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Chapter 1: Principles of Biochemistry
MULTIPLE CHOICE
1. The birth of modern biochemistry can be traced to the

a. end of the 19th century.
b. end of the 20th century.
c. beginning of the 19th century.
d. beginning of the 21st century.
ANS: A DIF: Easy REF: 1.1
OBJ: 1.1.a. List examples where biochemistry has made advancements in the lives of many
humans.
MSC: Remembering
2. The study of biochemistry attempts to explain

a. chemical processes at the atomic level.
b. biological processes at the molecular and cellular level.
c. the nature of life.
d. physical processes at the macromolecular level.
ANS: B DIF: Easy REF: 1.1
humans.
MSC: Remembering
3. Enzymes function as reaction catalysts in cells. If the enzymes were removed from a cell, the rate
of biochemical reactions would
a. increase.
b. remain the same.
c. decrease.
d. It is impossible to know.
ANS: C DIF: Medium REF: 1.1
humans.
MSC: Understanding
4. The essential ions calcium, magnesium, potassium, and sodium are all
a. anions.
b. cations.
c. transition metals.
d. halogens.
ANS: B DIF: Difficult REF: 1.2
OBJ: 1.2.a. List the elements that are most abundant in living organisms.
MSC: Analyzing
5. Look at the bond energies O-H, N-H, and P-H in the table below. O-H is the hardest bond to break
because it has the
a. greatest difference in relative affinities of the two atoms for electrons.

b. smallest difference in relative affinities of the two atoms for electrons.
c. smallest difference in atomic size.
d. largest difference in atomic size.
ANS: A DIF: Medium REF: 1.2
OBJ: 1.2.b. Identify the most abundant functional groups found in biomolecules.
MSC: Understanding
6. Given that methane (CH4) has a bond angle of and ethylene (C2H2) has a bond angle of
, what is the correct bond angle for acetylene (C2H2)?
a.
b.
c.
d.
MSC: Applying
7. Amino acids are the building blocks for which biomolecule(s)?

a. proteins
b. DNA
c. carbohydrates
d. micelles
OBJ: 1.2.c. Name the four major classes of biomolecules and the primary cellular functions
associated with each. MSC: Remembering
8. The __________ differentiates amino acids from one another.

a. number of silane groups
b. number of phosphoryl groups
c. side chains attached to the central carbon
d. number of hydroxyl groups
associated with each. MSC: Understanding
9. A nucleotide consists of which of the following?

a. nitrogenous base, four-membered sugar and phosphate groups
b. phosphate base, four-membered sugar and sulfate groups
c. nitrogenous base, five-membered sugar and phosphate groups
d. carboxylic acid, four-membered sugar and phosphate groups
10. Simple sugars are made of which of the following elements?
a. carbon, sulfur, and hydrogen
b. carbon, oxygen, and phosphate
c. carbon, oxygen, and helium
d. carbon, oxygen, and hydrogen
ANS: D DIF: Easy REF: 1.2
11. If energy in the form of ATP is required to make a polymeric macromolecule, which of the
following will happen if there is no ATP available?
a. The rate of polymer increases.

b. The polymer is broken down to release ATP.
c. The polymer continues to be made at the same rate.
d. The enzyme degrades to release ATP.
ANS: B DIF: Medium REF: 1.2
OBJ: 1.2.e. Explain the role of metabolic pathways in living systems.
MSC: Understanding
12. Why are fewer polypeptide sequences encountered biologically than are theoretically possible?
a. There is no way to make all the theoretical possibilities.
b. The phosphodiester linkages don’t allow for all the possibilities.
c. Not all have useful structural and functional properties.
d. Not all of the possibilities can be broken down.
OBJ: 1.2.d. Identify the biomolecules that form polymers. MSC: Applying
13. Humans do not have the enzyme cellulase. Is it likely that a human could survive on a plant-only
diet?
a. No, not enough ATP would be produced to generate energy.
b. No, not enough DNA would be produced to generate energy.
c. Yes, cellulase is not necessary to break down plant material.
d. Yes, ATP is not necessary to maintain life.
MSC: Analyzing
14. Which two functional groups are involved in producing a peptide bond?
a. alcohol and amino
b. amino and thiol
c. methyl and amino
d. amino and carboxyl
ANS: D DIF: Medium REF: 1.2
MSC: Understanding
15. Even though amylose and cellulose contain the same repeating unit of glucose, they are very
different in terms of function. Why?
a. A glycosidic bond cannot be cleaved.
b. The orientations of the glycosidic bond are different.
c. ATP cannot be generated from amylose.
d. There is no structural difference between the polymers.
OBJ: 1.2.d. Identify the biomolecules that form polymers. MSC: Applying
16. If the concentration of aspartate in the cell decreased, what would be the predicted outcome?
a. increased concentration of argininosuccinate

b. decreased concentration of citrulline
c. decreased concentration of fumarate
d. increased concentration of arginine
ANS: C DIF: Difficult REF: 1.2
MSC: Applying
17. The correct definition of a pathway intermediate is a molecule that

a. is both a product and a reactant in a pathway.
b. lowers the activation energy of a reaction.
c. increases the rate of a reaction.
d. is only a reactant in a pathway.
MSC: Understanding
18. If the concentration of F is high in a cell, the pathway will MOST likely shift to produce
a. more C.
b. less C.
c. more A.
d. less D.
ANS: D DIF: Difficult REF: 1.2
MSC: Applying
19. Plasmids are small, circular DNA molecules that are used in which of the following?
a. gene cloning
b. production of chromatin
c. cell movement
d. replication of nucleus
MSC: Remembering
20. What is the function of the chloroplast in a plant cell?
a. detoxification of macromolecules
b. degradation of macromolecules
c. conversion of light energy to chemical energy
d. conversion of glucose to ATP
OBJ: 1.2.f. Compare and contrast bacterial and eukaryotic cells.
MSC: Understanding
21. If a plasma membrane is hydrophobic, what kinds of amino acids are MOST likely to be found in
the membrane?
a. hydrophilic amino acids
b. hydrophobic amino acids
c. polar amino acids
d. charged amino acids
OBJ: 1.2.g. Name the key organelles found in a eukaryotic cell.
MSC: Applying
22. When a ligand binds to a receptor, it causes the receptor to

