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Chapter 10
From DNA to Proteins: Transcription and RNA
Processing
True/False
1. All RNAs are translated. (F)
2. In transcription, all parts of the DNA molecule are transcribed into RNA. (F)
3. In all organisms, all genes are transcribed from the same strand. (F)
4. Certain RNA molecules behave as enzymes that can catalyze activities such
as RNA processing, RNA replication, and peptide bond formation between amino
acids. (T)
5. The presence of consensus in a sequence of DNA nucleotides usually
indicates that the sequence is associated with an important function. (T)
6. Introns contain no sequence-specific information. (F)
7. Most eukaryotic genes contain more non-coding nucleotides than nucleotides
that encode amino acids in polypeptides. (T)
8. Initiation of transcription does not require a primer. (T)
9. Energy for phosphodiester bond formation comes from the removal of two
phosphates from each added nucleotide. (T)
10. RNA molecules have the same 5′ to 3′ orientation as the DNA template
strands to which they are complementary. (F)
11. Eukaryotic genes usually include a long stretch of T nucleotides that encode
a poly(A) tail at the 3’ end of the corresponding mRNA molecules. (F)
12. Typically, each human gene encodes a single polypeptide. (F)
13. Intron cleavage and exon splicing are both mediated by the spliceosome. (T)
14. Introns are degraded in the cytoplasm. (F)
15. Transcription and translation take place simultaneously in bacterial cells. (T)

From DNA to Proteins: Transcription and RNA Processing
Fill in the Blank
16. Whereas the nucleotide strand used for transcription is termed the template
strand, the nontranscribed strand is called the nontemplate strand.
17. In transcription, nucleotides are always added to the 3′ end of the elongating
strand.
18. In a transcription reaction, two phosphate groups are cleaved from the
incoming ribonucleoside triphosphate; the remaining phosphate group is
attached to the growing RNA molecule by a phosphodiester bond.
19. RNA polymerase must bind to a region of DNA called a(n) promoter in order
to begin transcription.
20. In eukaryotes, RNA polymerase II transcribes the genes that encode
proteins.
Multiple Choice
21. Which statement about RNA polymerase is not true?
a. RNA polymerase can initiate RNA synthesis from scratch and does not require
a primer.
b. RNA polymerase synthesizes RNA 5’ to 3’.
c. RNA polymerase reads DNA 3’ to 5’.
*d. RNA polymerase normally transcribes both strands of DNA during expression
of a particular gene.
e. RNA polymerase must bind to a promoter in order to initiate transcription.
22. Which is a mechanism that allows a single gene to encode more than one
polypeptide?
a. Regulation of mRNA stability
*b. Alternative RNA splicing
c. RNA interference
d. Reverse transcription
e. None of the above
23. If the sequence of an RNA molecule is 5’-GGCAUCGACG-3’, what is the
sequence of the template strand of DNA?
a. 5’-GGCATCGACG-3’
b. 3’-GGCATCGACG-5’
c. 5’-CCGTAGCTGC-3’
*d. 3’-CCGTAGCTGC-5’
24. During gene expression, which molecule(s) carries the information that
encodes polypeptides?
a. tRNA

Chapter 10
*b. mRNA
c. rRNA
d. snRNA
e. More than one of the above
25. If the sequence of a nontemplate strand of DNA is
5’-ACCGCATCCGAGTCAC-3’, what is the sequence of the primary product of
transcription?
a. 3’-UGGCGUAGGCUCACUG-5’
b. 3’-TGGCGTAGGCTCACTG-5’
*c. 5’-ACCGCAUCCGAGUCAC-3’
d. 5’-ACCGCATCCGAGTCAC-3’
26. An in vitro transcription system that contains a bacterial gene initiates
transcription, but from random points on the DNA. Which of the following proteins
most likely is missing from the reaction?
*a. sigma factor
b. rho factor
c. RNA polymerase II
d. TATA-binding protein
27. In eukaryotes, the 5′ cap on an mRNA is important for all the processes listed
below except for the a of an mRNA molecule.
*a. transcription
b. intron removal
c. stability
d. initiation of translation
28. An in vitro transcription system transcribes a bacterial gene but terminates
inefficiently. What is one possible problem?
a. There is a mutation in the –10 consensus sequence, which is required for
efficient termination.
*b. Rho factor has not been added.
c. Sigma factor has not been added.
d. Spliceosomes have not been added.
29. The DNA replication enzyme that most closely resembles RNA polymerase is
a. DNA polymerase I.
b. DNA polymerase III.
*c. primase.
d. telomerase.
e. helicase.
30. Which of the following is not necessary for RNA polymerase to recognize the
promoter of a bacterial gene?
a. sigma factor
*b. origin of replication

c. –10 consensus sequence
d. –35 consensus sequence
Use the following list for questions 31–35.
a. RNA only
b. DNA only
c. both RNA and DNA
d. neither RNA nor DNA
31. When this molecule is synthesized, both strands of a DNA molecule are used
as a template.
*b
32. This molecule is synthesized using triphosphate nucleotides as a substrate
for a polymerase enzyme that forms phosphodiester bonds.
*c
33. This molecule is synthesized using nucleotides containing the bases adenine,
guanine, cytosine, and uracil.
*a
34. The polymerase that synthesizes this molecule uses DNA as a template and
synthesizes new strands from 5′ to 3′.
*c
35. This molecule is made of nucleotides joined by phosphodiester bonds that
connect the 2′ OH to the 5′ phosphate.
*d
36. If the following DNA strand was used as a template, what would the
sequence of an RNA be?
5′ GTACCGTC 3′
a. 5′ GUACCGUC 3′
b. 5′ GACGGTAC 3′
c. 5′ CAUGGCAG 3′
*d. 5′ GACGGUAC 3′
e. 5′ GUCGGUAC 3′
37. What is the function of eukaryotic RNA polymerase I?
*a. transcription of rRNA genes
b. transcription of mRNA genes
c. transcription of tRNA genes
d. transcription of snRNAs
e. initiation of transcription (but not elongation)

