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Chapter Ten: DNA: The Chemical Nature of the Gene
COMPREHENSION QUESTIONS

Section 10.1

1. What four general characteristics must the genetic material possess?

Solution:
(1) The genetic material must contain complex information.
(2) The genetic material must replicate or be replicated faithfully.
(3) The genetic material must have the capacity to vary or mutate to generate
diversity.
(4) The genetic material must encode the phenotype or have the ability to code for
traits.

Section 10.2

2. Briefly outline the history of our knowledge of the structure of DNA until the time
of Watson and Crick. Which do you think were the principal contributions and
developments?

Solution:
1869: Johann Friedrich Miescher isolates nuclei from white blood cells and extracts
a substance that was slightly acidic and rich in phosphorous. He calls it
nuclein.
Late 1800s: Albrecht Kossel determines that DNA contains the four nitrogenous
bases: adenine, guanine, cytosine, and thymine.
1920s: Phoebus Aaron Levine discovers that DNA consists of repeating units, each
consisting of a sugar, a phosphate, and a nitrogenous base.
1950: Erwin Chargaff formulates Chargaff’s rules (A = T and G = C).
1947: William Ashbury begins studying DNA structure using X-ray diffraction.
1951–1953: Rosalind Franklin, working in Maurice Wilkins’ lab, obtains higher
resolution pictures of DNA structure using X-ray diffraction
techniques.
1953: Watson and Crick propose the model of DNA structure.
All of these scientists contributed information that helped Watson and Crick
determine the structure of the DNA double helix. Erwin Chargaff and Rosalind
Franklin made two important contributions that directly led to the discoveries by
Watson and Crick. By combining Chargaff’s rules with Rosalind Franklin’s X-ray
diffraction data, Watson and Crick were able to predict accurately the structure of
the DNA double helix.

3. What experiments demonstrated that DNA is the genetic material?


228 Chapter Ten: DNA: The Chemical Nature of the Gene

Solution:
Experiments by Hershey and Chase in the 1950s using the bacteriophage T2
demonstrated that DNA is the genetic material of the bacteriophage. Also, the
experiments by Avery, Macleod, and McCarty demonstrated that the transforming
material initially identified by Griffiths was DNA.

4. What is transformation? How did Avery and his colleagues demonstrate that the
transforming principle is DNA?

Solution:
Transformation occurs when a transforming material (or DNA) genetically alters
the bacterium that absorbs the transforming material. Avery and his colleagues
demonstrated that DNA is the transforming material by using enzymes that
destroyed the different classes of biological molecules. Enzymes that destroyed
proteins or RNA had no effect on the activity of the transforming material.
However, enzymes that destroyed DNA eliminated the biological activity of the
transforming material. Avery and his colleagues were also able to isolate the
transforming material and demonstrate that it had chemical properties similar to
DNA.

5. How did Hershey and Chase show that DNA is passed to new phages in phage
reproduction?

Solution:
Hershey and Chase used the radioactive isotope 32P to demonstrate that DNA is
passed to new phage particles during phage reproduction. The progeny phage
released from bacteria infected with 32P-labeled phages emitted radioactivity from
32
P. The presence of the 32P in the progeny phage indicated that the infecting phage
had passed DNA on to the progeny phage.

6. Why was the discovery of DNA structure so important?

Solution:
By deciphering the structure of the DNA molecule, Watson and Crick provided the
foundation for molecular studies of the genetic material or DNA, allowing scientists
to discern how genes function to produce phenotypes. Their model also suggested a
possible mechanism for the replication of DNA that would ensure the fidelity of the
replicated copies.

Section 10.3

7. Draw and identify the three parts of a DNA nucleotide.

Solution:
The three parts of a DNA nucleotide are phosphate, deoxyribose sugar, and a
nitrogenous base.
Chapter Ten: DNA: The Chemical Nature of the Gene 229

8. How does an RNA nucleotide differ from a DNA nucleotide?

Solution:
DNA nucleotides, or deoxyribonucleotides, have a deoxyribose sugar that lacks an
oxygen molecule at the 2′ carbon of the sugar molecule. Ribonucleotides, or RNA
nucleotides, have a ribose sugar with an oxygen linked to the 2′ carbon of the sugar
molecule. Ribonucleotides may contain the nitrogenous base uracil, but not
thymine. DNA nucleotides contain thymine, but not uracil.

9. How does a purine differ from a pyrimidine? What purines and pyrimidines are
found in DNA and RNA?

Solution:
A purine consists of a six-sided ring attached to a five-sided ring. A pyrimidine
consists of only a six-sided ring. In both DNA and RNA, the purines found are
adenine and guanine. DNA and RNA differ in their pyrimidine content. The
pyrimidine cytosine is found in both RNA and DNA. However, DNA contains the
pyrimidine thymine, whereas RNA contains the pyrimidine uracil but not thymine.

10. Draw a short segment of a single polynucleotide strand, including at least three
nucleotides. Indicate the polarity of the strand by labeling the 5′ end and the 3′ end.
230 Chapter Ten: DNA: The Chemical Nature of the Gene

Solution:

11. Which bases are capable of forming hydrogen bonds with each another?

Solution:
Adenine is capable of forming two hydrogen bonds with thymine. Guanine is
capable of forming three hydrogen bonds with cytosine.

12. What different types of chemical bonds are found in DNA and where are they
found?

Solution:
The deoxyribonucleotides in a single chain or strand of DNA are held by covalent
bonds called phosphodiester linkages between the 3′ end of the deoxyribose sugar
of a nucleotide and the 5′ end of the deoxyribose sugar of the next nucleotide in the
chain. Two chains of deoxyribonucleotides are held together by hydrogen bonds
between the complementary nitrogenous bases of the nucleotides in each chain.

13. What are some of the important genetic implications of the DNA structure?

Solution:
Referring back to Question 1, the structure of DNA gives insight into the four
fundamental genetic processes. The Watson and Crick model suggests that the
complex genetic information or instructions are encoded in the nucleotide
Chapter Ten: DNA: The Chemical Nature of the Gene 231

sequences. The complementary polynucleotide strands indicate how faithful


replication of the genetic material is possible. Finally, the arrangement of the
nucleotides is such that they specify the primary structure or amino acid sequence
of protein molecules.

14. What are the three major pathways of information that flow within the cell?

Solution:
The major transfers of genetic information are replication, transcription, and
translation. These are the components of the central dogma of molecular biology.

Section 10.4

15. What are hairpins and how do they form?

Solution:
Hairpins are a type of secondary structure found in single strands of nucleotides.
The formation of hairpins occurs when sequences of nucleotides on the single
strand are inverted complementary repeats of one another.

