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Reproduction
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Reproduction
and Cell Division
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Reproduction
and Cell Division
A. Malcolm Campbell, PhD

Christopher J. Paradise, PhD
Reproduction and Cell Division

Copyright © A. Malcolm Campbell and Christopher J. Paradise. 2016.
All rights reserved. No part of this publication may be reproduced, stored

in a retrieval system, or transmitted in any form or by any means—
electronic, mechanical, photocopy, recording, or any other except for
brief quotations, not to exceed 250 words, without the prior permission
of the publisher.
First published in 2016 by

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Abstract
Why do some children look more like one parent than another? How
can two parents with dark hair have a child with red hair? How can
two dark-skinned parents have a baby that has light skin? Everyone has
wondered these questions, but in order to understand such unexpected
outcomes, an understanding of what Gregor Mendel discovered—the
rules of genetics—is necessary. This book reproduces Mendel’s original
data that Mendel used to discover how traits are passed from one genera-
tion to the next. In addition to the rules governing DNA inheritance,
this book also examines how cells reproduce—all cells. Do bacterial cells
reproduce the same way animal cells do? And when a person has a cut that
needs to heal, do those cells reproduce the same way that sperm and egg
cells are produced? How do all these cells keep track of how much DNA is
needed in order to function properly? Data will be examined that explains
how reproduction works for every cell on the planet.
Keywords
haploid, gametes, diploid, genotypes, phenotypes, meiosis, dominant traits,
homozygous, heterozygous, central dogma, homologous chromosomes,
progeny, alleles, Punnett square, dihybrid cross, law of independent as-
sortment, law of segregation, random, centromere, chromatid, mitosis,
spindle fibers, microtubules, prophase, metaphase, anaphase, telophase,
interphase, cytokenesis, tetraploid
Contents
Preface...................................................................................................ix
Acknowledgments....................................................................................xi
Introduction.........................................................................................xiii
Chapter 1 Traits can Disappear and Reappear in a Later
Generation.........................................................................1
Mendel Discovers Inheritance Ratios.................................6
Mendel Discovers Variation from Ratios............................7
Mendel Finds Repeating Pattern of Inheritance..................7
Meiosis Contributes to Variation......................................10
Tool for Calculating Probabilities.....................................11
Chapter 2 Inheritance Patterns for Two Traits Simultaneously..........15
Mendel Discovers Two-Trait Ratios..................................15
Mendel Establishes Genetic Rules....................................19
Ethical, Legal, Social Implications:
The Scientific Meaning of “Random”............................21
Chapter 3 Prokaryotes Reproduce by Binary Fission.........................25
Chapter 4 Eukaryotes Reproduce Through Mitosis and
Cytokinesis......................................................................31
Chapter 5 Gametes are Formed After Meiosis...................................37
Meiosis I..........................................................................38
Meiosis II.........................................................................39
Ethical, Legal, Social Implications: The Ethics
of Genetic Engineering Humans...................................41
Conclusion............................................................................................45
Glossary................................................................................................47
Index....................................................................................................49
Preface
This book covers cell division and Mendelian genetics and is part of a
thirty book series that collectively surveys all of the major themes in
biology. Rather than just present information as a collection of facts, the
reader is treated more like a scientist, which means the data behind the
major themes are presented. Reading any of the thirty books by Campbell
and Paradise provides readers with biological context and comprehensive
perspective so that readers can learn important information from a single
book with the potential to see how the major themes span all size scales:
molecular, cellular, organismal, population and ecologic systems. The
major themes of biology encapsulate the entire discipline: information,
evolution, cells, homeostasis and emergent properties.
In the twentieth century, biology was taught with a heavy emphasis
on long lists of terms and many specific details. All of these details were
presented in a way that obscured a more comprehensive understanding.
In this book, readers will learn how genetically encoded traits are passed
on to the next generation as well as how cells replicate. With this informa-
tion, readers will also see for themselves some of the supporting evidence
behind our understanding of these important topics. The historic and
more recent experiments and data will be explored. Instead of believing
or simply accepting information, readers of this book will learn about the
science behind genetics and cellular reproduction the way professional
scientists do—with experimentation and data analysis. In short, data are
put back into the teaching of biological sciences.
Readers of this book who wish to see the textbook version of
this content can go to www.bio.davidson.edu/icb where they will find
pedagogically-designed and interactive Integrating Concepts in Biology for
introductory biology college courses or a high school AP Biology course.
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Acknowledgments
Publishing this book would not have been possible without the generous
gift of Dr. David Botstein who shared some of his Breakthrough Prize
with AMC. David’s gift allowed us to hire talented artists (Tom Webster
and his staff at Lineworks, Inc.) and copyeditor Laura Loveall. Thanks go
to Kristen Mandava for project management and guidance on the pub-
lishing process. In particular, we are indebted to Katie Noble and Melissa
Hayban for their many hours of help and attention to detail.
Kristen Eshleman, Paul Brantley, Bill Hatfield and Olivia Booker
helped us with technology at Davidson College. We are grateful to ad-
ministrators Tom Ross, Clark Ross, Carol Quillen, Wendy Raymond,
Verna Case, and Barbara Lom who had confidence in us and encouraged
us to persist despite setbacks along the way.
These books were the product of the shared labor of my two vision-
ary coauthors Laurie Heyer and Chris Paradise. We shared the dream
and the hardships and developed this book from scratch. My family has
been very supportive and I thank Susan, Celeste and Paulina for their
support and patience. I also want to thank Jan Serie, my pedagogical
mentor, who taught me so much about the art and science of helping stu-
dents learn. I benefited from the support of the Howard Hughes Medi-
cal Institute grant 52006292, the James G. Martin Genomics Program,
and Davidson College. This book would not have survived its first draft
without my students who endured the typos and the early versions of this
book. These undergraduates participated in a bold experiment to see if
beginners could construct their own knowledge, retain what they learned,
and transform the way they see themselves and the discipline of biology.
While many people said that beginning students were not up to the task,
my students proved them wrong.
Introduction
It is well known that DNA is the heritable material, but many people
have unanswered questions about how heredity works. For example, if
half of a child’s DNA is from its mother and half is from its father, why
don’t all siblings have exactly the same DNA and look like identical twins?
Why are some traits and diseases passed from one generation to the next,
but others are not? How can an inherited trait skip generations?
The key to understanding heredity is to understand how cells divide
and pass DNA information to the newly produced cells. Gregor Mendel,
a 19th century amateur geneticist, was able to explain many characteris-
tics of heredity that had mystified the professional biologists of his time.
In this book, the reader will learn how to predict patterns of inheritance
and how organisms pass their genetic information to future generations.
Cell division processes are different between eukaryotes, prokaryotes (bac-
teria) and, haploid gametes (eggs and sperm). Adults produce haploid
gametes, with half the diploid-amount of DNA, in a slightly different
way than they produce diploid cells. Yet all these cell division processes
use similar proteins, as would be expected given their common evolution-
ary history. The five chapters of this book focus on information at the
organismal level. There are two Ethical, Legal, Social Implication chapters
too that consider more than just the science of heredity.
CHAPTER 1
Traits can Disappear

and Reappear in a Later
Generation
For hundreds of years, people bred chickens, dogs, tomatoes, and roses to
maximize desired traits, but no one really understood the reason for the
patterns of inheritance that made these results possible. Then in 1865, an
Austrian monk named Gregor Mendel, who liked to grow vegetables in
his garden, started the field of genetics when he published a radical paper
that explained what had appeared to others as incomprehensible varia-
tion. Genetics is the study of how genotypes and phenotypes are propa-
gated through sexual reproduction. An English translation of Mendel’s
paper is available today at MendelWeb.org—all 41 pages of it.
Mendel studied many different plants and conducted thousands of
genetic experiments. His biggest insight was counting what he saw and
applying mathematical rigor and analysis to his biological data. Using
math to analyze biology is a prime example of the way genius sometimes
comes from having a new perspective rather than from knowing more
information.
Any time a biologist begins a new line of investigation, he or she must
select an appropriate organism for the questions under consideration.
Mendel was very careful about choosing which plant to study. He chose
the pea plant, because he had years of experience with it in his garden
and he knew that he could obtain thirty-four varieties of this one species.
Another key factor in his decision was the anatomy of the pea plant. Pea
flower pollination is less subject to chance because all the reproductive
parts of the plant are enclosed within the same flower. Pea flowers tend to
2 REPRODUCTION AND CELL DIVISION
Figure 1 Pea plant life cycle and parts of a seed. A, Diagram of pea
life cycle including self-fertilization. B, Like peas, peanuts contain
paired embryonic leaves and a root inside.
Source: A. Original art. B. Photo courtesy of A. Malcolm Campbell.
self-fertilize rather than being fertilized by other pea flowers. Peas offered
a very controlled experimental system for genetic experiments.
In the 1860s, a common definition for a species was that members of
the same species displayed “precisely similar characteristics.” It was not
clear to Mendel how many species he had with his thirty-four varieties of
pea plants, but as long as they could produce fertile offspring, he could
conduct his research. He worked for 2 years just to make sure that the
traits he observed were stable over many generations; 22 of the 34 varieties
he bred proved to be stable. When a line of individuals and their offspring
produce the same version of a trait, that line of individuals is described as
true-breeding. Mendel focused on 15 of the 22 true-breeding traits. Of
these 15, he chose 7 traits to study in-depth over the next 6 years.
Like most flowering plants, peas have a consistent life cycle (Figure 1).
Flowers produce gametes of pollen and eggs via meiosis. (Chapter 5 will
cover meiosis in detail.) The male reproductive organ, called the stamen,
produces pollen which is moved inside the flower to the stigma on top
Traits can Disappear and Reappear in a Later Generation 3
first generation peas (P) first generation peas (P)
A second generation peas (F1) B second generation peas (F1)
Figure 2 Mendel’s first breeding results. A, Pollen from true-breeding

