Biology for the Health Sciences Mechanisms of Disease - 1st

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 Contents

Preface ��������������������������������������������������������� vii SECTION 3: INFECTIOUS DISEASE

Student and Instructor e-Resources �����������ix Chapter 19. Infectious Diseases........ 359
Chapter 20. Respiratory Tract
SECTION 1: BASIC BIOLOGY AND Infections........................................ 387
BIOTECHNOLOGY Chapter 21. Diarrheal Diseases ......... 417
Chapter 1. Chemical Basis of Life ......... 3 Chapter 22. HIV/AIDS and Other
Chapter 2. Cell Structure and Sexually Transmitted Infections .... 441
Function............................................ 27 Chapter 23. Malaria and Other Vector-
Chapter 3. Protein Structure and transmitted Diseases ..................... 457
Function............................................ 49 Chapter 24: Parasitic Helminths........ 477
Chapter 4. Molecular Biology ............. 71 Index ��������������������������������������������������������� 493
Chapter 5. Genetics and Evolution ..... 89
Chapter 6. Genomics and the ’Omic
Revolution ...................................... 113
Chapter 7. Genetic Engineering ........ 133
Chapter 8. Cell Differentiation and
Stem Cells ...................................... 155
Chapter 9. The Immune System ........ 173

SECTION 2: NON-INFECTIOUS DISEASE

Chapter 10. Disease Overview and
Molecular Biomarkers.................... 207
Chapter 11. Genetic Diseases ........... 221
Chapter 12. Nutritional Disorders...... 239
Chapter 13. Diabetes ......................... 255
Chapter 14. Cardiovascular Disease. 267
Chapter 15. Chronic Kidney Disease
and Cardiorenal Syndrome............ 287
Chapter 16. Cancer ............................ 297
Chapter 17. Immunological
Disorders ........................................ 321
Chapter 18. Drug Misuse – Associated
Disorders ........................................ 339

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 Preface

Biology plays a central role in our understanding of health and disease. It

is therefore critical that health professionals have a solid foundation in
biology and especially the biological basis of disease. This is also true for
public health professionals in that a solid understanding of biological
principles will become increasingly important in disease management,
prevention, and control. Furthermore, biotechnology is being rapidly
applied to medicine as well as public health. So going forward, health
professionals will need a strong grounding in the fundamentals of the
biological basis of disease, as well as an understanding of biotechnol-
ogies such as stem cells, vaccines and other immunotherapies, genetic
engineering, and genomics.
The origins of this book are from a Biological Basis of Disease course
that was initially taught at the graduate level and then transformed into
an advanced undergraduate course within a school of public health.
However, the content of the book is also of interest to pre-medical
students as well as students pursuing other health-related professions,
such as pharmacy, veterinary medicine, or nursing. The book has three
major sections: Basic Biology and Biotechnology, Non-infectious Disease,
and Infectious Disease. The combining of these three topics into a single
book is unique and allows for a merger of the basic sciences with the
more applied biomedical sciences.
Section 1 consists of nine chapters that cover basic cell and molecular
biology, genetics, and immunology and vaccines, as well as recombinant
DNA technology and gene therapy, genomics, and stem cells. In advanced
courses the students may already be suffciently knowledgeable about
the basic aspects of cell structure and function and genetics, and these
chapters could serve as a review or reference within the same book. In
the later chapters there are numerous signposts referring the reader back
to the specifc pages in the earlier chapters where the basic concepts are
explained. In this way, readers who are unsure about the fundamental
concepts can quickly review basic biology as they pursue the biological
basis of disease. Conversely, there may be courses in which students do
not have a strong background in basic biology and the beginning of the
course needs to cover these topics. The book could even be used as part
of two sequential courses in which the frst course focused on basic
biology and biotechnology followed by a second course focusing on the
biological basis of disease.
The major non-infectious diseases of humans are covered in the nine
chapters of Section 2. The frst chapter of this section gives an overview
of disease and discusses aging, senescence, and degenerative diseases.

