Cell division: mitosis and meiosis

Learning Objectives

  1. Describe the chromosomal makeup of a cell using the terms chromosome, sister chromatid, homologous chromosome, diploid, haploid, and tetrad
  2. Recognize the function and products of mitosis and meiosis
  3. Compare and contrast the behaviors of chromosomes in mitosis and meiosis
  4. Recognize when cells are diploid vs. haploid
  5. Predict DNA content of cells in different phases of mitosis and meiosis
  6. Recall and describe the phases of the cell cycle
  7. Relate the cell cycle stages to changes in DNA content

The Cell Division Cycle

Cell_Cycle_2-2

Cell division cycle, figure from Wikipedia. Cells that stop dividing exit the G1 phase of the cell cycle into a so-called G0 state.

Cells reproduce genetically identical copies of themselves by cycles of cell growth and division. The cell cycle diagram on the left shows that a cell division cycle consists of 4 stages:

  • G1 is the period after cell division, and before the start of DNA replication. Cells grow and monitor their environment to determine whether they should initiate another round of cell division.
  • S is the period of DNA synthesis, where cells replicate their chromosomes.
  • G2 is the period between the end of DNA replication and the start of cell division. Cells check to make sure DNA replication has successfully completed, and make any necessary repairs.
  • M is the actual period of cell division, consisting of prophase, metaphase, anaphase, telophase, and cytokinesis.

Chromosomes

Chromosomes were first named by cytologists viewing dividing cells through a microscope. The modern definition of a chromosome now includes the function of heredity and the chemical composition. A chromosome is a DNA molecule that carries all or part of the hereditary information of an organism. In eukaryotic cells, the DNA is packaged with proteins in the nucleus, and varies in structure and appearance at different parts of the cell cycle.

Chromosomes condense and become visible by light microscopy as eukaryotic cells enter mitosis or meiosis. During interphase (G1 + S + G2), chromosomes are fully or partially decondensed, in the form of chromatin, which consists of DNA wound around histone proteins (nucleosomes).

chromatin_lg

DNA is packaged with histones to form nucleosomes, that resemble “beads on a string” when fully decondensed chromatin is visualized by electron microscopy. Chromatin can be wound into higher-order fibers and into fully condensed chromosomes visible by light microscopy during mitosis or meiosis. Figure from https://www.genome.gov/Glossary/index.cfm?id=32

In G1, each chromosome is a single chromatid. In G2, after DNA replication in S phase, as cell enter mitotic prophase, each chromosome consists of a pair of identical sister chromatids, where each chromatid contains a linear DNA molecule that is identical to the joined sister. The sister chromatids are joined at their centromeres, as shown in the image below. A pair of sister chromatids is a single replicated chromosome, a single package of hereditary information.

Human karyotype "painted" using fluorescent DNA probes. These mitotic chromosomes each consist of a pair of sister chromatids joined at their centromeres. The images of the homologous chromosome pairs (e.g., 2 copies of chromosome 1) have been lined up next to each other. Image from Bolzer et al., (2005) Three-Dimensional Maps of All Chromosomes in Human Male Fibroblast Nuclei and Prometaphase Rosettes. PLoS Biol 3(5): e157 DOI: 10.1371/journal.pbio.0030157

Human karyotype “painted” using fluorescent DNA probes. These mitotic chromosomes each consist of a pair of sister chromatids joined at their centromeres. The images of the homologous chromosome pairs (e.g., 2 copies of chromosome 1) have been lined up next to each other. Image from Bolzer et al., (2005) Three-Dimensional Maps of All Chromosomes in Human Male Fibroblast Nuclei and Prometaphase Rosettes. PLoS Biol 3(5): e157 DOI: 10.1371/journal.pbio.0030157

Ploidy

Humans are diploid, meaning we have two copies of each chromosome. We inherited one copy of each chromosome from other mother, and one copy of each from our father. Gametes (sperm cells or egg cells) are haploid, meaning that they have just one complete set of chromosomes.

Chromosomes that do not differ between males and females are called autosomes, and the chromosomes that differ between males and females are the sex chromosomes, X and Y for most mammals. Humans most commonly have 22 pairs of autosomes and 1 pair of sex chromosomes (XX or XY), for a total of 46 chromosomes. We say that humans have 2N = 46 chromosomes, where N = 23, or the haploid number of chromosomes.

Cells with complete sets of chromosomes are called euploid; cells with missing or extra chromosomes are called aneuploid. The most common aneuploid condition in people is variation in the number of sex chromosomes: XO (having just one copy of the X), XXX, or XYY. Having no X chromosome results in early embryonic death.