green (dark gray) pea plants and eggs from true-breeding yellow pea
(light gray) plants produced only yellow seeds. B, The reciprocal cross
also produced only yellow seeds.
Source: Original art.
of the elongated style, two parts of the female reproductive organ. When
pollen lands on the stigma, the pollen begins to grow a long tube that will
carry the haploid nucleus toward the egg, leading to fertilization of the
egg. The resulting seed, in this case a pea, can germinate and produce a
new plant, and the life cycle repeats itself again. Peas contain embryonic
plants and include a pair of simple leaves called cotyledons that emerge
immediately after germination, followed by roots and a shoot with com-
plex leaves (see Figure 1). When eating a peanut, it is possible to split it
in half and see the embryonic plant parts, which are similar in structure
to pea seeds.
At the time Mendel was conducting his amateur scientific research,
two prominent biologists had already published a study of how traits were
inherited, but their ideas did not agree with Mendel’s observations. They
stated that when true-breeding green pea plants were bred (crossed) with
true-breeding yellow pea plants, the offspring seeds (the peas) either were
intermediate colors of yellowish-green or looked like one parent and not
the other. Mendel was familiar with these published opinions, but he had
made different observations while working for more than two years to
obtain true-breeding plants. In his studies, Mendel observed that peas
could be either yellow or green, but he never saw any yellowish-green
peas (Figure 2). Similarly, Mendel had true-breeding plants that produced
smooth peas and others that produced only wrinkled peas, but his crosses
never produced the intermediate phenotype of mildly wrinkled peas.
Data from Mendel’s published work is available to analyze from the
figures in this book. For example, when Mendel crossed a true-breeding
yellow pea plant with a plant that produced only green peas (parental
generation = P), all of the peas (second generation = F1) were yellow,
regardless of whether the pollen came from a yellow pea plant or a green
pea plant. As Mendel discovered, dominant traits are apparent when
mixed with the DNA encoding a recessive trait. Based on the data in
Figure 2, it is possible to conclude that yellow is dominant to green pea
color. Therefore, green completely disappears in the second-generation
(F1) peas. Mendel knew nothing about DNA, but he understood that
each parent contributed something in equal proportions to the next gen-
eration. He deduced the main points of meiosis and gamete formation,
even though no one in the 1860s knew that the heritable material was
DNA. The processes of mitosis and meiosis were still undiscovered when
Mendel conducted his genetic experiments.
As Mendel began to understand inheritance patterns, his observation
that one trait was dominant to another trait contradicted the popular
understanding of breeding. However, he was certain which of his plants
had contributed heritable material to the next generation for two reasons.
First, he carefully removed the stamen and its pollen from the recipient
plant prior to the selective pollination he performed manually. Second,
the anatomy of the flowers effectively blocked the wind-driven entry of
pollen from the other pea plants.
Mendel’s next major insight came from allowing the second genera-
tion plants (F1) to self-pollinate. Because he needed to keep track of gen-
erations, Mendel invented a system for naming the generations that is
still used today. He called the original parental plants the P generation,
and the first offspring generation the F1 generation; F is for filial, which
is derived from the Latin word for child. Each subsequent generation
was numbered accordingly. Mendel planted the F1 peas from Figure 2
and allowed them to mature and produce F2 peas from self-pollination
(Figure 3). Contrary to the prediction of his peers, completely green peas
F1 peas (second generation)
F2 peas (third generation)
Figure 3 Mendel’s second experiments started with yellow F1 peas

from Figure 2. Plants derived from yellow F1 peas grew into plants
that were self-crossed, and the F2 peas were quantified for color.
Table 1 Results from mating true-breeding plants that produced

yellow or green peas.
generation green peas yellow peas
P 5 true-breeding green plants 5 true-breeding yellow plants
F1 0 green peas 273 yellow peas
F1 0 plants from green peas self-cross 258 plants from F1 yellow peas
F2 2,001 green peas 6,022 yellow peas
Source: Data from Mendel.
reappeared in the F2 generation after the mating of two true-breeding

plants in the P generation. The green that had disappeared in the F1 gen-
eration reappeared in the F2 generation, so Mendel called the green trait
recessive and the yellow trait dominant.
Mendel had another insight—gather lots of data to see the overall
trends (Table 1). As is apparent from the data, Mendel allowed 258 F1
plants from yellow peas to self-fertilize, and these plants produced a total
of 8,023 F2 peas. Of the F2 peas, 6,022 were yellow and 2,001 were green.
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Mendel Discovers Inheritance Ratios

Even though he conducted his research as a hobby, Mendel was an out-
standing scientist. Good scientists are often their own harshest critics,
and Mendel was no exception. He wondered whether the results that he
had observed would hold true for traits other than pea color. Although
he thoroughly tested all seven traits, this book considers only one more,
the wrinkled pea (Table 2).
Mendel started with two, true-breeding varieties of peas, crossed them,
and collected their F1 peas. The F1 seeds germinated into mature plants,
which Mendel allowed to self-pollinate. Mendel collected and counted these
F2 peas to determine how many were smooth and how many wrinkled. He
kept track of each generation of peas from each plant and measured them
individually (Table 3). Mendel analyzed the patterns of inheritance for ten
Table 2 Results from mating true-breeding plants that produced

smooth or wrinkled peas.
generation wrinkled peas smooth peas
P 5 true-breeding wrinkled plants 5 true-breeding smooth plants
F1 0 wrinkled peas 281 smooth peas
F1 0 plants from wrinkled peas self-cross 253 plants from F1 smooth peas
F2 1,850 wrinkled peas 5,474 smooth peas
Table 3 Variation in F2 pea phenotypes from individual matings for

two separate traits.
plant smooth wrinkled plant yellow green
number pea pea number pea pea
1 45 12 1 25 11
2 27 8 2 32 7
3 24 7 3 14 5
4 19 10 4 70 27
5 32 11 5 24 13
6 26 6 6 20 6
7 88 24 7 32 13
8 22 10 8 44 9
9 28 6 9 50 14
10 25 7 10 44 18
totals 336 101 totals 355 123
P-generation plants from two separate experiments and all their progeny.
He collected large enough numbers of F2 peas to detect trends that no one
else had noticed before. Mendel discovered that the dominant trait ap-
peared about three times more often than the recessive trait.
Mendel Discovers Variation from Ratios

Always an observant scientist, Mendel also noticed a subtlety in the data
in Table 3: Inheritance patterns were not consistent from plant to plant.
For example, the pea colors from mating #5 were in a ratio of about
2 yellow to 1 green, but mating #8 produced nearly 5 yellow to 1 green.
Mendel correctly deduced that plant-to-plant variation in phenotype
ratio was an indication of chance. Mendel’s observation was analogous
to flipping two coins one hundred times, and noticing that the number
of double heads varies from 17 to 33 out of 100. The long run average
number of double heads is 25 out of 100, but individual trials varied from
the average. In modern terms, color depended upon which pollen gamete
happened to fertilize a particular egg. His contemporaries had looked at
only one or two plants and could not detect the overall pattern. Further-
more, Mendel also detected variations within the pattern and was astute
enough to understand the role of chance and its biological consequences.
Whenever new information is discovered, there is a need for new
terms so the discovery can be discussed. Since Mendel was the first geneti-
cist, he needed terms and short-hand abbreviations to communicate his
research to others. He needed a way to describe the heritable material for
his peas depending if they were true-breeding or not. Mendel used capital
letters to represent dominance and lowercase letters to represent recessive
traits. The letter he chose for each trait was based on the characteristic he
first noticed. Mendel described his true-breeding yellow peas as YY, or
homozygous dominant. The peas that had yellow phenotype but were
not true-breeding were called heterozygous, and their genotype was Yy.
Mendel Finds Repeating Pattern of Inheritance

Mendel was curious about the F2 green and yellow peas that he had
produced (see Figure 3), and he carefully analyzed his results. All of the
all F1 peas =
1 3
4 4
F2 peas
2
3
1
100%
3 1 3
4 4
F3 peas
100% 100% 100% yellow

green yellow green
Figure 4 Mendel’s F3 results from F2 pea plants. For illustration

purposes, 24 peas were produced in each generation. Results from
self-pollinating F2 plants are shown as F3 peas.
plants that developed from F2 green peas produced only green F3 peas
(Figure 4). The F2 green peas were homozygous (yy) and were 1/4 of the
total F2 peas. The remaining 3/4 of the peas were yellow, and when 519
of the yellow F2 peas were allowed to grow and self-pollinate, about 1/3
of the 519 yellow F2 plants (homozygous YY) produced only yellow F3
peas. The remaining 2/3 of the 519 yellow F2 peas were heterozygous
and produced a mixture of yellow and green F3 peas in the same ratio
as the original F1 peas—that is, approximately 3 yellow peas for every
1 green pea. Mendel had finally discovered the repeating pattern of in-
heritance that had eluded all the so-called experts. With each subsequent
generation, half the peas returned to their true-breeding status (1/4 YY +
1/4 yy) of the P generation, while the other half were heterozygotes (Yy)
and each of these produced plants that continued to generate 3 yellow
peas for every 1 green pea, on average. As was seen in Table 3, any particu-
lar plant crossing might deviate from these precise ratios, but the overall
average was maintained for many generations.
Although Mendel did not have the knowledge about DNA as the
heritable material and how proteins are produced after transcription of
RNA, it is helpful to connect current understanding of central dogma
y y yy yy yy yy
25%
yy
Y Y YY YY YY YY
25%
YY
y Y YY Yy yY yy
50%
yY
Figure 5 Combining Mendel’s observations with DNA information.

From left to right: F2 seeds and their genotypes shown as homologous
chromosomes with colored alleles. The resulting F2 plants and the
F3 progeny from F2 self-pollination. F3 heterozygotes are shaded half
yellow and half green for illustration purposes, but the peas appear
fully yellow.
with his observations from the 1860s to see how genotype and pheno-
type are related (Figure 5). Mendel found that averaging data from a
large sample size revealed patterns by smoothing out the variability due
to chance. Figure 5 makes another simplification and follows only one
pair of chromosomes from the pea’s genome. This chapter will track only
the gene on a single chromosome that encodes for pea color. Peas have
seven pairs of chromosomes, and Mendel followed seven traits. As it turns
out, each of the seven traits Mendel followed was encoded on a different
chromosome.
Mendel had discovered a fundamental genetic principle, but he
wanted to test his conclusions, as any good scientist should. He expected
all seven traits to be inherited in the same way as pea color. The results of
his next round of experiments validated his previous conclusion, as shown
in Table 2 and Table 3. The smooth pea trait is dominant to wrinkled
peas, and the F2 ratio was about 3:1. Mendel detected the 3:1 pattern by
analyzing overall averages, even though individual plants deviated from

the average. The plant-to-plant variation must have confounded Mendel’s
contemporaries and led them to make incorrect conclusions about inheri-
tance patterns, such as the blending of colors to a yellowish-green pea that
did not really exist. Mendel’s contemporaries thought of pea color like
paint, something that could be mixed and blended to produce new col-
ors. Mendel’s breakthrough was discovering that pea color is determined
by bits of information that do not blend. Overall, F2 peas produced 25%
true-breeding recessive peas, 25% true-breeding dominant peas, and 50%
heterozygous peas. Mendel had uncovered a critical aspect of genetics,
even though he had never seen a chromosome and knew nothing about
the role of DNA in reproduction.
As diploid plants, all peas carry two alleles for pea color. Both alleles
are identical in true-breeding peas—that is, they are homozygous for the
color gene. Half of all the F2 progeny are true-breeding, even though it
is not possible to tell which yellow peas were true-breeding until their
offspring in Figure 4 were examined. It is possible to say with certainty
that the green peas have a genotype of yy, because they are homozygous
recessive.
Meiosis Contributes to Variation