a. degrade.
b. activate.
c. deactivate.
d. rapidly grow.
MSC: Understanding
23. How does the molecule adenosine monophosphate fit into the seven hierarchical levels that define
the chemical basis of life?
a. element/functional group
b. biomolecule
c. metabolism
d. organism
MSC: Understanding
24. The main difference between deoxyribonucleotides and ribonucleotides is that they have a
different
a. number of carbons in the sugar ring.
b. functional group on the carbon.
c. number of phosphates on the carbon.
d. functional group on the carbon.
OBJ: 1.3.b. Compare and contrast DNA and RNA. MSC: Understanding
25. A hydrogen bond can best be described as a

a. strong covalent interaction.
b. strong ionic interaction.
c. weak noncovalent interaction.
d. weak covalent interaction.
ANS: C DIF: Easy REF: 1.3
OBJ: 1.3.a. Identify the three components of a nucleotide. MSC: Remembering
26. Why can a guanine not be paired with adenine?

a. Guanine is only found in RNA and adenine is found only in DNA.
b. Guanine can form three hydrogen bonds and adenine can form two.
c. Guanine can form only two hydrogen bonds and adenine can form three.
d. Guanine can only pair with thymine.
OBJ: 1.3.b. Compare and contrast DNA and RNA. MSC: Analyzing
27. What structural feature of DNA is attributed to the fact that the two DNA strands are antiparallel?
a. DNA is a left-handed helix.
b. DNA has the phosphate groups on the interior.
c. DNA is a right-handed helix.
d. DNA forms a pleated sheet.
OBJ: 1.3.b. Compare and contrast DNA and RNA. MSC: Applying
28. The “central dogma of molecular biology” can best be described as the transfer of information
between
a. nucleic acids and proteins.
b. fatty acids and proteins.
c. carboxylic acids and proteins.
d. nucleic acids and DNA.
OBJ: 1.3.c. Define the central dogma of molecular biology. MSC: Remembering
29. A segment of DNA containing 20 base pairs includes 8 adenine residues. How many uracil
residues are present?
a. 12
b. 8
c. 0
d. 28
30. A genome is a set of

a. proteins.
b. transcripts.
c. genes.
d. genetic code.
OBJ: 1.3.b. Compare and contrast DNA and RNA. MSC: Remembering
31. Given a DNA sequence of -CAT- , what is the complementary sequence in mRNA?
a. -GUA-
b. -AUG-
c. -GUA-
d. -GTA-
ANS: A DIF: Difficult REF: 1.3
32. Who received the Nobel Prize in 1962 for elucidating the molecular structure of DNA?
a. Albert Einstein
b. Linus Pauling
c. John Kendrew and Max Perutz
d. Maurice Wilkins, James Watson, and Francis Crick
33. A single nucleotide base substitution in a wild-type DNA is an example of

a. random mutation.
b. transcription.
c. translation.
d. cloning.
OBJ: 1.4.a. Differentiate between germ-line cell mutations and somatic cell mutations.
MSC: Remembering
34. An inherited disease comes from the mutation of DNA in a __________ cell.
a. somatic
b. germ-line
c. adipose
d. blood
MSC: Understanding
35. The RNA world model is based on the hypothesis that __________ is stable.
a. RNA
b. DNA
c. uracil
d. adenine
MSC: Understanding
36. Bioinformatics shows that 98% of human DNA is identical to that of chimpanzees. If human DNA
contains 3.2 billion nucleotides, how many nucleotides are different between the two species?
a. 3.1 billion
b. 64 million
c. 3.1 million
d. 640 million
OBJ: 1.4.c. Identify the relationship between protein structure and function.
MSC: Applying
37. Highly conserved gene sequences that encode proteins with the same function in different
organisms are called __________ genes.
a. orthologous
b. conserved
c. parallel
d. antiparallel
OBJ: 1.4.b. Differentiate between orthologous genes and paralogous genes.
MSC: Understanding
38. If a mutation was made to the gene for glucose–6–phosphate dehydrogenase that prevented it from
functioning, a possible outcome would be the production of
a. more NADPH.
b. less NADPH.
c. more ATP.
d. less ATP.
MSC: Applying
39. The appearance of new gene speciation is an example of

a. gene multiplication.
b. gene singulation.
c. gene duplication.
d. random mutation.
MSC: Understanding
40. The amino acid sequence of a protein determines its structure. Which of the following statements
is true?
a. Two proteins with similar amino acid sequence should have similar structures.
b. Two proteins with different amino acid sequences will have identical structures.
c. Two proteins with similar amino acid sequences will always have the same function in a
cell.
d. It is impossible to determine how proteins will fold based on the amino acid sequence
alone.
MSC: Analyzing
41. Which of the following is NOT a common functional group?

a. COOH
b. CH3
c. SH
d. CHO
MSC: Understanding
42. By convention, nucleic acid chains are written starting at the __________ end.
a. amino
b. carboxyl
c.
d.
MSC: Understanding
43. The proposal that DNA is a double helix was based on what experimental evidence?
a. NMR
b. IR
c. HPLC
d. x-ray crystallography
MSC: Understanding
44. mRNA is used for what process in the cell?

a. as a template for protein synthesis
b. regulation of gene expression
c. regulation of RNA
d. replication of DNA
OBJ: 1.3.d. Define the terms transcriptome and proteome. MSC: Understanding
45. In DNA the phosphate groups are on the outside of the helix. Why does this stabilize the structure?
a. ionic interactions with the solvent
b. hydrogen bonding with itself
c. covalent binding to the solvent
d. It does not stabilize the structure.
46. What is the cause of the overall negative charge of a molecule of DNA?
a. hydrogen bonding between base pairs
b. the phosphate backbone
c. the sugars
d. the antiparallel orientation of the DNA
ANS: B DIF: Difficult REF: 1.3
OBJ: 1.3.b. Compare and contrast DNA and RNA. MSC: Evaluating
47. Mutations to proteins typically occur starting

a. at the protein itself.
b. with mRNA.
c. with tRNA.
d. with DNA.
OBJ: 1.3.c. Define the central dogma of molecular biology. MSC: Analyzing
48. Hydrogen bonds form between hydrogen and
a. oxygen.
b. helium.
c. carbon.
d. hydrogen.
49. The DNA double helix is stabilized by the interactions between nucleotides because of
__________ between nucleotides.
a. hydrogen bonding
b. pi–pi stacking
c. sigma bonds
d. ionic interactions
50. In structures of tRNA, base pairs form between

a. the same strand.
b. another strand of RNA.
c. a strand of DNA.
d. Base pairs do not form.
51. Ribose is a
a. four-carbon sugar.
b. five-carbon sugar.
c. six-carbon sugar.
d. five-carbon acid.
OBJ: 1.3.b. Compare and contrast DNA and RNA. MSC: Remembering
52. The process of fermentation uses sugar to produce which molecules?