Chapter 10
38–39. The poly(A) tails found in the 3′ end of an mRNA are important for all the
processes listed below except for c and d .
a. mRNA stability
b. translation
*c. intron splicing
*d. protein stability
Short Answer Discussion
40. What is an snRNP and what role does it play in the cell?
A small nuclear ribonucleoprotein is a complex of small nuclear RNAs
(snRNAs)—U1, U2, etc.—and proteins that catalyze the transesterification
reactions and splicing of exons during eukaryotic mRNA processing.
41. If you were asked to isolate total RNA from two unknown samples and then
were required to identify if the RNA was from prokaryotes or eukaryotes, what
aspects regarding the classes of RNA present would help you distinguish one
from the other?
RNA from prokaryotes will contain mRNA, tRNA, and rRNA. In addition to
these three types of RNA, eukaryotic samples will contain pre-mRNA,
snRNA, snoRNA, miRNA, and siRNA.
42. What are the three different eukaryotic RNA polymerases and what types of
genes do they transcribe?
(1) RNA polymerase I transcribes rRNA.
(2) RNA polymerase II transcribes pre-mRNA and some snRNA.
(3) RNA polymerase III transcribes tRNA, small rRNA, and some snRNAs.
43. What would you add to an in vitro transcription system that contains an E. coli
gene for glyceraldehyde 3-phosphate dehydrogenase, an enzyme in glycolysis,
in order to get transcription that begins from the normal transcription start site?
(1) RNA polymerase
(2) sigma factor
(3) ribonucleoside triphosphates with A, C, G, and U bases (rNTPs)
44. If you remove the TATA box and place it immediately upstream of a
transcription start site of a eukaryotic gene, and subsequently transcription of the
mRNA is assayed, will you still achieve transcription from the same start site?
No. The TATA box needs to be present a certain number of nucleotides
upstream of the transcription start site to allow enough space for the
assembly of TATA-binding protein and other transcription factors on the
core promoter. The RNA polymerase can then be placed appropriately over
the transcription start site.

45. What provides the energy for RNA synthesis?
Hydrolysis of PPi from each incoming ribonucleoside triphosphate (with
subsequent hydrolysis of PPi to two inorganic phosphates).
46. A new mutation in Saccharomyces cerevisiae, a eukaryotic yeast, causes the
cells to be unable to produce the amino acid histidine. Specifically, they cannot
catalyze the first reaction in the histidine biosynthesis pathway. When examined
closely, they are producing a completely wild-type enzyme for this reaction, but at
greatly reduced levels. Explain the mutation.
The mutation is in the promoter for the gene reducing transcription.
47. What is the RNA sequence transcribed from the DNA shown below?
promoter +1
5′ GTAACTATAATTAACGTAAGACTAT 3′
3′ CATTGATATTAATTGCATTCTGATA 5′
5′ GUAAGACUAU 3′
48. Explain at least two reasons why the following definition of a gene is
inadequate: “A gene consists of DNA sequences that are transcribed into a
single RNA molecule that encodes a single polypeptide.”
(1) Because of alternative splicing, a single gene can yield multiple mRNA
and protein products.
(2) Sometimes, as with ribosomal RNAs, a single transcript is made, but
then several RNA molecules are liberated from it.
(3) Splicing means that not all sequences end up in the mature RNA
(introns removed).
(4) RNA can be the functional product of a gene, so a gene doesn’t always
encode a polypeptide.
(5) Regulatory sequences (i.e., promoters, enhancers, etc.) that control
the timing, degree, and specificity of gene expression are required
elements for expression of any given gene, but they are not
transcribed.
49. In your own words, list a comprehensive definition for “gene” at the molecular
level.
A gene is a sequence of DNA, including all structural and regulatory
sequences, that encodes information dictating synthesis of one or more
functional polypeptides or RNA molecules. Note that this is one possible
definition; see if you can come up with an even better one.

Chapter 10
Extended Answer Discussion
50. The discovery of ribozymes led to the theory that the evolution of life on earth
began with an “RNA world.”
(a) Describe the chemical properties and functions of RNA that would allow it to
be the basis of the first self-replicating systems.
(1) As a nucleic acid, RNA can serve as a template for self-replication: an
RNA can be copied into a complementary strand that is a template for
generating more of the original RNA.
(2) As a nucleic acid, RNA can carry genetic information in its base
sequence.
(3) RNA possibly could have performed the reactions required of a self-
replicating system because RNA ribozymes have catalytic activity.
(b) Describe how the current cellular role of RNA supports this theory.
(1) RNA is an intermediate between DNA, the permanent genetic
information, and proteins.
(2) RNA has a role as a primer for DNA replication.
(3) RNA is required for protein synthesis as mRNA, rRNA, and tRNA.
(c) What properties of proteins and DNA make them more suitable than RNA as
enzymes and the cell’s genetic material?
(1) Proteins are made of 20 different amino acids; RNA is made of relatively
uniform nucleotides with only four different bases. Because they can be
more chemically diverse, proteins are better suited to catalyzing a
variety of cellular reactions.
(2) DNA is double-stranded, with a structure that protects the molecule
from degradation. The lack of 2′ OH makes the molecule less
susceptible to self-hydrolysis. Because it is more stable, DNA is a better
molecule for the cell’s genetic material.
51. List five different classes of RNA molecules found in eukaryotes and describe
their functions.
(1) Messenger RNA (mRNA): the molecular messenger that carries
functional genetic code, encoded in DNA, to the cytoplasm for
translation into polypeptide products
(2) Ribosomal RNA (rRNA): along with specific proteins, an integral
component of large and small ribosomal subunits
(3) Transfer RNA (tRNA): the “adaptor” molecule that couples the
functional genetic code, carried by mRNA, with ribosomes for
translation into specific polypeptides