16. What is DNA methylation?

Solution:
DNA methylation is the addition of methyl groups (–CH 3 ) to certain positions on
the nitrogenous bases on the nucleotide.

APPLICATION QUESTIONS AND PROBLEMS

Introduction

17. The introduction to this chapter, about the sequencing of 4000-year-old DNA,
emphasizes DNA’s extreme stability. What aspects of DNA’s structure contribute to
the stability of the molecule? Why is RNA less stable than DNA?

Solution:
Several aspects contribute to the stability of the DNA molecule. The relatively strong
phosphodiester linkages connect the nucleotides of a given strand of DNA. The
helical nature of the double-stranded DNA molecule results in the negatively
charged phosphates of each strand being arranged to the outside and away from each
other. The complementary nature of the nitrogenous bases of the nucleotides helps
hold the two strands of polynucleotides together. The stacking interactions of the
bases, which allow for any base to follow another in a given strand, also play a major
role in holding the two strands together. Finally, the ability of DNA to have local
variations in secondary structure contributes to its stability.
232 Chapter Ten: DNA: The Chemical Nature of the Gene

RNA nucleotides or ribonucleotides contain an extra oxygen at the 2′carbon of the


ribose sugar. This extra oxygen at each nucleotide makes RNA a less stable
molecule.

Section 10.2

*18. Match the researchers (a-j) with the discoveries listed.

a. Kossel
b. Fraenkel-Conrat
c. Watson and Crick
d. Levene
e. Miescher
f. Hershey and Chase
g. Avery, MacLeod, and McCarty
h. Griffith
i. Franklin and Wilkins
j. Chargaff

__ Took X-ray diffraction pictures used in constructing the structure of DNA.

__ Determined that DNA contains nitrogenous bases.

__ Identified DNA as the genetic material in bacteriophage.

__ Discovered regularity in the ratios of different bases in DNA.

__ Determined that DNA is responsible for transformation in bacteria.

__ Worked out the helical structure of DNA by building models.

__ Discovered that DNA consists of repeating nucleotides.

__ Determined that DNA is acidic and high in phosphorous.

__ Conducted experiments showing that RNA can serve as the genetic material in
some viruses.

__ Demonstrated that heat-killed material from bacteria could genetically transform


live bacteria.

Solution:
Took X-ray diffraction pictures used in constructing the structure of DNA:
i. Franklin and Wilkins
Chapter Ten: DNA: The Chemical Nature of the Gene 233

Determined that DNA contains nitrogenous bases:


a. Kossel

Identified DNA as the genetic material in bacteriophage:


f. Hershey and Chase

Discovered regularity in the ratios of different bases in DNA:


j. Chargaff

Determined that DNA is responsible for transformation in bacteria:


g. Avery, MacLeod, and McCarty

Worked out the helical structure of DNA by building models:


c. Watson and Crick

Discovered that DNA consists of repeating nucleotides:


d. Levene

Determined that DNA is acidic and high in phosphorous:


e. Miescher

Conducted experiments showing that RNA can serve as the genetic material in some
viruses:
b. Fraenkel-Conrat

Demonstrated that heat-killed material from bacteria could genetically transform live
bacteria:
h. Griffith

*19. A student mixes some heat-killed-type IIS Streptococcus pneumonia bacteria with
live type IIR bacteria and injects the mixture into a mouse. The mouse develops
pneumonia and dies. The student recovers some type IIS bacteria from the dead
mouse. If this is the only experiment conducted by the student, has the student
demonstrated that transformation has taken place? What other explanations might
explain the presence of the type IIS bacteria in the dead mouse?

Solution:
No, the student has not demonstrated that transformation has taken place. Unlike
Griffiths, who used strains IIR and IIIS to demonstrate transformation, the student is
using strains IIR and IIS. A mutation in the IIR strain injected into the mouse could
be sufficient to convert the IIR strain into the virulent IIS strain. By not conducting
the appropriate control of injecting IIR bacteria only, the student cannot determine
whether the conversion from IIR to IIS is due to transformation or to a mutation.
Although heat may have killed all the IIS bacteria, the student has not demonstrated
that the heat was sufficient to kill all the IIS bacteria. A second useful control
234 Chapter Ten: DNA: The Chemical Nature of the Gene

experiment would have been to inject the heat-killed IIS into mice and see if any of
the IIS bacteria survived the heat treatment.

20. Predict what would happen if Griffith had mixed some heat-killed type IIIS bacteria
and some heat-killed type IIR bacteria and injected these into a mouse. Would the
mouse have contracted pneumonia and died? Explain why or why not.

Solution:
The mouse would not have contracted pneumonia and died. Although the mouse
would have received IIIS DNA, which codes for virulent Streptococcus pneumoniae,
there are no live bacteria for this DNA to transform. Live bacteria are required for
pneumonia to develop.

21. Explain how heat-killed type IIIS bacteria in Griffith’s experiment genetically altered
the live type IIR bacteria. (Hint: See the discussion of transformation in Chapter 9.)

Solution:
The IIR strain of Streptococcus pneumonia must have been naturally competent or,
in other words, was capable of taking up DNA from the environment. The heat-killed
IIIS bacteria lysed releasing their DNA into the environment allowing for IIIS
chromosomal DNA fragments to come in contact with IIR cells. The IIIS DNA
responsible for the virulence of the IIIS strain was taken up by an IIR cell and
integrated into the IIR cell’s chromosome, thus “transforming” the IIR cell into a
virulent IIIS cell.

22. What results would you expect if the Hershey and Chase experiment were conducted
on tobacco mosaic virus?

Solution:
Infection by TMV results in the protein coat and RNA genome entering the host cell.
Inside the plant cell, the TMV protein coat unwinds, releasing the viral genome,
which initiates infection. If Hershey and Chase had used 32P and 35S to label TMV
particles, the RNA molecules would have been labeled with the 32P and the viral
proteins would have been labeled with 35S. However, the protein coat and the RNA
genome would have entered the cell, so “radioactive ghost proteins” would not have
been located outside the cell. Newly synthesized viral RNAs would have contained
measurable levels of 32P.

23. Which of the processes of information transfer illustrated in Figure 10.16 are
required for the T2 phage reproduction illustrated in Figure 10.4?

Solution:
DNA replication, transcription, and translation.

*24. Imagine you are a student in Alfred Hershey and Martha Chase’s lab in the late
1940s. You are given five test tubes containing E. coli bacteria that were infected
Chapter Ten: DNA: The Chemical Nature of the Gene 235

with T2 bacteriophage that have been labeled with either 32P or 35S. Unfortunately,
you forgot to mark the tubes and are now uncertain which were labeled with 32P and
which with 35S. You place the contents of the each tube in a blender and turn it on
for a few seconds to shear off the protein coats. You then centrifuge the contents to
separate the protein coats and the cells. You check for the presence of radioactivity
and obtain the following results. Which tubes contained E. coli infected with 32P-
labeled phage? Explain your answer.

