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 Ethnobotany
Forestry
Horticulture
Photosynthesis and Respiration
Plant Biotechnology
Plant Cells and Tissues
Plant Development
Plant Diversity
Plant Ecology
Plant Genetics
Plant Nutrition
 Carl-Erik Tornqvist

Series Editor
William G. Hopkins
Professor Emeritus of Biology
University of Western Ontario
Plant Genetics

Copyright © 2006 by Infobase Publishing

All rights reserved. No part of this book may be reproduced or utilized in any form
or by any means, electronic or mechanical, including photocopying, recording, or by
any information storage or retrieval systems, without permission in writing from the
publisher. For information contact:

Chelsea House
An imprint of Infobase Publishing
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New York NY 10001

Library of Congress Cataloging-in-Publication Data

Tornqvist, Carl-Erik.
Plant Genetics / Carl-Erik Tornqvist.
p. cm. — (The green world)
ISBN 0-7910-8563-5
1. Plant genetics—Juvenile literature. I. Title. II. Green world (Philadelphia, Pa.)
QK981.T67 2006
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 Introduction vii

1 History of Plant Genetics 2

2 Overview of Genetics 16

3 Meiosis, Mitosis, and Alternation

of Generations 28

4 Polyploid Plants 40

5 The Diversity of Plants:

Nature’s Palette of Genes 52

6 Genetically Controlled Processes 62

7 Identifying Functions of Genes 74

8 Agricultural Advances 82

9 Plant Genomics and Beyond 92

Glossary 102
Bibliography 114
Further Reading 115
Index 117
 By William G. Hopkins

“Have you thanked a green plant today?” reads a popular bumper sticker.
Indeed, we should thank green plants for providing the food we eat, fiber for
the clothing we wear, wood for building our houses, and the oxygen we breathe.
Without plants, humans and other animals simply could not exist. Psycholo-
gists tell us that plants also provide a sense of well-being and peace of mind,
which is why we preserve forested parks in our cities, surround our homes
with gardens, and install plants and flowers in our homes and workplaces. Gifts
of flowers are the most popular way to acknowledge weddings, funerals, and
other events of passage. Gardening is one of the fastest-growing hobbies in
North America and the production of ornamental plants contributes billions
of dollars annually to the economy.
Human history has been strongly influenced by plants. The rise of agricul-
ture in the Fertile Crescent of Mesopotamia brought previously scattered
hunter-gatherers together into villages. Ever since, the availability of land
and water for cultivating plants has been a major factor in determining the
location of human settlements. World exploration and discovery was driven
by the search for herbs and spices. The cultivation of New World crops—sugar,
vii
viii INTRODUCTION

cotton, and tobacco—was responsible for the introduction of slavery to

America, the human and social consequences of which are still with us. The
push westward by English colonists into the rich lands of the Ohio River
Valley in the mid-1700s was driven by the need to increase corn production
and was a factor in precipitating the French and Indian War. The Irish Potato
Famine in 1847 set in motion a wave of migration, mostly to North America,
that would reduce the population of Ireland by half over the next 50 years.
As a young university instructor directing biology tutorials in a classroom
that looked out over a wooded area, I would ask each group of students to
look out the window and tell me what they saw. More often than not, the
question would be met with a blank, questioning look. Plants are so much
a part of our environment and the fabric of our everyday lives that they
rarely register in our conscious thought. Yet today, faced with disappearing
rainforests, exploding population growth, urban sprawl, and concerns about
climate change, the productive capacity of global agricultural and forestry
ecosystems is put under increasing pressure. Understanding plants is
even more essential as we attempt to build a sustainable environment for
the future.
THE GREEN WORLD series opens doors to the world of plants. The series
describes what plants are, what plants do, and where plants fit into the
overall circle of life. Plant Genetics is a detailed look at the genetics of
plants, through historical scientific achievements, discussions of genetic
diversity, agricultural practices, and comparisons to animal and human
genetics. Readers will learn about the origins of important crops and
how plant breeders use genetics to improve crops. Details of some widely
used methods in biotechnology are also included to demystify genetic
engineering.

William G. Hopkins
Professor Emeritus of Biology
University of Western Ontario
 History of Plant Genetics

2
History of Plant Genetics

GREGOR MENDEL
Gregor Mendel, known as the father of genetics, grew up in the
early 1800s in Austria with a curiosity for math and science. At
an early age, Mendel showed interest in the way things worked
and how nature behaved. He worked on his family’s farm until
he was allowed to attend a special type of high school. Later,
after finishing his secondary education, Mendel attended the
University of Vienna. During his time in school, Mendel worked
various jobs to pay for his tuition and living expenses. Even-
tually, Gregor Mendel became a monk in the monastery of
St. Thomas in Brno, a city in what is now the Czech Republic.

Heredity and Garden Peas

During his time at the monastery, Mendel had a fascination with
how different characteristics of plants were passed on from
generation to generation—a process known as heredity. Heredity
is the passing down of genetic information from parent to off-
spring and is the basis for the science of genetics. Mendel’s
favorite plant to work with was the garden pea. By carefully
observing his garden, Mendel identified seven different charac-
teristics, or traits, of the peas, each of which came in two forms.
The seven traits Mendel observed were pod color, pod shape,
seed color, seed shape, flower color, flower position on the stems,
and plant height. On a given pea plant, the pod and seed color
were either yellow or green. The flower color was either white or
purple. The seed shape was either smooth or wrinkled. The pod
shape was either full or pinched. The plant height was either tall
or short. The flowers were found either at the tip of the branches
or all along the branches. Mendel looked at all of these traits
when describing the characteristics of the garden pea.
The seeds of Mendel’s pea plant varieties gave rise to plants
that had identical characteristics to the parents through each
generation. The plants that produced this type of seed were
called true breeding because the traits could be counted on to stay

4
 History of Plant Genetics 5

the same through each generation. The process of combining

hereditary information from flower parents to form seeds is
called fertilization. In fertilization, pollen, which carries the male
reproductive cells, is transferred to the stigma, the female repro-
ductive counterpart, in the flower and the creation of a seed is
initiated (Figure 1.1).
In both plants and humans, the first cell to be formed as
a result of fertilization is called a zygote. The first group of
multiple cells is called an embryo. In humans, an embryo grows
by adding cells, thereby becoming a fetus and then a baby. In
plants, the embryo also accumulates cells but then becomes a
dormant seed, which when germinated, starts to grow into a
seedling. The earliest stage of fertilization in plants can be
compared to the same process in humans. The human male
reproductive cells, called sperm, come in contact with the
female reproductive cell or ovum (egg), starting with zygote and
then embryo development.
The resulting seeds from fertilization will grow into a plant
that has genetic material from both of the parents. For the true
breeding plants, the flowers are on the same plant, with either
both male and female reproductive organs on one flower or on
two separate flowers. Mendel’s curiosity of the true breeding
pea plants led him to hypothesize what would happen if he
took pollen from one true breeding plant and applied it to the
stigma of a flower on a true breeding plant with different traits
than the male plant. The process Mendel used to combine the
hereditary material of one plant with another plant is called
cross-fertilization (or “crossing”).
When pollen from a plant that was true breeding was crossed
to another plant with different traits, Mendel found that certain
crosses would always lead to offspring (i.e., the plants that grew
from seeds made by the previous generation of pea plants) with
characteristics that were both like and unlike those of either
parent plant. These traits, he noticed, always appeared in the
6 Plant Genetics

