Concept


In sexual reproduction, offspring arise from the union of specialized sex cells — a female egg and a male sperm. Just before the rediscovery of Mendel's work, careful studies were made of chromosome behavior during the formation of sex cells (meiosis). First, homologous (like) chromosomes pair up at the cell equator where they actually exchange genetic information. Then, one chromosome from each pair is pulled toward each pole. At the end of this reduction division, each daughter cell receives only one homologous chromosome from each pair. Meiosis halves the chromosome number and randomly assorts homologous chromosomes into sex cells. The full chromosome number is restored when sperm and egg unite. This exactly mirrored the behavior of genes as deduced by Mendel three decades earlier.

Animation


Hello, I'm Theodor Boveri. In the late 1800's, I looked at chromosome behavior during meiosis. I also looked at how chromosomes affect the development of an organism. My colleagues and I believed that sex cells, produced during the process of meiosis, have a half set of chromosomes. Otherwise, each time a sperm fertilizes an egg, there would be a doubling of chromosome number. I showed that chromosome numbers are indeed reduced in meiosis. There are two rounds of division in meiosis. In meiosis I, chromosome number is reduced by half. Meiosis II is a cell division like mitosis. Let's first look at meiosis I, the reduction round of meiosis. Like in mitosis, a cell not undergoing meiosis is said to be in interphase. One of the things the cell does at this time is duplicate nuclear material. In prophase I, the chromosomes condense out of the nucleus. This cell has 4 chromosomes; each chromosome is visible as two sister chromatids joined at the centromere. Similar chromosomes — homologous chromosomes — pair up. Each member of the pair is called a homolog. In metaphase I, the homologous chromosomes move and line up at the cell equator. Let's zoom in and take a look. In anaphase I, the homologs separate and move to opposite poles of the cell. Telophase I ends the chromosome reduction stage of meiosis. Homologs collect at opposite poles of the mother cell, and the cytoplasm divides to produce two daughter cells. Chromosome number is halved in these daughter cells. In meiosis, these two daughter cells both undergo one more round of division. In meiosis II, the centromeres divide, and the daughter chromatids separate like in mitosis. Interphase II and prophase II are much shorter in meiosis II than in meiosis I. So, a cell with 4 chromosomes at the beginning of meiosis will divide twice to give 4 daughters cells, each with 2 chromosomes. Chromsomes must be important for heredity. Otherwise, it wouldn't matter how many chromosomes a cell gets during mitosis or meiosis. I used sea urchins to prove that the right number of chromosomes is needed for correct development. Sometimes sea urchin eggs can be fertilized by two sperm. When this happens, the first cellular division is uneven. Instead of dividing into two cells, the fertilized egg divides into three cells. Each of these three cells continue to divide, and the divisions all look normal, but these doubly fertilized eggs eventually die. They never become sea urchins. I realized that when two sperm fertilize one egg, an extra half set of chromosomes is introduced. The first division after fertilization is uneven, and each of these three cells receives an incomplete set of chromosomes. These cells then die. Chromosomes are the basis of heredity, and cells need a full set of chromosomes for proper development. I was not the only one who came to these conclusions about chromosomes. In 1902, Walter Sutton, an American student at Columbia University, published a paper on the number and shape of grasshopper chromosomes. In his paper, Sutton lined up the grasshopper chromosomes to show the differences in sizes and shapes. Sutton realized, as I did, that similar chromosomes pair during meiosis and segregation of the homologous pairs reduces chromosome number. Based on our work, Sutton and I established chromosomes as the physical basis of the Mendelian laws of heredity. The segregation of chromosomes during meiosis was what Herr Mendel had predicted for the segregation of factors.

Gallery


Excerpt of 1906 letter from W. Spillman to the American Breeders Association. The letter summarizes "recent" discoveries in genetics. Boxed text talks about meiosis in relation to Mendel's law of segregation.
A picture of a sperm fertilizing an egg.
Visual by www.PDImages.com.
A signed portrait of Theodor Boveri.
Theodor Boveri.

Audio/Video


Audio Glossary

Autosome, Cell, Chromosome, Diploid, Germ line, Haploid, Somatic cells

Video Interviews

Scott F. Gilbert

Scott Gilbert is professor of biology at Swarthmore College where he teaches developmental genetics, embryology, and the history and critiques of biology. He is the author of the college text Developmental Biology, and he works on a number of other projects having to do with developmental biology and the history and philosophy of biology.

Clip 1 (1:16)
Comments on Theodor Boveri and his experiments.

Clip 2 (1:39)
The relationships between Theodor Boveri, Edmund Wilson, Nettie Stevens and Thomas Hunt Morgan.

Biography


 

Theodor Boveri and Walter Sutton both described the process of meiosis. They also showed that although chromosomes may look similar, they have specific hereditary qualities.

THEODOR BOVERI (1862-1915)

Theodor Boveri was born in Bamberg, Germany, the son of a doctor. Boveri was artistically inclined. He initially enrolled to study the humanities, but in 1881, Boveri entered the University of Munich to study anatomy and biology. He graduated with his doctorate - summa cum laude - in 1885, with the thesis, Beiträge zur Kenntnis der Nervenfasern (On the Study of Nerve Fibers). He received a scholarship, and was able to continue doing independent research at the zoology department in Munich.

In the late 1880's and early 1890's, Boveri published some of his most significant findings. Oskar Hertwig had already discovered that sperm and egg nuclei fuse during fertilization. Boveri studied the maturation of egg cells of Ascaris megalocephala, the horse nematode. He saw that as egg cells matured, there comes a point where chromosome numbers are reduced in half. Boveri was one of the first to see evidence of the process of meiosis.