The two copies of a particular chromosome, such as chromosome 1, are called homologous. The karyotype image above shows the homologous pairs for all the autosomes. Homologous chromosomes are not identical to each other, unlike sister chromatids. They frequently have different variants of the same hereditary information – such as blue eye color vs brown eye color, or blood type A versus blood type B.

Mitosis

Mitosis produces two daughter cells that are genetically identical to each other, and to the parental cell. A diploid cell starts with 2N chromosomes and 2X DNA content. After DNA replication, the cells is still genetically diploid (2N chromosome number), but has 4X DNA content because each chromosome has replicated its DNA. Each chromosome now consists of a joined pair of identical sister chromatids. During mitosis the sister chromatids separate and go to opposite ends of the dividing cell. Mitosis ends with 2 identical cells, each with 2N chromosomes and 2X DNA content. All eukaryotic cells replicate via mitosis, except germline cells that undergo meiosis (see below) to produce gametes (eggs and sperm).

  • prophase – chromosomes condense; each chromosome consists of a pair of identical sister chromatids joined at the centromere.
  • metaphase – chromosomes line up at the middle of the cell, along the plane of cell division, pushed and pulled by microtubules of the spindle apparatus
  • anaphase – sister chromatids separate and migrate towards opposite ends of the cell
  • telophase – chromatids cluster at opposite ends of the cell and begin to decondense
  • cytokinesis – the membrane pinches in to divide the two daughter cells

Here is a simplified diagram illustrating the overall process and products of mitosis:

File:Major events in mitosis.svg

Source: Wikimedia Commons (https://commons.wikimedia.org/wiki/File:MajorEventsInMeiosis_variant_int.svg)

Questions or points to ponder or note about the figure above (answers at bottom of page):

  1. are the two daughter cells the same or different from each other, and from the parent cell at the start?
  2. why does the cartoon illustration of the chromosomes change (from a single rod to joined double rods) after DNA replication, and again (back to single rods) during mitosis?
  3. does the figure show 2 different chromosomes or a single pair of homologous chromosomes?
  4. can haploid cells undergo mitosis? what about triploid cells (cells that have 3N chromosomes)?

This animation below shows the packaging of DNA and condensation of chromosomes as a cell undergoes mitosis.

The video narration has a major error at time 1:22: chromosomes exist throughout the entire cell cycle (at all times in a cell’s life); they are visible in their condensed form only during mitosis and meiosis.

Meiosis

This is a special sequence of 2 cell divisions that produces haploid gametes from diploid germline cells. It starts with a diploid cell that has undergone chromosomal DNA replication: 2N chromosomes, 4X DNA content. Two successive divisions, with no additional DNA replication, results in 4 haploid gametes: 1N chromosomes, 1X DNA content.

NOVA has a good interactive side-by-side comparison of mitosis and meiosis on this page: How cells divide

Meiosis sets the stage for Mendelian genetics. Students need to know that most of the genetics action occurs in the first meiotic division:

  • homologous chromosomes pair up and align end-to-end (synapsis) in prophase I
  • crossing over occurs between homologous chromosomes in prophase I, before chromosomes line up at the metaphase plate
  • homologous chromosomes separate to daughter cells (sister chromatids do not separate) in the first division, creating haploid (1N) cells
  • the separation of each pair of homologous chromosomes occurs independently, so all possible combinations of maternal and paternal chromosomes are possible in the two daughter cells – this is the basis of Mendel’s Law of Independent Assortment
  • the first division is when daughter cells become functionally or genetically haploid

The last point appears to be the most difficult for students to grasp. Consider the X and Y chromosomes. They pair in prophase I, and then separate in the first division. The daughter cells of the first meiotic division have either an X or a Y; they don’t have both. Each cell now has only one sex chromosome, like a haploid cell.

One way of thinking about ploidy is the number of possible alleles for each gene a cell can have.  Right after meiosis I, the homologous chromosomes have separated into different cells.  Each homolog carries one copy of the gene, and each gene could be a different allele, but these two homologs are now in two different cells.  Though it looks like there are two of each chromosome in each cell, these are duplicated chromosomes; ie, it is one chromosome which has been copied, so there is only one possible allele in the cell (just two copies of it).

The second meiotic division is where sister (duplicated) chromatids separate. It resembles mitosis of a haploid cell. At the start of the second division, each cell contains 1N chromosomes, each consisting of a pair of sister chromatids joined at the centromere.

Here is a simplified diagram illustrating the overall process and products of meiosis:

Meiosis Overview from Wikipedia by Rdbickel. In prophase I, homologous chromosomes pair and separate in the first division (Meiosis I). In Meiosis II, sister chromatids separate.

And here is a video that walks through the steps of meiosis:

It is very important that you recognize how and why cells become haploid after meiosis I.