Figure 1.1 Fertilization is the process of combining hereditary information from

flower parents to form seeds. In this process, pollen is transferred to the stigma in
the flower and the creation of a seed is initiated.

same ratio, or relative numbers. Mendel wondered what would

happen if he were to mix the different types of pea plants
together. Because traits of offspring were believed to come about
by mixing of genetic information from the two parents, Mendel
wanted to test this hypothesis by seeing if pea plants resulting
from crosses would show new traits, meaning traits not seen in
either of the parent plants. If the progeny, or offspring, of this
cross-fertilization had traits such as lavender-colored flowers,
for example (mixing of white and purple), then Mendel could
conclude that the parental traits were passed on equally and
 History of Plant Genetics 7

blended in the offspring. Instead, Mendel found the offspring

had flowers that were always either white or purple, but never
lavender. Thus the traits were not blended.
This observation led Mendel to conclude that the trait of a
certain form, for example green seed color, was dominant to
another form and would mask, or cover, the characteristic of the
recessive form. The terms dominant and recessive refer to differ-
ent forms, called alleles, of a trait. In Mendel’s time, traits were
believed to come from unknown hereditary material. We now
know that this material is a gene. In humans, animals, and many
plants, there are two alleles for each gene. Through many crosses,
Mendel came to the conclusion that the seven traits could be
traced to one gene each, made up of two alleles, which are either
dominant or recessive. According to Mendel, if a plant has either
one allele or two alleles of the dominant form of a gene, the plant
shows the characteristic of the dominant allele and never of
the recessive allele. However, if the plant has two alleles of the
recessive type, then the plant shows the characteristic of the
recessive allele for the given trait because no other form of
the gene is present. The state of having both a dominant and
a recessive allele of the same gene is called heterozygous. The
state of having two of the same type of allele, either dominant
or recessive, is called homozygous. By analyzing the characteristics
of offspring and diagramming the crosses he made, Mendel was
able to determine which seven of the 14 traits were dominant and
which seven were recessive. Mendel concluded that tall plants,
yellow seed color, green pod color, round seed coat, smooth pod,
purple flower color, and flowers located all along the stem are the
dominant forms of the respective trait.
Based on his observations, Mendel could make two gener-
alizations about inheritance, or the passing down through
generations, of traits. These generalizations have come to be
known as Mendel’s laws of heredity. The first law, called the law
of segregation, states that traits are passed on through hereditary
8 Plant Genetics

material for which each trait has two forms and, during formation
of reproductive cells, each cell gets only one form of the heredi-
tary material. The second law, called the law of independent
assortment, states that alleles of genes segregate, or separate,
independently of other genes’ alleles when the parent plant
makes reproductive cells (Figure 1.2).
Even though Mendel made many brilliant observations, the
meaning of his work would not be known until 60 years later, in
1900. Carl Correns, Erich von Tschermak, and Hugo de Vries
rediscovered Mendel’s published articles on his pea plant exper-
iments. These European scientists repeated Mendel’s exact

Mendel’s Experiments with Crosses

Mendel followed seven different traits in his garden peas from generation
to generation. The genetic inventory of Mendel’s garden varied to the point
that pairs of plants had all seven to zero traits in common. In essence,
Mendel could pick and choose plants with precise traits with which to
perform crosses. Using this genetic resource to perform crosses, Gregor
Mendel discovered his laws of dominance, segregation, and independent
assortment.
The monohybrid crosses consisted of two parent plants with only one
different trait—height. This allowed Mendel to discover the principles of
dominance and segregation. The presence of one allele from a tall plant
and one from a small plant in the offspring of a cross between true-breed-
ing tall and small plants resulted in tall plants. Even though there was one
small allele in all of the heterozygous offspring, the phenotype was always
a tall plant. This observation led Mendel to his law of dominance: that some
alleles of genes could mask, or cover, the presence of the other allele. When
Mendel allowed the heterozygous peas to self-fertilize, the next generation
was comprised of both tall and small plants, but there were three times
more tall than small plants. This 3:1 phenotypic ratio is common in genetics.
 History of Plant Genetics 9

experiments, performing the same crosses he had performed

more than half a century earlier. The scientists found Mendel’s
results to be reproducible, and they published their own article
in a scientific journal, giving credit to Mendel for his discoveries
from decades before. The scientists brought attention to
Mendel’s work in a new scientific era, an era that would be
accepting of his ideas about heredity.

BARBARA MCCLINTOCK
Thirty years after the rediscovery of Mendel’s laws and a century
after Mendel carried out his experiments with peas, another

Self-fertilization of a plant that is heterozygous for a gene that singly

controls a trait, with one allele dominant and one recessive, will always
result in a segregating population of three-to-one dominant to recessive
phenotype. This last result led Mendel to his law of segregation in which
the two alleles of each gene in a heterozygote separate during production
of reproductive cells.
Mendel’s dihybrid crosses involved parent plants that differed in two
traits. Mendel selected the traits for seed shape and color to follow the move-
ment of the respective alleles during gamete formation. When pollen from
a plant that produced wrinkled, green seeds was germinated on plants that
produced round, yellow seeds, the next generation consisted of all plants hav-
ing round, yellow seeds. When Mendel allowed these plants to self-fertilize,
the result was a population with four different phenotypes segregating in a
ratio of 9:3:3:1. The four seed phenotypes were round/yellow, round/green,
wrinkled/yellow, and wrinkled/green. The new seed traits seen in the last
generation proved to Mendel that alleles of genes move independently of
each other during gamete formation because they made new combinations.
This became Mendel’s law of independent assortment.
10 Plant Genetics

Figure 1.2 Mendel’s experiments with peas evolved into Mendel’s laws
of heredity. Mendel crossbred peas that produced (A) yellow and (B)
green peas. This produced a generation in which the peas were (C) yellow.
By breeding the C generation peas together, Mendel discovered that the
next generation had a mixture of (D) yellow and green peas.
 History of Plant Genetics 11

young plant geneticist from the United States, named Barbara

McClintock, was studying chromosomes, microscopic structures
made of strands of genes from corn (known scientifically as
Zea mays). McClintock was interested in studying the structure
and behavior of the chromosomes in the nuclei of corn cells.
Using corn as a model plant, McClintock was the first to discover
transposons, which are genetic elements that can move about an
organism’s genome, a complete set of chromosomes containing
all of the genes. Transposons are not found in every organism,
but they are common in corn. If you have ever seen Indian corn,
with the mosaic of colored kernels, then you have seen trans-
posons at work (Figure 1.3).