(4) Small nuclear RNAs (snRNAs): integral components of the spliceosome,
which mediates precise processing of pre-mRNA molecules in the
nucleus
(5) Small nucleolar RNAs (snoRNAs): integral components for processing
(primarily) eukaryotic rRNAs; located in the nucleolus
53. In 1958, Francis Crick proposed that genes and their corresponding
polypeptides are “colinear.” Explain why the idea of colinearity must be qualified
for both prokaryotic and eukaryotic genes.
In this context, colinearity simply means that the linear nucleotide
sequence of a given gene corresponds directly to the linear amino acid
sequence in the corresponding polypeptide. This implies that the number
of nucleotides in a gene should be precisely proportional to the number of
amino acids present in the corresponding polypeptide. Colinearity
generally holds true for the coding regions of prokaryotic genes, which in
most cases don’t contain introns. However, intervening sequences
(introns) discovered during the mid-1970s disqualify this idea for the vast
majority of eukaryotic genes. In addition, another exception to colinearity
between genes and polypeptides is the presence of untranslated
sequences, or regions (called UTRs), in both prokaryotic and eukaryotic
genes. The protein-coding sequences in virtually all mRNA molecules
(prokaryotic and eukaryotic) are flanked by 5′ and 3′ UTRs of variable
length.
54. Devise a strategy to prove that splicing occurs in the nucleus.
Collect cells and fractionate them into nuclear and cytoplasmic fractions.
Isolate the proteins from each of the fractions. Add a pre-mRNA (made in
vitro) with introns to each of the protein fractions. Monitor the progressive
decrease in the size of pre-mRNA in the nuclear fraction by separating the
RNA products on a gel, which can resolve size differences between pre-
mRNA (longer) and spliced mRNA (shorter). The cytoplasmic fraction will
be incapable of splicing, and therefore the pre-mRNA remains intact over a
period of time without undergoing splicing. The difference in the ability to
splice suggests that the nuclear fraction contains the machinery essential
for splicing.
55. What are some of the different ways that the word “gene” is defined at the
molecular level? What are some of the advantages and disadvantages of the
different definitions?
(1) Definition of “gene”: A sequence of DNA-carrying information that
encodes a single polypeptide.
Advantages/disadvantages of this definition: This definition does not
take into account either the production of multiple mRNA transcripts via
alternative splicing or the transcription of RNA end products (e.g., tRNA,
rRNA, snRNA, etc.).

Chapter 10
(2) Definition of “gene”: A sequence of DNA that can be transcribed into an
RNA or protein product(s).
Advantages/disadvantages of this definition: This definition does not
take into account the influence of regulatory sequences (promoters,
termination signals, etc.) that are required for gene function and, in fact,
must be included in any definition of “gene.”
56. Explain how the transcription apparatus knows which strand of DNA on a
gene to transcribe and where to start and stop. Be sure to include in your
answer the names and descriptions of DNA sequence elements on the
transcription unit that are important to this process.
The promoter is a DNA sequence that RNA polymerase binds to and that
determines both the direction of transcription and which DNA strand is to
be transcribed. It also establishes where transcription will begin (the
transcription inititiation site). Promoters generally are found just upstream
from the transcription initiation site. Promoters vary in sequence from
gene to gene, but different promoters share certain conserved sequences
called consensus sequences that the transcription apparatus recognizes
and binds to. The terminator is a sequence near the downstream end of
the RNA-coding sequence that signals the end of transcription. Some
terminators require the action of an ancillary protein called rho factor, while
others do not require rho.
57. Explain why the nematode worm C. elegans has become an important model
system in genetics and developmental biology. Your answer should include
details regarding the anatomy, development, and life cycle of the organism, the
nature of its genome, and the genetic and molecular biology techniques that are
available to researchers.
C. elegans is small, easy to culture, and produces large numbers of
offspring. The generation time is only about 3 days. The worms are
transparent, allowing for easy observation of internal development at all
stages. The body structure is simple with fairly invariant patterns of cell
division leading to a predictable normal body plan. This has allowed
researchers to map the lineages of every cell in the body back to the
zygote. The worms occur as hermaphrodites and males, allowing for them
to fertilize themselves or to be crossed with other worms. The relatively
small size of the genome facilitates genomic analysis, and the genome has
been completely sequenced, allowing for identification of all genes.
Chemical and other mutagens have been used to generate and isolate
many mutations that affect all aspects of development, anatomy, and
physiology. C. elegans is particulary susceptible to RNA interference,
allowing for gene functions to be assayed fairly easily by shutting down
their expression at particular points of development.
Problems and Calculations

58. On the DNA strands shown below, two RNA polymerase enzymes are using
the top strand as a template. In the boxes, label the 5′ and 3′ ends of the DNA
molecules and the RNA molecules being made. With arrows, indicate the
directions, left to right or right to left, that the polymerases are moving.
59. Draw the structure of a typical eukaryotic pre-mRNA molecule transcribed
from a gene containing one intron and two exons. Indicate the introns and exon,
splice sites, branch point, poly(A) site, 5′ UTR, and 3′ UTR.
exon 1 exon 2
GU intron AG
AAUAAA
5′ UTR 3′ UTR
*5′ *5′
*5′
*5′
*3′
*3′
*3′
*3'
*
*

Exploring the Variety of Random
Documents with Different Content

out of ten cases a space between the title and the imprint of the
printer-publisher, so that this blank tends to be the strongest feature
on the page. When the device was first abandoned, the author,
printer or publisher took advantage of the leisure of the reader and
the blank at their disposal, to draft a tediously long title, subtitle and
list of the author's qualifications, designed to fill the entire page. The
present-day publisher goes to the other extreme, reducing the title
to as few short words as possible, followed with "by" and the
author's name. A professional writer may insert, e.g., "Author of The
Deluge" under his name or there may be incorporated a motto; but
apart from such exceptions, three and sometimes four inches of
space separate the author's name from the first line of the imprint.
The result is that unless the title is set in a size of type out of all
relation to that of the remainder of the book, this space is more
conspicuous than the chief line. It is more reasonable to lessen this
space by shortening the depth of the whole piece from title to
imprint. It is clear that a volume in 12-point does not require a 30-
point title unless it be a folio in double-column; and it is of no
consequence if the title page is a little shorter than the text pages.
There is no reason, other than a desire to be "different," for a title
page to bear any line of type larger than twice the size of the text
letter. If the book be set in 12-point, the title need be no larger than
24-point—and may decently enough be smaller. As lower-case is a
necessary evil, which we should do well to subordinate since we
cannot suppress, it should be avoided when it is at its least rational
and least attractive—in large sizes. The main line of a title should be
set in capitals; and, like all titling capitals, they should be spaced.
Whatever may happen to the rest of the composition, the author's
name, like all displayed proper names, should be in capitals.
V
Here we may pause to counter an objection. It will be contended
that whatever the value of our preceding conclusions, their adoption

