Cell Biol Educ 2(4): 214-219 2003
DOI: 10.1187/cbe.03-10-0043
© 2003 American Society for Cell Biology
WWW.Cell Biology Education
Robert Blystone
Department of Biology, Trinity University, San Antonio, Texas 78212
Submitted October 6, 2003;
Accepted October 16, 2003
Each quarter, Cell Biology Education calls attention to several
Web sites of educational interest to the life science community. The journal
does not endorse or guarantee the accuracy of the information at any of the
listed sites. The sites listed below were last accessed on October 5,
2003.
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THE HISTORY OF MITOSIS
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"Mitosis consists of four stages that are named in sequence:
prophase, metaphase, anaphase, and telophase. In between mitotic divisions is
interphase, the period of time during which DNA is replicated." The two
previous sentences along with their supporting material verge on a litany and
a catechism for biology. The column for this issue will focus on Web resources
for teaching and learning about mitosis.
Why do we call asexual cell division "mitosis"? Who created the
names "prophase," "metaphase," "anaphase,"
and "telophase?" Sir Henry Harris, a pathologist from the
University of Oxford, provides a wonderful accounting of the history of
mitosis in his book The Birth of the Cell (Yale University Press,
1999, ISBN 0-300-07384-4). Supplementing Harris' reference are two Web sites
using timelines that put these morphological descriptors of cell division in
their historical place.
Electronic Scholarly Publishing http://www.esp.org/timeline
Electronic Scholarly Publishing (ESP) is an effort established by Robert J.
Robbins. Two of the projects of ESP are a classical collection of genetics
papers and a genetics timeline. If you would like to have Mendel's 1866 paper,
it is here. ESP's genetics timeline places the discoveries leading to the
description of mitosis into chronological perspective. Beginning in 1750, the
timeline juxtapositions science with world events. For mitosis history, the
timeline between 1875 and 1895 is especially interesting.
Lasker Foundation http://www.laskerfoundation.org/news/gnn/timeline/timelinetop.html
The Lasker Foundation also has a nice timeline for genetics and genomics.
Not as ranging as the ESP timeline, the Lasker timeline provides background
information on some of the individuals including Walther Flemming, the person
who coined the term "mitosis" in 1882. The Lasker timeline can be
downloaded as a 211-page PDF file and includes a nice biography and picture of
the first person to recognize that chromosomes undergo longitudinal division
during mitosis, that is, Walther Flemming.
The various origins of the terms associated with mitosis are listed in
these timelines. W. Schleicher developed the term "karyokinesis"
in 1878. Eduard Strasburger created the terms "prophase,"
"metaphase," and "anaphase" in a paper published in
1884. Heinrich Wilhelm Gottfried Waldeyer gave us the term
"chromosome" in 1888. Surprisingly the ending phase of mitosis,
"telophase," was introduced by Martin Heidenhain in 1894. It is
curious that 10 years lapsed between prophase and telophase in terms of
naming. And finally an H. Lundegardh put "interphase" into the
mitotic lexicon in 1913. This history of cytology can be found in greater
depth in the Harris text and in the 1911 Encyclopedia Project.
The 1911 Encyclopedia Project http://93.1911encyclopedia.org/C/CY/CYTOLOGY.htm
This 1911 Encyclopedia article is interesting as a historical document
dealing with what was then the recent history of the development of cytology.
The text of the whole 1911 Encyclopedia is available at the above Web site. Of
a related nature, if you have an interest in structures that are named after
famous scientists, then you might what to check a medical eponym site.
Dictionary of Medical Eponyms http://www.whonamedit.com/
Almost 6500 medical eponyms are listed at this URL. This site is mentioned
because a nice biography of Waldeyer, the primary source for the term
"chromosome," may be found here.
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TEACHING MITOSIS TODAY
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A more recent history of mitosis research may be found in a Nature
article by T.J. Mitchison and E.D. Salmon. (2001. Mitosis: a history of
division. Nature Cell Biology 3(1): E17–21.) A PDF version of
this article may be found at the following URL.
T.J. Mitchison Lab Page http://mitchison.med.harvard.edu/Publications.htm
It was this paper in Nature Cell Biology that inspired this
column. The report provides an excellent pictorial view of the evolution of
thought concerning how the mitotic spindle is formed and generates force. This
five-page paper will bring you to the edge of contemporary ideas on
mitosis.
John Kimble's Electronic Biology Textbook http://users.rcn.com/jkimball.ma.ultranet/BiologyPages/M/Mitosis.html
A good place to start our tour of Internet mitosis sites is Kimble's
textbook description of mitosis. There are many hypertext links to other
locations that provide definitions and examples for key terms encountered in
the vocabulary of mitosis.
BioWeb at University of North Carolina at Charlotte http://www.bioWeb.uncc.edu/biol1110/Stages.htm
Biology 1110 at UNCC is the lab course that supports nonmajors biology. The
course Web site has an excellent traditional exercise on mitosis supported by
images such as found in Figure
1.
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Figure 1. A standard depiction of whitefish and onion mitosis. The chromosomes of
anaphase are clearly realized in each. Courtesy of Dr. Mark Clemens of the
Department of Biology, University of North Carolina at Charlotte.