The variation that Mendel documented in Table 3 can be explained by
what is known about gene the production of proteins via what is called
central dogma. Sexual reproduction will be covered in Chapter 5 of this
book. First, it is important to understand why the genotype of a pea results
from chance events (Figure 6). Plants produced pollen and egg gametes,
and each gamete contains only one chromosome of the homologous pair.
The plant in Figure 3.6 produced pollen and eggs, half with green alleles
(y) and half yellow alleles (Y). The fertilization of an egg by one pollen is
the consequence of a random event—which pollen grain happened to land
on the stigma and grow a pollen tube to fertilize a particular egg. Predict-
ing the outcome of any single mating is analogous to predicting the out-
come of flipping two coins, one for the pollen grain and one for the egg.
The genotype of the haploid pollen grain can be either Y or y, just as a coin
can be heads or tails. Similarly, the genotype of the egg can be either Y or y.
Figure 6 Chance events cause variations from the calculated ratio

of 3:1. Whether a particular pollen grain will land on the stigma is
random, as is the particular egg to be fertilized. Parental heterozygous
peas are shaded half yellow and half green for illustration purposes
but would appear fully yellow.
Chance events such as flipping coins or combining gametes are de-

scribed by probabilities, not certainties. For example, the probability of
flipping heads with two coins is 1/2 × 1/2 = 1/4. Likewise, the prob-
ability of producing a YY pea (Y allele from pollen and Y from egg) is 1/4.
Mendel’s understanding of probability was critical to his deciphering the
pattern of inheritance.
Tool for Calculating Probabilities

Mendel used symbols and mathematics to represent a complex pro-
cess that he did not understand at the molecular level. Based on
pollen Yy × Yy
genotypes maternal gamete genotypes
Y y
Y YY Yy
y Yy yy
Figure 7 A Punnett square. Genotype and phenotype outcomes for

mating two heterozygotes. Gamete genotypes are placed outside the
boxes, and pea genotypes are placed inside the boxes. Phenotypes are
represented by the yellow (light gray) or green (dark gray) colored
squares.
modern knowledge, it is possible to understand the mechanism shown in

Figure 7. On average, every time a heterozygote is self-pollinated, it pro-
duces 25% homozygous dominant peas, 50% heterozygous peas, and
25% homozygous recessive peas. A visual tool called a Punnett square
in Figure 7 is helpful for calculating ratios of progeny based on gamete
genotypes. For simplicity, it is easier to assume that each cross produces
four peas, although four is just a number of convenience. A Punnett
square lists all of the possible gamete genotypes on the outside of the
square (top and left), then combining all the columns and rows to
produce the progeny genotypes inside the square. It is easy to deduce
phenotypes from the genotypes. The ratio 1 YY:2 Yy:1 yy was one of the
famous ratios that Mendel published (Table 4). Using this ratio, Mendel
projected n generations into the future, which was easy because he made
a simplifying assumption that every plant produces only 4 peas and every
generation maintains the exact 1:2:1 ratio. Mendel reasoned that if he
started with a 1:2:1 ratio, the next generation will result in a ratio of
6:4:6, which simplifies to 3:2:3. Let’s examine this logic and use Table 4
to predict the future. Arrows connecting the first two generations show
how each number was deduced.
Look at the second generation data in Table 4. Each first generation
homozygous pea will produce only homozygous second generation peas.
Each Yy pea will produce one YY and one yy pea for a total of six YY
peas and six yy peas in the second generation. Four heterozygotes will be
present in the second generation, because each of the two heterozygous
Table 4 Ratio of pea genotypes for n future generations.

generations number of genotypes ratios
YY Yy yy YY Yy yy
Y 2 ×
1 1 × 1Y 2 1 yy 1 1 2 1
4 YY 2 4 yy
2 6 4 6 3 2 3
(2 × 2 Yy)
3 28 8 28 7 2 7
4 120 16 120 15 2 15
5 496 32 496 31 2 31
n ... ... ... 2n – 1 2 2n – 1
peas from the previous generation will produce two new heterozygotes;
2 × 2 = 4. Mendel repeated this pattern over and over and deduced a
formula to calculate the outcome for any generation, which is presented
in the bottom row of the ratio columns.
Generation #2 produced a YY ratio of 22 − 1 = 3, and the same
formula describes the yy ratio in the second generation. Note that the
heterozygote component of the ratio is a constant, which helps explain
why Mendel’s contemporaries thought heterozygotes “are inclined to re-
vert to the parental forms,” reconfirming their mistaken understanding
of inheritance patterns. Using Mendel’s formula, it is easily to predict the
ratio of peas ten generations into the future without having to write down
the outcomes of all the intermediate generations.
So far, this chapter has shown how eukaryotes transmit one trait at a
time. Like any good scientist, Mendel wondered if the patterns that he
had discovered scaled to more than one trait at a time. Chpater 2 examines
Mendel’s data when he followed pea color and texture at the same time.
Bibliography
Bateson W: Experiments in plant hybridization (1865) in translation form
Gregor Mendel’s orignal work: Versuche über Plflanzenhybriden,
Verhandlungen des natureforschenden Vereines in Brünn, Bd. IV für
das Jahr 1865. Abhandlungen, 3–47, 1901. Minor corrections by

Roger Blumberg.
Blumberg RB. Mendel’s paper in English: experiments in plant hybri
dization (1865) by Gregor Mendel, MendelWeb (website): http://
www.mendelweb.org/Mendel.html. Accessed April 1, 2014.
Judson HF, Hodgkin J, Relichova J. Mendel’s experiments and Mendel’s
law, Masaryk University Mendel Museum (website): http://www
.mendel-museum.com/eng/1online/room4.htm. Accessed April 1,
2014.
CHAPTER 2
Inheritance Patterns for

Two Traits Simultaneously
Having discovered the rule of inheritance for single traits, Mendel tried
some experiments in which he performed a dihybrid cross, which quan-
tified the results of two traits in the same mating—pea color and wrinkled
texture (Figure 8). Mendel used pollen from true-breeding smooth, yellow
pea plants with eggs from plants that were true-breeding wrinkled, green
peas. The results of the F1 generation were very clear, because 100% of
the peas were smooth and yellow. The F1 outcome should not be surpris-
ing, because it is easy to explain these results by following the dominant
alleles. A gamete contains one allele from its parent for each of the two
traits. Because one parent was homozygous dominant at both loci (pea
color and texture) and the other parent was homozygous recessive at both
loci, all diploid F1 peas had to contain exactly one dominant allele for
both traits and exhibit the dominant phenotypes. However, the hundreds
of F2 peas that resulted from self-fertilizing fifteen F1 plants produced a
more complex outcome that needed further investigation. Using his data
and his understanding of probability, Mendel was able to recognize the
pattern of seeds produced. From these results, he developed his second
famous ratio, one that is evident any time individuals are mated that are
heterozygous at two loci.
Mendel Discovers Two-Trait Ratios

Mendel extended his initial research by performing crosses of true-breeding
plants for combinations of two different traits over two generations.
His results uncovered one of the most famous ratios in biology, which
is 9:3:3:1. This ratio was verified by calculating the relative frequency of
YYSS yyss
F1
15 plants
YySs
F2
315 101 108 32
Figure 8 Two generations after mating YYSS 3 yyss plants. The

numbers under the F2 peas are Mendel’s published results. Y is
dominant yellow (light gray); S is dominant smooth.
each phenotype in Figure 8 and counting the different phenotype boxes

in the Punnett square in Figure 9. Probability rules can also be used to
calculate this ratio without making a Punnett square. For example, the
probability of getting at least one dominant S allele and at least one domi-
nant Y allele (smooth, yellow phenotype) is the probability of getting at
least one S allele (3/4) multiplied by the probability of getting at least one
Y allele (3/4), or 9/16. On average, nine of sixteen peas will display both
dominant traits, six will display only one dominant trait ([3/4 × 1/4] +
[3/4 × 1/4]), and one will display both recessive traits and be homozy-
gous recessive at both loci (1/4 × 1/4). Again, Mendel would never have
detected these ratios if he had crossed only a few plants and followed just
a few peas into the F1 generation. He detected the pattern of inheritance
because he averaged a lot of data, over multiple generations, which re-
duced the amount of variation caused by random pollination.
Inheritance Patterns for Two Traits Simultaneously 17
YySs × YySs
pollen egg genotypes
genotypes YS Ys yS ys
YS YYSS YYSs YySS YySs
Ys YYSs YYss YySs Yyss
yS YySS YySs yySS yySs
ys YySs Yyss yySs yyss
F2 seeds
315 101 108 32
Figure 9 Punnett square for F2 peas in Figure 8. Genotypes for

gametes and the Punnett square make it simple to predict the ratios of
genotypes and phenotypes from a dihybrid cross of YySs 3 YySs.
F2
315 101 108 32
301 plants
number of F3 plants
and seed types 38 65 60 138
Figure 10 F3 progeny from smooth, yellow (light gray) F2 peas.

Smooth, yellow F2 peas from Figure 9 were germinated, and the
flowers self-pollinated. Mendel’s F3 results are tabulated at the bottom.
Mendel wanted to be sure he had discovered fundamental rules in ge-

netics, so he examined more closely the outcomes from the experiment in
Figure 8. From the 315 smooth, yellow F2 peas, Mendel was able to raise
and self-pollinate 301 mature F2 plants that produced F3 peas (Figure 10).
Thirty-eight of the F2 self-pollinated plants produced only smooth, yel-
low peas; 65 F2 plants produced only smooth, yellow or smooth, green
peas; 60 F2 plants produced only wrinkled or smooth, yellow peas, and
138 F2 plants produced all four types of peas. These results demonstrated
96 plants 102 plants
28 68 35 67
A B
Figure 11 F3 progeny from self-pollenated F2 peas. A, 101 wrinkled,

yellow F2 peas from Figure 9 were germinated and self-pollinated.
Mendel’s F3 results are tabulated at the bottom as the number of
plants producing the peas shown above each number. B, Similar
experiment using 108 smooth, green F2 peas.
that both recessive alleles encoding two different traits remained hidden
for a generation, just as individual recessive alleles had done in earlier
experiments. The two traits were behaving independently of each other.
Mendel continued his genetics research by raising all 101 of the wrin-
kled, yellow peas from Figure 9 and allowing the plants to self-pollinate
(Figure 11A). Likewise, he raised and self-pollinated all 108 smooth,
green peas and counted the resulting peas (Figure 11B).
From the data in Figure 8 through Figure 11, it can be seen that
every combination of alleles is possible when two heterozygotes mate.
Since the pollen and egg cells in a dihybrid cross are produced with one of
four different genotypes, the multiplication principle indicates there are
4 × 4 = 16 entries in the Punnett square, as was seen in Figure 9, with
the ratio of phenotypes being 9 + 3 + 3 + 1 = 16. Note that there
are only nine different pea genotypes in the Punnett square in Figure 9,
because in biology, order of alleles does not matter. For example, Yy = yY.
All of these mathematical principles and the observed phenotypes led
Mendel to formulate two rules that are so fundamental, they are now
called laws.
meiosis gametes
Y
S S
s 25%
y Y Y
25% s
S s
y Y y
25% s
YySs YySs 25% y
Figure 12 Meiosis produces every possible gamete genotype. The laws
of independent assortment and segregation ensure that every possible
combination of chromosomes is produced. Centromeres are enlarged
for emphasis.
Mendel Establishes Genetic Rules