must mean an increase in standardization—all very well for those
who have an economic objective but very monotonous and dull for
those whose aim is that books shall possess more "life." This means
that the objectors want more variety, more "differentness," more
decoration. The craving to decorate is natural, and only if it is
allowed the freedom of the text pages shall we look upon it as a
passion to be resisted. The decoration of title pages is one thing—
that of a fount to be employed in books is another. Our contention,
in this respect, is that the necessities of a mass-production book and
the limited edition differ neither in kind nor in degree, since all
printing is essentially a means of the multiplication of a text set in an
alphabetical code of conventional symbols. To disallow "variety" in
the vital details of the composition is not to insist upon uniformity in
display. As already pointed out, the preliminary pages offer scope for
the utmost typographical ingenuity. Yet even here, a word of caution
may be in place, so soon do we forget, in arranging any piece of
display (above all, a title page), the supreme importance of sense.
Every character, every word, every line should be seen with
maximum clearness. Words should not be broken except
unavoidably, and in title pages and other compositions of centred
matter, lines should hardly begin with such feeble parts of speech as
prepositions and conjunctions. It is more reasonable, as assisting the
reader's immediacy of comprehension, to keep these to the ends of
lines or to centre them in smaller type and so bring out the salient
lines in a relatively conspicuous size.
No printer, in safeguarding himself from the charge of monotony in
his composition, should admit, against his better judgment, any
typographical distraction doing violence to logic and lucidity in the
supposed interests of decoration. To twist his text into a triangle,
squeeze it into a box, torture it into the shape of an hour-glass or a
diamond is an offence requiring greater justification than the
existence either of Italian and French precedents of the fifteenth and
sixteenth centuries, or of an ambition to do something new in the
twentieth. In truth, these are the easiest tricks of all, and we have
seen so much of them during the late "revival of printing" that we

now need rather a revival of restraint. In all permanent forms of
typography, whether publicly or privately printed, the typographer's
only purpose is to express, not himself, but his author. There are,
admittedly, other purposes which enter into the composition of
advertisement, publicity and sales matter; and there is, of course, a
very great deal common to both book and advertisement
composition. But it is not allowable to the printer to relax his zeal for
the reader's comfort in order to satisfy an ambition to decorate or to
illustrate. Rather than run this risk the printer should strive to
express himself by the use of this or that small decorative unit,
either of common design supplied by the type founders or drawn for
his office by an artist. It is quite true that to an inventive printer
decoration is not often necessary. In commercial printing, however, it
seems to be a necessity, because the complexity of our civilization
demands an infinite number of styles and characters. Publishers and
other buyers of printing, by insisting upon a setting which shall
express their business, their goods, their books and nobody else's
business or goods or books, demand an individuality which pure
typography can never hope to supply. But book-printers, concerned
with the permanently convenient rather than with the transiently
sensational or the merely fashionable, should be on their guard
against title-page borders, vignettes and devices invented to ease
their difficulties. There is no easy way with most title pages; and the
printer's task is rendered more difficult by the average publisher's
and author's incompetence to draft a title or to organize the
preliminaries in reasonable sequence.
VI
Those who would like to lessen or vary the tendency towards
standardization in day-today book production have a field for their
activity in the last-mentioned pages. The position on the page of the
half-title, title, dedication, etc., and their relation to each other, are
not essentially invariable. Nevertheless, as it is well for printers and

publishers to have rules, and the same rules, it may be suggested
that the headings to Preface, Table of Contents, Introduction, etc.,
should be in the same size and fount as the chapter heads; and
should be dropped if they are dropped. The order of the
preliminaries remains to be settled. With the exception of the
copyright notice, which may be set on the verso of the title page, all
should begin on a recto. The logical order of the preliminary pages is
Half-title or Dedication (I see no reason for including both), Title,
Contents, Preface, Introduction. The certificate of "limitation," in the
case of books of that class, may face the title where there is no
frontispiece, be incorporated with the half-title, or be taken to the
end of the volume. This order is applicable to most categories of
books. Novels need neither Table of Contents nor List of Chapters,
though one or the other is too often printed. If it is decided to retain
either, it would be reasonable to print it on the back of the half-title
and facing the title page, so that the structure, scope and nature of
the book will be almost completely indicated to the reader at a single
opening. Where the volume is made up of a few short stories, their
titles can be listed in the otherwise blank centre of the title page.
VII
Fiction, Belles-Lettres and Educational books are habitually first
published in portable, but not pocketable formats; crown octavo (5
by 7-1/2 in.) being the invariable rule for novels published as such.
The novel in the form of Biography will be published as a Biography,
demy octavo (5-5/8 by 8-3/4 in.), the size also for History, Political
Study, Archaeology, Science, Art and almost everything but Fiction.
Novels are only promoted to this format when they have become
famous and "standard"; when they are popular rather than famous
they are composed in pocket (4-1/2 by 6-3/4 in.) editions. Size,
therefore, is the most manifest difference between the categories of
books.

