Chapter 4: The Chromosome Theory of Inheritance
When available, the links under the Suggested Readings section will take you to websites
that either provide free electronic full-text versions of these references, or links to online
abstracts. All of these abstracts are themselves free. Many of the abstract sites contain
links to online full-text versions of the references; some of these full-text versions are
free, while others require personal or institutional subscriptions. None of the sites that can
be accessed through the links below are affiliated with McGraw-Hill Publishers.
More on the History of Chromosome Inheritance
Suggested Readings
 Bridges, Calvin. 1916. Non-disjunction as proof of the chromosomal theory of
inheritance. Part I. Genetics 1: 1-52 [full text link]
Bridges was a researcher in the “Fly Lab” of Thomas Hunt Morgan at the
beginning of the 20th century. This paper describes some of his initial work on
sex-linked inheritance and how it supports the chromosomal theory of inheritance.
 Bridges, Calvin. 1916. Non-disjunction as proof of the chromosomal theory of
inheritance. Part II. Genetics 1: 107-163 [full-text link]
The second part of the manuscript by Bridges that describes his work on sexlinked inheritance and how it supports the chromosomal theory of inheritance.
 Morgan, T.H. 1910. Chromosomes and heredity. The American Naturalist 44:
449-496. [full-text link]
This paper describes how chromosomes may be the factor associated with the
particulate theory of inheritance. Proof of this theory was provided by Calvin
Bridges in 1916 (see links above).
 Morgan, T.H, 1910. Sex-limited inheritance in Drosophila. Science 32: 120-122.
[full-text link].
The discovery of a white-eyed mutant in the Drosophila stocks in Morgan’s lab
opened up the opportunity to how sex-linked inheritance differed from predicted
Mendelian ratios. This paper describes some of the initial crosses conducted by
Morgan that led to proof of the particulate and chromosome theories of
inheritance.
 Sutton, W.S. 1902. On the morphology of the chromosome group in Brachystola
magna. Biological Bulletin 4: 24-39 [full-text link]
This paper describes Sutton’s work on the grasshopper, Brachystola magna . In
this study Sutton confirms the Mendelian principle that chromosomes exist in
pairs and that meiosis serves to separate the chromosome pairs.
 Sutton, W.S. 1903. Chromosomes in heredity. Biological Bulletin 4: 231-251.
[full-text link].
In this follow-up to his 1902 publication, Sutton discusses how his research on the
grasshopper Brachystola magna supports Mendel’s work.
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Websites
 Biography of Thomas Hunt Morgan, Nobelprize.org
http://nobelprize.org/medicine/laureates/1933/morgan-bio.html
This site provides a brief biography of Morgan’s life and his pivotal research in
Drosophila genetics at the start of the 20th century. Links on the sit provide access
to additional resources on Morgan as well as a transcript of his Nobel Prize
acceptance speech.

