14 15 19 21 22 1  2  3  4  5  6  7  8  9  10  11  12  13  14  15  16  17  18  19  20  21  22  X
14 1 11 12 13 14 15 17 18 21 22 X 1  2  3  4  5  6  7  8  9  10  11  12  13  14  15  16  17  18  19  20  21  22  X  Y
15 1 4 5 9 11 12 13 14 16 17 19 20 21 22 1 2 3 5 6 7 8 9 10 11 12 13 14 15 16 1719 21 22 X
16 1 2 3 7 8 9 10 11 12 15 17 20 21 22 1 7 10 11 12 13 14 16 17 18 21 22
17 1  3  4  5  7  8  9  10  11  12  13  14  15  16  17  18  20  21  22 1  2  3  4  5  6  8  9  10  11  12  13  14  15  16  17  18  19  20  21  22  X
18 1 2 6 9 12 13 16 17 20 22 1  2  3  5  8  9  10  11  12  13  14  17  19  20  21  22  X
19 1 3 5 6 7 8 11 12 13 14 15 17 20 21 22 1 2 3 4 5 6 7 8 11 12 13 14 15 17 18 19 22
20 1 4 8 9 11 13 14 17 18 20 21 22 1  3  7  9  11  12  14  15  16  17  18  19  20  22  X
21 1 6 7 11 12 13 14 1518 20 21 22 1 3 5 7 8 9 10 11 12 13 14 15 16 17 18 19 21 22
22 1 7 8 9 11 13 14 15 16 20 21 22 1 2 3 4 5 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 X
X 1 4 10 12 14 15 16 17 18 20 22 1 2 6 11 13 14 17 19
Y 14 1 2 3

Table 3: Chromosome long arm and short arm activity.

S. No P Q
1 633 1834
2 470 179
3 489 1080
4 75 788
5 76 507
6 381 213
7 612 354
8 274 3520
9 961 4000
10 278 152
11 344 3598
12 928 394
13 53 313
14 44 5022
15 106 1389
16 587 194
17 202 1647
18 52 2072
19 991 302
20 88 237
21 83 2538
22 80 4780
X 295 24
Y 21 10
Summary 8123 35147

Table 4: Data comparison for long and short arms of chromosome.

Karyotype analysis of the long and short arm of chromosomes

It was found from our analysis that there is no obvious regularity in the data from the long or short arms of the single chromosome and in the other single chromosomes, but in summary of karyotyping the long arm is more obvious than the short arm. The ratio of the number of genes that dominate the exchange of chromosome material is 22: 2, except for two, chromosomes 6 and 16 (Table 5). In the short arm of the chromosome, the exchange of chromosomes materials between the short arm and the long arm is found to be higher than that on the short arm and short arm, however in long arm the material exchanged between chromosomes occurring on long arms is greater than the number of chromosomal material exchanges occurring on long arms and short arms. Karyotyping data elucidates that the long arms have a definite advantage, and this may be due to the arm length of the chromosome. In the analysis of data, in order to eliminate the odd single data while analyzing the whole data, we sorted out the frequency of mass exchange between chromosomes, we analyzed the karyotype of each chromosome frequency by size, arranged in the first 24 calculation, and so on, (X, Y chromosome were calculated) (Figures 1a - 1x).

Long vs short arms
 S. No P Q Overlapping
1 22 24 21
2 17 18 14
3 19 19 15
4 8 15 6
5 10 17 8
6 18 14 12
7 18 19 17
8 13 22 13
9 16 19 15
10 8 17 8
11 20 23 20
12 23 23 21
13 11 23 11
14 11 24 11
15 14 20 12
16 14 12 8
17 19 23 18
18 10 17 8
19 15 17 13
20 12 15 8
21 12 18 10
22 12 22 12
X 11 8 3
Y 1 3 0

Table 5: Comparison of long arms and short arms (karyotype analysis of chromosomes).

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Figure 1a: Genetic material exchange frequency of P and Q arm of chromosome-1 and rest of chromosomes.

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Figure 1b: Genetic material exchange frequency of P and Q arm of chromosome-2 and rest of chromosomes.