Another obvious difference is bulk, calculated in accordance with the
publisher's notion, first, of the general sense of trade expectation
and, secondly, of the purchasing psychology of a public habituated
to certain selling prices vaguely related to number of pages and
thickness of volume (inconsistently enough, weight does not enter
into these expectations). These habits of mind have consequences in
the typography; they affect the choice of fount and size of type, and
may necessitate the adoption of devices for "driving out," i.e.,
making the setting take up as much room as possible. By putting the
running headline between rules or rows of ornaments; introducing
unnecessary blanks between chapters; contracting the measure;
exaggerating the spaces between the words and the lines;
excessively indenting paragraphs; isolating quoted matter with areas
of white space: inserting wholly unnecessary sectional titles in the
text and surrounding them with space; contriving to drive a chapter
ending to the top of a recto page so that the rest of it and its verso
may be blank; using thick paper; increasing the depth of chapter
beginnings and inserting very large versals thereto; and so on, the
volume can be inflated to an extra sixteen pages and sometimes
more—which is a feat the able typographer is expected to
accomplish without showing his hand.
Limited editions of standard authors, or of authors whose publishers
desire them to rank as such, are commonly given a rubricated title
or some other feature not strictly necessary. A dreadful example of
overdone rubrication is to be found in an edition of Thomas Hardy's
verse, in which the running heads throughout the book are in red—
the production of a firm which desired to make an impression on the
purchaser in view of the price asked for the edition. This could have
been better done by reserving colour for the initial letters.
Handmade paper is generally used for éditions de luxe, and none but
the brave among publishers will disregard the superstitious love of
the book-buying classes for its untrimmed, ugly and dirt-gathering
edges. That most of the public prefer to have it so is because a
trimmed book looks "ordinary" to them. Any book which is
"different" from the "ordinary" in one superficial way or another is

apt to impress those lacking trade experience. And there has been a
notable increase during recent years in the category of books,
generally illustrated, known to the trade as fine printing, éditions de
luxe, press-books, limited editions, collectors' books, etc. Hence, it is
hoped that the above setting out of the first principles of typography
may give the discriminating reader some sort of yardstick which he
can apply not only to the entries catalogued by the book-sellers as
limited editions, but to the output of publishers responsible for
printing the literary and scientific books which are more necessary to
society, and are often designed with greater intelligence.

COMPOSED IN NEW TIMES ROMAN TYPES

CARL PURINGTON ROLLINS
American Type Designers and Their Work
Published by The Lakeside Press 1947-1948. Reprinted by permission of the
publisher.
A piece of paper about two inches square, originally pinned to the
manuscript of the Rev. Ezra Stiles's diary in the Yale University
Library is all that remains of the first original American type design.
It is a proof of letters made by Abel Buell, a Connecticut Yankee, in
1769.[35] Buell was his own designer, punch cutter, and caster, since
in his day, as for many years after, the making of type was entirely a
hand operation. Not the least exacting part of the work was the
cutting of the punch on the end of a short bar of softened steel. It
was not until the invention of the Benton pantograph punch-cutting
machine in 1885 that any other method was known. All type made
before 1885 was therefore dependent on hand punch cutting, and
the designer of the type was almost always the same man who cut
the punches.
Who these type designers were after Buell is a matter of uncertainty
and obscurity. The first type specimen book in America was that of
Binny & Ronaldson of Philadelphia, issued in 1812; and from then
almost to our own day the type foundries have taken the credit for
the type designs which they have offered for sale. Type designers,
like architects, got no credit; possibly Modesty, with a backward
glance at old specimen books, raised a warning finger, and the

designers were willing to let the foundries have whatever glory there
was.
The Punch Cutting Machine. Courtesy George Macy
Publications.
Abel Buell and his contemporaries and successors followed the
general trends in design in the arts as a whole. The Greek Revival
and the Victorian Age, marked by the two great expositions at
London and Philadelphia with their crudities and extravagancies in
design, found echoes in our imitative craft of printing. So it is not
surprising that type design began to improve, along with the other
arts, with the advent of the '90's. We have always followed European
and especially English models, and it is natural that the upheaval in
type design in England under Morris's influence had immediate
repercussions here. But while imitations of Kelmscott types were

soon on the market, two surprisingly original American designs
appeared at the same time as the imitations. About 1894 or 1895
the Central Type Foundry of St. Louis introduced a face which
became widely used, called (for no better reason than attends the
christening of most type faces) "De Vinne." It is of unknown
parentage, though there is some reason to suppose that it
descended from the Elzevirs; but it was a face of character and
distinction. At the same time the same foundry brought out another
design which had an acknowledged father—Will Bradley. Of this face
it has been said that it has "remarkably bold letters, with
peculiarities of form never before attempted." Thus we have in the
De Vinne and the Bradley faces two fresh and distinctively American
types, destined to be the forerunners of many others. And in one
case the name of the designer was definitely attached.
With the invention of the pantograph punch cutter, type design
became an "art" rather than a craft, and as might be expected the
personality of the designer became for various reasons more
important. It is not without interest that the chief designer of the
American Type Founders Company—a man responsible for almost
the whole type output of that foundry for many years—Morris Fuller
Benton, was the son of the man whose machines were responsible
for this revolution in type design. For it was the two basic machines
invented and developed by Linn Boyd Benton which made it possible
for those unskilled in the intricacies of type making to provide the
basic designs for type. The machines were very ingenious, and the
designs partook of the "faultily faultless, icily regular" perfection of
the mechanical device. This method of making type faces involved
the drawing of the design and the making of two or three patterns in
thin brass of the outline of the letter—each pattern good for several
sizes of type, and slightly modified for another group of sizes. This is
the way in which modern type is designed. It is the reason why such
a type series as "Cheltenham," designed by the architect Bertram G.
Goodhue in 1900 for the Mergenthaler Linotype Company, while very
expertly handled in the details, seems monotonous in mass; whereas