a. alcohol and carbon monoxide
b. alcohol and carbon dioxide
c. acid and carbon monoxide
d. acid and carbon dioxide
humans.
MSC: Understanding
53. An example of experimental biochemistry is trying an experiment and

a. quitting after it fails to prove your hypothesis.
b. continuing to try with no changes to protocol.
c. optimizing experimental design.
d. successfully proving your hypothesis after the first attempt.
humans.
MSC: Applying
54. What side products of pyruvate are being converted into alcohol and carbon dioxide by yeast?
a. CO and NADH
b. CO2 and NADH
c. CO2 and NAD+
d. CO and NAD+
MSC: Applying
55. Pyruvate decarboxylase converts pyruvate into

a. carbon dioxide and acetaldehyde.
b. carbon monoxide and acetaldehyde.
c. water and glucose.
d. ethanol and carbon dioxide.
humans.
MSC: Understanding
56. Which of the following is an example of a metabolic pathway?

a. mitochondria
b. organelles
c. glycolysis
d. plasma membrane
MSC: Understanding
57. Which of the following is an example of an ecosystem?

a. mammals
b. plasma membrane
c. insects
d. forest
MSC: Understanding
58. A requirement for a covalent bond to form between two atoms is that
a. there are unpaired electrons on each atom.
b. the atoms are ions.
c. both the atoms must be metals.
d. one of the atoms must be a halogen.
MSC: Applying
59. What is the maximum number of covalent bonds a carbon atom can make?
a. 2
b. 4
c. 6
d. 8
MSC: Applying
60. The correct name for the VSEPR arrangement around a carbon in methane is
a. linear.
b. trigonal bipyrimidal.
c. tetrahedral.
d. octahedral.
MSC: Analyzing
61. Cell signaling and cell membranes are examples of functions performed by which biomolecule?
a. amino acid
b. nucleotide
c. simple sugar
d. fatty acid
associated with each. MSC: Applying
62. The figure below shows an example of which functional group?
a. amino
b. hydroxyl
c. phosphoryl
d. methyl
63. ATP is an abbreviation for which energy currency molecule?

a. adenosine triphosphate
b. adenosine thymine
c. adenosine dinucleotide
d. amino triphosphate
64. Which of the following is the correct formula for glucose?

a. C5H10O5
b. C6H12O6
c. C5H12O6
d. C6H10O6
65. What is another way to describe the amphipathic nature of a fatty acid?
a. polar head and nonpolar tail
b. nonpolar head and polar tail
c. polar head and double bonds in tail
d. nonpolar head and double bonds in tail
66. Triacylglycerols are neutral molecules made of

a. three fatty acid esters covalently linked to glycine.
b. three fatty acid esters covalently linked to glycerol.
c. two fatty acid esters covalently linked to glycine.
d. three acyls covalently linked to glycerol.
67. Proteins are a polymer of __________, whereas nucleic acids are polymers of __________.
a. nucleotides; fatty acid esters
b. nucleotides; amino acids
c. amino acids; nucleotides
d. sugars; nucleotides
OBJ: 1.2.d. Identify the biomolecules that form polymers. MSC: Understanding
68. Vitamin B2 is a metabolite. Lack of vitamin B2 can lead to blurred vision and a swollen tongue.
Vitamin B2 has such a strong effect on health because metabolites
a. are the catalysts that drive biochemical reactions necessary for life-sustaining processes.
b. are complex chemical reactions in cells.
c. process essential genetic information needed for life.
d. are needed for construction of microtubules.
MSC: Analyzing
69. Cyclic pathways contain several metabolites that regenerate during each turn of the cycle. Another
way to describe a metabolite is that it functions as a
a. reactant or product.
b. catalysis.
c. transition state.
d. enzyme.
MSC: Applying
70. Enzymes and chromosome are found where in the bacterial cell?
a. nucleoid
b. pili
c. cytoplasm
d. capsule
MSC: Understanding
71. What does chromatin in eukaryotic nucleus contain?

a. DNA packaged with proteins.
b. RNA packaged with proteins.
c. ATP packaged with proteins.
d. DNA packaged with ATP.
MSC: Understanding
72. The function of chloroplasts in plant cells is to convert

a. heat energy to light energy.
b. heat energy to chemical energy.
c. light energy to chemical energy.
d. chemical energy to heat energy.
MSC: Applying
73. What process replicates DNA to make more DNA?

a. DNA transcription
b. DNA replication
c. DNA translation
d. RNA transcription
OBJ: 1.3.c. Define the central dogma of molecular biology. MSC: Understanding
74. What process converts DNA to RNA?

b. DNA replication
c. DNA translation
d. RNA transcription
75. What process describes using mRNA templates to produce proteins?

b. DNA replication
c. DNA translation
d. mRNA translation
SHORT ANSWER
1. What are the three biochemical principles that together provide a framework for understanding life
at the molecular level? How are they interrelated?
ANS:
Hierarchical organization of biochemical processes within cells, organisms, and ecosystems
underlies the chemical basis of life on Earth. DNA is the chemical basis for heredity and encodes
the structural information for RNA and protein molecules, which mediate biochemical processes in
cells. The function of a biomolecule is determined by its molecular structure, which is fine-tuned
by evolution through random DNA mutations and natural selection. These processes cannot
function without each other.
DIF: Easy REF: Introduction

humans.
MSC: Understanding
2. List three examples of how biochemistry has made advancements in the lives of many humans.
ANS:
A list can be found in Section 1.1 of the text. Examples include developing new pharmaceutical
drugs, diagnostic tests, improved detergents, and faster ripening of fruit and vegetables.
DIF: Easy REF: 1.1

humans.
MSC: Remembering
3. Which six elements make up 97% of the weight of most organisms?
ANS:
Hydrogen, oxygen, carbon, nitrogen, phosphorus, and sulfur
DIF: Easy REF: 1.2

MSC: Remembering
4. Identify the following functional groups.
ANS:
Amino, hydroxyl, sulfhydryl, phosphoryl, carboxyl, methyl
DIF: Easy REF: 1.2

MSC: Understanding
5. Identify the following biomolecules.