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Figure 1c: Genetic material exchange frequency of P and Q arm of chromosome-3 and rest of chromosomes.

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Figure 1d: Genetic material exchange frequency of P and Q arm of chromosome-4 and rest of chromosomes.

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Figure 1e: Genetic material exchange frequency of P and Q arm of chromosome-5 and rest of chromosomes.

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Figure 1f: Genetic material exchange frequency of P and Q arm of chromosome-6 and rest of chromosomes.

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Figure 1g: Genetic material exchange frequency of P and Q arm of chromosome-7 and rest of chromosomes.

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Figure 1h: Genetic material exchange frequency of P and Q arm of chromosome-8 and rest of chromosomes.

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Figure 1i: Genetic material exchange frequency of P and Q arm of chromosome-9 and rest of chromosomes.

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Figure 1j: Genetic material exchange frequency of P and Q arm of chromosome-10 and rest of chromosomes.

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Figure 1k: Genetic material exchange frequency of P and Q arm of chromosome-11 and rest of chromosomes.

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Figure 1l: Genetic material exchange frequency of P and Q arm of chromosome-12 and rest of chromosomes.

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Figure 1m: The 1st value of Q is 44, the second is 56, the 8th is 23, the 12th is 25, and the 14th is 39.

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Figure 1n: Genetic material exchange frequency of P and Q arm of chromosome-14 and rest of chromosomes.

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Figure 1o: Genetic material exchange frequency of P and Q arm of chromosome-15 and rest of chromosomes.

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Figure 1p: Genetic material exchange frequency of P and Q arm of chromosome-16 and rest of chromosomes.

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Figure 1q: Genetic material exchange frequency of P and Q arm of chromosome-17 and rest of chromosomes.

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Figure 1r: Genetic material exchange frequency of P and Q arm of chromosome-18 and rest of chromosomes.

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Figure 1s: Genetic material exchange frequency of P and Q arm of chromosome-19 and rest of chromosomes.

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Figure 1t: Genetic material exchange frequency of P and Q arm of chromosome-20 and rest of chromosomes.

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Figure 1u: Genetic material exchange frequency of P and Q arm of chromosome-21 and rest of chromosomes.

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Figure 1v: Genetic material exchange frequency of P and Q arm of chromosome-22 and rest of chromosomes.

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Figure 1w: Genetic material exchange frequency of P and Q arm of chromosome-23 and rest of chromosomes.

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Figure 1x: Genetic material exchange frequency of P and Q arm of chromosome-24 and rest of chromosomes.

Chromosome segregation statistics between the long arms

From the chromosomes statistical data as shown in Tables 6A6D karyotype occurring between the short arms is relatively small, because the short arms of many chromosomes have no non-1 region. In the comparison of region 1 and non-region 1, it can be seen that the number of karyotypes covered by non-region 1 is more than that of region 1, and the specific data is in more than 1 zone, suggesting that the activity of the non 1 region is greater than the region 1, and they also have the same coverage. Data analysis of the short & long arm, long & short arm, long and long arm of the chromosomes are more obvious. It is further suggested that the coverage of non-1-region is much greater than the rest (Figure 2).

Time Zone
  Non 1 zone 1 zone
1PP 2 3 6 8 9 X 2 6 7 9
2PP 1 7 9 1 9
3PP 1 3 6
4PP   1 3 7 
5PP   1 3 7 
6PP 1 7 9 1 3
7PP 2 1 9
8PP 1 9  
9PP 1 2 6 8 9 1 2
10PP   3
11PP   1 3 7 8 9 
12PP   1 2 3 6 7 8 9 X
13PP   6 9
14PP   X
15PP   1 9 
16PP   1 2
17PP   1 7 8 9 
18PP   2 6 9 
19PP   1 3 6 7 
20PP   9
21PP    
22PP   8 9 
XPP 1 4
YPP    