Tube number Presence of radioactivity in


1 cells
2 protein coats
3 protein coats
4 cells
5 cells

Solution:
Tubes 1, 4, and 5. The DNA of the bacteriophage contains phosphorous and the
protein contains sulfur. When the bacteriophages infect the cell, they inject their
DNA into the cell, but the protein coats stay on the surface of the cell. The protein
coats are sheared off in the blender, while the cells with the DNA pellet at the
bottom of the tube. Thus, cells infected with 35S -labeled bacteriophage will have
radioactivity associated with the protein coats, whereas those cells infected with 32P-
labeled bacteriophage will have radioactivity associated with the cells.

25. Figure 10.8 illustrates Fraenkel-Conrat and Singer’s experiment on the genetic
material of TMV. What results would you expect in this experiment if protein
carried the genetic information of TMV instead of RNA?

Solution:
When type A RNA was combined with type B protein (left side of Figure 10.8), the
resulting progeny viruses would have type B RNA and protein. When type B RNA
was combined with type A protein (right side of Figure 10.8), the resulting progeny
viruses would have type A RNA and protein.

Section 10.3

*26. DNA molecules of different sizes are often separated with the use of a technique
called electrophoresis (see Chapter 19). With this technique, DNA molecules are
placed in a gel, an electrical current is applied to the gel, and the DNA molecules
migrate toward the positive (+) pole of the current. What aspect of its structure
causes a DNA molecule to migrate toward to the positive pole?

Solution:
The phosphate backbone of DNA molecules typically carries a negative charge, thus
making the DNA molecules attractive to the positive pole of the current.
236 Chapter Ten: DNA: The Chemical Nature of the Gene

*27. Each nucleotide pair of a DNA double helix weighs about 1 × 10–21 g. The human
body contains approximately 0.5 g of DNA. How many nucleotide pairs of DNA are
in the human body? If you assume that all the DNA in human cells is in the B-DNA
form, how far would the DNA reach if stretched end to end?

Solution:
If each nucleotide pair of a DNA double helix weighs approximately 1 × 10–21 g and
the human body contains 0.5 grams of DNA, then the number of nucleotide pairs can
be estimated as: (0.5 g DNA / human)/(1 × 10–21 g / nucleotide) = 5 × 1020
nucleotides pairs/human.
DNA that is in B form has an average distance of 0.34 nm between each nucleotide
pair. If a human possesses 5 × 1020 nucleotide pairs, then that DNA stretched end to
end would reach: (5 × 1020 nucleotides / human) × (0.34 nm/nucleotide pair) = 1.7 ×
1020 nm, or 1.7 × 108 km.

28. One nucleotide strand of DNA molecule has the base sequence illustrated below.

5′—ATTGCTACGG—3′

Give the base sequence and label the 5′ and 3′ends of the complementary DNA
nucleotide strand.

Solution:
Answer: 3′—TAACGATGCC—5′

*29. Erwin Chargaff collected data on the proportions of nucleotide bases from the DNA
of a variety of different organisms and tissues (E. Chargaff, in The Nucleic Acids:
Chemistry and Biology, vol. 1, E. Chargaff and J. N. Davidson, Eds. New York:
Academic Press, 1955). The following data are from the DNA of several organisms
analyzed by Chargaff.
Percent
Organism and tissue A G C T
Sheep thymus 29.3 21.4 21.0 28.3
Pig liver 29.4 20.5 20.5 29.7
Human thymus 30.9 19.9 19.8 29.4
Rat bone marrow 28.6 21.4 20.4 28.4
Hen erythrocytes 28.8 20.5 21.5 29.2
Yeast 31.7 18.3 17.4 32.6
E. coli 26.0 24.9 25.2 23.9
Human sperm 30.9 19.1 18.4 31.6
Salmon sperm 29.7 20.8 20.4 29.1
Herring sperm 27.8 22.1 20.7 27.5

a. For each organism, compute the ratio of (A + G)/(T + C) and the ratio of
(A + T)/(C + G).
Chapter Ten: DNA: The Chemical Nature of the Gene 237

Solution:
Organism (A + G)/(C + T) (A + T)/(C + G)
Sheep Thymus 1.03 1.36
Pig liver 0.99 1.44
Human thymus 1.03 1.52
Rat bone marrow 1.02 1.36
Hen erythrocytes 0.97 1.38
Yeast 1.00 1.80
E. coli 1.04 1.00
Human sperm 1.00 1.67
Salmon sperm 1.02 1.43
Herring sperm 1.04 1.29

b. Are these ratios constant or do they vary among the organisms? Explain why.

Solution:
The ratios for the (A + G)/(T + C) are constant at approximately 1.0 for the
different organisms. Each of these organisms contains a double-stranded
genome. The percentages of guanine and cytosine are almost equal to each other
and the percentages of adenine and thymine are almost equal to each other as
well. In other words, the percentage of purines should be equal to the percentage
of pyrimidines for double-stranded DNA. This means that (A + G) = (C + T).
The (A + T)/(C + G) ratios are not constant. The relative numbers of AT base
pairs and GC base pairs are unique to each organism and can vary between the
different species.

c. Is the (A + G)/(T + C) ratio different for the sperm samples? Would you expect
it to be? Why or why not?

Solution:
The ratios for the two sperm samples are essentially the same. The equal ratio
should be expected. As stated in the answer to part b. of this question, the
percentage of purines should equal the percentage of pyrimindines.

30. Boris Magasanik collected data on the amounts of the bases of RNA isolated from a
number of sources, expressed relative to a value of 10 for adenine (B. Magasanik, in
The Nucleic Acids: Chemistry and Biology, vol. 1, E Chargaff and J. N. Davidson,
Eds. New York: Academic Press, 1955).

Percent
Organism and tissue A G C U
Rat liver nuclei 10 14.8 14.3 12.9
Rabbit liver nuclei 10 13.6 13.1 14.0
Cat brain 10 14.7 12.0 9.5
Carp muscle 10 21.0 19.0 11.0
Yeast 10 12.0 8.0 9.8
238 Chapter Ten: DNA: The Chemical Nature of the Gene

a. For each organism, compute the ratio of (A + G)/(U + C).

Solution:
Organism and tissue (A + G)/ (U + C)
Rat liver nuclei 0.91
Rabbit liver nuclei 0.87
Cat brain 1.15
Carp muscle 1.03
Yeast 1.24

b. How do these ratios compare with the (A + G)/(T + C) ratio found in DNA
(see Problem 29)? Explain.