vii

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 viii Preface

This is followed by a chapter on monogenic genetic diseases. Sickle cell

disease, cystic fbrosis, and Huntington’s disease are described in detail
as examples of how genetic defects lead to pathogenesis at the molecular
and cellular levels. The next chapter deals with the role of nutrition in
disease with a focus on overnutrition and obesity since obesity is a factor
in many diseases. Thereafter, there are chapters on diabetes,
cardiovascular disease, chronic kidney disease, cancer, immunological
disorders, and addictive disorders. Each of the disease-specifc chapters
gives an overview of the disease and describes disease etiology and
pathogenesis. The molecular and cellular aspects of etiology and
pathogenesis are also discussed for each of the diseases. Clinical
manifestations, as well as diagnosis, treatment, and disease management,
are also described for these various diseases. The clinical aspects of the
disease are described at a level suitable for non-physicians and pre-
professional students.
The fnal six chapters of the book make up Section 3 and are devoted
to infectious diseases. Infectious diseases tend to have superfcial
coverage in most biology of disease books. However, the COVID-19
pandemic has certainly awakened the world to the importance of
infectious diseases in both resource-rich regions and resource-poor
regions. Section 3 starts with an overview of pathogens and infectious
disease. Included in the frst chapter of this section is a discussion of drug
resistance and the mechanisms by which pathogens become resistant to
drugs. The subsequent chapters focus on a particular type of infectious
disease, which includes chapters on respiratory tract infections,
gastrointestinal infections and diarrheal diseases, HIV and other sexually
transmitted infections, vector-borne disease with a focus on malaria, and
parasitic worms. In each of these chapters the biologies of the various
pathogens are extensively covered, as well as the biology of the host–
pathogen interaction, including immunity. As in the chapters on
non-infectious diseases, the pathogenesis associated with infectious
diseases is described, as well as clinical manifestations, diagnosis, and
treatment. Examples of interactions between the pathogen and human
host at the molecular and cellular levels are also described. In regard to
disease etiology, modes of pathogen transmission are thoroughly
discussed including methods to prevent transmission.
In some respects, this book is three books in one. And that was some
of the impetus for writing the book since there was not a single book that
comprehensively covered both non-infectious and infectious diseases as
well as the applications of modern biotechnology. This book could serve
as the primary textbook in many public health biology courses or courses
in pre-medical or pre-professional curricula. In addition, many pre-
medical or pre-professional students may fnd this book helpful in
preparing for qualifying exams and interviews. At the end of each chapter
are a summary of the key points, defnitions of the key terms, and review
questions that can assist students in studying for exams.

Mark F. Wiser

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 Student and Instructor
Resources

STUDENT RESOURCES
In addition to the review questions found at the end of each chapter in
the book, Biology for the Health Sciences also comes with a number of
interactive questions, hosted online, which the student can use to test
their understanding of the chapter material.

This interactive resource makes use of several different question types:

multiple choice, true or false, fll-in-the-blanks, and matching-type ques-
tions. Where necessary, feedback is also provided to help the student to
understand why a particular answer is correct.

The students can access these resources by visiting

https://routledge.com/cw/wiser

INSTRUCTOR RESOURCES
For instructors, all the fgures found in the textbook have been compiled
into Figure Slides, which the instructors may wish to use in their lectures.
These are available in two convenient formats: PowerPoint and PDF.
To access these, please visit https://routledge.com/cw/wiser

ix

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 SECTION
Basic Biology and
Biotechnology 1
Chapter 1. Chemical Basis of Life
Chapter 2. Cell Structure and Function
Chapter 3. Protein Structure and Function
Chapter 4. Molecular Biology
Chapter 5. Genetics and Evolution
Chapter 6. Genomics and the ’Omic Revolution
Chapter 7. Genetic Engineering
Chapter 8. Cell Differentiation and Stem Cells
Chapter 9. The Immune System