ANS:
Amino acid, nucleotide, sugar, fatty acid
DIF: Easy REF: 1.2

6. What functions can polysaccharides perform?
ANS:
They provide structural support to cells and energy storage.
DIF: Difficult REF: 1.2

7. What is an advantage of using polypeptides for storage and information transmission?
ANS:
The complexity of monomeric units forms a stable arrangement that is perfect for information
storage and transmission.

OBJ: 1.2.d. Identify the biomolecules that form polymers. MSC: Evaluating
8. Explain how a linear pathway is different from a forked pathway.
ANS:
In a linear pathway, a reaction generates only one product that is then used in the next reaction. In
a forked pathway, two products are produced that each enter a different metabolic pathway.

MSC: Analyzing
9. Compare and contrast at least three different characteristics of prokaryotic and eukaryotic cells.
ANS:
Prokaryotes are often 1 in diameter, their cytoplasm contains all the enzymes and
chromosomes, and they have flagella and pili. Eukaryotic cells are 10 to 100 in diameter,
chromatin is contained in the nucleus, and they have a cytoskeleton.

MSC: Analyzing
10. Name four organelles found in a eukaryotic cell.
ANS:
Many answers are correct including the following: lysosome, smooth endoplasmic reticulum,
rough endoplasmic reticulum, ribosomes, peroxisome, mitochondria, nucleus, Golgi apparatus, and
cytoskeleton.
DIF: Easy REF: 1.2
MSC: Remembering
11. Describe how a receptor is activated and inactivated by a ligand in the figure below.
ANS:
The receptor ligand is bound to the receptor, which causes a conformation change to the active site
on the protein. The ligand is then released and returns the receptor to an inactive conformation.

OBJ: 1.2.h. Explain the role of receptors in multicellular organisms.
MSC: Analyzing
12. Name the three components of a nucleotide.
ANS:
Nucleotide base, five-carbon ribose, and one or more phosphate groups
DIF: Easy REF: 1.3

OBJ: 1.3.a. Identify the three components of a nucleotide. MSC: Remembering
13. Compare and contrast the bases found in DNA and RNA.
ANS:
DNA is composed of the deoxyribonucleotides (lacking an hydroxyl group on the position of
ribose) guanine, cytosine, adenine, and thymine. RNA is composed of ribonucleotides (containing
an hydroxyl group on the position of ribose) guanine, cytosine, adenine,
and uracil.
DIF: Medium REF: 1.3 OBJ: 1.3.b. Compare and contrast DNA and RNA.
MSC: Analyzing
14. Define the central dogma of molecular biology.
ANS:
The central dogma of molecular biology describes how genetic information stored in DNA is used
to direct the biological processes in cells.
DIF: Medium REF: 1.3

15. Compare and contrast a transcriptome and a proteome.
ANS:
Although both transcriptomes and proteomes are collections of genetic material, a collection of
DNA transcripts (RNA products) generated by DNA transcription is called a transcriptome,
whereas a proteome is the collection of proteins produced by mRNA translation.

OBJ: 1.3.d. Define the terms transcriptome and proteome. MSC: Analyzing
16. Differentiate between germ-line cell mutations and somatic cell mutations.
ANS:
Although both are mutations in DNA, if the mutation is passed from the parent to the offspring,
then the mutation is contained within the DNA of a germ-line cell. If the DNA mutation occurs
during the lifetime of the organism in a somatic cell, then this disease is limited to that individual
organism.

MSC: Analyzing
17. Differentiate between orthologous genes and paralogous genes.
ANS:
Orthologous genes are highly conserved gene sequences that encode proteins with the same
function in different organisms and are believed to have arisen from a common ancestral gene.
Paralogous genes are related genes within a species. Paralogous genes have orthologous genes in
other species.

OBJ: 1.4.b. Differentiate between orthologous genes and paralogous genes.
MSC: Analyzing
18. Evaluate the following statement: Two amino acid sequences with high sequence conservation
must have the same function in an organism.
ANS:
Although two amino acid sequences with high sequence conservation may have the same structure,
it does not mean that those two proteins will perform the same function in a body. As is shown in
Figure 1.30, similar structures may have different function.

MSC: Evaluating
19. For the figure below, label the and ends for each strand, the hydrogen bonds, and the
pyrimidine and purine molecules, as well as the direction of polarity.
ANS:
MSC: Remembering
20. Describe the difference between a deoxyribonucleotide and a ribonucleotide.
ANS:
Deoxyribonucleotides are monomeric units of DNA and lack a hydroxyl group on the carbon on
the position, whereas a ribonucleotide does have a hydroxyl group on the carbon on the on the
position.
MSC: Applying
21. Give an example of each of the following: element, biomolecule, macromolecule, metabolism,
cell, organism, and ecosystem.
ANS:
Any element from the periodic table is acceptable;
Biomolecules: amino acids, nucleotides, simple sugars, fatty acids;
Macromolecules: DNA/RNA, proteins, carbohydrates;
Metabolism: glycolysis, citrate cycle, urea cycle;
Cells: cell wall, plasma membrane, organelles;
Organisms: trees, mammals, fish, bird, insects;
Ecosystems: rivers, islands, forest, desert.
DIF: Medium REF: 1.2

MSC: Remembering
22. Amino acids are the building blocks of proteins. There are 20 amino acids; how do these amino
acids differ from one another?
ANS:
Amino acids differ from one another in the side chain attached to the central carbon.
DIF: Easy REF: 1.2

23. Describe the differences between the structures of pyrimidine and a purine.
ANS:
A pyrimidine is an aromatic molecule with nitrogen at positions 1 and 3 on the ring, along with a
carbonyl at position 4. Examples of pyrimidines are cytosine, thymine, and uracil. A purine is a
heterocyclic aromatic molecule with nitrogen at positions 1, 3, 7, and 9. Examples of purines are
guanine and adenine.