Transposons and Corn

Transposons are small genetic elements that may contain an
entire gene. Transposons get their name from their ability to
undergo transposition, or movement, to new locations in the
genome. Whether containing an entire gene or not, transposons
have the ability to insert themselves in the genome in a random
way. Therefore, a transposition event may result in a transposon
inserting itself within a gene, thereby disrupting the gene’s abil-
ity to reveal a trait. Some transposons can move on their own
with an accompanying enzyme, or catalyst, called transposase.
Others, called disabled transposons, require the presence and
activity of an active transposon to be mobilized. Not all trans-
posons are removed from the genome before moving to another
location; retrotransposons are copied first and the copy is inserted
in a new location, leaving the original copy in its place. So, there
are two methods of transposing: the cut-and-paste method and
the copy-and-paste method.
McClintock discovered a mechanism of mobilizing genes
within a genome, but, since transposons didn’t follow the
Mendelian laws of inheritance, there were many skeptics of
McClintock’s work. Making McClintock’s findings harder to
12 Plant Genetics

Figure 1.3 Transposons are genetic elements that can move about an organism’s
genome. Transposons are not found in every organism, but they are common in corn.
This mosaic of colored kernels in Indian corn shows transposons at work.

impress upon the male-dominated scientific community was the

discrimination against women that characterized the entire
nation throughout most of the twentieth century. Although she
received the highest award from the Genetics Society of America
and served on the executive boards of many scientific organiza-
tions, McClintock’s work was not taken seriously by the larger
scientific community until decades later. Social barriers that kept
women from reaching tenured positions at universities fueled
the scientific community’s rejection of McClintock’s ideas.
It was not until later, when male scientists studying bacteria
also found evidence of transposable elements, that McClintock’s
work on transposons in maize was given the attention it
deserved. Since research on bacteria was deemed important,
 History of Plant Genetics 13

the fact that transposons could be found in this organism

was extremely exciting to the general scientific community.
McClintock soon received awards and became elected to several
high posts in prestigious scientific societies. It was not until 1983,
50 years after her pioneering research on transposable elements
in maize, that McClintock was given her highest honor: she was
chosen as the recipient for the 1983 Nobel Prize in Physiology
or Medicine.

OTHER MODELS OF GENETIC TRANSFER

There are other models of genetic transfer through generations
that do not follow Mendelian principles. Incomplete dominance
gives rise to alleles that are neither dominant nor recessive.
Rather, the combination of the alleles that make a heterozygote
leads to a physical appearance, or phenotype, showing the char-
acteristics of both alleles. Codominance occurs when both alleles
of a gene are dominant and the heterozygous genotype yields a
phenotype that is a blending of the two traits (i.e., unlike that of
either homozygous form). The carnation, for example, is a plant
that expresses codominant alleles in flower color. Both the phe-
notypes of white and red flower color come from homozygous
dominant genotypes. These can be designated WW for white and
RR for red. The genotype of heterozygous dominant, or WR,
leads to a pink flower. Because the genes for producing proteins
that create white pigment are being expressed at the same level
as the genes producing proteins for red color, the resulting flower
looks pink—the color you get when you mix red and white.
These genotypes and the associated phenotypes can be dia-
grammed using a Punnett square, named after Reginald Punnett,
who was also an early geneticist (Figure 1.4).
In 2005, Dr. Susan J. Lolle at Purdue University published a
research article in the journal Nature that described the inheri-
tance of an allele in an Arabidopsis thaliana plant (a small plant in
the mustard family) that had skipped the parent generation; in
14 Plant Genetics

Figure 1.4 Genotypes and the associated phenotypes can be diagrammed

using a Punnett square. This Punnett square shows the possible combi-
nations of offspring that can be found from pure and mixed pea pods.
 History of Plant Genetics 15

other words, the allele was from the grandparent plant. The
fact that the genetic makeup of the current generation of the
Arabidopsis plant in Dr. Lolle’s hands had alleles that could not
have been inherited from either parent, according to Mendel’s
principles, was mysterious. Dr. Lolle and her colleagues are cur-
rently working to decipher this genetic puzzle. Mendel’s genetic
principles have stood the test of time, but with advanced scientific
research, more exceptions to his rules are being discovered.
Gregor Mendel and Barbara McClintock are alike in that the
impacts of both scientists on the scientific community were not
recognized until long after they did their research. Mendel was
far ahead of his time, and his observations have been funda-
mental to our understanding of genetics today. McClintock’s
pioneering work on maize chromosomes and transposition has
been invaluable to the field of cytogenetics.
Mendel’s observations on the genetics of peas explained how
alleles of genes can recombine in different combinations upon
fertilization. The assortment of the alleles among the resulting
gametes, or sex cells, is random, but the location of the alleles is
always at the same position on the chromosome. Using corn
chromosomes, McClintock studied a mechanism in which
genetic elements move about the genome in a random fashion,
leading to new chromosomal compositions with genes in
completely different locations than those found in the parent
or previous generations.
 Overview of Genetics

16
Overview of Genetics

DNA: THE BUILDING BLOCKS OF LIFE

The basic building blocks of life that hold the genetic information
are nucleic acids, which are chemicals found in the cells of all
organisms, from bacteria to sunflowers to humans. Nucleic acids
can be described as having three distinct chemical parts: (1) a
phosphate group, which is a phosphate atom surrounded by
four oxygen atoms; (2) a type of sugar, known as pentose; and
(3) a ring-like molecule, called a base. Two different types of
ring structures characterize the bases: a single five-sided ring
(a pentagon shape) and a double ring consisting of a five- and a
six-sided ring joined at one side (Figure 2.1). The single-ring base
is called a pyrimidine, and the double-ring base is called a purine.
There are two types of nucleic acid molecules crucial for life;
these are deoxyribonucleic acid (DNA) and ribonucleic acid (RNA).
The prefixes deoxyribo- and ribo- describe the type of sugar
molecule attached to the acid. The deoxy- in DNA means that the
sugar in this nucleic acid is missing an oxygen atom (de- means
“without”); the sugar in RNA has that oxygen atom (Figure 2.2).
This simple difference in the chemical composition of DNA
and RNA is important to the structures and characteristics of
these molecules.
DNA and RNA each come in four different types, called
nucleotides, based on how additional atoms are attached at differ-
ent points on the base-ring structure. For DNA, the names of the
four nucleotides are adenine, thymine, cytosine, and guanine.
Adenine and guanine are purines, while cytosine and thymine are
pyrimidines. RNA also has four nucleotides; all but one are the
same as in DNA. Instead of thymine, the fourth RNA nucleotide
is uracil, which is also a pyrimidine. The difference between
thymine and uracil is the presence of an extra chemical compo-
nent called a methyl group, attached to the uracil ring. In order for
multiple nucleotide molecules to fuse together to make a DNA or
RNA chain, the phosphate group from each nucleotide bonds to
the sugar component of the neighboring nucleotide. When all of

18
 Overview of Genetics 19

Figure 2.1 Two different types of ring structures characterize the bases:
a single five-sided ring (pyramidine) and a double ring consisting of a
five- and a six-sided ring joined at one side. This cytosine-guanine
base pair are the nucleic acids that bond in a DNA chain. The atoms
are carbon (green), nitrogen (blue), hydrogen (white), and oxygen (red).
Cytosine is a pyrimidine (bottom), and guanine is a purine (top).

the nucleotides are connected in a chain, the nucleic-acid molecule

forms a sugar-phosphate backbone (Figure 2.3). The properties of
DNA and RNA are not identical, but they share common
nucleotides. Because the nucleotides of DNA and RNA are similar,
we will focus the rest of the discussion in terms of DNA.