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VIEWING MITOSIS
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A search of the Internet can provide excellent illustrations and
photographs of mitosis that move beyond the traditional text and lab
materials. Below is a brief sampling of some of these wonderful and often
esthetic views of cell division.
Biology 304 Homepage http://www.micro.utexas.edu/courses/levin/bio304/genetics/celldiv.html
This Web site supports a majors-based course at the University of Texas at
Austin titled Biology 304—Ecology and Evolutionary Biology. The first
image on this Web page has a traditional view of plant mitosis and that figure
is reproduced here as Figure
2.
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Figure 2. The four stages of plant mitosis are seen in this composite with the
chromosomes staining blue and the spindles red. Courtesy of Dr. Andrew Bajer,
Professor Emeritus of the University of Oregon.
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Biology Teaching Organisation http://www.bto.ed.ac.uk/courses/year1/mac1h/gallery/index.html
This site is located at the University of Edinburgh. The image gallery
supports many courses including a first-year course in Molecules and Cells.
Figure 3 is an example of a
beautiful mitotic figure fluorescently stained and found at the Edinburgh
site.
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Figure 3. An early anaphase with red stained chromosomes and green spindle
microtubules of a mammalian kidney cell is illustrated. Courtesy of Professor
William C. Earnshaw, Institute of Cell and Molecular Biology, University of
Edinburgh.
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The Vivek Malhotra Lab http://www-biology.ucsd.edu/labs/malhotra/SciencePictures.htm
The Web site listed here has been developed by Dr. Vivek Malhotra's Lab in
the Division of Biological Sciences at the University of California, San
Diego. There are images at this site that answer a question which students
often ask: what happens to the cell organelles during mitosis.
Figure 4 reveals Golgi being
partitioned during mitosis.
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Figure 4. This photomicrograph appeared on the cover of the April 17, 2000 issue of
the Journal of Cell Biology. The caption states: "Laser
scanning confocal fluorescence image showing the organization of the Golgi
membranes during various stages of the cell cycle. Normal rat kidney cells
(courtesy of Dr. Lippincott-Schwartz) expressing galactosyl transferase-YFP
(shown green) were costained with the DNA-specific stain propidium iodide
(shown blue) and antitubulin antibody (shown red). In the nondividing cells
the Golgi membranes are found in the perinuclear area." The article
associated with the cover image may be found at
http://www.jcb.org/cgi/content/short/149/2/331.
Courtesy of Vivek Malhotra of UCSD and the Rockefeller University Press.
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John Runions' Home Page http://www.plantsci.cam.ac.uk/Haseloff/JohnRunions/Web/Lineage.html http://www.plantsci.cam.ac.uk/Haseloff/JohnRunions
John Runions was a post-doctoral fellow in the lab of Jim Haseloff at the
University of Cambridge when the images found at the URL above were made.
Accompanying the beautiful pictures of Arabidopsis root, Runions'
site has explanations of the techniques employed to produce the wonderful
pictures. The images here help a student understand the lineage of cells that
are rapidly dividing in succession as illustrated in
Figure 5.
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Figure 5. Cell lineage analysis of Arabidopsis root apical meristem is
demonstrated here. Yellow fluorescent protein is bound to histone 2B and
reveals all the cell nuclei that are related to one another. Courtesy of Dr.
C. John Runions of the Department of Biological and Molecular Sciences at
Oxford Brooks University.
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Control of Cell Division, Course BS313 http://www.teaching-biomed.man.ac.uk/ramsay/Overv.htm
The School of Biological Sciences at the University of Manchester offers a
course titled Control of Cell Division. The course has a supporting
Web site that offers information about the control of mitosis at nine
different time points. By clicking on one of the time points arrayed in a
diagram, clear organizational illustrations appear that indicate the
relationships of such compounds as cdc2, cyclin B, wee1, and cdc25 in the
molecular control of mitosis.
Cande Lab http://mcb.berkeley.edu/labs/cande/mitosis.html
Dr. W. Zacheus Cande is a member of the Department of Molecular and Cell
Biology at the University of California at Berkeley. His lab site has some
wonderful images of mitosis including rotating three-dimensional
reconstructions of mitotic chromosomes and represented by
Figure 6.
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Figure 6. This still image was taken from a set of five rotating movies of yeast cell
chromosomes in division as seen from reconstructed laser confocal microscopy
data. Courtesy of Zacheus Cande of the University of California, Berkeley.
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Cyr Lab http://www.bio.psu.edu/People/Faculty/Cyr/Lab/coolmovies/coolmovies.htm
The lab of Dr. Richard J. Cyr of the Department of Biology, Penn State
University, explores questions concerning the cellular basis of plant
morphogenesis. At the URL above are a number of movies that show Golgi
movement, cycle cell movement, and mitosis. The movies found there are in a
variety of Web formats. Figure
7 is representative of the imagery found at this site.