Mendel did not know about DNA or chromosomes, but he correctly de-
duced their behavior based on the matings of pea plants and their result-
ing offspring. It is easier to understand his conclusions if his terminology
is changed to a modern understanding of how alleles and chromosomes
behave during meiosis when pollen and eggs are formed (Figure 12).
Mendel noticed that plants which contained two traits passed on either
one or the other trait, but never both traits in the same fertilization. He
also determined that two different traits (such as color and texture) are
transmitted independently of each other. Based on his conclusions, Men-
del is credited for discovering two of only a handful of laws in all of biology.
The law of segregation simply states that homologous chromo-
somes move to opposite poles during meiosis I (covered in Chapter 5),
which is half the mechanism that generates every possible combination of
gamete genotype. The fact that many pairs of homologous chromosomes
migrate during meiosis independently of each other is the foundation for
the law of independent assortment, which accounts for the other half of
diversity in gamete genotypes. Without knowing anything about meio-

sis, chromosomes, or DNA as the heritable material, Mendel was able
to deduce these two laws that proved to be the basis for genetics and the
diversity of life in every organism that sexually reproduces.
Mendel developed a sophisticated understanding that explained the
pattern of inheritance, and his rules remain intact after 150 years. A good
model should allow one to make testable predictions to verify the model’s
accuracy. Mendel tested his models with predictions about what would
happen when he cross plants that were heterozygous at three loci. He
mated two generations of plants and collected the peas from 639 individual
plants and tabulated the frequencies of 27 different combinations of phe-
notypes. Mendel tested his understanding on many different traits in peas,
as well as traits in other fruiting plants. His rules were always validated.
He overturned previous incorrect explanations of inheritance patterns
and wrote one of the most important papers in all of biology.
One of Mendel’s contemporaries confessed that in his own experi-
ments, he had found it very difficult to determine which parent a particular
pea resembled more closely. Many biologists in the 1860s were trying to
make the data fit their hypotheses rather than using data to challenge their
hypotheses. Mendel’s data were clean because he spent two years generating
true-breeding varieties, one trait at a time. Mendel calculated the number
of peas he would have needed to count if his original parental plants varied
at all seven traits. If he had raised F1 seeds and allowed all the flowers to
self-pollinate, Mendel would have needed to count peas with 128 different
phenotype combinations resulting from 2,187 different genotype combina-
tions. It is no wonder that others had missed the patterns; they had not sim-
plified their starting plants to isolate single traits that were true-breeding.
It seems reasonable that anyone who single-handedly invented the
discipline of genetics and who published this work as the solo author
would have become a world-wide celebrity. However, the monk’s work
was not fully appreciated at the time of publication; maybe this was in
part because of the Civil War in the United States and its repercussions
in Europe. Mendel’s work remained obscure for 15 years until three bot-
anists, Hugo Marie de Vries, Erich von Tschermak, and Carl Correns,
made the same discoveries as Mendel. Once they realized Mendel had al-
ready published the same conclusions, they must have been disappointed.
www.Ebook777.com
Fortunately, these three botanists retold Mendel’s story and gave him full
credit for his pioneering work and assured themselves obscurity instead of
fame. Mendel understood how genotypes and phenotype were produced
in subsequent generations with an equal understanding of variation pro-
duced during individual matings. His work gave mathematical structure
to inheritance patterns by explaining the phenotype and genotype ratios
when one or many traits were analyzed. Working in isolation, Mendel
documented in exquisite detail how genetic information is recombined
during sexual reproduction, not consumed or destroyed, despite recessive
traits hiding for one generation. Mendel was not able to win a Nobel Prize
because he died before prizes in physiology or medicine were awarded,
and Nobel Prizes are never given to dead people. Perhaps it is inconse-
quential, but if Mendel had been working under today’s tenure system,
he would probably have been denied tenure because his research took too
long, and he only published one paper that was not cited for 15 years.
Ethical, Legal, Social Implications: The Scientific

Meaning of “Random”
The term “random” has become the latest victim to casual slang that de-
values its true meaning. The problem with devaluing “random” is that it
has a specific meaning and important uses in mathematics and biology.
To be random, the outcome of a particular event is not knowable, but its
overall probability can be calculated. When looking at Figure 13, many
people mistakenly think panels A or B are more random than panel C,
which illustrates a lack of understanding of what random really is. Is it
possible to have a random mechanism produce one outcome 75% of the
A B C
Figure 13 Which pattern of dots best represents randomness?

time and a different outcome 25% of the time? Or do random mecha-

nisms always produce outcomes with equal frequency? Rolling dice and
flipping coins are typical examples of random events, because we cannot
predict a specific outcome, even though we can predict the probability
of an outcome. It is common knowledge that each fill has a 50% chance
of getting heads, and a 1 in 6 chance of rolling a six on a six-sided die.
However, if the coin was significantly heavier on one side, it might land
on heads 60% of the time, and yet the process would still be random. In
short, outcomes can produce asymmetrical ratios such as 3:1 as a result of
a random process, such as sexual reproduction.
Another example of how people fail to understand “random” is gam-
bling on the lottery. If a coin is tossed, is it possible to flip heads ten times
in a row? By the multiplication rule, the probability of flipping heads 10
times in a row is (1/2)10 = 1/1024, about a 0.1% chance, which is unlikely
but not impossible. Should someone bet money on flipping ten heads in a
row? This type of bet might make sense if the cost was low and the reward
was high, which is what casinos bank on. Lotteries, horse races, and casino
games are designed to make the payoff appealing enough that people are
willing to lose a little money on the slight chance that they might win big.
Lotto is the best example of people voluntarily giving their money away
on the very small chance that they will win millions. Go to the PowerBall
website as search for prizes to read the odds of winning the grand prize and
the odds of winning the smallest prize of $3 when the ticket costs $1. Note
that PowerBall uses “odds” instead of probability, but the probability can
be calculated by dividing the two numbers. For example, the probability of
winning any amount of money is approximately 1/35.11, or about 0.028.
If the odds of winning $3 were 1 in 3, then over time, a person would break
even if he or she gambled frequently. Is it a good idea to give a neighbor one
dollar every day for the opportunity to flip heads 27 times in a row? The
neighbor would benefit, as do the lottery companies, because the probability
of winning a Lotto grand prize is as likely as flipping heads 27 times straight.
Another area of everyday life where people do not understand ran-
domness is in the interpretation of statistics about safety. If an action
doubles a person’s odds of getting hurt, how bad is that? Well, it depends
on the original odds. The National Safety Council has published the odds
of dying from a range of accidents (Table 5).
Table 5 Odds of dying by different causes.

cause of death lifetime odds (USA)
any accident 1 in 36
motor vehicle accident 1 in 81
firearm 1 in 202
poisoning 1 in 344
falling object (excluding objects from space) 1 in 4,873
drowning in bathtub 1 in 10,455
suffocation by plastic bag 1 in 130,498
Source: The National Safety Council.
If the odds of being suffocating by a plastic bag were to double, it

might be reasonable to accept the doubled risk. But if the odds of being
in a car accident doubled, to 2 in 81, then a person should probably take
more care. Wearing a seat belt can reduce a person’s risk of dying in a car
accident to 1 in 140, for instance, and yet many people don’t wear seat
belts. Some drivers talk on a cell phone despite studies showing that even
a hands-free phone conversation increases the driver’s chances of causing a
car accident to the same level as driving while intoxicated. The probability
of getting in an accident when talking on a cell phone can be calculated,
but it is unknown whether the driver will have an accident on any particu-
lar trip. However, the more a person drives while talking on the phone, the
higher the probability that the driver will eventually be in a car accident.
One final example of failure to understand random is related to genetic
testing. The costs of health and life insurance are based on the probability of
the insured getting sick or dying within a given period of time. At the mo-
lecular level, getting sick is not random, but since we cannot predict when
a particular person will get sick or die, we treat these conditions as random
events and set insurance prices according to their probabilities. If an insur-
ance company adjusts the cost of insurance based on a positive genetic test,
randomness has been reduced and the cost for insurance will go up.
Since insurance companies have to cover their own expenses and they
are for-profit companies, the average person will pay more for his or her
insurance than the average person will receive payments, just as casinos
use probability to make sure they make more money per person than
the average customer wins. However, results from genetic testing can be
wrong, in which case a person might be charged a higher rate even though
he or she is no more likely than anyone else to suffer what the test pre-
dicted. In short, the more genetic testing insurance companies require,
the higher the probability the tested person will pay more for insurance
than he or she will collect from the company.
In closing, the next time someone says something was “totally ran-
dom,” keep in mind that random events cannot be predicted with cer-
tainty but have a knowable probability of happening. The next time
someone says something that seems unusual or out of the ordinary, it
would be more accurate to say, “That was strange and unexpected.” By
casually using the word “random,” people are contributing to the abuse of
a mathematically, biologically, and socially important word.
Bibliography
Bateson W. Experiments in plant hybridization (1865) in translation form
Gregor Mendel’s orignal work: Versuche über Plflanzenhybriden,
Verhandlungen des natureforschenden Vereines in Brünn, Bd. IV für
das Jahr 1865. Abhandlungen, 3–47, 1901. Minor corrections by
Roger Blumberg.
Blumberg RB. Mendel’s paper in English: experiments in plant hybri
dization (1865) by Gregor Mendel, MendelWeb (website): http://
www.mendelweb.org/Mendel.html. Accessed April 1, 2014.
Ethical, Legal, Social Implications: The Scientific Meaning

of “Random”
Crandall CS, Olson LM, Sklar DP. Mortality reduction with air bag and
seat belt use in head-on passenger car collisions. Am J Epidemiol
153(3):219–224, 2001.
Popular Science. The sky is falling! the sky is falling! Popular Science
(website): http://www.popsci.com/scitech/article/2002-01/sky-falling-
sky-falling. Accessed April 1, 2014.
Multi-State Lottery Association. Powerball: prizes and odds, Powerball
(website): http://www.powerball.com/powerball/pb_prizes.asp. Ac-
cessed April 1, 2014.
Strayer DL, Drews FA, Crouch DJ. A comparison of the cell phone driver
and the drunk driver. Hum Factors 48(2): 381–391, 2006.
CHAPTER 3
Prokaryotes Reproduce by
Binary Fission
DNA is an ancient molecule and is the genetic material inside every