Table 6A: Statistics of chromosome segregation between short arm and short arm

Comparison table between non 1 zone and 1 zone
  Non 1 zone 1 zone
1PQ 1 2 3 5 6 7 8 9 10 11 12 13 14 16 17 21 7 8 9 10 12 13 14 15 16 17
2PQ 1 3 4 5 7 8 10 11 14 18 8 11 14
3PQ 1 2 3 5 6 7 8 10 12 13 14 15 17 21 5 8 9
4PQ 1 2 3 6 14 1 3 17
5PQ 1 5 11 14 1 3 7 8 15 17
6PQ 2 3 4 5 6 7 8 9 10 11 12 13 14 15 17 1 3 8 17
7PQ 1 3 6 7 11 12 14 17 21 1 5 9 11 14 17
8PQ 18 9 12 14 21 1 3 8 12 13 14 17
9PQ 1 2 8 9 11 12 13 14 15 21 1 7 9 12 13 15 18
10PQ 111 13 14 17 1 11 12
11PQ 1 2 3 4 5 7 11 14 17 2 3 6 7 8 9 10 11 12 14
12PQ 1 2 3 5 6 7 8 9 10 14 15 16 17 21 1 2 4 5 6 7 8 9 11 12 13 14 15 17 18
13PQ   12 15
14PQ 18 1 11 12 15 17
15PQ   1
16PQ 1 3 7 9 11 12 21 1 7 8 11 12 15 17 21
17PQ 1 10 11 16 1 3 4 5 8 9 11 13 15 17 18 21
18PQ 1 1 2 13 16 17
19PQ 1 7 11 14 17 1 3 5 8 11 12 13 14 15 17 21
20PQ 1 8 1 4 8 11 13 14 17 18
21PQ 1 1 6 7 14 18 21
22PQ 1 7 9 1 8 13 14 16
XPQ 1 12 14 16 17 15 18
YPQ 14  

Table 6B: Statistics of chromosome segregation between short arm and long arm.

   Non 1 zone  1 zone
 1 QP  1 2 3 7 8 9 X  6 7 8 9
 2 QP  1 3 6 9  1 9 
 3 QP  1 2 3 6 7 6 8
 4 QP  2 6  
 5 QP  1 2 3 6  37
 6 QP  1 3 6 X  
 7 QP  1 2 3 6 7  1 9
 8 QP  1 2 3 6 7 8 9  1 2 3 6 8
 9 QP  1 6 7 8 9 1 3 7 8 9
 10 QP  1 2 3 6  c
 11 QP  1 2 6 7 9  1 2 7 
 12 QP  1 3 6 7 8 9 X  1 8 9 
 13 QP  1 3 6 7 9  1 8 9 
 14 QP  1 2 3 6 7 8 9 X 1 2 7 8
 15 QP  3 6 9   1 9
 16 QP 1 1
 17 QP  1 2 3 6 7 8 X 1678
 18 QP 12 39X
 19 QP  1 3 6 8  2 5 6 8
 20 QP 9 9 X
 21 QP  1 3 7 8 9 9
 22 QP  1 2 3 7 X 12 8
 X QP 126  
 YQP   2

Table 6C: Chromosome segregation statistics between the long arm and short arm data.