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Figure 7. "The image is of a tobacco cell (BY-2) that has been stably
transformed with a Golgi marker in the red channel (nactylglucosamine:dsRed)
and a microtubule marker in the green (microtubule binding domain of
MAP4:GFP). Plant cells have a barrel shaped anastral spindle and so it looks a
bit different (from) animals." The movie represented by this anaphase
figure will form a phragmoplast, which plant cells use in cytokinesis to lay
down a new plate. Description and figure courtesy of Dr. Richard Cyr,
Department of Biology, Pennsylvania State University.
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Cytographics http://www.cytographics.com/gallery/gal.html http://www.cytographics.com/gallery/clips/prophase.mpg http://www.cytographics.com/gallery/clips/Celldiv.mov http://www.cytographics.com/gallery/clips/Sickcell.mov
Dr. Jeremy Pickett-Heaps has been producing awe-inspiring views of cells
for years. He and his colleagues have formed a distribution company of
educational materials called Cytographics. The first URL takes one to a
gallery of video resources. The last three sites listed above link to direct
downloads of short videoclips of cells in division; the clips may download
slowly. The fourth site provides a view of misdivision in mitosis and shows a
chromosome going astray during division.
Figure 8 is typical of the
quality of the images found at the site.
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Figure 8. A still image from the videoclip of newt lung cells in division. The
chromosomes are on the metaphase plate. Courtesy of Julianne Pickett-Heaps of
Cytographics.
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MBC Online Sampler http://www.molbiolcell.org/misc/video1.shtml
The American Society for Cell Biology publication Molecular Biology of
the Cell has an online sampler page of videos. One of the videos in the
collection represents mitosis in newt lung cell using a new type of
microscope. The video represents the work of Shinya Inoue and Rudolf
Oldenbourg and the paper supporting Figure
9 seen below appeared in Molecular Biology of the Cell
9(7): 1603–07, July 1998. The text of the article may be found at
<http://www.molbiolcell.org/cgi/content/full/9/7/1603>.
The video images are of exceptional quality.
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Figure 9. The figure caption from the article above states: "Mitosis in
tissue-cultured lung cell of a newt, Taricha granulosa, recorded with
the new Pol-Scope." Courtesy of the American Society for Cell
Biology.
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The Kevin Sullivan Lab http://www.scripps.edu/cb/sullivan/movies/ks701.mov http://pingu.salk.edu/~wahl/ks701.mov
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Figure 10. This still image comes from a 3-D reconstruction and rotation where the
centromeres of a living human cell can be seen during interphase. Courtesy of
Kevin Sullivan of Scripps Research Institute.
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Dr. Kevin Sullivan of the Department of Biology at Scripps Research
Institute, working in conjunction with the lab of Geoffrey Wahl at the Salk
Institute, has developed a technique to follow the centromeres during
Interphase. Using GFP probes and centromere binding protein, the centromere
satellite DNA can be visualized as in
Figure 10 and provide insight
into the activities of the cell before the mitotic process begins.
Mitosis World: the Lab of Ted Salmon http://www.bio.unc.edu/faculty/salmon/lab/mitosis/prometspinfixed.mov http://www.bio.unc.edu/faculty/salmon/lab/mitosis/mitosis.html
Dr. Edward D. Salmon of the Department of Biology at the University of
North Carolina at Chapel Hill and his lab work on chromosome movement. They
have created a meta-site for mitosis. The movie section of the site named
Mitosis World has ten excellent movies depicting different aspects of mitotic
division and an image from one of the movies may be seen in
Figure 11.
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Figure 11. A Differential Interference Contrast Microscopy still image was taken from
a 43-min time-lapse movie of newt lung cell mitosis. Courtesy of Ted Salmon of
the University of North Carolina at Chapel Hill.
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PEER INSTRUCTIONAL MATERIALS
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Two student mitosis Web sites were especially interesting and are found
below.
Troy High School http://www.troy.k12.ny.us/thsbiology/skinny/skinnymitosis.html
This site is maintained by Troy High School in Troy, New York. The image
seen in Figure 12 is taken
from an attractive and understandable animation of mitosis. This high school
also has online labs to perform onion root tip preparations.
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Figure 12. The pink chromosomes move with the green spindle apparatus. Courtesy of
Joseph Dibari, a Biology Instructor at Troy High School, Troy, New York.
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Haverford College Biology 300 http://www.haverford.edu/biology/Courses/bio300/BioGallery02/presentations/CrozierJohnson.htm
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Figure 13. Jurkat cells to which DNA has been stained with DAPI. Three stages of
mitosis are seen. Met = metaphase; Ana = anaphase; and Telo = Telophase.
Images courtesy of Dr. Karl Johnson, Department of Biology, Haverford
College.
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Haverford College biology juniors work on special research topics during
one quarter. For their project Kathryn Crozier and Brandon Johnson produced
some immunofluorescence images of cells in mitosis and examples may be seen in
Figure 13.
It is heartening to consider that undergraduate students can perform such
techniques to visualize mitotic events when 120 years ago Flemming,
Schleicher, and Strasburger were just trying to name the stages of
mitosis.
Thank you for taking this tour through Web resources for studying mitosis.
If you want to comment on the selections or suggest future inclusions, please
send a message to
rblyston{at}trinity.edu.
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THE HISTORY OF MITOSIS
TEACHING MITOSIS TODAY