Solution:
The ratios are not as similar to each other or as close to the value of 1.0 as
found for the (A + G)/(T + C) ratio in DNA. Many RNA molecules are
single-stranded and do not have large regions of complementary sequences as
we would expect to find in DNA.

31. Which of the following relations or ratios would be true for a double-stranded DNA
molecule?

a. A + T = G + C

Solution:
No

b. A + T = T + C

Solution:
No

c. A + C = G + T

Solution:
Yes

A +T
= 1.0
d. C + G

Solution:
No

A +G
e. = 1.0
C+T
Chapter Ten: DNA: The Chemical Nature of the Gene 239

Solution:
Yes

A G
f. =
C T

Solution:
No

g.

Solution:
No

A G
h. =
T C

Solution:
Yes

A double-stranded DNA molecule will contain equal percentages of A and T


nucleotides and equal percentages of G and C nucleotides. The combined
percentage of A and T bases added to the combined percentage of the G and C
bases should equal 100.

*32. If a double-stranded DNA molecule is 15% thymine, what are the percentages of all
the other bases?

Solution:
The percentage of thymine (15%) should be approximately equal to the percentage
of adenine (15%). The remaining percentage of DNA bases will consist of cytosine
and guanine bases (100% – 15% – 15% = 70%); these should be in equal amounts
(70%/2 = 35%). Therefore, the percentages of each of the other bases if the thymine
content is 15% are adenine = 15%; guanine = 35%; and cytosine = 35%.

33. Suppose that each of the bases in DNA were capable of pairing with any other base.
What effect would this capability have on DNA’s capacity to serve as the source of
genetic information?

Solution:
DNA’s ability to be replicated faithfully and to encode phenotypes would be
destroyed. If each base could pair with any other base, the result during replication
would be changes in the DNA sequences of the newly replicated strands. The two
new molecules of DNA would not be identical to the original molecule or to each
other because different bases would be inserted in each newly synthesized strand. If
240 Chapter Ten: DNA: The Chemical Nature of the Gene

the DNA base sequence was constantly changing due to the random pairing of
bases, then no consistent “code” could be maintained. This lack of a code would
inhibit the ability of a DNA molecule to faithfully code for any particular protein.

34. Heinz Shuster collected the following data on the base composition of ribgrass virus
(H. Shuster, in The Nucleic Acids: Chemistry and Biology, vol. 3, E. Chargaff and J.
N. Davidson, Eds. New York: Academic Press, 1955). On the basis of this
information, is the hereditary information of the ribgrass virus RNA or DNA? Is it
likely to be single stranded or double stranded?

Percent
A G C T U
Ribgrass virus 29.3 25.8 18.0 0.0 27.0

Solution:
Most likely, the ribgrass viral genome is a single-stranded RNA. The presence of
uracil indicates that the viral genome is RNA. For the molecule to be double-
stranded RNA, we would predict equal percentages of adenine and uracil bases and
equal percentages of guanine and cytosine bases. Neither the percentages of adenine
and uracil bases nor the percentages of guanine and cytosine bases are equal,
indicating that the viral genome is likely single stranded.

*35. The relative amounts of each nucleotide base are tabulated here for four different
viruses. For each virus listed in the following table, indicate whether its genetic
material is DNA or RNA and whether it is single stranded or double stranded.
Explain your reasoning.

Virus T C U G A
I 0 12 9 12 9
II 23 16 0 16 23
III 34 42 0 18 39
IV 0 24 35 27 17

Solution:
Virus I is a double-stranded RNA virus. Uracil is present indicating an RNA
genome and we see equal percentages of adenine and uracil and equal percentages
of guanine and cytosine, which we would expect if it is a double-stranded genome.

Virus II is a double-stranded DNA virus. The presence of thymine indicates that the
viral genome is DNA. As expected for a double-stranded DNA molecule, we see
equal percentages of adenine and thymine bases and equal percentages of guanine
and cytosine bases.

Virus III is a single-stranded DNA virus. Thymine is present suggesting a DNA


genome. However, we see unequal percentages of thymine and adenine and unequal
Chapter Ten: DNA: The Chemical Nature of the Gene 241

percentages of guanine and cytosine, which suggest a single-stranded DNA


molecule.

Virus IV is a single-stranded RNA virus. Uracil is present indicating an RNA


genome. However, the percentage of adenine does not equal the percentage of
uracil, and the percentage of guanine does not equal the percentage of cytosine.
These unequal amounts suggest a single-stranded genome.

*36. A B-DNA molecule has 1 million nucleotide pairs. How many complete turns are
there in this molecule?

Solution:
B-form DNA contains approximately 10 nucleotides per turn of the helix. A
B-DNA molecule of 1 million nucleotide pairs will have about the following
number of complete turns: (1,000,000 nucleotides) / 10 nucleotides/turn) = 100,000
complete turns.

*37. For entertainment on a Friday night, a genetics professor proposed that his children
diagram a polynucleotide strand of DNA. Having learned about DNA in preschool,
his 5-year-old daughter was able to draw a polynucleotide strand, but she made a
few mistakes. The daughter’s diagram (represented here) contained at least 10
mistakes.
O

O— P—O–

OH— CH C base

H H
H OH
H O


O— P —O
OH— CH C base

H H
H OH
H OH

a. Make a list of all the mistakes in the structure of this DNA polynucleotide
strand.

Solution:
(1) The 5′ carbon for each of the two sugars is directly linked to phosphorous.
(2) Neither 5′ carbon of the two sugars should have an OH group attached.
242 Chapter Ten: DNA: The Chemical Nature of the Gene

(3) Neither sugar molecule has oxygen in its ring structure between the 1′ and
4′ carbons.
(4) In one sugar, the 2′ carbon has an –OH group attached, which does not
occur in deoxyribonucleotides.
(5) At the 3′ position in both sugars, only hydrogen is attached, as opposed to
an –OH group.
(6) The 1′ carbon for each of the two sugars has an –OH group, as opposed to
just a hydrogen attached.
(7) The two nucleotides are connected at the 2′ to 5′ position instead of the 3′
to 5′.
(8) The two nucleotides are not connected by a phosphodiester bond.
(9) Neither of the two phosphates are double-bonded to an oxygen. There
should be one oxygen-phosphate double bond in each phosphate.
(10) The phosphate connecting the two sugars shows two negative charges
instead of only one negative charge, which would be present if the
phosphorous double-bond to oxygen was present.

b. Draw the correct structure for the polynucleotide strand.

Solution:
O
P O
O
O
C CHO CH Base
H2
HC C
O H2
O
O P
O
C O Base
H2 CH CH
HC C
HO H2

38. Chapter 1 considered the theory of the inheritance of acquired characteristics and
noted that this theory is no longer accepted. Is the central dogma consistent with the
theory of the inheritance of acquired characteristics? Why or why not?