the Caslon type of the original cutting shows all the inevitable
variations of hand work.
A survey of the types of the first quarter of the present century,
made by the Editor of the Inland Printer in 1927, displays 161 type
faces brought out by seven or eight of the leading foundries
between 1900 and 1925. Of these, it was possible to name the
designers of 72, almost all from the Barnhart Brothers & Spindler
foundry of Chicago, whose records seem to have been in better
shape, or whose generosity was more spontaneous. Oswald Cooper,
Sydney Gaunt, Will Ransom, Robert Wiebking, and George Trenholm
were the chief names. It is unfortunate that the names of the
designers of the types put out by the American Type Founders
Company have not been preserved except in rare instances. Of
course, Benton was responsible for the greater portion, and on the
aesthetic side they occasionally scored a triumph as in the case of
the "Cloister" face.
The list included in the Inland Printer's survey fails to include some
of the outstanding designs of the period. Goodhue's "Merrymount"
was done in 1894, but after 1900 we have Mr. Rogers's "Centaur,"
Mr. Hunter's odd but forceful types (properly cut on punches by the
designer), the output of the rapidly growing composing-machine
industry, and Frederic W. Goudy's fifty designs completed in that
quarter century. Goudy's output of six score type designs in fifty
years is an amazing record, one probably never equalled. Such
designs as those for "Goudy Modern," "Goudy Text," and "Hadrian"
would establish his reputation. He had his limitations as a designer—
most of his designs lack a certain crispness—but his versatility was
extraordinary.
In the years since 1925 new designers have come to the fore:
Blumenthal with his "Emerson," Dwiggins with his "Electra" and
"Caledonia," Ruzicka with his "Fairfield," and Chappell with his
"Lydian." This brief survey cannot hope to mention all types or
designs which American designers have contributed, but it is well to
see if any tendencies can be detected.

The type which Buell made in 1759, as well as the type of his
immediate successors into the first decades of the nineteenth
century, were mainly variations on the so-called "modern romans" of
Didot, Bodoni, Austen, and Thorowgood. As the artistic styles in
design in general, not alone in type, gradually lost the evolutionary
force which has developed letter forms through the centuries,
eccentricity and anarchy came into play. The nineteenth-century
types as shown in the specimen books of Bruce, Connor, Farmer,
etc., and exhibited in all their grotesque horror in Fred Phillips' "Old-
fashioned Type Book," had no legitimate parentage, and they are as
well relegated to the bizarre and pseudo-nostalgic advertisement.
The result of the Kelmscott "revival" was to turn attention to type
forms of the past which could be revived for modern use, and the
type designers after 1900 did a remarkable piece of work in
introducing good type faces. The advertisers have been eager to use
new and novel faces, and have greatly stimulated this activity, even
in many cases over-exciting it. The most interesting result has been
the renewed interest in calligraphy. First directed toward new forms
of script, the truer form of broad pen lettering is now beginning to
influence type design, to free it from too slavish a devotion on the
one hand to the serif, and on the other to a too-free rejection of the
serif altogether. Such a face as Mr. Chappell's "Lydian" is an example
of real advance in design, and if one could adduce European
examples, more could be cited.
American designers have not developed many new or good book
faces; such types as Oxford, Centaur, Emerson, Fairfield, Electra, are
the exception. Their efforts have been given to the drawing of
display and advertising types—too often not to the enrichment of the
printer's repertory. It is quite as true now as in the past that
distortions of the normal Roman letter form in the direction of extra
condensed or extra heavy or very light mono-line letters result in
eccentricities which have no permanent value. On the other hand
such novel type designs as Garamond Bold Italic, Hadriano, the
newspaper Ionics, and Lydian are meritorious additions to the
printer's fonts. When it is realized that eccentricity and originality are

not the same thing, we may expect from our increasingly intelligent
designers indigenous types of usefulness and charm.

FOOTNOTES:
[35] Reprinted in Lawrence C. Wroth's "The First Work with
American Types," page 65.

TYPOGRAPHY—ERIC GILL
From Printing & Piety, An Essay on Typography by Eric Gill. Copyright 1931 by J.
M. Dent and Sons, Ltd., London. Reprinted by permission of the publisher.
One of the most alluring enthusiasms that can occupy the mind of
the letterer is that of inventing a really logical and consistent
alphabet having a distinct sign for every distinct sound. This is
especially the case for English speaking people: for the letters we
use only inadequately symbolize the sounds of our language. We
need many new letters and a revaluation of existing ones. But this
enthusiasm has no practical value for the typographer; we must take
the alphabets we have got, and we must take these alphabets in all
essentials as we have inherited them.
First of all, then, we have the ROMAN ALPHABET of CAPITAL letters
(Upper-case), and second the alphabet which printers call ROMAN
LOWER-CASE. The latter, tho' derived from the Capitals, is a distinct
alphabet. Third we have the alphabet called ITALIC, also derived
from the Capitals but through different channels. These are the
three alphabets in common use for English people.
Are there no others? It might be held that there are several; there
are, for example, the alphabet called Black Letter, and that called
Lombardic. But these are only partial survivals, and very few people
could, without reference to ancient books, write down even a
complete alphabet of either. As far as we are concerned in modern
England, Roman Capitals, Lower-case and Italics are three different
alphabets, and all are current "coin." But however familiar we are

with them, their essential differences are not always easily
discovered. It is not a matter of slope or of serifs or of thickness or
thinness. These qualities, though one or other of them may be
commonly associated with one alphabet more than another, are not
essential marks of difference. A Roman Capital A does not cease to
be a Roman Capital A because it is sloped backwards or forwards,
because it is made thicker or thinner, or because serifs are added or
omitted; and the same applies to Lower-case and Italics (see Fig. 1).

Figure 1 illustrates the contention that slope in either
direction does not deprive Capitals, Lower-case or Italics
of their essential differences.
Figure 2 in which the upper line of letters is essentially
"Roman Lower-case"; the lower essentially "Italic."
The essential differences are obviously between the forms of the
letters. The following letters, abdefghklmnqrtu and y, are not Roman
Capitals, and that is all about it. The letters shown in the lower line
of Fig. 2 are neither Capitals nor Lower-case. The conclusion is
obvious: there is a complete alphabet of Capital letters, but the
Lower-case takes ten letters from the Capital alphabet, and the Italic
takes ten from the Capitals and twelve from the Lower-case. Figure
3 shows the three alphabets completed, and it will be seen that
CIJOPSVWX and Z are common to all three, that bdhklmnqrtu and y
are common to Lower-case and Italics; that ABDEFGHKLMNQRTU
and Y are always Capitals; and that aef and g are always Lower-
case.