DNA Double Helix

In the cell, DNA is in the form of a double helix, which is like a
ladder that has been twisted lengthwise (Figure 2.4). Each side of
the ladder is a chain of connected nucleotides. The helix is double
because the nucleotides are found in pairs. The bases of a pair
are always the same: cytosine pairs with guanine and adenine
pairs with thymine. The bases that pair with each other are called
20 Plant Genetics

Figure 2.2 The structural component of a nucleic acid includes a type of sugar called
a pentose. A pentose has a single five-sided ring structure. An RNA ribose molecule
includes a hydroxyl group (OH on each side of the bottom sides), whereas a DNA
2-deoxyribose molecule does not. The deoxy- in DNA means that the sugar in this
nucleic acid is missing an oxygen atom; the sugar in RNA has that oxygen atom.

complementary bases, which form the rungs of the ladder. The

nucleotides that make up the sides of the ladder are called com-
plementary strands. Because each nucleotide has only one
complementary partner, one can figure out the order, or
sequence, of the second strand if only the sequence of the first
strand is known. When geneticists talk of DNA sequence, they
are referring to the order of the nucleotides on the strand. The
association of the nucleotide pairs is based on hydrogen bonds.
Although hydrogen bonds are not as strong as the phosphate-
sugar bonds of the DNA backbone, this base pairing makes the
double helix of DNA a very strong structure.
 Overview of Genetics 21

Figure 2.3 Multiple nucleotide molecules fuse to make a DNA or RNA

chain by bonding the phosphate group from each nucleotide to the sugar
component of the neighboring nucleotide. When all of the nucleotides
are connected in a chain, the nucleic acid molecule forms a sugar-
phosphate backbone.

Blueprint for Proteins

The nucleotide bases of DNA can be written in shorthand as A,
C, G, and T, for adenine, cytosine, guanine, and thymine, respec-
tively. Geneticists use this notation to write a DNA sequence.
Protein products are made according to the DNA sequence,
22 Plant Genetics

Figure 2.4 DNA is in the form of a double helix, which is like a ladder that has
been twisted lengthwise. Each rung of the ladder is a chain of connected
nucleotides. The helix is double because the nucleotides are found in pairs. The
bases of a pair are always the same: cytosine pairs with guanine and adenine pairs
with thymine. The nucleotides that make up the sides of the ladder are called com-
plementary strands–nucleotide bases (blue-red) project from the outer phosphate
backbone (yellow).

which is transcribed to messenger RNA (mRNA) and then, by way

of transfer RNA (tRNA), translated into amino acids, which are the
building blocks of proteins. There are only 20 amino acids in
nature that combine to make proteins found in all organisms.
The series of amino acids needed to make a particular protein
is determined by codons, the triplet language of RNA. One can
think of codons as “words” that are three letters long; the letters
of the nucleic acid “alphabet” being A, C, G, and T (or U for uracil
in RNA). Since amino acids are translated from RNA, the codon
 Overview of Genetics 23

alphabet that represents them is made up of the letters A, C, G,

and U. Some amino acids have only one codon that represents
them, while others may be represented by up to six different
codons. In addition to amino acids, there are three codons, called
stop codons, that signal the end of translation and one codon that
signifies the start of translation (AUG, which codes for the amino
acid methionine). Stop codons do not code for an amino acid; they
simply signal the translational machinery to stop.

Chromosomes
Mendel thought of genes as pieces of material with information
about how to create certain traits. These genetic elements could be
inherited by offspring, and the traits would appear in certain ratios
that Mendel could predict by knowing the genetic makeup of the
parent plants. It was not until after Mendel’s time that the word
gene would be used to describe the genetic elements, or alleles, that
Mendel had been studying. Geneticists during Mendel’s period
could observe the results of crosses and could determine pheno-
types of plants, but that was as far as they could go. During
McClintock’s days as a young researcher, great strides were taking
place in the field of genetics. Scientists discovered that the genetic
elements Mendel described were genes located on thread-like
structures, called chromosomes, in the nucleus of a cell.
The nucleus, a membrane-bound structure within a cell,
contains all of a cell’s genetic material, or genome. Chromo-
somes are found in twisted bundles that are held together by
nuclear proteins called nucleosomes. Nucleosomes can be thought
of as having a barrel shape, around which DNA is wound, like
string around a spool. Together, the chromosome strands and
the nucleosomes make chromatin, the fundamental structure of
chromosomes (Figure 2.5). It is from these thread-like bundles
that all of the genes, in the form of DNA, are transcribed into
mRNA, which, in turn, is translated into amino acids that join
together to form proteins.
24 Plant Genetics

Figure 2.5 Chromosome strands and nucleosomes make chromatin, the funda-
mental structure of chromosomes. Through these thread-like bundles, all genes in
the form of DNA are transcribed into mRNA, which is then translated into amino
acids that join to form proteins.

Transcription and Gene Expression

Genes are transcribed with the help of DNA-specific and RNA-
specific enzymes and then are translated into amino acids by
specialized enzymes that form complexes, referred to as ribosomal
complexes. When the transcriptional enzymes encounter a gene
to be expressed, the usual binding point of the enzyme complex
on the chromosome is at a site before the actual start of the gene.
The area before the gene is called the promoter, which is a stretch
of sequence that is recognized by the transcriptional machinery
of the cell as the site onto which the RNA polymerase enzyme
should attach. There may be additional sequences before or after
the gene or within introns (the noncoding regions of the gene) that
regulate the expression of the associated gene. These additional
 Overview of Genetics 25

DNA sequences are called enhancers or silencers, depending on

whether gene expression is increased or decreased. The regu-
latory region has a unique sequence that is recognized by
transcription factors. Most transcription factors have two
domains, or sections, that affect the expression of the target
gene. A DNA-binding domain allows the transcription factor
to bind to the actual DNA of the enhancer or silencer. The other
domain is a transcriptional activator that associates with RNA
polymerase enzymes to either activate or suppress transcrip-
tion of the gene. Once the enhancer or silencer region of a gene
is recognized, the appropriate action will occur: either more or
less expression of the gene.

Spotlight on Cytogenetics
The study of cells is called cytology and the study of genes is called genetics.
Hence, the study of genes at the cellular level, or scale, is called cytoge-
netics. Since chromosomes are large clusters of DNA strands that encode
genes, the focus of cytogenetics is chromosome structure and number.
Cytogeneticists use many special skills and tools to study the character of
chromosomes. The techniques used by cytogeneticists who research plants
are the same as those who study animals. Regardless of the organism, a
technique called chromosome painting is used to identify the different
chromosomes in a set. Chromosome painting uses special chemical dyes to
“paint” a pattern on a chromosome. Different dyes “stick” to different parts
of a chromosome, creating a unique pattern for each chromosome in the set.
A cytogeneticist studies the condition of a chromosome by viewing it under a
microscope. Chromosome maps are used to identify the placement of genes
along chromosomes. An ordered display of all of the chromosomes in a cell
is called a karyotype. Cytogeneticists may karyotype the amniotic fluid of a
pregnant woman to make sure that the baby will be healthy. A plant may be
karyotyped to investigate its lineage or ancestral origins.
26 Plant Genetics

Transposons
What makes transposable elements (transposons) unique
enough to be singled out by transposase enzymes for excision, or
cutting-out, is the presence of an inverse-repeat sequence at each
end of the transposon. An inverse-repeat sequence is a series of
about nine to 15 nucleotides (units of DNA represented by the
letters A, T, G, and C) that are immediately followed by the same
sequence but inverted so that the sequence looks like the reverse
of the preceding sequence. These short stretches of DNA are
strictly conserved, meaning they have survived through evolu-
tion, allowing for enzymatic recognition and excision of the
targeted transposon. Once reinserted into the genome, the
genetic information encoded by the transposon is transcribed
into mRNA and translated into amino acids that form a peptide
chain. However, if the transposon is inside the protein-coding
region of a gene, it may disrupt the gene’s expression, leading to
loss of that protein’s function.