Boveri was also one of the first to do experiments in the field of cytology. He chronicled the development of sea urchin eggs, when one egg was fertilized by two sperm. He concluded that male sperm nuclei and female egg nuclei were equivalent in the amount of hereditary information. They each had a half set (haploid number) of chromosomes. As long as there was a set of both (diploid number of chromosomes), there was fairly normal development of the sea urchin larvae. Any more or any less and there was abnormal development. When Mendel's laws were rediscovered in 1900, Boveri recognized the correlation between Mendel's factors and the cytology work being done on chromosomes.

In 1893, Boveri was appointed Professor of Zoology and Comparative Anatomy at the University of Würzburg. In 1902 and 1908, Nettie Stevens spent time in Boveri's lab, and likely was influenced by his cytology work. Boveri was not overly fond of Stevens, and complained about how she was a "bloodsucker," learning much but contributing little.

Boveri remained at the University of Würzburg until his death in 1915.

WALTER STANBOROUGH SUTTON (1877-1916)

Walter Sutton was born in Kansas City, and graduated from Kansas University. He was E. B. Wilson's graduate student in the Department of Zoology at Columbia University. In the spring of 1902, when he was only 25, Sutton deduced that chromosomes are the basis of heredity, and that the reduction of chromosomes in meiosis is directly related to Mendel's laws of inheritance.

The behavior of chromosomes and its importance in heredity was a "hot topic" at the turn of the century. Many scientists, including Sutton's supervisor, E. B. Wilson, were working on this problem. Theodor Boveri made the connection between chromosomes and heredity by doing his own observations and experiments. Sutton, working independently in Wilson's lab, came to the same conclusions. Wilson admitted later that when Sutton first explained his theory to him, he "did not at once fully comprehend his conception or realize its entire weight."

Sutton did his observations using grasshopper cells. His paper, in 1902, clearly showed that each chromosome is different, and meiosis reduces chromosome number in the gametes. Sutton's 1903 paper, The Chromosomes in Heredity, summarized and discussed the importance of his conclusions. The paper even more strongly drew the connection between Mendel's laws of heredity and chromosomes.

Wilson was very impressed with Sutton's abilities as an investigator. Unfortunately, Sutton never finished his doctorate. Sutton left research and entered medical school. He graduated from the College of Physicians and Surgeons at New York, and became a surgeon. Sutton served in France during World War I, and distinguished himself in treatments of wounded soldiers. Sutton died following an operation for appendicitis. He was only 39.

Factoid

Links


 

Links

The Biology Project at the University of Arizona

This site is a good resource in all areas of biology. It has a tutorial on meiosis.

UrchiNet Database

This site is an online database of information about sea urchins. It has a simulator that models early development of sea urchin in 3-D.

Bibliography

  • Moore, John, A., 1985, Science as a Way of Knowing, American Society of Zoologists.

  • Portugal, Franklin, H., and Cohen, Jack, S., 1977, A Century of DNA: A History of the Structure and Function of the Genetic Substance, The Massachusetts Institute of Technology, U. S. A.

  • Stubbe, Hans, 1972 (English Translation), History of Genetics, The Massachusetts Institute of Technology, Cambridge, Massachusetts.

  • Sturtevant, A. H., 1965, A History of Genetics, Harper & Row, Publishers, New York.

Glossary


Autosome - Any chromosome other than a sex chromosome. Humans have 22 pairs of autosomes.
Cell - The basic unit of any living organism. It is a small, watery, compartment filled with chemicals and a complete copy of the organism's genome.
Chromosome - One of the threadlike "packages" of genes and other DNA in the nucleus of a cell. Different kinds of organisms have different numbers of chromosomes. Humans have 23 pairs of chromosomes, 46 in all: 44 autosomes and two sex chromosomes. Each parent contributes one chromosome to each pair, so children get half of their chromosomes from their mothers and half from their fathers.
Diploid - Refers to a cell that has two sets of chromosomes. In humans, almost all the cells, except the gametes are diploid. Because there are 23 chromosomes in a single human set, most human cells contain 46 chromosomes.
Germ line - A sequence of cells, each descended from earlier cells in the lineage, which will develop into new sperm and egg cells for the subsequent generation.
Haploid - The number of chromosomes in a sperm or egg cell, half the diploid number.
Somatic cells - All body cells, except the reproductive cells.

Children resemble their parents.
Genes come in pairs.
Genes don't blend.
Some genes are dominant.
Genetic inheritance follows rules.
Genes are real things.
All cells arise from pre-existing cells.
Specialized chromosomes determine gender.
Chromosomes carry genes.
Genes get shuffled when chromosomes exchange pieces.
Evolution begins with the inheritance of gene variation.
Mendelian laws apply to human beings.
Mendelian genetics cannot fully explain human health and behavior.
DNA and proteins are the molecules of the cell nucleus.
One gene makes one protein.
A gene is made of DNA.
Bacteria and viruses have DNA too.
The DNA molecule is shaped like a twisted ladder.
A half DNA ladder is a template for copying the whole.
RNA is an intermediary between DNA and protein.
DNA words are three letters long.
A gene is a discrete sequence of DNA nucleotides.
The RNA message is sometimes edited.
Some viruses store genetic information in RNA.
RNA was the first genetic molecule.
Mutations are changes in genetic information.
Some types of mutations are automatically repaired.
A chromosome is a package for DNA.
Higher cells incorporate an ancient chromosome.
Some DNA does not encode protein.
Some DNA can jump.
Genes can be turned on and off.
Genes can be moved between species.
DNA responds to signals from outside the cell.
Different genes are active in different kinds of cells.
Master genes control basic body plans.
Development balances cell growth and death.
A genome is an entire set of genes.
Living things share common genes.
DNA is only the starting point for understanding human biology.
adi_at_dnaftb