Figure 3 shows the differences and similarities between
the three "current" alphabets. Note: the curve of the Italic
y's tail is due to exuberance, and not to necessity.
But tho' this is a true account of the essential differences between
the three alphabets, there are customary differences which seem
almost as important. It is customary to make Roman Capitals
upright. It is customary to make Lower-case smaller than Capitals
when the two are used together; and it is customary to make Italics
narrower than Lower-case, sloping towards the right and with
certain details reminiscent of the cursive hand-writing from which
they are derived. Fig. 4 shows the three alphabets with their
customary as well as their essential differences.
Figure 4 shows the Capitals, Roman Lower-case and
Italics with their customary as well as their essential
differences.

Properly speaking there is no such thing as an alphabet of Italic
Capitals, and where upright or nearly upright Italics are used
ordinary upright Roman Capitals go perfectly well with them. But as
Italics are commonly made with a considerable slope and cursive
freedom, various sorts of sloping and quasi-cursive Roman Capitals
have been designed to match. This practice has, however, been
carried to excess; the slope of Italics and their cursiveness have
been much overdone. In the absence of punch cutters with any
personal sensibility as letter designers, with punch cutting almost
entirely done by machine, the obvious remedy is a much more
nearly upright and non-cursive Italic, and for Capitals the ordinary
upright Roman. Even with a nearly upright Italic, the mere presence
of the Italic aef and g alters the whole character of a page, and with
a slight narrowness as well as a slight slope, the effect is quite
different from that of a page of Lower-case.
The common practice of using Italics to emphasize single words
should be abandoned in favour of the use of the ordinary Lower-
case with spaces between the letters (l e t t e r - s p a c e d). The
proper use of Italics is for quotations and footnotes, and for books in
which it is or seems desirable to use a lighter and less formal style of
letter. In a book printed in Italics upright Capitals may well be used,
but if sloping Capitals be used they should only be used as initials—
they go well enough with Italic Lower-case, but they do not go with
one another.
We have, then, the three alphabets, and these are the printer's main
outfit; all other sorts of letters are in the nature of fancy letters,
useful in inverse proportion to the importance and quantity of his
output. The more serious the class of book he prints, the wider the
public to whom he appeals, so much the more solemn and
impersonal and normal will be and should be his typography. But he
will not call that book serious which is merely widely bought, and he
will not call that a wide appeal which is made simply to a mob of
forcibly educated proletarians. A serious book is one which is good in
itself according to standards of goodness set by infallible authority,

and a wide appeal is one made to intelligent people of all times and
nations.
The invention of printing and the breakdown of the medieval world
happened at the same time; and that breakdown, tho' hastened by
corruption in the Church, was chiefly caused by the recrudescence of
a commercialism which had not had a proper chance since the time
of the Romans. The invention of double-entry book-keeping also
happened about the same time, and though, as with modern
mechanical invention, the work was done by men of brains rather
than men of business, it was the latter who gained the chief
advantage.
Printing, a cheaper method of reproducing books than hand-writing,
came therefore just at the right moment. Since its first fine careless
rapture, and in spite of the genuinely disinterested efforts of
ecclesiastical presses, University presses and the work of many
notable individual printers and type-founders, the history of printing
has been the history of its commercial exploitation. As is natural with
men of business, the worse appears the better reason. Financial
success is, rightly, their only aim, and technical perfection the only
criterion they know how to apply to their works.
TYPOGRAPHY (the reproduction of lettering by means of movable
letter types) was originally done by pressing the inked surface or
"face" of a letter made of wood or metal against a surface of paper
or vellum. The unevenness and hardness of paper, the irregularities
of types (both in respect of their printing faces and the dimensions
of their "bodies") and the mechanical imperfections of presses and
printing methods made the work of early printers notable for
corresponding unevennesses, irregularities and mechanical
imperfections. To ensure that every letter left its mark more or less
completely and evenly, considerable and noticeable impression was
made in the paper. The printed letter was a coloured letter at the
bottom of a ditch.
The subsequent development of typography was chiefly die
development of technical improvements, more accurately cast types,

smoother paper, mechanically perfect presses. Apart from the history
of its commercial exploitation, the history of printing has been the
history of the abolition of the impression. A print is properly a dent
made by pressing; the history of letter-press printing has been the
history of the abolition of that dent.
But the very smooth paper and the mechanically very perfect
presses required for printing which shall show no "impression" can
only be produced in a world which cares for such things, and such a
world is of its nature inhuman. The industrial world of today is such,
and it has the printing it desires and deserves. In the industrial world
Typography, like house building and sanitary engineering, is one of
the necessary arts—a thing to be done in working hours, those
during which one is buoyed up by the knowledge that one is serving
one's fellow men and neither enjoying oneself like an artist nor
praising God like a man of prudence. In such a world the only
excuse for anything is that it is of service.
Printing which makes any claim on its own account, printers who
give themselves the status of poets or painters, are to be
condemned; they are not serving; they are shirking. Such is the tone
of the more romantic among men of commerce; and the
consequence is a pseudo-asceticism and a bastard aesthetics. The
asceticism is only a sham because the test of service is the profits
shown in the accounts; and the aesthetics is bastard because it is
not founded upon the reasonable pleasure of the mind of the
workman and of his customer, but upon the snobbery of museum
students employed by men of commerce to give a saleable
appearance to articles too dull otherwise to please even the readers
of The Daily Mail.
Nevertheless, as we have already shown, commercial printing,
machine printing, industrial printing would have its own proper
goodness if it were studiously plain and starkly efficient. Our quarrel
is not with such a thing but only with the thing that is neither one
nor the other—neither really mechanically perfect and physically
serviceable nor really a work of art, i.e., a thing made by a man