Comparison table between non 1 zone and 1 zone
  Non 1 Zone 1 Zone
1QQ 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 21 X 2 3 5 6 8 9 11 12 13 14 16 17 18 21
2QQ 1 3 4 5 9 11 12 13 14 15 X 8 11 13 X
3QQ 1 2 3 4 5 6 7 8 11 12 13 14 17 21 1 6 7 8 11 12 13 17 18 
4QQ 1 2 3 5 6 9 11 13 17 8 14 
5QQ 1 2 3 4 5 7 10 11 12 1417 18 21 1 3 11 12 13 14 17 21
6QQ 1 3 7 8 10 11 12 14 X 1 3 11 13 14 15
7QQ 1 3 6 8 9 10 11 12 13 14 15 16 3 9 11 12 13 14 17 18 
8QQ 1 3 7 9 10 11 1213 14 15 21 1 3 4 9 12 13 14 15 17 21
9QQ 1 2 4 7 8 9 10 11 12 15 17 21 1  7 8 11 12 13 14 15 17
10QQ 1 5 6 7 8 9 11 12 13 14 15 17 18 21 8 12 13 14 15 17
11QQ 1 2 3 4 5 6 7 8 9 10 12 1 15 16 17 18 21 X 1 5 6 7 9 12 13 14 15 17
12QQ 1 2 3 4 5 6 7 8 9 10 11 14 17 1 3 5 7 8 9 10 11 12 13 15 17 18 
13QQ 1 2 3 4 7 8 9 10 13 14 15 16 18 X 1 3 5 6 7 8 9 10 11 12 13 14 15 17 21 
14QQ 1 2 3 5 6 7 8 10 11 12 13 14 15 16 17 18 21 X 1 4 5 6 7 8 9 10 11 17 21
15QQ 1 2 7 8 9 10 11 13 14 17 21 1 6 8 9 10 11 12 13 14 15 17 21 
16QQ 1 7 11 13 14 17 21 1 17
17QQ 1 3 4 5 9 10 11 12 14 15 16 17 21 X 1 3 5 7 8 9 10 11 12 13 15 16 18 21
18QQ 1 5 8 10 11 14 21 1 3 12 13 17 21
19QQ   1 3 4 5 7 11 12 14 15 17 18 
20QQ   1 3 7 9 11 14 X
21QQ 1 3 5 8 9 10 11 14 15 16 17 18 1 5 8 13 14 15 17 18 21 
22QQ   1 2 3 4 5 7 8 9 10 11 12 13 14 15 16 17 18 21 
XQQ    
YQQ   1 3

Table 6D: Chromosome segregation statistics between the long arm and long arm of the data.

cell-science-therapy-chromosome-activity

Figure 2: Aggregation of chromosome activity.

Aggregation of chromosome activity

In the karyotype’s analysis for chromosomal aggregation, it was observed that long- short arm of the chromosomes overlap with each other in a certain range, meaning there is overlap between homologues and non-homologous chromosomal material exchanges, which strongly supports the concept of Boveri chromosome domain (chromosome territories) (Figures 3 and Figures 4a - 4x).

cell-science-therapy-Chromosomes-aggregation

Figure 3: Chromosomes aggregation.

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Figure 4a: Summary of karyotype analysis of chromosome no.1.

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Figure 4b: Summary of karyotype analysis of chromosome no.2.

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Figure 4c: Summary of karyotype analysis of chromosome no.3.

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Figure 4d: Summary of karyotype analysis of chromosome no.4.

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Figure 4e: Summary of karyotype analysis of chromosome no.5.

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Figure 4f: Summary of karyotype analysis of chromosome no.6.

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Figure 4g: Summary of karyotype analysis of chromosome no.7.

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Figure 4h: Summary of karyotype analysis of chromosome no. 8.

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Figure 4i: Summary of karyotype analysis of chromosome no.9

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Figure 4j: Summary of karyotype analysis of chromosome no.10.

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Figure 4k: Summary of karyotype analysis of chromosome no.11.

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Figure 4l: Summary of karyotype analysis of chromosome no.12.

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Figure 4m: Summary of karyotype analysis of chromosome no.13.

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Figure 4n: Summary of karyotype analysis of chromosome no.14.

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Figure 4o: Summary of karyotype analysis of chromosome no.15

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Figure 4p: Summary of karyotype analysis of chromosome.

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Figure 4q: Value of 15 is 1230.

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Figure 4r: Summary of karyotype analysis of chromosome no.18.

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Figure 4s: Summary of karyotype analysis of chromosome no.19.

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Figure 4t: Summary of karyotype analysis of chromosome no.20.

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Figure 4u: Summary of karyotype analysis of chromosome no.21.

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Figure 4v: Summary of karyotype analysis of chromosome no.2.

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Figure 4w: Summary of karyotype analysis of chromosome X.

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Figure 4x: Summary of karyotype analysis of chromosome Y.

Intensity of chromosome activity

Since karyotype analysis is a clinical activity and we conduct karyotyping analysis of patients, mainly concerned on hematological tumors. Karyotype analysis was performed in patients with greater clinical relevance, such as deformity and infertility, those who have no special clinical significance of deformity were exempted. It was also governed by various factors of the patient profile. For example, the common karyotype of chronic myelogenous leukemia is t (9; 22) (q34; QLL), and the common karyotype of acute promyelocytic leukemia is t (15; 17) (q22; Q12), the data is surprisingly high in certain cases. We calculated the intensity of karyotype analysis, which can effectively reflect on chromosome activity and cumulative frequency of exchange between chromosome exchange intensity (Figure 5).

cell-science-therapy-Intensity-chromosome

Figure 5: Intensity of chromosome activity.