cell. The mechanism of DNA replication is highly conserved across all
domains of life, as shown by the existence of very similar DNA poly-
merase molecules in every species. However, once cells have synthesized
new chromosomes, how do prokaryotes (bacteria and archaea) partition
their twin DNA molecules into the next generation of organisms? To fully
understand bacterial propagation of genetic information, it is helpful ex-
amine real data from published papers to deduce the mechanisms that all
prokaryotes use to transmit their genetic information to the next genera-
tion. Through case studies, the reader can construct an understanding of
bacterial reproduction.
Before analyzing data using Escherichia coli as a model (well-studied)
bacterial species, having an understanding of the physical dimensions of a
single cell is helpful. Because cells are three-dimensional, scientists need to
take into account their volume as well as their height and width. Biologists
assessed the range of volumes of E. coli bacteria. Millions of E. coli cells
were grown at 378 C and then harvested so that their cell volumes could
be calculated. After carefully measuring the length and radius of each tu-
bular cell, investigators calculated the volume of each cell. They plugged
the cellular dimensions into the formula for the volume of a cylinder and
plotted a frequency distribution of the volume. Even though all cells in
the culture were genetically equivalent, they did not all have the same vol-
ume. Even equivalent genotypes can produce variable phenotypes, such
as cell volume. Using all of these values, the microbiologists calculated
an average E. coli cell volume, under their experimental conditions, of
0.83 µm3. But just knowing the average does not tell the whole story.
www.Ebook777.com
The range of volumes for E. coli provides evidence that allowed biolo-
gists to deduce how bacteria produce new cells. For example, do parental
cells produce little baby cells or two cells of equal volume? Rather than a
normal distribution of sizes, with a symmetrical curve around the average
size, the graph displayed a “tail” on the right side indicating some cells
were exceptionally larger than most. This indicated there were some very
large cells but no cells that were tiny. Another question was about the
effect of temperature on their growth. E. coli often lives in the guts of
animals. To survive, they must grow at a variety of temperatures beyond
their control. As one step in discovering how bacteria reproduce, a team
of microbiologists working in the 1960s wanted to know whether the
growth rate of bacteria was determined genetically or environmentally.
In their experiment, the investigators grew genetically equivalent popula-
tions of bacteria in two separate flasks. The independent variable was the
temperature of the two different cultures. Instead of counting the num-
ber of cells produced in each flask, the microbiologists measured cellular
reproduction, the dependent variable, by drying and weighing aliquots
of cells. After 2.5 hours at 258 C or 308 C, cells from both flasks were di-
luted with fresh media in their separate flasks and grown at 378 C. While
the cells grew at 378 C for another 90 minutes, aliquots were periodically
removed for drying and weighing. Prior research had shown that volume
of individual cells varies within a narrow range, so the number of cells
would proportional to the mass. From this experiment, they determined
the rate of bacterial reproduction was controlled by genetic information
and the environmental factor of temperature.
The microbiologists determined that E. coli cells have an average vol-
ume of 0.83 µm3 with some variation in the population. A small percent-
age of cells were much larger than the average as revealed by the extended
“tail” on the right side of the graph’s major peak, but even these larger cells
are less than twice the average size. They also concluded that bacteria grow
very rapidly, doubling their population in less than an hour, but the exact
rate depends on the temperature. Cells grow faster at 308 C than at 258 C,
as indicated by the different slopes of the plotted lines showing cell mass
as a function of time. However, the rate of growth was not genetically set,
because the cells were able to adjust their growth rate rapidly when shifted
to 378 C. When the two populations were shifted to 378 C, they grew
Prokaryotes Reproduce by Binary Fission 27
faster than before but at identical rates, as indicated by the parallel lines
in the graphs of mass as a function of time. These scientists knew that
chemical reactions accelerate when they are heated because the molecules
vibrate faster, allowing more frequent random interactions between them.
The experiment showed the rate of bacterial growth was controlled by
temperature and DNA, which indicates chemical reactions are involved
bacterial division. From these experiments, they learned that prokaryotes
such as E. coli grow as a result of biochemical activity, the growth rate is
sensitive to temperature, and the cells do not all grow at the same rate.
If they had compared different species at the same temperatures, they
would have been able to determine that each species has a genetically
constrained growth rate that differs between the species. Therefore DNA
and environmental factors govern growth rates.
Biologists wanted to learn more about how bacteria reproduce. They
hypothesized that proteins and their chemical interactions were at the
heart of bacteria reproduction. If proteins were involved, there had to
be a genetic component to bacterial reproduction, even though the rate
was not determined exclusively by DNA. To probe further, microbiolo-
gists measured the average volume of mutant E. coli cells grown either
in the presence or absence of the DNA base thymidine (Figure 14). The
20 m
Figure 14 Effect of DNA replication on cell size. Mutant E. coli

cells that required the addition of thymidine to replicate their DNA.
Mutants could be grown either with or without added thymidine.
Micrographs show thyA mutant E. coli (top) compared to wild-type
E. coli (bottom).
Source: Courtesy of Elizabeth Brunner and Malcolm Campbell, Davidson, NC.
bacterial cells they used in this experiment had been mutated prior to the
experiment, so the mutant cells could not produce their own thymidine
base, which is required for DNA synthesis. Therefore, these cells had all
the nutrients that they needed for cell growth, but they could not repli-
cate their DNA. The researchers reported only the average cell volume
from a population, not individual cells. They repeatedly sampled aliquots
of cells over seven hours and plotted the “average volume of cells for each
time point.” Cells lacking thymidine continued to grow in volume, even
though they were unable to make new chromosomes (Figure 14, top).
Cells in a second flask were given thymidine and exhibited volumes that
were either large or small. Approximately every two hours, the population
of cells exhibited two volume categories—approximately 0.4 µm3 and
0.8 µm3. Seeing the dual size distribution for cells given thymidine, the
biologists understood how bacteria reproduce.
The experiment illustrated in Figure 14 is a great example of how sci-
entists can deduce many conclusions from data that appear rather simple.
It is hard to imagine how a cell could grow in size over time and then
abruptly decrease in volume, only to repeat the entire process over again
every 2 hours. However, if the parental cells reproduce by generating two
equivalent cells, the new cells would be about half the volume that the pa-
rental cell had been just before cell division. By reporting two cell volumes
at a single time point, the investigators did not average all cell volumes into
a single number but split the cells into two categories based on averaged
cell volume within each category. Based on the periodic overlapping of
volume categories, they estimated that cells with thymidine in this experi-
ment divided about once every 2 hours, which is slower than the 378 C
doubling time of wild-type cells. The data in Figure 14 led biologists to
pose a testable hypothesis—cells can divide into two equal-sized daughter
cells once a parental cell has reached a certain volume, and the parental cell
has replicated its DNA. Cells lacking thymidine continued to grow in vol-
ume but detected the lack of a replicated chromosome and therefore did
not divide. To test their hypothesis that bacteria reproduce by dividing in
half, biologists used time-lapse photography, which was invented in 1965.
To perform their time-lapse photography experiment on bacterial
reproduction, microbiologists connected a camera to a microscope and
Prokaryotes Reproduce by Binary Fission 29
took photos every 15 seconds over several minutes. There are many mov-
ies of bacteria growth which can be found on YouTube. E. coli cells elon-
gate and then become constricted in the middle; they eventually form
two new cells as the last cellular connection breaks. DNA is partitioned
into each of the new cells during the process. Notice that the parental
cells in the movie do not divide synchronously, which is consistent with
the data in showing variation in cell size and volume at any given time
for a population. What was not clear from movies of cells dividing was
how a bacterium could sense whether its chromosome had replicated or
not. How cells sense chromosome replication is beyond the scope of this
book, but it is known from Figure 14 that these unicellular organisms
must be able to respond to the presence of two chromosomes within the
cytoplasm of a single cell.
Prokaryotes grow until they reach a certain volume and have two cop-
ies of their DNA present in the same cytoplasm. Over a narrow period
of time, the cells constrict near the middle of the elongated cell to pro-
duce two equivalent cells. The process of prokaryotic replication is called
cellular fission. The two new cells are about half the mother cell’s size,
but it is impossible to predict with certainty what a new cell’s volume will
be because volume varies within a range that is constrained genetically.
However, the rate of growth is not determined exclusively by the DNA;
the proteins involved behave like chemical reactants so that warmer tem-
peratures lead to faster growth. Proteins are responsible for the mecha-
nism of cellular fission because each species divides within its own genetic
range of division rate.
This chapter presented the data that informed microbiologists that
prokaryotes replicate their genetic information and then partition the
two chromosomes into new cells that are genetically equivalent. Passing
DNA from one generation to the next defines each species and provides
continuity of information as individuals reproduce. DNA information is
used (transcription and translation) by each cell and is neither lost nor
consumed in the process. The highly regulated process of cellular fission is
an example of reproducing information at the organismal level. Chapter
4 analyzes data to determine whether eukaryotes undergo the same cell
division process as prokaryotes.
Bibliography
Ducret A, Maisonneuve E, Notareschi P, et al. A microscope automated
fluidic system to study bacterial processes in real time. PLoS One
4(9):1–8, 2009.
Edgar BA, Kim KJ. Cell biology: sizing up the cell. Science 325(5937):
158–159, 2009.
Harvey RJ, Marr AG, Painter PR. Kinetics of growth of individual cells
of Escherichia coli and Azotobacter agilis. J Bacteriol 93(2):605–617,
1967.
Hoffman H, Frank ME. Time-lapse photomicrography of cell growth
and division in Escherichia coli. J Bacteriol 89(1):221–216, 1965.
Kubitschek HE. Control of cell growth in bacteria: experiments with thy-
mine starvation. J Bacteriol 105(2):472–476, 1971.
Leavitt RW, Wohlhieter JA, Johnson EM, et al. Isolation of circular
deoxyribonucleic acid from Salmonella typhosa hybrids obtained
from matings with Escherichia coli Hfr donors. J Bacteriol 108(3):
1357–1365, 1971.
Ng H, Ingraham JL, Marr AG. Damage and depression in Escherichia coli
resulting from growth at low temperatures. J Bacteriol 84:331–339,
1962.
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CHAPTER 4
Eukaryotes Reproduce
Through Mitosis and
Cytokinesis
Human cells divide millions of times a day. Hair cells, cheek cells, blood
cells, skin cells, and so on divide every day of a person’s life. When a child
scrapes his or her knee, skin cells respond and make replacement cells
in just a few days. How do eukaryotic cells like ours partition their ge-
netic information evenly between the two new cells? The simplest answer
would be for all cells to divide the same way E. coli does, but do the data
support this answer?
Consider some facts, and then see whether prokaryotic cellular fission
could work for eukaryotic cells. E. coli has a single circular chromosome
but humans have 23 pairs of linear chromosomes (46 total). Given that
humans have forty-six times the number of DNA fragments to partition
between daughter cells, it seems unlikely that the two types of cells could
divide the same way. If human cells merely constricted in the middle, how
would the nucleus and the chromosomes partition evenly?
Some aspects of passing information on to the next generation are
extremely similar between prokaryotes and eukaryotes. Like bacteria, eu-
karyotic cells must replicate their DNA before they divide. Replicating
DNA first makes sense, because only after duplicating every chromosome
will the human parental cell have enough DNA to provide each new cell
with the full 46 chromosomes. As displayed in Figure 15, the 24 types
of human chromosomes are not identical (numbers 1–22 of matched
pairs plus X and Y = 24 in men; mitochondrial genome not consid-
ered here). Our chromosome pairs range in size from largest (number 1)
to smallest (number 22) plus the two sex chromosomes X and Y. Each
Figure 15 Photograph of a man’s chromosomes, organized by size

(number 1 in top left corner). Twenty-three pairs are shown, with the
only uneven pair being the larger X and smaller Y chromosomes in the
bottom right corner.
Source: Image courtesy of the National Human Genome Research Institute, from www.genome
.gov/glossary/resources/karyotype.pdf.
chromosome carries a different amount and sequence of double-stranded