Solution:
The central dogma of molecular biology is not consistent with the theory of
inheritance of acquired characteristics. The flow of information predicted by the
central dogma is:
DNA RNA Protein
One exception to the central dogma is reverse transcription, whereby RNA codes
for DNA. However, biologists currently do not know of a process that will allow
Chapter Ten: DNA: The Chemical Nature of the Gene 243

for the flow of information from proteins back to DNA. The theory of inheritance
of acquired characteristics necessitates such a flow of information from proteins
back to the DNA.

Section 10.4

*39. Write a sequence of bases in an RNA molecule that will produce a hairpin structure.

Solution:
For a hairpin structure to form in an RNA molecule, an inverted complementary
RNA sequence separated by a region of noncomplementary sequence is necessary.
The inverted complements form the stem structure, and the loop of the hairpin is
formed by the noncomplementary sequences.

5′ —UGCAU—3′ …unpaired nucleotides…5′ —AUGCA—3′

UA
AU
CG
GC
5′ ----------U A---------3′

40. Write a sequence of nucleotides on a strand of DNA that will form a hairpin structure.

Solution:
For a hairpin structure to form in a strand of DNA, the DNA strand must contain
inverted complementary DNA sequences separated by a region of
noncomplementary sequence. The inverted complements form the stem structure,
and the loop of the hairpin is formed by the noncomplementary sequences:

5'—TGCATTACTCAATGCA—3'

or C T
A C
TA
GC
CG
AT
TA
5' T A 3'
244 Chapter Ten: DNA: The Chemical Nature of the Gene

CHALLENGE QUESTIONS

Section 10.1

*41. Suppose that an automated, unmanned probe is sent into deep space to search for
extraterrestrial life. After wandering for many light-years among the far reaches of
the universe, this probe arrives on a distance planet and detects life. The chemical
composition of life on this planet is completely different from that of life on Earth,
and its genetic material is not composed of nucleic acids. What predictions can you
make about the chemical properties of the genetic material on this planet?

Solution:
Although the chemical composition of the genetic material may be different DNA, it
more than likely will have similar properties to DNA. As discussed earlier in the
chapter, the genetic material must possess three general characteristics:
(1) It must contain complex information.
(2) It must replicate or be replicated faithfully.
(3) It must encode the phenotype.
(4) It must have the ability to vary.
Even if the material is not DNA, it must meet these criteria. For instance, if the
material could not be replicated or duplicated faithfully, then life on that planet could
not continue because ultimately no offspring could be produced. A lack of fidelity
would result in the loss of information. Genetic material from any lifeform has to
store the information necessary for the survival of that organism. Also, the genetic
material will need to be stable. Unstable molecules will not allow for long-term
storage of information, resulting in the loss of information and change in phenotype.
In addition, this material must be able to vary or mutate, and the mutant form must
be as stable as the original form, or else species will not be able to evolve.

Section 10.2

42. How might 32P and 35S be used to demonstrate that the transforming principle is
DNA? Briefly outline an experiment that would show that DNA rather than protein
is the transforming principle.

Solution:
The first step would be to label the DNA and proteins of the donor bacteria cells with
35
S and 32P. The DNA could be labeled by growing a culture of bacteria in the
presence of 32P. The cells as they replicate ultimately will incorporate radioactive
phosphorous into their DNA. A second culture of bacteria should be grown in the
presence of 35S, which ultimately will be incorporated into proteins.
Material from each culture should be used to transform bacteria cells that previously
had not been exposed to the radioactive isotopes. Transformed cells (or colonies)
that would be identified by the acquisition of a new phenotype should contain low
levels of the radioactive material due to the uptake of the labeled molecules. If the
Chapter Ten: DNA: The Chemical Nature of the Gene 245

transforming material were protein, then cells transformed by the material from the
35
S exposed bacterial cultures would also contain 35S. If the transforming material
were DNA, then the cells transformed by the material from the 32P exposed bacterial
cultures would also contain 32P.

Section 10.3

43. Researchers have proposed that early life on Earth used RNA as its source of genetic
information and that DNA eventually replaced RNA as the source of genetic information.
What aspects of DNA structure might make it better suited than RNA to be the genetic
material?

Solution:
Due to the lack of an attached oxygen molecule at the 2' carbon position of the sugar
molecule in deoxynucleotides, DNA molecules are more stable and less reactive than
RNA molecules. The double-helical nature of the DNA molecule provides a greater
opportunity for DNA repair and fidelity during replication. If mistakes occur in one
strand, the complementary strand can serve as a template for corrections.

44. Scientists have reportedly isolated short fragments of DNA from fossilized dinosaur
bones hundreds of millions of years old. The technique used to isolate this DNA is
the polymerase chain reaction, which is capable of amplifying very small amounts of
DNA a million fold (see Chapter 19). Critics have claimed that the DNA isolated
from dinosaur bones is not purely of ancient origin but instead has been
contaminated by DNA from present-day organisms such as bacteria, mold, or
humans. What precautions, analyses, and control experiments could be carried out to
ensure that DNA recovered from fossils is truly of ancient origin?