The Mechanism of Heredity
http://www.cshl.edu/
This site provides a narrative history of the work being done at the beginning of
the 20th century that led to the proof of the chromosome theory of inheritance.
Photos of Thomas Hunt Morgan, Calvin Bridges and Alfred Sturtevant are
provided.
Mechanisms of Sex-determination in Various Organisms
Suggested Readings
 Cline T.W. 1993. The Drosophila sex-determination system: how do flies count to
two? Trends in Genetics 9(11): 385-390. [Entrez-PubMed link]
This article describes the genetic mechanism of sex-determination in Drosophila.
Specifically it examines some of role of genes in determining the X:A ratio.
 Marx, J. 1995. Tracing How the Sexes Develop. Science 269(5232):1822-1824.
[Entrez-PubMed link]
This article examines how sex-determination occurs in the nematode C. elegans
and D. melanogaster.
 Bull, J.J. 1980. Sex-determination in reptiles. Quarterly Review of Biology 55(1):
3-20. [JSTOR link]
This paper represents an early review of the major sex-determination mechanisms
found in reptiles.
 Manolakou, P., G. Lavranos and R. Angelopoulou. 2006. Molecular Patterns of
Sex Determination in the Animal Kingdom: A Comparative Study of the Biology
of Reproduction. Reprod. Biol. Endocrinol. 4(1):59. [Entrez-PubMed link]
This recent manuscript provides a review of what is known about the molecular
mechanisms of the major sex-determination mechanisms in animals. Discussions
include the ZZ-ZW system, X:A ratio system, and the XX:XY system.
Recent Research into the Biochemical Mechanisms Underlying Mitosis and Meiosis
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Suggested Readings
 Crittenden, SL, CR Eckmann, L. Wang, DS Bernstein, M. Wickens and J.
Kimble. 2003. Regulation of Mitosis/Meiosis in the Caenorhabditis elegans
germline. Philos Trans R Soc Lond Biol Sci. 358(1436):1359-1362. [EntrezPubMed link]
This paper examines how RNA-binding proteins help determine whether a cell
undergoes mitosis or meiosis. It focuses on the use of C. elegans and D.
melanogaster as model organisms for the study of mitosis/meiosis regulation.

Bernstein, E. and SB Hake. 2006. The Nucleosome: A Little Variation Goes a
Long Way. Biochem Cell Biol. 84(4): 505-517 [Entrez-PubMed link]
This paper presents evidence that the histones of the nucleosome may be involved
in the regulation of how chromosomes segregate during meiosis and mitosis.
Websites
 The Cell Cycle & Mitosis Tutorial, The Biology Project
http://www.biology.arizona.edu/cell_bio/tutorials/cell_cycle/main.html
This site provides an overview of the cell cycle and mitosis. Animations are
provided, as well as a series of problems to test your comprehension of the
material.

Meiosis Tutorial, The Biology Project
http://www.biology.arizona.edu/cell_bio/tutorials/meiosis/main.html
This site provides an overview of meiosis. Animations are provided, as well as a
series of problems to test your comprehension of the material.
Further Examples of Sex-linked Inheritance in Humans
Suggested Readings
 Okulicz, JF and RA Schwartz. Hereditary and Acquired Icthyosis vulgaris. Int J.
Dermatology. 42(2): 95-98. [Entrez-PubMed link]
This article examines what is known of icthyosis vulgaris, a disease of the skin
that causes the epidermis to form scale-like structures.
 Tuchman M., N. Jaleel, H. Morizono, L. Sheehy and MG Lynch. 2002. Mutations
and Polymorphisms in the Human Orinithine Transcarbamylase Gene. Hum
Mutat. 19(2): 93-107. [Entrez-PubMed link]
Orninthine decarbamylase deficiency is a disease that results in the accumulation
of ammonia in the blood, frequently resulting in mental retardation. The gene for
this enzyme is located on the X chromosome. This paper examines what is known
about the numerous mutations (244 to date) that are associated with this gene.
 Kriaucionis S and A. Bird. 2003. DNA Methylation and Rett Syndrome. Hum
Mol Genet. 12(2):R221-227. [Entreez-Pubmed link]
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Rett syndrome is an X-linked dominant condition characterized by
neurodegeneration. Using the mouse as a model, this paper examines the
relationship mutations and DNA methylation of the MeCP2 gene.
Websites
 National Hemophilia Foundation
http://www.hemophilia.org
This organization is dedicated to the study of all forms of blood-clotting diseases,
including those associated with the sex-linked disease hemophilia.

International Rett Syndrome
http://www.rettsyndrome.org/
The Medical Reference link on this site contains useful information on how
individuals can get tested for the disease and how to interpret genetic testing
results.

Severe Combined Immune Deficiency (SCID)
http://www.scid.net/
This site provides information on SCID, a disease characterized by a lack of
immune system cells. The research tab at the top of the page provides links to
scientific journals that contain more information on this disease.
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