DNA (dsDNA). Males have one X and one Y chromosome (the smallest
one), but females have a pair of X chromosomes. Humans do not have
the most chromosomes of all mammals with several species carrying over
40 pairs. One species of fern has 52 pairs of chromosomes!
Having more than one chromosome inside a nucleus presents a prob-
lem for diploid cells. How do parent cells make sure that the two new
cells get exactly two copies of each chromosome? Cells do not have the
capacity to count, nor can they see the location of each chromosome
stuffed inside their nuclei. In fact, the nucleus presents yet another chal-
lenge that E. coli cells do not have to face. How can a human cell divide its
membrane-bound nucleus in two equivalent parts, each with 23 pairs of
matched chromosomes? To answer these questions, it is easier to observe
microscopic data in the form of an original movie made in the middle of
the twentieth century (search mitosis on YouTube). The movie shows the
smooth and continuous process of chromosomes separating inside cells of
a eukaryotic cell.
Eukaryotes Reproduce Through Mitosis and Cytokinesis 33
From the movie of DNA partitioning some of the major steps can
be seen in the process of dividing genetic information into the new pair
of nuclei. Notice that the whole cell did not divide in this process; only
the chromosomes separated and partitioned into two equivalent halves.
In the first phase of nuclear division, the chromosomes condense, which
helps to keep them from getting tangled when they separate later. Within
a few minutes, the chromosomes migrate one by one to the midline and
jostle around until all the other chromosomes reach the center. Moving
to the center of the cell is not a process left to random diffusion. Long
molecular tubes called spindles extend from the two cleared zones, and
attach to each chromosome at its constricted area. In a very coordinated
manner, all the chromosomes appear to split in half, which finally reveals
the results of DNA replication. The newly divided chromosomes migrate
in opposite directions and toward the cleared areas because of the physical
forces applied to them via the spindles.
Most of the time, chromosomes are composed of one long molecule
of dsDNA. But after replication, each chromosome contains two dsDNA
equivalent copies of itself that are attached to one another at the area of
constriction called a centromere. While the two equivalent copies of a
chromosome are attached at the centromere, each copy is called a chro-
matid. The difference between a chromatid and a chromosome is physical
placement; two chromatids are attached to one another, but once they
separate, the same pieces of DNA are referred to as two chromosomes.
Keep in mind that DNA polymerases make mistakes during replication,
so the freshly separated chromatids may not be 100% identical due to
new mutations. Joining the two chromatids to each other in the first por-
tion of nuclear division ensures that the two chromatids will separate and
the equivalent chromosomes will move to opposite poles of the nucleus.
In the movie of mitosis, the chromatids did not get tangled at this stage,
which is both impressive and efficient.
Upon separation, the previously joined chromatids are referred to as
chromosomes. The process of DNA separation concludes with the slow
movement of chromosomes to opposite poles, where they will eventually
become less condensed and the nuclear membranes will re-form a pair of
nuclei. By the end of the chromosomal separation process, the cell has
two equivalent nuclei each with the diploid number of chromosomes.
The partitioning of replicated chromosomes to the newly formed

nuclei is called mitosis, which was orchestrated successfully more than
50 trillion times to form the cells of a typical human body. All eukary-
otic cells undergo a similar process, though details, such as the number of
chromosomes, varies by species. The movie available online was filmed to
highlight the chromosomes over any other aspect of the cells. However,
others have used fluorescent tags to label two different parts of the mitotic
machinery. The long fluorescent orange molecules are the spindle fibers,
which are composed of microtubules. The fluorescent green dots are the
centromere attachment sites of microtubules to the paired chromatids.
Chromosomes appear as dark shapes in a light green background. These
images of live cells undergoing mitosis with labeled spindles helped biolo-
gists understand how the chromosomes are able to gather at the midpoint of
each cell and then move away from each other into the new pair of nuclei.
Once mitosis is complete, most cells divide the contents of their cy-
toplasm in a process called cytokenesis to produce two equivalent cells.
Cell biologists have divided mitosis into four discrete steps, but as is clear
from watching the movie online, the distinctions between the steps are
not clearly demarcated.
Step 1. Cells condense their chromosomes and begin to move to the

midline during prophase. The prefix “pro” means coming
before the main event, as in prologue.
Step 2. When chromosomes align along the midline, cells are said to
be in metaphase.
Step 3. At a coordinated time during metaphase, sister chromatids
separate from each other and metaphase has given way to
anaphase.
Step 4. Finally, as the freshly separated chromosomes finish their
migration and the nuclear membrane reforms, cells are in
telophase. Cytokenesis follows telophase.
At the end of mitosis, one human cell contains two nuclei total-
ing 92 chromosomes. During cytokenesis, cell division establishes new
membranes to divide the cytoplasm into two similar parts, permanently
separating the equivalent nuclei. After DNA replication, mitosis, and
Eukaryotes Reproduce Through Mitosis and Cytokinesis 35
cytokenesis, one cell has produced two new cells that are nearly identi-
cal in genetic information. Chromosomes gather at the middle during
metaphase because they are pushed there by the microtubules, and they
continue to bounce around there until all the chromosomes are gathered.
Generally, cells sense when all the replicated chromosomes are aligned
in the midline and do not progress into anaphase prematurely. When
the microtubules draw the chromatids away from each other in normal
cells, all of the chromosomes are drawn back to the clear place where the
microtubules are anchored. Finally, each new cell must reestablish nuclear
membranes around the two sets of chromosomes, relax the compact state
of the chromosomes, and subdivide all the cellular contents that used to
be inside the original parent cell.
In addition to the processes of mitosis and cytokinesis, eukaryotic
cells share an evolutionarily conserved cell cycle in which cells prepare for
cell division (Table 6). When a cell is not undergoing mitosis, it is in a
stage called interphase. Each species and cell type varies somewhat, but
the general trends and proportion of time are typical. Rapidly dividing
cells, such as cancer cells, repeat the loop of cell cycle from M to G1, S,
G2, and so on, faster than shown in Table 6. Cytokinesis overlaps with the
end of mitosis and the beginning of G1. Some cells, such as neurons, can
exit the cell cycle and are described as in G0. As in prokaryote replication,
the two new cells are half the volume of the parental cell.
There are two microtubule organizing centers, seen as the clear zones,
inside a parental cell. Radiating out of these two organizing centers, mi-
crotubules attach to the paired chromatids at their centromeres and move
them toward the midline of the cell. Chromosomes jostle back and forth
during metaphase, because another microtubule from the other side binds
to the other chromatid and the DNA is in the middle of a tug-o-war.
After a while, proteins holding each of the paired chromatids together
Table 6 Steps in typical eukaryotic cell cycle.

name description duration (hours)
G1 growth and normal cellular functions 10
S synthesis of DNA 8
G2 growth and normal cellular functions 4
mitosis separation of chromosomes 2
www.Ebook777.com
simultaneously separate and the sister chromatids are pulled away from
each other. The newly liberated chromatids, now called chromosomes,
migrate toward opposite organizing centers to produce two equivalent,
but not identical, nuclei. Perhaps the back and forth chromosomal mov-
ing is one way that the cell senses whether all chromosomes have been
connected to opposing microtubules. Normally, once the cell senses that
the chromosomes are attached to opposing spindles, the chromatids are
permitted to separate.
The end product of mitosis is analogous to the end product of bac-
terial cell fission, and it also illustrates the unifying theme of biological
information transmission at the organism level. Mitosis uses information
encoded in the DNA but does not consume or significantly alter the ge-
netic information. Eukaryotic cells transmit the heritable information
from one parent cell to two new cells via asexual reproduction. Mitosis
and cytokinesis provides continuity of the species and ensures that nearly
every cell in a person’s body is genetically equivalent. Given that the cell
cycle is so important to the function of all organisms, it should not be
surprising to know many proteins (such as cyclin kinases) tightly control
progression of cell division. Gametes of sperm and eggs, however, are
different from all the other cells in a person’s body. The sperm and egg
cells that fused to become each person were not the products of mitosis.
Chapter 5 examines how eukaryotes that sexually reproduce are able to
produce haploid sperm and egg cells without mitosis.
Bibliography
Cimini D, Cameron LA, Salmon ED. August 2006b. Mitosis Movie
Series: 2. Meeting in the Middle before Parting: Metaphase-to-
Anaphase. American Society for Cell Biology. <http://www.cellimage
library.org/images/8078>. Accessed 5 April, 2014. <http://www.cell
imagelibrary.org/images/8079>. Accessed 5 April, 2014. <http://
www.cellimagelibrary.org/images/12020>. Accessed 5 April, 2014.
Rieder C, Cole R, Waters J. November, 1995. CELLebration: Part IV:
Cell Division 17. Mitosis and Nondisjunction. American Society for
Cell Biology. Washington, DC.
CHAPTER 5
Gametes are Formed After

Meiosis
Most multicellular organisms in the world are the products of sexual re-
production. Animals produce gametes of sperm and egg, which fuse to
form a new individual. Plants reproduce via pollen and eggs, which pro-
duce seeds that germinate into new organisms. Chapter 5 investigates
how each parent can pass on genetic information and produce a new
organism with the same number of chromosomes. Note that this chapter
will follow only the nuclear genes since mitochondrial genes are always
passed from mother to child since the sperm rarely contributes any mito-
chondria to the next generation. If human sperm and eggs were diploid
products of mitosis, then the fusion of equal gametes would result in tet-
raploid cells with 92 chromosomes, which would not produce a human.
Therefore, gametes need to be haploid and contain half the usual number
of chromosomes before fusing.
Over the course of evolution, living organisms rarely develop complex
processes from scratch. Typically, a preexisting complex process becomes
slightly modified to produce a new outcome. For example, all vertebrates
have eyes, but some can see in color and others cannot. How did color
vision evolve from black and white vision? One possible route would have
been for color vision to develop from a completely different visual organ
than the existing eye that could see only black and white. Alternatively,
a black and white-seeing eye could have been slightly modified to gain
color vision, which is what happened in vertebrate evolution. Similarly,
gamete formation uses a mechanism comparable to mitosis, but with a
slight modification that cuts in half the amount of DNA in the result-
ing nuclei. The formation of gametes is the outcome of a process called
meiosis. Because meiosis is more complex than mitosis, the best way to
see meiosis is to watch an animation online (search “meiosis cell division”