Solution:
An initial precaution would be to handle all the material in the most sterile manner
possible. People handling the samples should wear gloves and masks to help keep
the area as devoid of extraneous DNA as possible. Instruments used in the sampling
should be sterilized to eliminate any contamination by bacteria, viruses, fungi, and so
on. They should also be treated to remove trace DNAs. In addition, the source
material surrounding the bones should be treated to remove contaminating DNAs.
Controls also need to be conducted. The DNA from people involved in the procedure
should be tested to see if amplification occurs. Material at various locations around
the site and isolated bugs and microorganisms from the area should be sampled to
see if similar amplification patterns emerge. The design of the primers used for
amplification should be considered carefully and should be executed considering the
sequences of potential dinosaur descendants, such as birds or reptiles, in an attempt
to limit random amplifications. Furthermore, every experiment should be
reproducible.
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[26] Rooms are concealed by this talus, the walls of which
project in places out of the ground.
[27] Laguna creek is entered at this point on the right by a
stream bifurcating into the Cataract and East tributaries, which
flow through canyons of the same names. In or near East canyon
are four large ruins: Ladder House, Cradle House, Forest-glen
House, and Pine-tree House. The largest ruin in Cataract canyon
is Kitsiel. The Navaho sometimes speak of the East canyon as
the Salt, or Alkaline, bokho.
[28] Another geological feature of the sites of the large cliff-
dwellings of the Navaho Monument is the almost constant
presence of a vertical cliff-wall below the cave floor, the talus
rarely extending to the base of the lowest rooms.
[29] According to Hopi legends, the Horn clans (animals with
horns) are kin to the Snake, and formerly lived with the Snake
clans at Tokónabi. Later they united with the Flute clans at
Lengyanobi, and still later joined the Snake clans at Walpi.
Lengyanobi (“Pueblo of the Flute”) is a large ruin north of the Hopi
mesas.
[30] “Adobe bricks” with straw, according to Mr. W. B. Douglass,
are found at Inscription House near the end of the White mesa.
The writer has found adobe cubes in some of the walls of Cliff
Palace, but these contain no straw.
[31] Although circular kivas are found in several ruins in the
Navaho National Monument, as Kitsiel, Inscription House,
Scaffold House, and others, they were not seen in Betatakin,
which has the rectangular ceremonial room with side entrance
above mentioned. Although such rooms possess some of the
features of kivas, it is perhaps better to restrict that term to the
circular chambers and adopt the word kihu to designate the
rectangular rooms above ground. The ceremonial chambers of
Betatakin suggest the Flute room at Walpi. This fact and the
discovery of a flute in one of the rooms make it appear that
Betatakin was inhabited by Flute clans, which, according to Hopi
legends, lived in this region.
[32] For the accompanying view of the ruin (pl. 1), from
photographs taken by Mr. William B. Douglass, the writer is
indebted to the General Land Office.
[33] The kivas appear to be circular; one of them has the large
banquette, like kiva M in Cliff Palace. No pilasters for supporting
roofs have yet been reported.
[34] The two ruins Kitsiel and Betatakin are those about which
extravagant statements as to size and character were made
about two years ago by newspapers and otherwise reliable
magazines.
[35] Like all ruins in East canyon, Cradle House is situated in a
small side canyon on the left bank.
[36] Trickling-spring House is not located on the accompanying
map and, so far as could be ascertained, had not been visited by
archeologists previously to the writer’s visit. A young Navaho
guided the writer to it a short time before he left the region.
[37] Of course some of the rooms in Cliff Palace, especially
those at the western extension of the northern end, are
dependent, the cliff forming their rear walls.
[38] Both kinds of circular kivas are found in the cliff-ruins at
Casa Blanca and in Mummy cave in the Canyon de Chelly.
[39] These rites in all the Hopi pueblos are performed, as in
ancient times, in rectangular rooms not called kivas. The Snake
rites are performed now, as when the clan lived at Tokónabi in
subterranean rooms (kivas), the present form of which is
rectangular instead of circular, as at Tokónabi.
[40] It appears that in some of the ruins of the Navaho National
Monument there were both circular subterranean kivas and
rectangular rooms used for ceremonial purposes. At Wukóki the
former do not exist, but two of the latter can be recognized, one of
which has a construction like a ventilator.
[41] None of the five Walpi kivas is older than 1680, and one or
two are of later construction.
[42] Haus und Dorf bei den Eingeborenen Nordamerikas, in
Arch. für Anthr., n. f., Bd. vii, Heft 2 and 3, 1908.
[43] The circular kivas of Kükütcomo, the twin ruins on the
mesa above Sikyatki, near Walpi, are the only ceremonial rooms
of this form known from the Hopi mesas. These were the work of
the Coyote clan and are of Eastern origin.
[44] There are two types of cavate ruins, or rooms artificially
excavated in the tops or faces of cliffs, near Flagstaff. In one type,
Old Caves, the entrance to the subterranean rooms is vertical; in
the other, New Caves, it is from the side. In one type the walls of
masonry are built above the caves; in the other in front of them.
The common feature is the existence of chambers artificially
excavated in the cliff. Both types differ essentially from pueblos
built in the open or in natural caverns, although some of the kivas
of the latter are excavated in the solid rock.
MINOR ANTIQUITIES

Notwithstanding the limited duration of the writer’s visit to the


Navaho National Monument, a few specimens of stone, wood,
pottery, and other objects were collected. The whole pieces of
pottery, numbering 14 specimens (pls. 15-18), the majority of which
came probably from Inscription House and other ruins near Red
Lake, were presented to the Smithsonian Institution by Mr. Stephen
Janus, Navaho agent at Tuba, who accompanied the writer on the
trip to the Marsh Pass ruins. Fragments of pottery were picked up on
the surface at Betatakin, Kitsiel, and several other ruins, and the
most characteristic of these were brought back to Washington. No
excavations were attempted, nor could all objects that were seen be
brought away. Although up to within a few years these ruins were
practically in the condition they were when abandoned, unfortunately
of late they have been despoiled and many beautiful specimens
have been taken from them. Many objects still remain which should
be removed lest they fall into improper hands.

BUREAU OF AMERICAN ETHNOLOGY BULLETIN 50 PLATE 16


a. rough vase of corrugated ware
Cat. No. 257777. Height, 7 inches.
b. vase with constricted neck
Cat. No. 257778. Height, 8 inches.
POTTERY FROM NAVAHO NATIONAL
MONUMENT

Pottery

The pottery collected consists of jars, vases, food bowls, and


circular disks with a row of perforations about the margin. There are
also dipper handles and broken ladles of the usual shape. Some of
these specimens are of corrugated ware, others have smooth
surfaces with painted decoration. The proportion of corrugated and
indented ware found in the Navaho Monument ruins is about the
same as in the Mesa Verde National Park. The finest coiled ware
was obtained from the latter locality. Several fragments of flat dishes,
perforated on their margins (pl. 15, b), or colanders having holes in
the middle, form part of the collection.[45]
The most instructive form of pottery in the collection brought back
from northern Arizona is a decorated globular vase of black-and-
white ware (pl. 16, b). The decoration on this specimen is not
confined to the exterior but is found also on the inner surface of the
lip; it consists mainly of triangles so united as to form hour-glass
figures. A unique design on this vessel consists of two parallel lines,
each with dots on one side, suggesting similar bands in red on the
inner wall of the third story of the square tower of Cliff Palace.
Three small bowls of crude ware are fluted on the outside, the
ridge, or fluting, being raised somewhat above the surface of the
bowl and having a zigzag course. One of the best of these unique
ceramic forms has this fluting broken into S-shaped figures, as
shown in the accompanying illustration (pl. 17, a).
The writer collected also several perforated clay disks which were
possibly used as spindle whorls, although they may have been
gaming implements. A similar disk made of mountain-sheep horn
was found at Kitsiel.
The largest and one of the finest vases (pl. 18, a) from the
neighborhood of Red Lake is also of black-and-white ware. The
decoration is external and consists of black figures covering the neck
and upper body. The base is rounded and the lip slightly flaring. This
vase may have been used for containing water or possibly as a
receptacle for prayer (corn) meal. The food bowls from Red Lake are
chiefly of black-and-white ware, the red and yellow varieties being
less numerous. A common feature in food bowls of this region is a
handle on one side, as shown in plate 15, d. Some of these vessels,
although of smooth ware, are without decoration on either the
exterior or the interior.
The shallow, slightly concave clay disk[46] shown in plate 15, b, is
characteristic in possessing a row of holes near the rim. This disk
seems to represent a common type, as several fragments with
similar holes were found on the surface of the ruins. The same or
related forms appear to have been common in ruins near the Hopi
pueblos. These are found in the collection of votive offerings now in
the Peabody Museum at Cambridge, from Jedito spring, near
Awatobi, and the writer has discovered specimens elsewhere in Hopi
ruins, a brief mention of which occurs in a report on the archeological
results of his expedition to Arizona in 1895.[47]
Several fragments of deep bowls, each having a handle (pl. 18, b)
on the surface, were obtained in the sands below cliff-house B; these
are commonly of red ware and have reddish-brown and black
decorations. A small dish of black-and-white ware (pl. 15, a) has the
rim slightly elevated and rounded on one side. The cups or mugs
from this region are shaped unlike those from the Mesa Verde. Mugs
from the latter region are cylindrical in form or the walls incline
slightly inward so that the diameter of the opening is somewhat less
than that of the base. The lip is thick and decorated. One of these
cups, here figured, has a constricted neck, and a slightly flaring rim
which is thin and undecorated. The decoration of another cup (pl. 15,
c) suggests the designs on several mugs from the Little Colorado
ruins. So far as form and decoration are concerned, this cup, or
handled vase, might have come from Homolobi, Chevlon, or Chaves
pass.[48]