GENETIC INFORMATION IN RNA

All organisms hold their genetic code in the form of nucleic
acids, but the meaning of the genetic information can be seen in
the final product: the protein. While all eukaryotes including
plants and humans have DNA genomes, some types of viruses
store their genetic material in the form of RNA. The genes of
these viruses must go through a reverse transcription step to
replicate, which means converting the RNA sequence first to a
DNA template and then back to RNA. Most genes encode for a
protein product, but there are some genes that code for special
RNA molecules that serve specific purposes, such as moving
through the cell or organism, carrying a signal, or interacting
with proteins, DNA, or other RNA molecules.

Alleles
Alleles can be thought of as different protein forms. Mendel
 Overview of Genetics 27

looked at each gene as a package, and the product of that

package could be thought of as a protein. For instance, the
purple flower and white flower situation can be explained by the
fact that the protein for a purple flower allele is more successful
at causing pigmentation than the protein of the white flower
allele. Thus, one can better understand the concept of dominant
alleles because the expression of the trait is due to a dominant,
or overpowering, form of the protein.
Mutations can cause disruptions in the alleles of genes by
changing the DNA sequence. Some mutations cause chemical
changes in nucleotides which, upon replication, result in differ-
ent DNA sequences than on the template strand. The new DNA
strands then pass on the altered genetic information to mRNA.
The sequence of the resulting mRNA may encode different
amino acids, introduce new start or stop codons, or have no
effect on the amino acid sequence. The last outcome is possible
because some amino acids are represented by more than one
codon. For the first two results, novel protein products may arise
or the protein may not be formed. Mutations can change the
form of a gene from dominant to recessive or vice versa. For
example, in humans, the gene for eye color could have one
mutated allele that could lead to the translation of an incorrect
protein or no protein at all. This could cause the mutant form
of the protein to be recessive because the normal allele would
produce the normal, more effective protein. Since the normal
protein is the functioning form, this would mask the effect of the
recessive form and thus would be dominant over the mutant
form. However, if a mutated allele were to become the dominant
form of the gene, then the mutant phenotype would be seen in a
plant that is heterozygous for the wild type, or normal form of a
gene. This type of mutation is called a dominant negative because
the disrupted gene form is expressed over the wild type form.
 Meiosis, Mitosis, and
Alternation of Generations

28
 Meiosis, Mitosis, and
Alternation of Generations

MEIOSIS
The basis of Mendel’s law of independent assortment can be
understood by examining the cellular process of meiosis. Meiosis
is the double cell division event that results in four daughter cells,
each with half of the genetic content of the original mother cell.
The cells of the male and female gametophytes—the pollen
grain and embryo sac—result from the cell divisions of meiosis.
Cell division by way of meiosis is best described by break-
ing down the two divisions into their four stages or phases
(Figure 3.1). The Roman numeral after each phase represents the
division of meiosis.

Figure 3.1 Meiosis is the basis of Mendel’s law of independent assortment. Meiosis is the
double cell division event that results in four daughter cells, each with half of the genetic
content of the original mother cell. Meiosis divides cells through phases—prophase (not
pictured), metaphase, anaphase, and telophase.

30
Meiosis, Mitosis, and Alternation of Generations 31
32 Plant Genetics

The initial state of the cell is prophase I: the chromosomes

have doubled but are not yet distinguishable from each other.
Then, in late prophase, the duplicated sister chromatids pair
up and can be identified because the chromatid strands are
stretched. At this point, the phenomenon known as crossing
over occurs. A crossover event occurs when homologous sis-
ter chromatids attach to each other at points other than the
centromere. Where the sister chromatids join, a break occurs,
allowing chromosomal segments to be swapped between the two
chromatids. Crossing over is a key step in the creation of gametes
with unique genotypes. Next, in metaphase I, the chromosome
pairs, which are joined at the centromere, are centrally aligned
in the middle of the cell, and spindle fibers attach to each chro-
mosome of each pair. In the third phase of meiosis, anaphase I,
the chromosome pairs are pulled apart by the spindle fibers and
are drawn to opposite ends of the cell. In telophase I, the last stage
in meiosis I, the chromosomes are completely at opposite ends
of the cell and cytokinesis, or division of the cytoplasm, follows.
The two resulting cells, each with two copies of the chromosome,
begin the second division of meiosis, or meiosis II.
In meiosis II, the chromosomes go through the same four
stages as in meiosis I, except the chromosomes are not doubled
in prophase II, as they were in prophase I. This difference is what
leads to the final formation of four daughter cells with only half
the chromosome number as the original mother cell. Also, in
telophase II, nuclear membranes form around the chromosomes
in each new daughter cell prior to cytokinesis.

Cytokinesis
Up to the point of cytokinesis, every step in cell division is the
same in animals as it is in plants. The division of the cell’s cyto-
plasm, called cytokinesis, differs between plants and animals.
This difference is most likely due to the presence of both a cell
wall and a plasma membrane around plant cells, and not just a
 Meiosis, Mitosis, and Alternation of Generations 33

plasma membrane, as is the case in animal cells. In human cells,

division occurs by a pinching method that gradually narrows the
cytoplasmic space between the two opposite sides on the division
plane. Eventually, the plasma membranes from either side of the
cell meet in the middle and fuse together, creating two separate
cells. In plant cells, the division process is very different. Follow-
ing telophase, when the chromosomes are at either end of the
mother cell, a new cell wall begins to grow along the division
plane from opposite sides of the cell; ultimately, this cell wall
comes together to produce a solid wall separating the two daugh-
ter cells. When the chromosome pairs are pulled apart, after
crossing over, different pieces of the chromosome pairs can stay
attached to each other, resulting in a break in the chromosome
strand and in more chromosome material on one side and less
on the other. This means that some of the daughter cells would
be deficient in the genes that were on the lost chromosome
pieces. This event would result in the newly arising cells having
abnormal genetic content, as proposed by McClintock. Mendel
saw, without the aid of a microscope, that the process of creat-
ing gametes somehow resulted in new traits found in the progeny
pea plants. It is the resulting cells, or gametes, of meiosis that
merge from both parent plants to form the zygote, the first cell of
the offspring plants.

MITOSIS
Sexual Reproduction and Gamete Formation
The first cell of the progeny is actually a fusion of a haploid
nucleus from the female and from the male plant. Therefore,
the new zygote that will form into an embryo and, in most
plants, eventually a seed has a mixture of the two parents’
genetic makeup. The special cells for reproduction, or
gametes, contain only half of the chromosome number
(haploid) as the rest of the plant. These cells reside in a
specialized part of the plant called the sporangium, within
34 Plant Genetics

Theory of the Breakage-Fusion-Bridge Cycle

Barbara McClintock’s first major pioneering theory was the breakage-
fusion-bridge cycle. By describing the chromosome structure, the concept
of the breakage-fusion-bridge cycle was McClintock’s way of explaining
the presence or absence of genes in the offspring of corn whose reproductive
cells had been mutated by X-rays. X-ray mutations caused whole chromosome
segments to be chopped off and reattached in different ways. Often, broken
chromosomes would attach to homologous partners at the breakage site.
This led to chromosomes that were dicentric, or had two centromeres.
McClintock proposed that, due to their structure and to crossover events,
dicentric chromosomes could be torn apart during meiosis and become rear-
ranged in other ways in the resulting gametes (See Figure).