OBJ: 1.3.a. Identify the three components of a nucleotide. MSC: Analyzing
24. What is the function of mRNA in the cell?
ANS:
mRNA is used as templates for protein synthesis in a process referred to as mRNA translation.
DIF: Easy REF: 1.3 OBJ: 1.3.b. Compare and contrast DNA and RNA.
MSC: Analyzing
25. If you were a biochemist who just discovered a new protein, how would you gain insight into the
function of the protein?
ANS:
One way to gain insight into the function of a protein is to compare its amino acid sequence to
those of other proteins to see if conserved regions appear that might suggest an important function.
This is done using the genetic code to convert the DNA sequence in the coding stand of a gene into
the inferred amino acid sequence of the encoded protein.

MSC: Evaluating
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postulate and the recent development of atomic
theory
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Title: The quantum postulate and the recent development of atomic

theory
Author: Niels Bohr
Release date: January 29, 2024 [eBook #72800]
Language: English
Original publication: London: The London, Edinburgh, and Dublin

Philosophical Magazine and Journal of Science, 1915
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*** START OF THE PROJECT GUTENBERG EBOOK THE

QUANTUM POSTULATE AND THE RECENT DEVELOPMENT OF
ATOMIC THEORY ***
THE QUANTUM POSTULATE
AND THE RECENT
DEVELOPMENT OF ATOMIC
THEORY [1].
By Prof. N. BOHR, For. Mem. R.S.
Nature 121, 580-590 (1928)

Supplement to “Nature,” April 14, 1928.
CONTENTS
1. QUANTUM POSTULATE AND CAUSALITY.
2. QUANTUM OF ACTION AND KINEMATICS.
3. MEASUREMENTS IN THE QUANTUM THEORY.
4. CORRESPONDENCE PRINCIPLE AND MATRIX THEORY.
5. WAVE MECHANICS AND QUANTUM POSTULATE.
6. REALITY OF STATIONARY STATES.
7. THE PROBLEM OF THE ELEMENTARY PARTICLES.
ABSTRACT
IN connexion with the discussion of the physical interpretation of the

quantum theoretical methods developed during recent years, I
should like to make the following general remarks regarding the
principles underlying the description of atomic phenomena, which I
hope may help to harmonise the different views, apparently so
divergent, concerning this subject.
[1] The content of this paper is essentially the same as that of a
lecture on the present state of the quantum theory delivered on
Sept. 16, 1927, at the Volta celebration in Como. For a summary
of the theory just previous to the development of the new methods
the reader is referred to a lecture of the author, “Atomic Theory
and Mechanics,” published in this periodical (NATURE, 116, 809;
1925). The rapid development which has taken place since has
given rise to a considerable number of publications. The present
paper is confined to a few references to recent articles which have
a special bearing on the subject now under discussion.
1. quantum postulate and causality.
The quantum theory is characterised by the acknowledgment of a
fundamental limitation in the classical physical ideas when applied to
atomic phenomena. The situation thus created is of a peculiar
nature, since our interpretation of the experimental material rests
essentially upon the classical concepts. Notwithstanding the
difficulties which hence are involved in the formulation of the
quantum theory, it seems, as we shall see, that its essence may be
expressed in the so-called quantum postulate, which attributes to
any atomic process an essential discontinuity, or rather individuality,
completely foreign to the classical theories and symbolised by
Planck’s quantum of action.
This postulate implies a renunciation as regards the causal
space-time co-ordination of atomic processes. Indeed, our usual
description of physical phenomena is based entirely on the idea that
the phenomena concerned may be observed without disturbing them
appreciably. This appears, for example, clearly in the theory of
relativity, which has been so fruitful for the elucidation of the classical
theories. As emphasised by Einstein, every observation or
measurement ultimately rests on the coincidence of two independent
events at the same space-time point. Just these coincidences will
not be affected by any differences which the space-time co-
ordination of different observers otherwise may exhibit. Now the
quantum postulate implies that any observation of atomic
phenomena will involve an interaction with the agency of observation
not to be neglected. Accordingly, an independent reality in the
ordinary physical sense can neither be ascribed to the phenomena
nor to the agencies of observation. After all, the concept of
observation is in so far arbitrary as it depends upon which objects
are included in the system to be observed. Ultimately every
observation can of course be reduced to our sense perceptions. The
circumstance, however, that in interpreting observations use has
always to be made of theoretical notions, entails that for every
particular case it is a question of convenience at what point the
concept of observation involving the quantum postulate with its
inherent ‘irrationality’ is brought in.
This situation has far-reaching consequences. On one hand, the
definition of the state of a physical system, as ordinarily understood,
claims the elimination of all external disturbances. But in that case,
according to the quantum postulate, any observation will be
impossible, and, above all, the concepts of space and time lose their
immediate sense. On the other hand, if in order to make observation
possible we permit certain interactions with suitable agencies of
measurement, not belonging to the system, an unambiguous
definition of the state of the system is naturally no longer possible,
and there can be no question of causality in the ordinary sense of
the word. The very nature of the quantum theory thus forces us to
regard the space-time co-ordination and the claim of causality, the
union of which characterises the classical theories, as
complementary but exclusive features of the description, symbolising
the idealisation of observation and definition respectively. Just as the
relativity theory has taught us that the convenience of distinguishing
sharply between space and time rests solely on the smallness of the
velocities ordinarily met with compared to the velocity of light, we
learn from the quantum theory that the appropriateness of our usual
causal space-time description depends entirely upon the small value
of the quantum of action as compared to the actions involved in
ordinary sense perceptions. Indeed, in the description of atomic
phenomena, the quantum postulate presents us with the task of
developing a ‘complementarity’ theory the consistency of which can
be judged only by weighing the possibilities of definition and
observation.
This view is already clearly brought out by the much-discussed
question of the nature of light and the ultimate constituents of matter.
As regards light, its propagation in space and time is adequately
expressed by the electromagnetic theory. Especially the interference
phenomena in vacuo and the optical properties of material media are
completely governed by the wave theory superposition principle.
Nevertheless, the conservation of energy and momentum during the
interaction between radiation and matter, as evident in the
photoelectric and Compton effect, finds its adequate expression just
in the light quantum idea put forward by Einstein. As is well known,
the doubts regarding the validity of the superposition principle on one
hand and of the conservation laws on the other, which were
suggested by this apparent contradiction, have been definitely
disproved through direct experiments. This situation would seem
clearly to indicate the impossibility of a causal space-time description
of the light phenomena. On one hand, in attempting to trace the laws
of the time-spatial propagation of light according to the quantum
postulate, we are confined to statistical considerations. On the other
hand, the fulfilment of the claim of causality for the individual light
processes, characterised by the quantum of action, entails a
renunciation as regards the space-time description. Of course, there
can be no question of a quite independent application of the ideas of
space and time and of causality. The two views of the nature of light
are rather to be considered as different attempts at an interpretation
of experimental evidence in which the limitation of the classical
concepts is expressed in complementary ways.
The problem of the nature of the constituents of matter presents
us with an analogous situation. The individuality of the elementary
electrical corpuscles is forced upon us by general evidence.
Nevertheless, recent experience, above all the discovery of the
selective reflection of electrons from metal crystals, requires the use
of the wave theory superposition principle in accordance with the
original ideas of L. de Broglie. Just as in the case of light, we have
consequently in the question of the nature of matter, so far as we
adhere to classical concepts, to face an inevitable dilemma, which
has to be regarded as the very expression of experimental evidence.
In fact, here again we are not dealing with contradictory but with
complementary pictures of the phenomena, which only together offer
a natural generalisation of the classical mode of description. In the
discussion of these questions, it must be kept in mind that, according
to the view taken above, radiation in free space as well as isolated
material particles are abstractions, their properties on the quantum
theory being definable and observable only through their interaction
with other systems. Nevertheless, these abstractions are, as we
shall see, indispensable for a description of experience in connexion
with our ordinary space-time view.
The difficulties with which a causal space-time description is
confronted in the quantum theory, and which have been the subject
of repeated discussions, are now placed into the foreground by the
recent development of the symbolic methods. An important
contribution to the problem of a consistent application of these
methods has been made lately by Heisenberg (Zeitschr. f. Phys., 43,
172; 1927). In particular, he has stressed the peculiar reciprocal
uncertainty which affects all measurements of atomic quantities.
Before we enter upon his results it will be advantageous to show
how the complementary nature of the description appearing in this
uncertainty is unavoidable already in an analysis of the most
elementary concepts employed in interpreting experience.
2. quantum of action and
kinematics.
The fundamental contrast between the quantum of action and the
classical concepts is immediately apparent from the simple formulæ
which form the common foundation of the theory of light quanta and
of the wave theory of material particles. If Planck’s constant be
denoted by , as is well known,
where and are energy and momentum respectively, and the