BUREAU OF AMERICAN ETHNOLOGY BULLETIN 50 PLATE 17


a. bowls bearing relief ornaments
(From left to right): Cat. Nos. 257783, 257784, 257782. Heights, 2½ inches, 2
inches, 2¾ inches.

b. handles of food-bowls
Cat. No. 258326.
c. stone implements
(From left to right): Cat. Nos. 258334, 258335, 258336, 258337. Dimensions, 6 x 4
x 1¾ inches; 5¼ x 3¾ x 2½ inches; 4½ x 3¾ x 2 inches; 4¾ x 2¾ x 2½ inches.
POTTERY AND STONE IMPLEMENTS FROM NAVAHO NATIONAL
MONUMENT

The designs on fragments of pottery found in ruins in northern


Arizona are identical with or related to those from the Black Falls
ruins, but differ somewhat from those on pottery from ruins higher up
the Little Colorado river. If the history of the modification of ceramic
symbols in any of the large composite pueblos of the Southwest be
studied, it will be noticed that there are often radical changes, the
later symbols not being modifications of earlier ones. Thus modern
Zuñi pottery designs differ materially from those found in ruins in the
same valley. The modern pottery from East mesa is wholly different
from that of Sikyatki, a few miles away. Again, in so-called modern
Hopi pottery, Tewa symbols derived from the Rio Grande have
replaced old Hopi symbols dominant before the advent of Tewa
clans. The changes in pottery symbols in every large composite
pueblo are not due to evolution of the modern from the ancient, but
reflect the history of the advent of new clans, powerful enough to
substitute their designs for those formerly existing. One of the
problems of the ethnologist is to determine symbols associated with
certain clans, and by means of legends to identify clans with ruins.
Having determined the symbols introduced by certain clans and the
places where these clans halted in their migrations and built pueblos,
the course of these prehistoric movements may be followed.
Comparison of symbols on pottery from northern Arizona with those
from Black Falls ruins support, so far as they go, the legends that the
Snake people, who once lived at Wukóki near the Black Falls, lived
also in cliff-houses now ruins near Marsh pass or the White mesa.
The symbolism indicates the presence of the same clans, and
tradition is thereby supported.

Cliff-dwellers Cradle

One of the most instructive specimens collected in the Navaho


National Monument was found by Mr. W. B. Douglass in a ruin
designated as Cradle House. This object is a cradle made of basket
ware, open at one end and continued at the opposite end into a
biped extension to serve for the legs. It is decorated on the outside
with an archaic geometric ornamentation, the unit design of which is
shown in the accompanying illustration. This specimen (pls. 19-21)
may be regarded as one of the finest examples of prehistoric
basketry from the Southwest; moreover, with one exception, it is the
only known cradle of this form. A pair of infant’s sandals found with
the cradle leaves no doubt as to its use, while the character and
symbolism of the decoration refer it to the ancient cliff-house culture.
The design (fig. 3) suggests that which characterizes certain
specimens of the well-known black-and-white pottery found in the
San Juan drainage. Evidences of long use and repair appear,
especially on one side. Unfortunately, the specimen, although entire
when found, later was broken across its middle.
Fig. 3. Design on cliff-dwellers cradle.

The only other known cradle of this type was brought to the
attention of ethnologists by Dr. W. J. McGee when in charge of the
anthropological exhibit at the St. Louis Exposition. This was found in
San Juan county, Utah, not far from the Colorado river.[49] This
specimen is better preserved than that here figured, but the
decoration is practically identical; so near, in fact, that the two might
have been made by the same woman.

Miscellaneous Objects

The stone implements (pl. 17, c) consist of axes, pounding stones,


[50]and hatchets. On one of the roofs at Kitsiel there was picked up
a curved stick[51] identical with those placed by the Walpi Snake
priests about the sand-painting of their altar. A good specimen of a
planting stick and a rod formerly used as a spindle were found near
by; the latter is a perforated disk made of horn. A flute identical with
those used at the present day by Flute priests at Walpi was found at
Betatakin, thus tending to support the legend that the Flute clan once
lived at the latter pueblo.
FOOTNOTES:

[45] These dishes resemble those sometimes used by the Hopi


for sprinkling water on their altars as a prayer for rain. They may
have been used also in sifting sand on the kiva floor, to form a
layer upon which the sand picture is later drawn with sands of
different colors.
[46] Small perforated clay disks are not rare here, as in other
ruins. They were used in the same way as the horn disk
mentioned on page 30.
[47] In Seventeenth Annual Report of the Bureau of American
Ethnology, pt. 2.
[48] Compare figures from these ruins, in the Twenty-second
Annual Report of the Bureau of American Ethnology.
[49] The finder was Mr. E. B. Wallace. This specimen was
owned at one time by Mr. J. T. Zeller, an architect of St. Louis.
The writer has been informed that Mr. Zeller sold the cradle and
that it is now in the Field Museum of Natural History in Chicago.
[50] A common feature of stone mauls is a raised ferrule above
and one below the groove to which the handle is attached.
[51] These sticks, or “crooks” (gnela), found on the Antelope
altar in the Walpi Snake ceremony are reported to have been
brought to Walpi from Tokónabi.
SUMMARY AND CONCLUSIONS