To eliminate the chromosomal bias caused by the clinical factors of a single disease it is seen that the exchange intensities of chromosomes are different, but the active chromosomes are still relatively high in data, while some chromosomes are relatively conserved such as sex chromosomes X, the exchange rate of Y is relatively low, may be consistent with the relative sex chromosomes to maintain their genetic stability.

Possible arrangement of chromosomes in metaphase cells

Based on the frequency of chromosomal material exchange, the frequency is considered as a pair of chromosomes that are closer to each other. Patterns for arrangement can be considered as circular arrangement, or honeycomb arrangement (Figures 6 and 7). From the karyotyping data analysis X and Y chromosome expression is very minute and the rules are very difficult, but we found out in comparing their respective high-frequency data, Both X, Y and chromosome 14 have higher frequency, while the X chromosome and chromosome 18 has the highest frequency. Therefore, the sex chromosomes X and Y may be closer to chromosomes 14 and 18 in the cell as presented in Table 7.

cell-science-therapy-honeycomb-arrangement

Figure 6: The arrangement of chromosomes in the cell equatorial plate honeycomb arrangement.

cell-science-therapy-equatorial-plate

Figure 7: The arrangement of chromosomes in the cell equatorial plate honeycomb arrangement.

1 19 7 16 3 1 22 14 11 15 17  
    12 6 9 2 8 13 x 21 4 5
  20 10 18 y            
2 8 5 3 14 13 11 1 18 7 12  
    22 9 16 4 19 x y 15 6  
  10 17 2 20 21          
3 3 5 14 21 8 12 2 1 22 7 6
    16 17 4 11 13 19 10 15 9  
  18 20 y x            
  11 14 1                
    7 8 9 13 15 20 4 10 16  
  18 21 x y            
5 3 2 17 12 11 14 1 7 15 19 8
    5 10 18 21 22 4 6 13 9  
6 9 11 14 1 3 12 8 10 7 19  
    13 17 x 4 6 15 2 5 18  
  21 16 20 22 y          
7 1 11 9 12 7 14 10 2 17 3  
    16 22 6 5 8 19 13 21 4  
  15 20 18 x y          
8 21 14 22 3 16 2 9 1 8 12  
    17 13 6 5 15 7 11 19 20  
  10 4 18 x y          
9 22 11 6 20 7 12 8 1 14 17  
    15 9 2 18 13 21 16 3 4  
  10 5 19 x y          
10 11 14 17 1 7 22 6 12 13 5 8
    3 15 21 2 9 18 4 10 16  
  19 20 x y            
11 14 4 9 19 22 10 6 7 17 1 5
    2 18 16 12 20 3 15 8 11  
  21 13 x y            
12 21 16 9 3 22 14 1 5 7 17  
    13 8 4 2 15 6 11 19 10  
  18 12 x 20 y          
13 2 1 14 12 15 8 13 22 21 17 3
    9 10 6 7 5 18 19 4 11  
  16 20 x y            
14 18 8 11 19 3 1 6 12 13 14 2
  17 x 9 4 7 22 5 21 20 y  
  10 16 17 15            
15 1 14 13 21 9 12 22 5 11 8
    19 3 6 10 15 16 2 7 4  
  20 x 18 y            
16 1 8 12 21 11 16 7 17 14 10  
    22 2 15 3 9 13 18 20 4 5
  6 19 x y            
17 15 1 11 5 12 18 10 8 9 19 x
    22 7 3 13 16 20 21 4 17  
  14 6 2 y            
18 14 x 17 22 11 2 1 21 9 12  
    20 3 5 13 6 8 10 16 19 4
  7 15 18 y            
19 1 11 14 17 22 5 12 6 3 4 7
    8 19 15 2 13 18 20 21 9  
  10 16 x y            
20 9 20 1 14 11 17 22 18 8 x 3
    4 7 12 13 15 16 19 21 2 5
  6 10 y              
21 8 12 3 16 1 21 15 13 22 17  
    14 18 9 7 11 5 10 6 19  
  20 2 4 x y          
22 9 11 8 1 12 3 18 14 21 17  
    22 7 13 15 4 2 10 16 19  
  20 5 x 6 y          
X 18 1 14 17 2 10 11 12 16 4 6
    13 15 19 20 22 3 5 7 8 9
  21 x y              
Y 14 1 2 3 4 5 6 7 8 9  
    10 11 12 13 15 16 17 18 19  
  20 21 22 X   Y        