corresponding period of vibration and wave-length. In these formulæ
the two notions of light and also of matter enter in sharp contrast.
While energy and momentum are associated with the concept of
particles, and hence may be characterised according to the classical
point of view by definite space-time co-ordinates, the period of
vibration and wave-length refer to a plane harmonic wave train of
unlimited extent in space and time. Only with the aid of the
superposition principle does it become possible to attain a connexion
with the ordinary mode of description. Indeed, a limitation of the
extent of the wave-fields in space and time can always be regarded
as resulting from the interference of a group of elementary harmonic
waves. As shown by de Broglie (Thèse, Paris, 1924), the
translational velocity of the individuals associated with the waves can
be represented by just the so-called group-velocity. Let us denote a
plane elementary wave by
where and are constants determining respectively the amplitude

and the phase. The quantity is the frequency, , , , the
wave numbers in the direction of the co-ordinate axes, which may be
regarded as vector components of the wave number in the
direction of propagation. While the wave or phase velocity is given
by , the group-velocity is defined by . Now according to the
relativity theory we have for a particle with the velocity :
where denotes the velocity of light. Hence by equation (1) the

phase velocity is and the group-velocity . The circumstance that
the former is in general greater than the velocity of light emphasises
the symbolic character of these considerations. At the same time, the
possibility of identifying the velocity of the particle with the group-
velocity indicates the field of application of space-time pictures in the
quantum theory. Here the complementary character of the
description appears, since the use of wave-groups is necessarily
accompanied by a lack of sharpness in the definition of period and
wave-length, and hence also in the definition of the corresponding
energy and momentum as given by relation (1).
Rigorously speaking, a limited wave-field can only be obtained by
the superposition of a manifold of elementary waves corresponding
to all values of and , , . But the order of magnitude of the
mean difference between these values for two elementary waves in
the group is given in the most favourable case by the condition
where , , , denote the extension of the wave-field in

time and in the directions of space corresponding to the co-ordinate
axes. These relations—well known from the theory of optical
instruments, especially from Rayleigh’s investigation of the resolving
power of spectral apparatus—express the condition that the wave-
trains extinguish each other by interference at the space-time
boundary of the wave-field. They may be regarded also as signifying
that the group as a whole has no phase in the same sense as the
elementary waves. From equation (1) we find thus:
as determining the highest possible accuracy in the definition of the