animation on YouTube).
Like mitosis, meiosis begins after DNA replication has occurred so
that each chromosome is composed of two chromatids. By definition,
each chromatid is attached to its sister chromatid but becomes a full-
fledged chromosome once the two chromatids are separated. Cells that
will give rise to gametes must go through two rounds of DNA separation.
Any photographs are static moments frozen in time from a process that is
smooth and continuous. The online animation gives a better sense for the
fluid and continuous transition between the stages that are named but not
really as distinct as many mistakenly believe.
While watching the animation, several differences between mitosis
and meiosis should be noticed, besides the different final outcomes of
haploid nuclei. Read through the following steps of meiosis, and look
for differences between mitosis and meiosis. Cell biologists have divided
meiosis into two phases. Both of the phases are subdivided into four steps,
because each phase of meiosis is very similar to the four steps of mitosis.
The first phase of meiosis is called meiosis I, followed by meiosis II, which
is more similar to mitosis than is meiosis I.
Meiosis I
Step 1. Cells condense their conjoined chromatids and, unlike mito-
sis, similar chromosomes form pairs and become intertwined
during prophase I. For example, maternal chromosome
13 pairs with the paternal chromosome 13, bringing to-
gether four equivalent chromatids. Paired chromosomes are
called homologs or homologous chromosomes.
Step 2. Homologous chromosomes, composed of four chromatids,
align along the midline during metaphase I.
Step 3. Homologous chromosomes separate in anaphase I, but the
chromatids remain attached. For example, the two copies of
chromosome 13, each composed of two chromatids, move
away from each other. The fact that chromatids remain at-
tached is a very important difference between meiosis I and
mitosis.
Gametes are Formed After Meiosis 39
Step 4. Meiosis I is completed after telophase I, when the separated

chromosomes, as paired chromatids, finish their migration
and the nuclear membrane reforms.
Cytokinesis separates the newly formed nuclei and partitions the

cytoplasm and organelles as well. Not all species undergo cytokinesis at
this step.
The two new cells contain haploid nuclei, because each nucleus has
only one copy of each chromosome. In our example, the two copies of
chromosome 13 moved to opposite poles and each chromosome is still
composed of two chromatids. Be sure to think through this stage care-
fully, because these haploid nuclei still contain as much DNA (23 chro-
mosomes × 2 chromatids) as a diploid cell with 46 chromosomes.
Meiosis II
Meiosis II is similar to meiosis I, but more closely resembles mitosis.
Meiosis II happens in both of the newly formed haploid nuclei:
Step 1. In prophase II, chromosomes condense again but the paired

chromatids do not become intertwined as happened dur-
ing prophase I. Prophase II is more similar to prophase in
mitosis.
Step 2. Each chromosome aligns along the midline in metaphase II,
which prepares the paired chromatids for separation as seen
in mitosis. Microtubules are attached to the centromeres and
move them back and forth as seen in the mitosis movie.
Step 3. Paired chromatids separate and begin migrating in opposite
directions in anaphase II. Note that once the chromatids
have become separated, they are called chromosomes again,
even though the DNA in each structure is exactly the same
as it was when it was referred to as a chromatid. This is very
similar to anaphase in mitosis.
Step 4. Telophase II concludes meiosis and produces two pairs of
haploid nuclei that are separated into four haploid cells once
cytokenesis is complete.
Meiosis produces haploid gamete cells with half the number of chro-
mosomes of a typical cell, and half the amount of DNA. Eggs and sperm
go through the same meiotic process to produce haploid nuclei, but their
size, shape, and cytoplasmic content differ. However, an important part of
meiosis has not been considered—the functional consequence of the four
equivalent chromatids becoming intertwined during prophase I. By just
watching the process of meiosis, the subtle recombination, or cutting and
pasting, of DNA that takes place during prophase I cannot be detected. To
understand what happens during recombination, it is necessary to zoom
in to see each of the four individual chromatids. In high-magnification
images, individual chromatids can be seen crossing over each other, which
happens only when the four chromatids are interwoven during prophase I.
Everywhere there is crossing over of chromatids, two dsDNA molecules
are being cut and they are swapping segments. This was shown in the
meiosis animation mentioned above. No chromatid remains unchanged.
Recombination is a random process, and afterward, no two chromatids
will be identical. Prophase I is one reason why non-twin siblings look
similar but not identical. With this level of recombination for each chro-
matid, multiplied it by the number of chromosomes in a species, it is
easy to see why every gamete is unique and a population always expresses
variation in genotype and phenotype. Recombination during prophase I
is a crucial aspect of meiosis and contributes to more diversity of gametes.
It is difficult to see evolutionarily shared traits when organisms like
prokaryotes and eukaryotes diverged hundreds of millions of years ago,
but difficult is not the same as impossible. Very old processes, such as
passing genetic information to new cells, display evidence of our ances-
tral heritage that unites all cells to their shared origins of life. Mitosis
and meiosis share many structural features. They both use microtubules
that originate from organizing centers at opposite ends of the cell. These
microtubules attach to the chromatids at the centromere where the chro-
matids attach to each other. Centromeres contain unique proteins dis-
tinct from the remaining portions of each chromosome. Chromosomes
in both processes condense after replication and prior to chromosomal
movement. Both processes are divided into four nearly identical steps, but
mitosis undergoes only one round of DNA separation, whereas meiosis
has two. Mitosis separates its chromatids immediately, whereas chromatid
separation takes place only in anaphase II of meiosis. Mitosis and meiosis

share so many features that they most certainly have similar evolutionary
origins. Their shared evolutionary history has been confirmed many times
by examination of the common proteins involved in mitosis and meiosis
in every eukaryote species.
During mitosis, chromosomes do not recombine, which is good be-
cause cells are able to reproduce equivalent copies of themselves and not
introduce genetic variation. However, in meiosis, chromatids must cross
over during prophase I before a cell can progress to metaphase I. At the
end of meiosis, each chromosome is a mosaic of the four homologous
chromatids from the parent, which means every chromosome in a gamete
is different from every parental chromosome. Recombination in gamete
formation produces variation in the population even if only two parents
produced all of the progeny. Remember, crossing over during prophase I
happens only among homologous chromatids and not between different
chromosomes, such as #13 and #14.
Haploid gametes from the same species unite during fertilization. It is
worth the time it takes to watch another movie online that shows the fusion
of sea urchin gametes (two eggs are immediately visible) to produce new
individuals with similar but not identical DNA to their parents. Sea urchins
have been a favorite model organism of developmental biologists for over
100 years, because they are easy to manipulate and visualize. Meiosis in
sea urchins, pea plants, yeast, flies, worms, and humans is nearly identical
in every way because all of these species evolved from a common ancestor
and all organisms are connected through an unbroken chain of inherited
genetic information. The information in DNA remains equivalent in nearly
every cell within an individual, except in gametes, where chromatids recom-
bine to produce new chromosomes. Recombination is the genetic source of
genotype and phenotype variation in a population. Variation within the
population is a key component of biological information and evolution.
Ethical, Legal, Social Implications: The Ethics

of Genetic Engineering Humans
Having learned about gamete formation and cell division, perhaps it
would be appropriate to consider the consequences of wanting to have
healthy children. To date, the Human Genome Project has identified over
2,000 genetic diseases with more discovered every week. There are now
over 1,000 genetic tests for all sorts of traits, not just diseases. There is a
genetic test that claims to determine if a child will “be a good athlete” be-
fore he or she is even born. It is not reasonable to think that an Olympic
gold medal winner will be determined by a single gene. It seems more
reasonable to take into consideration training, nutrition, and determina-
tion. All of these non-genetic aspects also play a role in determining who
will be the next superstar athlete.
In the 1950s, people were convinced that a child’s social environment
determined most of his or her traits, but since 2000, popular opinion has
swung to the other extreme—a belief in genetic determinism or the idea
that our DNA determines our fate. Given the rising costs of healthcare
and the economic savings of disease-free children, those who believe in
genetic determinism might argue that society should practice disease pre-
vention at the genetic level. For example, spina bifida, a potentially lethal
birth defect of the central nervous system, is caused by both genetic and
environmental factors. Because so many people accept genetic determin-
ism, parents tend to move toward gene screening when, in fact, eating
green vegetables during pregnancy can reduce the risk of spina bifida by
sixty percent.
Currently, three genetic diseases are commonly tested in America:
Down syndrome (caused by trisomy 21), cystic fibrosis, and spina bifida.
While cystic fibrosis and spina bifida are often fatal at an early age, Down
syndrome results in different kinds of people, implying that society de-
fines this difference as a disease. Should society eliminate these forms of
individual diversity? Science gives us the tools needed to eradicate these
people from the next generation. Increasingly, parents are aborting fe-
tuses if Down syndrome is detected. As the number of Down syndrome
children decreases, parents may be considered genetically irresponsible
if they do not abort fetuses diagnosed prior to birth. What is the right
thing to do?
In 2002, most states in the United States screened newborns for a
handful of disorders. As of 2016, most states typically screen for at least
29 different disorders. Not only are these DNA tests increasing in num-
ber, but they are becoming mandatory in more states, although only a few
of the tests have been approved by the Food and Drug Administration
(FDA). The remaining tests have not been validated by anyone except the
companies that market the tests, yet many parents in the United States
mistakenly believe genetic tests are always correct.
Cystic fibrosis is the most common genetic disease in Caucasians and
is the phenotype produced by any one of over 900 different mutant al-
leles. A positive result using fetal cells and a typical genetic test for cystic
fibrosis will be correct less than 30% of the time, but very few parents
know the test's accuracy. Mandating genetic testing may put society in
financial conflict with parents who choose not to abort. It is in the best
financial interest of society not to support children with special needs that
cost a lot of money.
Perhaps some people agree that genetic diseases should be eliminated
whenever possible. How far should society be willing to go to reach per-
fection? The composer Beethoven had deteriorating hearing, and the im-
pressionist painter Van Gogh had glaucoma along with mental illnesses.
Should we forego creative genius as a tradeoff for fewer diseases? If other
nations start engineering designer babies, perhaps super warriors, are all
countries obliged to defend themselves with similar genetic manipula-
tions? Some men have been described as genetically violent and pre-
disposed to criminal acts. Should preemptive incarceration and forced
sterilization be used to prevent crime to protect our way of life? Is it ap-
propriate to protect our lives by aborting fetuses that might grow up to
become terrorists?
Many of these questions have been addressed in books such as Brave
New World (1931) by Aldous Huxley and movies such as Gattaca (1997)
and Minority Report (2002). The reader may not have formed an opin-
ion yet because having children is not a major concern. But decisions are
being made today that will affect everyone’s reproductive freedom and
options. Twenty years ago, there were a few genetic tests. Now there are
many genetic tests. Student of biology should be part of the public discus-
sions that help society make these important decisions.
Bibliography
Hartwell, Leland, et al. 2000. Genetics: from genes to genomes. McGraw
Hill. Boston, MA. 813 pages.
Ethical, Legal, Social Implications: The Ethics of Genetic

Engineering Humans
Written with the assistance of Kyri Bye-Nagel, Davidson, NC.