Table 7: Chromosome high frequency data arrangement.

Discussion

Chromosomes are considered natural unit of subdivision of complete genome. During mitosis chromosomes can easily be seen in cells as a highly condensed structures but for many years their morphological nature during interphase remained undefined [16]. Thus, it is particularly important to elucidate the arrangement of chromosome during metaphase based on karyotype expression.

Chromosome structure and arrangement in cells are only well observed in metaphase of cells. Karyotype analysis is a prompt examination of clinical diseases (O’Connor, 2008). In our results we evaluated that the exchange of material between chromosomes follows a certain rule that the exchange may occur only in chromosomes adjacent to or close to each other. Individual chromosomes are different, some are more active, some relatively quiet; the long arm and short arm of chromosome is free in the cytoplasm of the cells; activity of the long arm of chromosome is greater and dominant than short arm of chromosome (Tables 1, 3, 4) and (Figures 1a - 1x).

We also found that segregation far from the centromere in different regions of the arm is greater than the segregation near the centromere (many chromosomes do not have region other than zone 1, and these chromosomes are not mentioned) (Tables 6A - 6D). In our work we found both sex chromosomes (X and Y) to be very close to chromosome number 14 and 18 (Table 7). Previously Boyle et al. have revealed that the chromosome which is gene-rich (human chromosome 19) is positioned more central in nucleus than the equally sized chromosome 18 which is gene-poor. Boyle et al. also report that the chromosomes of different functional characteristics and having similar physical size adopt different positions nucleus [17].

So, from our work we conclude that, Arrangement of chromosomes in the cell occurred in a regular pattern. Through the massive karyotyping data, we found an exchange of chromosomal materials for adjacent chromosomes only. The present study suggested that patterns for arrangement of chromosome can be considered as circular arrangement, or honeycomb arrangement (Figures 6 and 7). According to the data analysis in Figures 4A to 4X) it is clear that chromosome No-1 show grater exchange of genetic material with chromosome No-7, 19, and 16. Chromosome No-1 also show low genetic material exchange with Chromosome No-X, Y and 15. Chromosome No-2 show grater exchange of genetic material with Chromosome No-8 and 13. Chromosome No-3 and 5 show grater exchange of genetic material for each other. Chromosome No-4 show grater exchange of genetic material with Chromosome No-11 but chromosome No-11 show grater exchange of genetic material wit chromosome No-10 and 14 in addition chromosome No-14 show grater exchange of genetic material with chromosome No-18 and Y-chromosome. Chromosome No-8, 21 and 15, 17 show grater exchange of genetic material for each other. Chromosome-X and 18 show grater exchange of genetic material.

Conclusion

The above discussed pattern of genetic material exchange is not possible in other form of arrangement than honeycomb arrangement. Due to honeycomb arrangements of chromosome it may be possible to exchange genetic material in such a way. In addition to the location of chromosomes, karyotyping may have some relationship with the structure of chromosomes, which needs further investigation. Through mathematical and statistical analysis of chromosome karyotype analysis, we re-recognized the value of karyotyping.

References

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Citation: Tao J, Chunxiao W, Ali W, Donyun J, Han H, Zhiren Z , et al. (2019) Arrangement of Human Cell Metaphase Chromosomes on the Equatorial Plate. J Cell Sci Ther, 10: 289. doi: 10.35248/2157-7013.19.10.289

Copyright: © 2019 Jiang T, et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
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