energy and momentum of the individuals associated with the wave-
field. In general, the conditions for attributing an energy and a
momentum value to a wave-field by means of formula (1) are much
less favourable. Even if the composition of the wave-group
corresponds in the beginning to the relations (2), it will in the course
of time be subject to such changes that it becomes less and less
suitable for representing an individual. It is this very circumstance
which gives rise to the paradoxical character of the problem of the
nature of light and of material particles. The limitation in the classical
concepts expressed through relation (2) is, besides, closely
connected with the limited validity of classical mechanics, which in
the wave theory of matter corresponds to the geometrical optics, in
which the propagation of waves is depicted through ‘rays.’ Only in
this limit can energy and momentum be unambiguously defined on
the basis of space-time pictures. For a general definition of these
concepts we are confined to the conservation laws, the rational
formulation of which has been a fundamental problem for the
symbolical methods to be mentioned below.
In the language of the relativity theory, the content of the relations
(2) may be summarised in the statement that according to the
quantum theory a general reciprocal relation exists between the
maximum sharpness of definition of the space-time and energy-
momentum vectors associated with the individuals. This
circumstance may be regarded as a simple symbolical expression
for the complementary nature of the space-time description and the
claims of causality. At the same time, however, the general character
of this relation makes it possible to a certain extent to reconcile the
conservation laws with the space-time co-ordination of observations,
the idea of a coincidence of well-defined events in a space-time point
being replaced by that of unsharply defined individuals within finite
space-time regions.
This circumstance permits us to avoid the well-known paradoxes
which are encountered in attempting to describe the scattering of
radiation by free electrical particles as well as the collision of two
such particles. According to the classical concepts, the description of
the scattering requires a finite extent of the radiation in space and
time, while in the change of the motion of the electron demanded by
the quantum postulate one seemingly is dealing with an
instantaneous effect taking place at a definite point in space. Just as
in the case of radiation, however, it is impossible to define
momentum and energy for an electron without considering a finite
space-time region. Furthermore, an application of the conservation
laws to the process implies that the accuracy of definition of the
energy momentum vector is the same for the radiation and the
electron. In consequence, according to relation (2), the associated
space-time regions can be given the same size for both individuals in
interaction.
A similar remark applies to the collision between two material
particles, although the significance of the quantum postulate for this
phenomenon was disregarded before the necessity of the wave
concept was realised. Here this postulate does indeed represent the
idea of the individuality of the particles which, transcending the
space-time description, meets the claim of causality. While the
physical content of the light quantum idea is wholly connected with
the conservation theorems for energy and momentum, in the case of
the electrical particles the electric charge has to be taken into
account in this connexion. It is scarcely necessary to mention that for
a more detailed description of the interaction between individuals we
cannot restrict ourselves to the facts expressed by formulæ (1) and
(2), but must resort to a procedure which allows us to take into
account the coupling of the individuals, characterising the interaction
in question, where just the importance of the electric charge
appears. As we shall see, such a procedure necessitates a further
departure from visualisation in the usual sense.
3. measurements in the quantum
theory.
In his investigations already mentioned on the consistency of the
quantum theoretical methods, Heisenberg has given the relation (2)
as an expression for the maximum precision with which the space-
time co-ordinates and momentum-energy components of a particle
can be measured simultaneously. His view was based on the
following consideration: On one hand, the co-ordinates of a particle
can be measured with any desired degree of accuracy by using, for
example, an optical instrument, provided radiation of sufficiently
short wave-length is used for illumination. According to the quantum
theory, however, the scattering of radiation from the object is always
connected with a finite change in momentum, which is the larger the
smaller the wave-length of the radiation used. The momentum of a
particle, on the other hand, can be determined with any desired
degree of accuracy by measuring, for example, the Doppler effect of
the scattered radiation, provided the wave-length of the radiation is
so large that the effect of recoil can be neglected, but then the
determination of the space co-ordinates of the particle becomes
correspondingly less accurate.
The essence of this consideration is the inevitability of the
quantum postulate in the estimation of the possibilities of
measurement. A closer investigation of the possibilities of definition
would still seem necessary in order to bring out the general
complementary character of the description. Indeed, a discontinuous
change of energy and momentum during observation could not
prevent us from ascribing accurate values to the space-time co-
ordinates, as well as to the momentum-energy components before
and after the process. The reciprocal uncertainty which always
affects the values of these quantities is, as will be clear from the
preceding analysis, essentially an outcome of the limited accuracy
with which changes in energy and momentum can be defined, when
the wave-fields used for the determination of the space-time co-
ordinates of the particle are sufficiently small.
In using an optical instrument for determinations of position, it is
necessary to remember that the formation of the image always
requires a convergent beam of light. Denoting by the wave-length
of the radiation used, and by the so-called numerical aperture, that
is, the sine of half the angle of convergence, the resolving power of a
microscope is given by the well-known expression . Even if the
object is illuminated by parallel light, so that the momentum of the

incident light quantum is known both as regards magnitude and
direction, the finite value of the aperture will prevent an exact
knowledge of the recoil accompanying the scattering. Also, even if
the momentum of the particle were accurately known before the
scattering process, our knowledge of the component of momentum
parallel to the focal plane after the observation would be affected by
an uncertainty amounting to . The product of the least
accuracies with which the positional co-ordinate and the component
of momentum in a definite direction can be ascertained is therefore
just given by formula (2). One might perhaps expect that in
estimating the accuracy of determining the position, not only the
convergence but also the length of the wave-train has to be taken
into account, because the particle could change its place during the
finite time of illumination. Due to the fact, however, that the exact
knowledge of the wave-length is immaterial for the above estimate, it
will be realised that for any value of the aperture the wave-train can
always be taken so short that a change of position of the particle
during the time of observation may be neglected in comparison to
the lack of sharpness inherent in the determination of position due to
the finite resolving power of the microscope.
In measuring momentum with the aid of the Doppler effect—with
due regard to the Compton effect—one will employ a parallel wave-
train. For the accuracy, however, with which the change in wave-
length of the scattered radiation can be measured the extent of the
wave-train in the direction of propagation is essential. If we assume
that the directions of the incident and scattered radiation are parallel
and opposite respectively to the direction of the position co-ordinate
and momentum component to be measured, then can be taken
as a measure of the accuracy in the determination of the velocity,
where denotes the length of the wave-train. For simplicity, we here
have regarded the velocity of light as large compared to the velocity
of the particle. If represents the mass of the particle, then the
uncertainty attached to the value of the momentum after observation
is . In this case the magnitude of the recoil, , is sufficiently
well defined in order not to give rise to an appreciable uncertainty in
the value of the momentum of the particle after observation. Indeed,
the general theory of the Compton effect allows us to compute the
momentum components in the direction of the radiation before and
after the recoil from the wave-lengths of the incident and scattered
radiation. Even if the positional co-ordinates of the particle were
accurately known in the beginning, our knowledge of the position
after observation nevertheless will be affected by an uncertainty.
Indeed, on account of the impossibility of attributing a definite instant
to the recoil, we know the mean velocity in the direction of
observation during the scattering process only with an accuracy
. The uncertainty in the position after observation hence is
. Here, too, the product of the inaccuracies in the