To confirm for yourself that you understand meiosis, work through one or more of these interactive tutorials:

  • The U. Arizona Cell Biology Project’s Meiosis tutorial has a click-through animation of meiosis, with 10 thought-provoking problem questions.
  • Jung Choi’s interactive flash tutorial, programmed by Pearson, uses human chromosome 7, with wild type and cystic fibrosis alleles for CFTR, to track segregation through meiosis, with and without crossing over: Meiotic Segregation tutorial

Chromosomes, chromatids, what is the difference and how many chromosomes are there at different times of the cell cycle and after mitosis and meiosis?

Chromosomes by definition contain the DNA that makes up the fundamental genome of the cell. In a prokaryote, the genome is usually packaged into one circular chromosome consisting of a circular DNA molecule of a few million base pairs (Mbp). In eukaryotes, the genome is packaged into multiple linear chromosomes, each consisting of a linear DNA molecule of tens or hundreds of Mbp. Chromosomes exist at all different phases of the cell cycle. They condense and become visible to light microscopy in prophase of mitosis or meiosis, and they decondense during interphase, in the form of chromatin (DNA wrapped around nucleosomes, like “beads on a string”).

The chromosome number, N, in eukaryotes, refers to the number of chromosomes in a haploid cell, or gamete (sperm or egg cell). Diploid cells (all the cells in our body except our gametes) have 2N chromosomes, because a diploid organism is created by union of 2 gametes each containing 1N chromosomes. In terms of chromosome number (ploidy), it’s useful to think of chromosomes as packages of genetic information. A pair of sister chromatids is one chromosome because it has genetic information (alleles) inherited from only one parent. A pair of homologous chromosomes, each consisting of a single chromatid in a daughter cell at the end of mitosis, has alleles from the father and from the mother, and counts as 2 chromosomes.

This chromosome number stays the same after chromosome replication during S phase: each chromosome entering cell division now consists of a pair of sister chromatids joined together at the centromere. Then in mitosis, the sister chromatids of each chromosome separate, so each daughter cell receives one chromatid from each chromosome. The result of mitosis is two identical daughter cells, genetically identical to the original cell, all having 2N chromosomes. So during a mitotic cell cycle, the DNA content per chromosome doubles during S phase (each chromosome starts as one chromatid, then becomes a pair of identical sister chromatids during S phase), but the chromosome number stays the same.

A chromatid, then, is a single chromosomal DNA molecule. The number of chromatids changes from 2X in G1 to 4X in G2 and back to 2X, but the number of chromosomes stays the same.

The chromosome number is reduced from 2N to 1N in the first meiotic division, and stays at 1N in the second meiotic division. Because homologous chromosomes separate in the first division, the daughter cells no longer have copies of each chromosome from both parents, so they have haploid genetic information, and a 1N chromosome number. The second meiotic division, where sister chromatids separate, is like mitosis. Chromosome number stays the same when sister chromatids separate.

Using the information above, compare these two simplified diagrams of mitosis and meiosis to visualize why cells are haploid after meiosis I. Specifically, compare the chromosomes in cells at the end of mitosis vs the end of meiosis I, recognizing that the diagram of mitosis tracks just a single pair of homologous chromosomes, whereas the diagram of meiosis tracks two pairs of homologous chromosomes (one long chromosome and short chromosome):

File:Major events in mitosis.svg

Meiosis Overview from Wikipedia by Rdbickel

The video below is geared toward a high school audience, but it does present a helpful way for recognizing how many chromosomes are present in a cell (and thus the ploidy level of that cell).  While watching, see if you can recognize why the products of meiosis 1 are haploid cells:

Answers to questions about the mitosis figure:

  1. The two daughter cells are the same as each other, and same as the parental cell
  2. Each rod represents a chromatid, and DNA replication results in two sister chromatids joined at their centromeres. Mitosis separates the sister chromatids.
  3. A single pair of homologous chromosomes. Red and blue are chromosomes inherited from the male and female parents.
  4. Any cell can divide by mitosis – haploid, triploid, even aneuploid cells.
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7 Responses to Cell division: mitosis and meiosis

  1. Rohan Raman says:

    Is everything we have to know for midterm four on the blog?

    • Jung Choi says:

      This blog does not yet address basic genetics, so no.
      The answer is yes as of Fall 2014; we consider these pages ready for student use. If you feel the page is missing something important, please comment. – response edited by J. Choi 3/27/2015

  2. Julia Fraser says:

    Why are the daughter cells of Meiosis I and of Meiosis II both haploid when they have a different number of chromosomes?

  3. Jung Choi says:

    Here’s a nifty video of mitosis using fluorescence microscopy images. The condensed chromosomes glow red, and the microtubules of the mitotic spindle glow green.
    https://twitter.com/BlackFinch/status/670338456309080065

  4. Jung Choi says:

  5. Pingback: Revision – bbkamodules

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