Figure: The broken end of the chromosome is caused by X-ray bombardment and
is where the two homologous partners fuse. When spindle fibers attach to the two
centromeres and pull to opposite poles of the cell, an anaphase bridge is formed
that eventually breaks under the tension created by the contracting spindle
fibers. The broken sister chromatids are each pulled to opposite poles where
they become part of the chromosome set of the new daughter cells. The broken
ends fuse to homologous chromosomes in the daughter cell in subsequent cell
divisions and the cycle continues.
 Meiosis, Mitosis, and Alternation of Generations 35

which the haploid cells are produced. These haploid cells,

which have only one complete set of chromosomes in the
nucleus, are generated by two consecutive meiotic events and
one round of mitosis. Mitosis is another type of cell division,
but instead of ending up with half of the chromosome number,
as in meiosis, the daughter cells have exactly the same number
of chromosomes as the mother cell. Mitosis is involved in division
of cells that are not reproductive, like in the shoots, roots, and
leaves (Figure 3.2).
In male flowers or in male parts of bisexual flowers, the
microsporocyte is the diploid cell that, through meiosis, gives rise
to four microspores. The four cells complete mitosis to form two
pollen grains each, resulting in a total of eight pollen grains for
each microsporocyte. Each pollen grain consists of two cells: one
called the generative cell and the other called the tube cell.
In the female flower or flower parts, the diploid cell respon-
sible for gamete production is called the megasporocyte. The
megasporocyte goes through meiosis I and II, resulting in four
haploid cells called megaspores. Only one of the four resulting
cells survives to undergo three mitotic divisions. The resulting
female gametophyte, created by three cell divisions of one
megaspore, is multicellular.
Upon pollination of a female flower, the generative cell
divides to become two sperm cells. When a pollen grain lands on
a female flower, the tube cell will elongate and grow down into
the ovary, where the mature, female gametophyte can be
reached. In a process called double fertilization, each sperm cell
releases its nucleus into the embryo sac; one sperm fertilizes
the egg cell that will become the diploid zygote, while the other
fertilizes the two nuclei of the large central cell to form triploid
(3n) endosperm tissue (Figure 3.3). The endosperm tissue serves
as the nutrient source for the plant embryo during seed germi-
nation. The yellow part of a corn kernel is the endosperm, which,
in this case, makes up the majority of the seed.
36 Plant Genetics

Figure 3.2 In mitosis, the daughter cells have exactly the same num-
ber of chromosomes as the mother cell. Mitosis is involved in division
of cells that are not reproductive, like in the shoots, roots, and leaves.
Like meiosis, mitosis has four phases—prophase, metaphase,
anaphase, and telophase.

ALTERNATION OF GENERATIONS
In flowering plants, the gametophyte is in the flower, which
produces the seed; the rest of the plant, which is called the
 Meiosis, Mitosis, and Alternation of Generations 37

Figure 3.3 The life cycle of corn is shown here. Each sperm cell releases
its nucleus into the embryo sac; one sperm fertilizes the egg cell that
will become the diploid zygote, while the other fertilizes two nuclei of
the large central cell to form triploid (3n) endosperm tissue. The
endosperm tissue serves as the nutrient source for the plant embryo
during seed germination. The yellow part of a corn kernel is the
endosperm, which makes up the majority of the seed.
38 Plant Genetics

sporophyte, has a diploid chromosome number. However, in

nonvascular plants such as mosses, the haploid part of the plant
is most of what we see. This transition from haploid to diploid
and again to haploid state is unique to plants and is called
alternation of generations. Alternation of generation means that all
plants have both a gametophytic phase and a sporophytic phase
as part of their life cycle. In flowering plants, the sporophytic
phase can be thought of as the vegetative state, or the state in
which the plant grows branches and produces a lot of leaves. The
gametophytic generation of vascular plants is confined to the
small inconspicuous organs dedicated to the purpose of repro-
duction. In nonvascular plants such as mosses, liverworts, and
hornworts, the gametophytic generation is the green part that we
see; the sporophytic generation is much smaller and less visible
(Figure 3.4).

Other Modes of Reproduction

Plants do not have to undergo fertilization in order to reproduce.
They can also reproduce, or propagate, asexually (i.e., without
fertilization), which is a characteristic known as vegetative repro-
duction. All plant cells have the ability to become any specialized
cell of the plant. This characteristic of plant cells to become any type
of cell is called totipotency. The act of becoming a specialized cell
is called differentiation, and plant cells become differentiated just
after forming in the cell-producing area called the meristem.
The undifferentiated cells found in the meristem are a type of
stem cell, like the stem cells of humans. Recent media coverage
has brought attention to research conducted on two different
forms of human stem cells: those found in embryos and those
found in the bone marrow, or the center of the bone, of adults. The
meristematic cells of plants are more like the human stem cells
found in embryos because, like this type of human stem cell,
plant stem cells can become any type of specialized cell and, thus,
any type of organ. Adult stem cells found in bone marrow, on
 Meiosis, Mitosis, and Alternation of Generations 39

Figure 3.4 Alternation of generation means that all plants have both a gametophytic
phase and a sporophytic phase as part of their life cycle. This carpet moss has
distinct sporophytes growing on gametophytes (the visible green part). Sporophytes
reproduce asexually whereas gametophytes reproduce sexually.

the other hand, can only be conditioned to become certain types

of cells. Unlike human cells, a plant cell can dedifferentiate and
become a cell that is completely different than the type of cell
it was before. Dedifferentiation is the process in which a plant
cell loses all of its specialized structures and organelles, which are
the membrane-bound structures within the cell, and becomes a
simple cell containing only the nucleus and the essential
organelles needed for survival. Scientists can manipulate a
dedifferentiated cell to become a particular cell by exposing it to
various hormones that induce gene expression changes and affect
development of the cell. The genetic basis of these physiological
processes will be discussed further in chapter 6.
 Polyploid Plants

40
Other documents randomly have
different content
 The Project Gutenberg eBook of Souvenir
Map and Guide for Tourists in the Black Hills
of South Dakota
 This ebook is for the use of anyone anywhere in the United
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eBook.

Title: Souvenir Map and Guide for Tourists in the Black Hills of
South Dakota

Creator: S.D. Mitchell Dakota Engineering Company

Release date: September 3, 2019 [eBook #60217]

Most recently updated: October 17, 2024

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*** START OF THE PROJECT GUTENBERG EBOOK SOUVENIR MAP

AND GUIDE FOR TOURISTS IN THE BLACK HILLS OF SOUTH
DAKOTA ***
 EDWARD K. MATHER, C. E. ORIN L. KIPP, C. E.
Managing Engineer Associate Engineer

Dakota Engineering Company

Automobile Map and Guide Department

MAP AND GUIDE OF HIGHWAYS IN THE BLACK HILLS, 50c

MAP AND GUIDE OF THE SCENIC HIGHWAY THROUGH SOUTH

DAKOTA, 50c
SIOUX FALLS-MITCHELL TO RAPID CITY

In Preparation

MAP AND GUIDE OF THE “CAPITOL CITY TRAIL”

SOUTH DAKOTA SECTION OF CHICAGO-YELLOWSTONE PARK
HIGHWAY

And Other Cross State Road Maps and County Road Maps
311 Western National Bank Building
MITCHELL, S. D.
 Souvenir Map and Guide for Tourists in the
BLACK HILLS
of South Dakota