The route chosen by the author for visiting the ruins of the Navaho
National Monument is via Flagstaff and Tuba, the distance being not
far from 200 miles to Marsh pass and 10 miles beyond to the largest
cliff-dwellings. Although the wagon road is long, requiring a journey
of at least five days, it may be traversed with carriage or buckboard,
the sandy stretch between Tuba and Red Lake being the most
difficult. The trail from Marsh pass to the great cliff-dwellings,
although now passable only on horseback, could be made into a
wagon road at small expense.
The nature of the cliffs in which the ruins of the Navaho Monument
are situated favored the construction of cliff-dwellings rather than of
open pueblos in this region. These cliffs are full of caverns, large and
small, presenting much the same condition as the cliffs of the red
sandstone elsewhere in the Southwest, as the Mesa Verde, Canyon
de Chelly, the Red Rocks south of Flagstaff, and other sections
where caverns abound. Fragments of fallen rocks present good
plane surfaces for walls of masonry, and there is abundant clay for
plastering. Trees suitable for rafters and beams are not wanting. In
short, all conditions are favorable for stone and adobe houses in the
cliffs. The neighboring Sethlagini mesa is of different geological
formation; in it are no caverns, the mesa top is broad, and ruins
thereon are necessarily open pueblos. The effect of difference in
geological structure is nowhere more evident than in these adjacent
formations.

BUREAU OF AMERICAN ETHNOLOGY BULLETIN 50 PLATE 18


a. large black-and-white vase
Cat. No. 257774. Height, 17 inches.
b. large vase with handle
Cat. No. 257787. Height, 8½ inches.
POTTERY FROM NAVAHO NATIONAL
MONUMENT

If environment has had so marked an influence on the character of


building, we can readily see how it has affected arts and crafts. We
can hardly imagine a people living any length of time in this region
without being mentally influenced by the precipitous cliffs that rise on
all sides. The summits of these heights are eroded into fantastic
shapes resembling animals or grotesque human forms. The constant
presence of these marvelous forms, of awe-inspiring size and weird
appearance, exerted a profound influence on the supernatural ideas
of the inhabitants. Here were born many conceptions of earth gods
and the like, survivals of which still remain among the Hopi.
As a rule the cliff-houses are not situated in sight of the main
stream, but are hidden away in secluded side canyons, approached
by narrow entrances, their sites having been determined no doubt by
the position of the springs with their constant water supply.
Almost every side canyon, even in a dry season, has its spring of
water which, trickling out of the rocks, follows the canyon bed until it
is finally drunk up by the thirsty sands. Often water seeps out of a
soft stratum of rock in the cave itself, where it was gathered in
artificial reservoirs that in ancient times furnished an adequate
supply for the inhabitants. One feature of these side canyons is that
they enlarge into basins surrounded on all sides by lofty cliffs. Many
of these basins are so hidden that they can be discovered only by
following dry stream-beds from their junction with the creeks. How
many of these basins are still undiscovered no one can yet tell. In
these basins now covered with bushes the aboriginal farms were
probably situated.
As the width of the valley of Laguna creek from Marsh pass to the
point where the stream receives its largest branches on the left bank
varies, the amount of arable land is greater in some places than in
others. In stretches where the stream almost washes the bases of
the ruins there could have been no extensive farming lands. The
creek meanders through the soft clay and sand which fill the valley to
the depth of many feet, forming treacherous banks that are
continually falling and changing the course of the stream, so it is
quite possible that the present configuration of the valley is very
different from what it was when the cliff-dwellings were inhabited. If
the occupants once had farms within its limits all traces of them
would have long since been obliterated. Although too much
credence should not be given to Navaho traditions, it is not
unreasonable to believe that in one particular at least they are
correct. These state that, before the introduction of sheep, grass was
much higher in the level part of the valley than at present, and
formerly game (at least the mountain sheep and the antelope) may
have been more abundant. This condition would have exerted a
marked influence on the life of the cliff-dwellers. Pictographs show
that the ancient people, either here or in their former homes, were
familiar with these animals, and various objects of bone and horn are
significant in this connection.
The Navaho National Monument (see sketch map, pl. 22) contains
two kinds of ruins,[52] cliff-dwellings and pueblos. Most of the latter
are situated on promontories or on low hills. The structural features
of the cliff-dwellings are characteristic, their walls being constructed
of stone or adobe built against, rarely free from, vertical faces of the
cliff.
There are two types of kivas, one circular and subterranean, allied
to those of the Mesa Verde, the other rectangular, above ground,
entered from the sides.
The masonry of these northern ruins is crude, resembling that of
modern Walpi. The component stones are neither dressed nor
smoothed, but the walls are sometimes plastered. There is a great
similarity in architecture. No round towers[53] relieve the monotony or
impart picturesqueness to the buildings. The walls of ruined pueblos
in this region and the ceramic remains closely resemble those at
Black Falls on the Little Colorado. A prominent feature of the walls is
a jacal construction in which the mud is plastered on wattling
between upright poles. The ends of many of these supports project
high above the ground, constituting a characteristic feature of the
ruins. This method of wall construction is unknown at Black Falls or
at Walpi, but survives in modified form in one or more Oraibi kivas
and in one at least of the Mesa Verde ruins.[54] It has been described
by Mr. Cosmos Mindeleff as common to several ruins in the Canyon
de Chelly.
The key to the culture of the people from which the cliff-dweller
culture was derived is probably the kiva, which furnishes also a good
basis for the classification of the Pueblos and cliff-dwellers into
subordinate groups.
Architecturally the kiva reached its highest development in the
Mesa Verde region, where it is a circular subterranean room with
pilasters and banquettes, ventilators and deflectors, fireplaces and
ceremonial openings, the features of which have been described
elsewhere. As we follow the San Juan down to its junction with the
Colorado we find a gradual simplification of the circular type of kiva
by the elimination of pilasters, ventilators, and other features, the
round kiva being here represented by rooms in which almost the only
architectural feature remaining is the large banquette. The question
naturally arising in this connection is, whether the circular kiva in the
eastern region is a development of that simpler form existing in the
western or whether the latter is a degenerate form of the eastern. In
other words, does the evidence show that this particular modification
spread from the east down the San Juan or from the west up the
river to the east? In this connection it may be urged that originally the
form of circular kiva lacking pilasters extended along the entire
course of the San Juan and that the kivas of the Mesa Verde
became highly specialized forms in which pilasters were developed,
while those lower down the river remained the same. We can not
definitely answer either of these questions, but taken with other
evidence it would seem that the circular form of kiva originated in the
eastern section and gradually extended westward.

BUREAU OF AMERICAN ETHNOLOGY BULLETIN 50 PLATE 19


CLIFF-DWELLERS CRADLE—FRONT

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