who, however laughable it may seem to men of business, loves God
and does what he likes, who serves his fellow men because he is
wrapped up in serving God—to whom the service of God is so
commonplace that it is as much bad form to mention it as among
men of business it is bad form to mention profits.
There are, then, two typographies, as there are two worlds; and,
apart from God or profits, the test of one is mechanical perfection,
and of the other sanctity—the commercial article at its best is simply
physically serviceable and, per accidens, beautiful in its efficiency;
the work of art at its best is beautiful in its very substance and, per
accidens, as serviceable as an article of commerce.
The typography of Industrialism, when it is not deliberately diabolical
and designed to deceive, will be plain; and in spite of the wealth of
its resources—a thousand varieties of inks, papers, presses and
mechanical processes for the reproduction of the designs of tame
designers—it will be entirely free from exuberance and fancy. Every
sort of ornament will be omitted; for printers' flowers will not spring
in such a soil, and fancy lettering is nauseating when it is not the
fancy of type-founders and printers but simply of those who desire
to make something appear better than it is. Paradoxical though it be,
the greater the wealth of appliances, the less is the power of using
it. All the while that the technical and mechanical good quality is
increasing, the de-humanizing of the workmen is also increasing. As
we become more and more able to print finer and more elaborate
and delicate types of letter it becomes more and more intellectually
imperative to standardize all forms and obliterate all elaborations
and fancifulness. It becomes easier and easier to print any kind of
thing, but more and more imperative to print only one kind.
On the other hand, those who use humane methods can never
achieve mechanical perfection, because the slaveries and
standardizations of Industrialism are incompatible with the nature of
men. Humane Typography will often be comparatively rough and
even uncouth; but while a certain uncouthness does not seriously
matter in humane works, lack of uncouthness is the only possible

excuse for the productions of the machine. So while in an
industrialist society it is technically easy to print any kind of thing, in
a humane society only one kind of thing is easy to print, but there is
every scope for variety and experiment in the work itself. The more
elaborate and fanciful the industrial article becomes, the more
nauseating it becomes—elaboration and fancifulness in such things
are inexcusable. But there is every excuse for elaboration and fancy
in the works of human beings, provided that they work and live
according to reason; and it is instructive to note that in the early
days of printing, when human exuberance had full scope, printing
was characterized by simplicity and decency; but that now, when
such exuberance no longer exists in the workman (except when he
is not at work), printing is characterized by every kind of vulgarity of
display and complicated indecency.
But, alas for humanity, there is the thing called compromise; and the
man of business who is also the man of taste, and he of taste also
who is also man of business will, in their blameless efforts to earn a
living (for using one's wits is blameless, and earning a living is
necessary), find many ways of giving a humane look to machine-
made things or of using machinery and the factory to turn out, more
quickly and cheaply, things whose proper nature is derived from
human labor. Thus we have imitation "period" furniture in Wardour
Street, and we have imitation "arts and crafts" in Tottenham Court
Road. The-man-of-business-who-is-also-man-of-taste will tend to the
"period" work, the-man-of-taste-who-is-also-man-of-business will
tend to the imitation handicrafts. And, in the printing world, there
are business houses whose reputation is founded on their
resuscitations of the eighteenth century, and private presses whose
speed of output is increased by machine-setting and gas engines.
These things are more deplorable than blameworthy. Their chief
objectionableness lies in the fact that they confuse the issue for the
ordinary uncritical person, and they turn out work which is neither
very good nor very bad. "Period" printing looks better than the usual
vulgar products of unrestrained commercialism, and there is no
visible difference, except to the expert, between machine-setting

and hand-setting, or between sheets worked on a hand-press and
those turned out on a power-driven Platen.
Nevertheless, even if these things be difficult to decide in individual
instances, there can be no sort of doubt but that as industrialism
requires a different sort of workman so it also turns out a different
kind of work—a workman sub-human in his irresponsibility, and work
inhuman in its mechanical perfection. The imitation of the work of
pre-industrial periods cannot make any important ultimate
difference; the introduction of industrial methods and appliances into
small workshops cannot make such workshops capable of
competition with "big business." But while false standards of good
taste may be set up by "period" work, this "good taste" is entirely
that of the man of business and his customers; it is not at all that of
the hands—they are in no way responsible for it or affected by it; on
the other hand, the introduction of mechanical methods into small
workshops has an immediate effect on the workmen. Inevitably they
tend to take more interest in the machine and less in the work, to
become machine-minders and to regard wages as the only reward.
And good taste ceases to be the result of the restraint put upon his
conscience by the workman himself; it becomes a thing imposed
upon him by his employer. You cannot see the difference between a
machine-set page and one set by hand. No, but you can see the
difference between Cornwall before and after it became "the English
Riviera"; you can see the difference between riding in a hansom and
in a motor-cab—between a "cabby" and a "taxi-man"; you can see
the difference between the ordinary issue of The Times today and its
ordinary issue a hundred years ago; you can see the difference
between an ordinary modern book and an ordinary book of the
sixteenth century. And it is not a question of better or worse; it is a
question of difference simply. Our argument here is not that
Industrialism has made things worse, but that it has inevitably made
them different; and that whereas before Industrialism there was one
world, now there are two. The nineteenth century attempt to
combine Industrialism with the Humane was necessarily doomed,
and the failure is now evident. To get the best out of the situation

we must admit the impossibility of compromise; we must, in as
much as we are industrialists, glory in Industrialism and its powers
of mass-production, seeing that good taste in its products depends
upon their absolute plainness and serviceableness; and in so much
as we remain outside Industrialism, as doctors, lawyers, priests and
poets of all kinds must necessarily be, we may glory in the fact that
we are responsible workmen and can produce only one thing at a
time.
That if you look after goodness and truth beauty will take care of
itself, is true in both worlds. The beauty that Industrialism properly
produces is the beauty of bones; the beauty that radiates from the
work of men is the beauty of the living face.
COMPOSED IN PERPETUA TYPES

FREDERIC W. GOUDY
TYPES AND TYPE DESIGN
The Syracuse University School of Journalism awarded its first medal of honor to F.
W. G. in 1936, "for distinctive achievement in typographic design." His address
then, reflecting the typographic philosophy and practice of two-score years, is
reprinted as published by the University in 1936.
It would be mere affectation on my part were I to pretend not to be
touched by the signal honor you extend to me this evening, and I
would be ungrateful indeed if I neglected to voice my very great
appreciation of your kindness. I wish that I might express that
appreciation in words that would leave no shadow of doubt in your
minds as to the depth and sincerity of my feeling.
I am not conscious of any outstanding reasons for the kind words
spoken here tonight of my work. At the same time I am under no
illusions as to the ultimate value of the work I have attempted to do,
although it is, after all, merely the every-day work of an earnest
craftsman who endeavors to perform each task well and the next

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