Maps prepared especially for the use of Automobile Tourists

BY THE

DAKOTA ENGINEERING COMPANY

MITCHELL, SOUTH DAKOTA

The Mitchell Pub. Co., Mitchell, S. D. Copyright, 1913, by Edward K. Mather, C. E.
 he possibility of motoring through various sections of the
Black Hills for pleasure and business has long been
known. The idea is new, however, of selecting good
automobile routes to reach the larger cities and points of
scenic interest in this region, and of platting the same as
a guide to the stranger who may be easily lost amid the windings and
branchings of the roads in forest, mining camp or city. The strenuous
efforts of various localities and counties along these routes to
improve them during the last two years has made them now such as
may be travelled with pleasure.
The natural scenery on a scale majestic in size, yet of readily
appreciated magnitude; the proximity of the primitive forest to the
noise and rush of the mining or industrial center; the strange
thoughts on realizing that here lies the richest tract one hundred
miles square on earth, all serve to make the trip through the Black
Hills a memorable one; even to him who has traveled far and near
across ocean, desert sands or broad plains in search of nature’s
beauty spots.
From a personal standpoint, the renewed vigor, and quieted
nerves that result from rest among such pleasant surroundings,
breathing pure air, laden with the odor of the pines, bathing in or
drinking mineral waters all make it a trip well worth while.
It is hoped that to the stranger to the Black Hills region this
book will bring some idea of the pleasure and profit to be found amid
such surroundings, and that it will be an aid to him in the proper
enjoyment of the same.
 Moderate temperature,
winter and summer, abundance of
sunshine and clear, dry air make
CLIMATE the Black Hills particularly
desirable from a tourist’s
standpoint. The altitude ranging from 3500 to 8000 feet above sea
level. Absolute freedom from mosquitoes in many parts of the hills
and cool, dry nights make it an ideal country for camping.

In the wilder portions,

bears, wild cats, wolves, silver
foxes, and other large game may
GAME be found. Special provision has
been made for re-stocking the
Forest Reserve at frequent intervals with deer and elk which,
together with legal protection for these animals from December 1st
to November 1st makes them very plentiful. Plover, partridges,
pheasants, grouse, ducks and geese may be easily had in season.
Although many kinds of fish are found in the mountain streams,
trout are the most common. The streams are restocked at frequent
intervals from the Government Fishery at Spearfish with Brook,
Locklaven, Speckled and Rainbow trout, so that fishing will continue
to be one of the leading diversions of the pleasure seeker. From
November 1st to May 1st when trout fishing is prohibited in South
Dakota, the sport may be continued across the line in Wyoming
where no such restriction exists.

Residents of other states

using their cars in South Dakota
will not be required to re-register
SOUTH their cars in South Dakota
DAKOTA providing they have complied
LAWS with all laws regarding
registration of automobiles in the
CONCERNING state from which they came, and
AUTOMOBILE providing such state extends a
similar privilege to South
S Dakotans.
Automobiles must be
driven in a careful and prudent manner, at a rate of speed such as
not to endanger the property, life or limb of any person, provided
that a rate of speed in excess of 25 miles per hour is presumptive
evidence of driving at a rate of speed which is not careful and
prudent, in case of injury to the person or property of another. Local
authorities may limit speed to a rate of 10 miles per hour provided
proper notices or warnings are placed in the public highway
indicating such limitation of speed.
The raising of the hand, by anyone who is leading, driving or
riding a horse or other draft animal, as a signal to stop shall cause
the driver of any automobile thus signalled to, to stop his car until
the danger of frightening such animal is past.

Perhaps the best known city in the Northern Hills is

Deadwood, the center of the industrial and mining interests of this
 section. Most excellent hotel and
garage accommodations, its
convenience to surrounding
DEADWOOD points of scenic interest, and a
general spirit of genuine western
hospitality toward all will continue to make it a most popular tourist
center. Although high in the Hills it is connected with the plains
outside by a good road of easy gradient.

At the point of entrance to

the Black Hills from the east,
Rapid City has become a leading
RAPID CITY industrial and railway center.
Good hotel and garage
accommodations make this a pleasant stopping place in the foothills.
The trip westward into the Hills along Rapid Creek to Pactola by
auto, or by rail to Mystic, should not be overlooked by any tourist in
this territory.

Spearfish is located on the

northern margin of the Hills,
close to the mouth of Spearfish
SPEARFISH canyon, to Sand Creek, to Higgins
Gulch, Crow Peak and other
fishing and hunting resorts. It has good hotel and garage
accommodations, and is visited by thousands of tourists each year.

Twenty-eight miles north of Deadwood and close to the

100,000 acre government irrigation project is a thriving county seat
town of 3000 population. It has been a noted shipping point for
those interested in cattle and sheep raising in years gone by and is
 now becoming the principal
center of business for the adjacent
irrigated districts.
BELLE
FOURCHE

A thriving village at the

point of entrance into the Hills
which will be remembered by the
WHITEWOOD tourist going by rail or auto to
Deadwood, Lead, Spearfish or
other points in the northern Hills.

This has been one of the

leading towns in the heart of the
Black Hills district since the early
HILL CITY days of the Black Hills history.
Numerous tin, mica and other
mines and considerable rich agricultural land are found near here.
About 45 miles from Deadwood, Rapid City and Hot Springs it is a
convenient stopping place for tourists travelling through this
territory.

The largest gold mine in

the world, employing an average
of 3500 men and producing from
THE $5,000,000 to $8,000,000 worth
HOMESTAKE of gold bullion per year, has its
MINE principal properties and mines at
Lead.
The Elliston Hoist, the
largest now operated by the company, is 1850 feet deep. The 1200 H.
P. hoisting engine with its hoisting rope, a 7¾ inch by ⅝ inch flat
laced steel cable weighing 7 pounds to the foot, lifting a load of 10
tons, not including the weight of the cable from the bottom of the
mine to the surface in a few seconds, is an interesting sight. Nearby
are two air compressors, one delivering air at 80 pounds pressure for
the use of rock drills and other machinery; the other compressing air
to 850 pounds pressure for use of compressed air haulage motors.
Across the gulch the drill sharpening shop where 3000 drills
are sharpened each day, and the foundry and machine shop are
points of interest. The noise of operation of the stamp mill night and
day, 365 days in the year make its location easily determinable. Here
240 stamps each weighing 900 pounds and dropping at the rate of
90 times per minute crush to a powder 1000 tons of ore each 24
hours. To each ton of powdered rock is added 2400 gallons of water
which wash it over the silver plated copper plates. In another
building it is re-ground, and the coarsest portion or sands sent to the
Cyanide Plant. Here solution is accomplished by the addition of
potassium cyanide, and the gold then precipitated by the addition of
zinc dust. The finer portion of the powder or the slime is piped to the
Slime Plant at Deadwood, where the gold bearing powder is caught
in filter presses and the gold then extracted as before.
The Homestake Pumping Plant at Hanna, 6 miles from Lead,
and the Hydro-electric plant at Spearfish, 15 miles from Lead,
furnish the water and power used in the Homestake properties. The
Hydro-electric Plant is the largest of its kind in the Hills. The water is
carried in a tunnel 5 miles long from the river in Spearfish Canyon, at
a point 8 miles from Spearfish, to the reservoir on top of the ridge
overlooking and directly south of the city. The three surge towers, on
the pipe lines leading from the reservoir down the 700 foot drop to
the power plant, may be seen for many miles.