measurement of position and momentum is thus given by the
general formula (2).
Just as in the case of the determination of position, the time of
the process of observation for the determination of momentum may
be made as short as is desired if only the wave-length of the
radiation used is sufficiently small. The fact that the recoil then gets
larger does not, as we have seen, affect the accuracy of
measurement. It should further be mentioned, that in referring to the
velocity of a particle as we have here done repeatedly, the purpose
has only been to obtain a connexion with the ordinary space-time
description convenient in this case. As it appears already from the
considerations of de Broglie mentioned above, the concept of
velocity must always in the quantum theory be handled with caution.
It will also be seen that an unambiguous definition of this concept is
excluded by the quantum postulate. This is particularly to be
remembered when comparing the results of successive
observations. Indeed, the position of an individual at two given
moments can be measured with any desired degree of accuracy; but
if, from such measurements, we would calculate the velocity of the
individual in the ordinary way, it must be clearly realised that we are
dealing with an abstraction, from which no unambiguous information
concerning the previous or future behaviour of the individual can be
obtained.
According to the above considerations regarding the possibilities
of definition of the properties of individuals, it will obviously make no
difference in the discussion of the accuracy of measurements of
position and momentum of a particle if collisions with other material
particles are considered instead of scattering of radiation. In both
cases we see that the uncertainty in question equally affects the
description of the agency of measurement and of the object. In fact,
this uncertainty cannot be avoided in a description of the behaviour
of individuals with respect to a co-ordinate system fixed in the
ordinary way by means of solid bodies and unperturbable clocks.
The experimental devices—opening and closing of apertures, etc.—
are seen to permit only conclusions regarding the space-time
extension of the associated wave-fields.
In tracing observations back to our sensations, once more regard
has to be taken to the quantum postulate in connexion with the
perception of the agency of observation, be it through its direct
action upon the eye or by means of suitable auxiliaries such as
photographic plates, Wilson clouds, etc. It is easily seen, however,
that the resulting additional statistical element will not influence the
uncertainty in the description of the object. It might even be
conjectured that the arbitrariness in what is regarded as object and
what as agency of observation would open up a possibility of
avoiding this uncertainty altogether. In connexion with the
measurement of the position of a particle, one might, for example,
ask whether the momentum transmitted by the scattering could not
be determined by means of the conservation theorem from a
measurement of the change of momentum of the microscope—
including light source and photographic plate—during the process of
observation. A closer investigation shows, however, that such a
measurement is impossible, if at the same time one wants to know
the position of the microscope with sufficient accuracy. In fact, it
follows from the experiences which have found expression in the
wave theory of matter, that the position of the centre of gravity of a
body and its total momentum can only be defined within the limits of
reciprocal accuracy given by relation (2).
Strictly speaking, the idea of observation belongs to the causal
space-time way of description. Due to the general character of
relation (2), however, this idea can be consistently utilised also in the
quantum theory, if only the uncertainty expressed through this
relation is taken into account. As remarked by Heisenberg, one may
even obtain an instructive illustration to the quantum theoretical
description of atomic (microscopic) phenomena by comparing this
uncertainty with the uncertainty, due to imperfect measurements,
inherently contained in any observation as considered in the ordinary
description of natural phenomena. He remarks on that occasion that
even in the case of macroscopic phenomena we may say, in a
certain sense, that they are created by repeated observations. It
must not be forgotten, however, that in the classical theories any
succeeding observation permits a prediction of future events with
ever-increasing accuracy, because it improves our knowledge of the
initial state of the system. According to the quantum theory, just the
impossibility of neglecting the interaction with the agency of
measurement means that every observation introduces a new
uncontrollable element. Indeed, it follows from the above
considerations that the measurement of the positional co-ordinates
of a particle is accompanied not only by a finite change in the
dynamical variables, but also the fixation of its position means a
complete rupture in the causal description of its dynamical
behaviour, while the determination of its momentum always implies a
gap in the knowledge of its spatial propagation. Just this situation
brings out most strikingly the complementary character of the
description of atomic phenomena which appears as an inevitable
consequence of the contrast between the quantum postulate and the
distinction between object and agency of measurement, inherent in
our very idea of observation.

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Chapter 1: Principles of Biochemistry

1. The birth of modern biochemistry can be traced to the

2. The study of biochemistry attempts to explain

a. greatest difference in relative affinities of the two atoms for electrons.

7. Amino acids are the building blocks for which biomolecule(s)?

8. The __________ differentiates amino acids from one another.

9. A nucleotide consists of which of the following?

a. The rate of polymer increases.

a. increased concentration of argininosuccinate

17. The correct definition of a pathway intermediate is a molecule that

22. When a ligand binds to a receptor, it causes the receptor to

25. A hydrogen bond can best be described as a

26. Why can a guanine not be paired with adenine?

30. A genome is a set of

33. A single nucleotide base substitution in a wild-type DNA is an example of

39. The appearance of new gene speciation is an example of

41. Which of the following is NOT a common functional group?

44. mRNA is used for what process in the cell?

47. Mutations to proteins typically occur starting

50. In structures of tRNA, base pairs form between

52. The process of fermentation uses sugar to produce which molecules?

53. An example of experimental biochemistry is trying an experiment and

55. Pyruvate decarboxylase converts pyruvate into

56. Which of the following is an example of a metabolic pathway?

57. Which of the following is an example of an ecosystem?

62. The figure below shows an example of which functional group?

63. ATP is an abbreviation for which energy currency molecule?

64. Which of the following is the correct formula for glucose?

66. Triacylglycerols are neutral molecules made of

71. What does chromatin in eukaryotic nucleus contain?

72. The function of chloroplasts in plant cells is to convert

73. What process replicates DNA to make more DNA?

74. What process converts DNA to RNA?

75. What process describes using mRNA templates to produce proteins?

DIF: Easy REF: Introduction

DIF: Easy REF: 1.1

3. Which six elements make up 97% of the weight of most organisms?

DIF: Easy REF: 1.2

4. Identify the following functional groups.

DIF: Easy REF: 1.2

5. Identify the following biomolecules.

DIF: Easy REF: 1.2

6. What functions can polysaccharides perform?

DIF: Difficult REF: 1.2

7. What is an advantage of using polypeptides for storage and information transmission?

DIF: Difficult REF: 1.2

8. Explain how a linear pathway is different from a forked pathway.

DIF: Difficult REF: 1.2

DIF: Difficult REF: 1.2

10. Name four organelles found in a eukaryotic cell.

DIF: Difficult REF: 1.2

12. Name the three components of a nucleotide.

DIF: Easy REF: 1.3

14. Define the central dogma of molecular biology.