Conclusion
All cells come from preexisting cells; we know this to be true. Prokary-
otic cells reproduce primarily through cellular fission and can grow ex-
ponentially in a very short period of time. Prokaryotes and eukaryotes
contain many of the same genes and proteins involved in DNA repli-
cation and chromosomal segregation. However, cytokinesis and mitosis
are much more complex processes found only in eukaryotes. Evolution
tends to modify existing structures and mechanisms rather than produce
completely novel processes. Therefore, it is not surprising to see many
similarities between mitosis and meiosis, though the latter produces hap-
loid nuclei instead of diploid. Gametes use random separation of chro-
mosomes (Mendel’s laws of independent assortment and segregation) as
well as recombination of chromatids to produce nearly infinite combina-
tions of alleles in gametes. Studying patterns of inheritance takes time and
mathematics training, which the monk Gregor Mendel had in abundance.
He discovered the famous ratios of 3:1 and 9:3:3:1 as well as the laws of
independent assortment and segregation. This book presented Mendel’s
150-year-old data to uncover the fundamentals of genetics that he used
deduced the steps required for eukaryotic and prokaryotic reproduction.
www.Ebook777.com
Glossary
alleles. alleles are different versions of the same gene.
anaphase. when chromosomes separate during mitosis.
anaphase I. when homologous chromosomes separate during meiosis I.
anaphase II. when sister chromatids separate during meiosis II.
cellular fission. the process of cellular replication whereby a mother cell divides
into two equivalent daughter cells.
central dogma. it refers to the critical role of converting DNA information into
RNA and then into protein.
centromere. the narrowed portion of a chromosome, which leads the way during
chromosome separation.
chance. it indicates a random event whose precise outcome is unknowable, but
whose probabilities can be calculated.
chromatid. replicated DNA molecules in a single chromosome still attached to
each other at the centromere.
cotyledons. embryonic leaves that emerge immediately upon germination.
crossed. synonym for mated.
cytokenesis. the process whereby the entire cell is partitioned into equivalent
halves that become new cells with a complete set of chromosomes.
dependent variable. it is measured in response to the independent variable.
dihybrid cross. any mating where two traits are being observed simultaneously.
diploid. cells and organisms that carry two copies of almost every chromosome.
dominant traits. traits that are always detectable in the phenotype of an individual.
dsDNA. abbreviation for double-stranded DNA.
F1 generation. the first generation derived from the P generation.
gametes. the egg and sperm, are haploid and are required for sexual reproduction.
genotypes. the genetic composition that determines an organism’s appearance or
behavior.
germination. the process of a new plant sprouting from a seed.
haploid. it contains only one of each chromosome, half as many as a diploid cell.
heterozygous. heterozygous individuals have two different versions of a gene.
homologous chromosomes. the maternal and paternal copies of similar
chromosomes.
homozygous. homozygous individuals have two copies of the same version of a
gene.
independent variable. the variable manipulated by the investigator.
interphase. the bulk a eukaryotic cell’s life, outside of mitosis or meiosis, when
the cell is performing its typical functions.
www.Ebook777.com
48 GLOSSARY
law of independent assortment. it states that non-homologous chromosomes

migrate independently of each other.
law of segregation. it states that paired chromosomes move to opposite nuclei.
meiosis. the process of partitioning nuclear DNA that produces haploid gametes
from diploid parental eukaryotic cells.
meiosis I. the first half of meiosis and results in two haploid nuclei with homolo-
gous chromosomes partitioned into different cells.
meiosis II. the second half when chromatids separate and four haploid nuclei are
formed.
metaphase. during mitosis, when all the chromosomes align in the middle, just
before chromatids are separated to the two poles.
metaphase I. during meiosis I when paired homologous chromosomes align
along the middle of the cell prior to anaphase I.
metaphase II. during meiosis II when paired chromatids align along the middle
of the cell prior to anaphase II.
microtubules. protein hollow poles that form the spindles used to separate chro-
matids during mitosis.
mitosis. the division of eukaryotic chromosomes after replication into a pair of
equivalent nuclei.
model. a model organism is one that is studied extensively because its biology is
amenable to investigation.
P generation. the parental generation, which is the first cross in a multi-
generational series of matings.
phenotypes. the way an organism appears or behaves due to its genetic makeup.
progeny. synonym for offspring or babies; the next generation.
prophase. the first phase of mitosis, when chromosomes condense prior to metaphase.
prophase I. during meiosis I when paired homologous chromosomes condense
and coalesce all four chromatids; chromatids recombine or cross over to generate
novel chromatids prior to metaphase I.
prophase II. during meiosis II when paired chromatids are connected to spindle
fibers that push them into metaphase II.
Punnett square. a tool to help calculate the probability of outcomes from matings.
random. random dispersion occurs when the probability of finding an individual
at any point in the area is the same for all points.
recessive trait. traits that are only detectable in the absence of a dominant allele.
spindle fibers. microtubules that connect to chromosomes and pull them apart
during anaphase of either mitosis or meiosis.
telophase. the final portion of mitosis, when separation of paired chromatids is
complete and nuclei reform.
telophase I. the final portion of meiosis I, when separation of homologous
chromosomes is complete and immediately prior to prophase II of meiosis II.
telophase II. the final portion of meiosis II, when separation of paired chromatids
is complete and nuclei reform.
tetraploid. cells have four copies of each chromosome instead of two copies, as
in diploids.
true-breeding. organisms always produce the same trait in every generation.
Index
Alleles, 10 Gametes, xiii, 2
Anaphase, 34 formation of, 37–43
Anaphase I, 38 ethical, legal, social implications
Anaphase II, 39 of, 41–43
meiosis I, 38–39
Bacteria reproduction, 27 meiosis II, 39–41
Binary fission, and prokaryotes Genetic engineering humans,
reproduction, 25–29 ethics of, 41–43
Genetic rules, 19–21
Cellular fission, 29 Genetics, 1
Central dogma, 8, 10 Genotypes, 1
Centromere, 33, 40 Germination, 3
Chance, 7
Chromatid, 33, 36, 38 Haploid, xiii, 39
Chromosome, 31–36 Heterozygous, 7
Cotyledons, 3 Homologous chromosomes
Crossed, 3 (homologs), 19, 38
Cystic fibrosis, 42, 43 Homozygous, 7
Cytokinesis, 34, 35, 39 Human Genome Project, 42
Dependent variable, 25 Independent variable, 25

Dihybrid cross, 15 Inheritance patterns
Diploid, xiii, 32 repeating, 7–10
DNA, 25 for two-traits
double-stranded, 40, 32, 33 ethical, legal, social implications
formation, 9 of, 21–24
replication, 27–29, 31, 33, 38 genetic rules, 19–21
Dominant traits, 4, 5 ratios, 15–18
Double-stranded DNA (dsDNA), 32, Inheritance ratios, 6–7
33, 40 variation in, 7
Down Syndrome, 42 Interphase, 35
Escherichia coli (E coli), 25–29, 31, 32 Law of independent

Eukaryotes reproduction, assortment, 19–20
through mitosis and Law of segregation, 19
cytokinesis, 31–36
Meiosis, 19, 2
F1 generation, 4–7 gametes formation after, 37–43
F2 generation, 4–5, 6, 7, 17, 18 ethical, legal, social implications
F3 generation, 17, 18 of, 41–43
Food and Drug Administration meiosis I, 38–39
(FDA), 43 meiosis II, 39–41
50 INDEX
and mitosis, similarities between, Random, scientific meaning of, 21–24

40–41 Recessive trait, 4, 5
variation in, 10–11
Meiosis I, 38–39 Spina bifida, 42
Meiosis II, 39–41 Spindle fibers, 34
Metaphase, 34 Spindles, 33
Metaphase I, 38 Stamen, 2–3
Metaphase II, 39
Microtubules, 34, 40 Telophase, 34
Mitosis, 34, 36 Telophase I, 39
and meiosis, similarities Telophase II, 39
between, 40–41 Tetraploid, 37
Traits
National Safety Council, 22 appearance in later generation,
1–13
Pea plant, life cycle of, 2 dominant, 4, 5
P generation, 4, 5 recessive, 4, 5
Phenotypes, 1 true-breeding, 2–8, 10
Probabilities, tool for calculation two-traits, inheritance patterns
of, 11–13 for, 15–18
Prokaryotes reproduction, binary True-breeding, 2–8, 10
fission and, 25–29 Two-traits, inheritance patterns
Prophase, 34 for, 15–18
Prophase I, 38, 40 ethical, legal, social implications
Prophase II, 39 of, 21–24
Proteins, 29 genetic rules, 19–21
Punnett square, 12, 17 ratios, 15–18
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Reproduction and Cell Division

All rights reserved. No part of this publication may be reproduced, stored

First published in 2016 by

ISBN-13: 978-1-94474-913-2 (print)

Momentum Press Biology Collection

Cover and interior design by S4Carlisle Publishing Services Private Ltd.,

Printed in the United States of America

Traits can Disappear

first generation peas (P) first generation peas (P)

A second generation peas (F1) B second generation peas (F1)

Figure 2 Mendel’s first breeding results. A, Pollen from true-breeding

Traits can Disappear and Reappear in a Later Generation 5

F1 peas (second generation)

F2 peas (third generation)

Figure 3 Mendel’s second experiments started with yellow F1 peas

Table 1 Results from mating true-breeding plants that produced

Source: Data from Mendel.

reappeared in the F2 generation after the mating of two true-breeding

Mendel Discovers Inheritance Ratios

Table 2 Results from mating true-breeding plants that produced

Source: Data from Mendel.

Table 3 Variation in F2 pea phenotypes from individual matings for

Mendel Discovers Variation from Ratios

Mendel Finds Repeating Pattern of Inheritance

100% 100% 100% yellow

Figure 4 Mendel’s F3 results from F2 pea plants. For illustration

Figure 5 Combining Mendel’s observations with DNA information.

analyzing overall averages, even though individual plants deviated from

Meiosis Contributes to Variation

Figure 6 Chance events cause variations from the calculated ratio

Chance events such as flipping coins or combining gametes are de-

Tool for Calculating Probabilities

Figure 7 A Punnett square. Genotype and phenotype outcomes for

modern knowledge, it is possible to understand the mechanism shown in

Table 4 Ratio of pea genotypes for n future generations.

Source: Original art.

das Jahr 1865. Abhandlungen, 3–47, 1901. Minor corrections by

Inheritance Patterns for

Mendel Discovers Two-Trait Ratios

Figure 8 Two generations after mating YYSS 3 yyss plants. The

each phenotype in Figure 8 and counting the different phenotype boxes

Figure 9 Punnett square for F2 peas in Figure 8. Genotypes for

Figure 10 F3 progeny from smooth, yellow (light gray) F2 peas.

Mendel wanted to be sure he had discovered fundamental rules in ge-

96 plants 102 plants

Figure 11 F3 progeny from self-pollenated F2 peas. A, 101 wrinkled,

YySs YySs 25% y

Mendel Establishes Genetic Rules

diversity in gamete genotypes. Without knowing anything about meio-

Ethical, Legal, Social Implications: The Scientific

Figure 13 Which pattern of dots best represents randomness?

time and a different outcome 25% of the time? Or do random mecha-

Table 5 Odds of dying by different causes.

If the odds of being suffocating by a plastic bag were to double, it

Ethical, Legal, Social Implications: The Scientific Meaning

DNA is an ancient molecule and is the genetic material inside every

Figure 14 Effect of DNA replication on cell size. Mutant E. coli

Figure 15 Photograph of a man’s chromosomes, organized by size

chromosome carries a different amount and sequence of double-stranded

The partitioning of replicated chromosomes to the newly formed

Step 1. Cells condense their chromosomes and begin to move to the