There is probably no other

scenic place in the Black Hills
which has been given as much
SPEARFISH attention by tourists,
CANYON photographers, and magazine
writers as Spearfish Canyon. The
Royal Gorge in Colorado, the Palisades of the Hudson and many
other points of national scenic wonder may be found reproduced
here on a scale perhaps less grand, but making up in beauty all that
is lost in magnitude. Good automobile roads traverse long stretches
of this canyon, entering it from Deadwood, and from Spearfish as
platted herein. It is anticipated that in a few years the road will be
made suitable for auto traffic the entire length of the canyon.

Crystal Cave, in the

Northern Hills, is resplendent
with stalactite and stalagmite
CRYSTAL formations and is quite different
CAVE in general structure from Wind
Cave. During 1913 it is expected
that the construction of a new
road from Deadwood will make this cave accessible to automobile
tourists.

One of the most interesting

places in the Hills from a scenic
point of view is located about
SYLVAN LAKE midway between Deadwood and
THE NEEDLES Hot Springs. Harney Peak, the
HARNEY highest point in the Hills from
which can be seen four states, the
PEAK hundreds of Needles, some rising
as much as 500 feet in cathedral
tower like grandeur, the great varied vistas, and the quiet beauty of
Sylvan Lake surrounded by monstrous rock walls and pine clad
heights, all bring to one the realization of the real wonder and beauty
of nature. Excellent hotel accommodations at Sylvan Lake makes this
region justly popular as a tourist resort.
 This is a typical Black Hills
stream in Wyoming close to
Deadwood and Spearfish. The fact
SAND CREEK that Wyoming laws allow trout
fishing at all seasons of the year
makes this stream the rendezvous of the fisher both summer and
winter.

One of the great sights of

this region, in many respects
excelling the famous Mammoth
WIND CAVE Cave of Kentucky, is Wind Cave.
Over 200 miles of passageway
and many thousands of rooms have been explored, some of these
over 600 feet below the entrance. Yet, apparently neither the bottom
nor the sides of this cave have been reached. The great abundance of
“frost work” and “box work” in many forms make this cave unusually
attractive and interesting. Guides will take parties into the Cave each
day at 9:00 A. M. and 2:00 P. M. and occasionally at other times.

As a health resort the

Southern Hills have gained a
National reputation. A sanitarium
HEALTH for treatment of tuberculosis
RESORTS patients has been established by
the State near Custer, 29 miles
north of Hot Springs. At the latter
city is the State Home for Old Soldiers, and the Government Hospital
for Disabled Volunteer Soldiers and Sailors. Also numerous private
sanitariums, for the treatment of various maladies, which are filled
throughout the year.
 The city gets its name from
the hundred springs near there
which give out large quantities of
HOT SPRINGS water at a temperature of 98
degrees F. or warmer. The water
from these and from certain cold springs have been found to contain
unusual mineral elements very valuable in the treatment of
rheumatism of all kinds, nervous complaints, indigestion and
intestinal disorders, pulmonary affections, diseases of the urinary
organs, gout and skin diseases. Particularly noticeable have been the
curing of parties affected with rheumatism.
Besides the small swimming pools at the various sanitariums,
two immense pools have been constructed which are centers of
attraction during the summer months. These are the “Evans” and the
“Mammoth Springs Plunge,” both of which are enclosed in large
buildings, and contain individual dressing rooms and all other
possible conveniences.
One of the largest and finest equipped hotels in the state is
located here.

Edgemont is a busy railway

division point on the C. B. & Q.
Ry. It is located on the Cheyenne
EDGEMONT River southwest of the Black Hills.
To those who have seen the
prairies of eastern South Dakota, the Bad Land and Black Hills
formations, a new type of topography is presented extending to the
south and west of Edgemont.

East of the Black Hills, and within easy reach, is scenery,

somewhat mountainous in nature, yet very different from anything
in the Black Hills or in other mountainous regions. To anyone who
 has not travelled the “Scenic
Highway through South Dakota,”
a two days trip eastward from
THE BAD Rapid City over this road is well
LANDS worth while.
The tourist enters the Bad
Lands near the town of Scenic, 45 miles east of Rapid City. From this
point the next 50 miles is in the midst of some of the strangest and
most wonderful scenery known. Broad grass covered basins are
dotted with bare rounded hillocks or great irregular shaped buttes
and ridges. Along the northern border of this region and much of the
time within sight of the road is “The Great Wall” several hundred feet
in height, appearing against the sky line to be a succession of domes,
towers, pinnacles and precipitous walls and gulches. The soil varies
in color through the shades of white, buff, yellow, red and green. The
valleys and flat plateaus on the tops of the ridges or buttes are
usually grass covered while the slopes stand out most prominent
because of their sheer height and nakedness. It is a scene that cannot
be properly described; strange and wonderful in the extreme. At first
seemingly grotesque, then strangely beautiful, impressions are left
on the mind that will never be forgotten.
Pictures and more detailed description of this region, also a
complete guide of the road from Rapid City eastward will be found in
our “_Map and Guide of the Scenic Highway through South
Dakota_.”
No tourist should leave the Black Hills until this trip has been
taken either by automobile or rail. In either case he should go as far
east as the towns of Interior or Kadoka, and actually go through
Cedar Pass which is five miles northeast of Interior.
 RAPID CITY

apid City, the gateway to the Hills, lies on Rapid Creek

for which it is named, midway between its source in the
Western Black Hills and its mouth, where it empties into
the Cheyenne river. Its location is ideal from the
standpoint of natural environment embracing, as it
does, the rugged backbone of the hills and a foreground of rolling
prairie. In early days pioneers were quick to see the natural
advantages of the location for a town and their selection proved the
later choice of railroad engineers, who have made it the central point
for two great systems in western South Dakota. Four distinct lines
radiate from Rapid City, viz: The Omaha division of the
Northwestern providing connection with Deadwood on the North
and the great South and Southeast. The Pierre division running to
the state capital and Chicago. The Milwaukee has its terminal here. It
runs southeast through the Bad Lands, the only railroad traversing
this Wonderland of Nature, and on to Chicago. The Rapid City, Black
Hills & Western has its headquarters in Rapid City, and is one of the
most wonderfully constructed railroads in the country. It follows
Rapid Creek west to Mystic, about thirty-five miles where it connects
with the Burlington system. This is called the “Scenic” route of the
Hills and affords tourists one of the grandest of pictueresque views to
be had in the world. It is Colorado and the Grand Canyon in
miniature.
 Located as it is in a country blessed by nature Rapid City
provides more amusement to lovers of the out door life than most
cities. With a climate ofttimes as equitable in January as that of
Southern California and with rainfall usually confined to the spring
months out of door life finds here its perfect environment. The
nights, owing to the altitude of nearly thirty-two hundred feet, are
always cool, there being few in the hottest of the summer months
when blankets are not needed. The absence of dew makes camping a
delightful pastime and there is trout fishing, for those who enjoy the
gentle art of Isaac Walton, as good as the country affords anywhere.
Deer are plentiful in season and game birds, including grouse,
partridge, quail and duck, are plentiful enough to make it worth the
while of any follower of Nimrod.
Rapid City itself being thus favorably located is the center of
business activity radiating east as far as the Missouri river and west,
north and south to the boundaries of the Black Hills. It has the three
requisites of a progressive and up-to-date growing city—first-class
hotels, a new up-to-date theatre and a live newspaper. From the city
all of the interesting points in the Hills can be reached by railroads,
or by automobiles over roads which are being constantly improved.
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