As we better understand disease at the molecular, cellular, genetic, and

immunological levels, it is important to have a strong foundation in basic
molecular and cellular biology, genetics and evolution, and immunology.
Furthermore, recombinant DNA technology, genomics, stem cells, and
other technologies are increasingly being applied to the diagnosis,
treatment, and management of disease. The frst four chapters of the frst
section provide an overview of cellular and molecular biology. Genetics
also plays a role in all disease and, accordingly, a chapter on genetics
that covers the basic principles of inheritance follows the chapters on
basic cellular and molecular biology. Also included in this chapter is a
brief discussion of evolution and our current understanding of human
evolution. These opening chapters can serve as a review or a reference
for students who have prior knowledge of these topics. Students who do
not have a suffcient background can use these chapters to acquire basic
knowledge that will be needed later in the book.
Understanding basic molecular biology and genetics is also important
to understanding the biotechnologies that are being applied at a rapid
rate to medicine and related felds. Following the chapter on genetics is

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 a chapter on genomics, which describes genomics and its derivatives,
such as transcriptomics, proteomics, and metabolomics. Also included
in this chapter are discussions on how genomics can be used to improve
human health through the development of better diagnostics and
therapies and the prospect of personalized medicine. Genome-wide
association studies are also described. The chapter following genomics
describes recombinant DNA technology including gene therapy and how
recombinant DNA technology leads to the more effcient production
of biopharmaceuticals and its applications to agriculture. Chapter 8
discusses the organization of cells into tissues and organs and gives an
overview of the cell differentiation process. Also included in this chapter
is a discussion of stem cells and cell-based therapies. The fnal chapter in
this section provides an overview of the immune system and immunology.
Discussions of serology and vaccines are also included.

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 Chemical Basis of Life 1

All living organisms, at the most fundamental level, are composed of basic ATOMIC STRUCTURE
chemical elements. Chemical elements form the molecules that make
up cellular structures and participate in chemical reactions necessary to
sustain life. These molecules and chemical reactions occur in an aqueous INTERACTIONS BETWEEN
environment and the unique properties of water play an essential role in ATOMS
the existence and sustenance of life. Biologically important molecules
consist of a carbon backbone and fall into four chemically distinct classes LIFE-AFFIRMING
of molecules: carbohydrates, lipids, proteins, and nucleic acids.
PROPERTIES OF WATER

ATOMIC STRUCTURE IONIZATION AND PH

Atoms are composed of subatomic particles called electrons, protons, and
neutrons. Atoms are typically depicted as a central nucleus, composed of THE CARBON ATOM
protons and neutrons, being orbited by electrons (FIGURE 1.1). Protons
(p+) are positively charged and the attraction between negative and CLASSES OF BIOLOGICAL
positive charges maintains the negatively charged electrons (e-) in this
MOLECULES
orbit. Neutrons have no charge since they are composed of a proton and
an electron. Atoms containing the same number of protons and electrons
have no net charge. A gain or loss of electrons orbiting the nucleus results
in the ionization of the atom that imparts either a negative or a positive
charge on that atom. nucleus

Atoms defne the basic chemical elements that make up – –

all of nature
Atoms are defned as substances that cannot be broken down into
another substance by ordinary means. The atomic nuclei provide stability
to atoms and ordinary forces, such as heat, electricity, and light, do not + ++
– + –
adversely affect or change the atomic structure. There are 92 elements +
that occur naturally and each element corresponds to a distinct type of –
atom. These basic elements are defned by the number of protons making –
up the atom. For example, hydrogen, the simplest atom, is composed of
one proton and its corresponding electron. The next element, which is
helium (He), is composed of two protons and the corresponding electrons,
as well as two neutrons. All 92 naturally occurring elements are defned + proton neutron – electron
by the number of protons ranging from 1 to 92 and the number of protons
is referred to as the atomic number.
FIGURE 1.1 Model of atomic structure. Atoms
Protons and neutrons have mass and atoms can also be defned by
are typically depicted as a core nucleus composed
the total number of neutrons and protons. The mass of the electron is of neutrons and protons being circled by electrons.
negligible compared with the masses of the protons and neutrons and The number of protons defnes an element.
is therefore not included in the atomic mass. So, in addition to defning

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 4 CHAPTER 1 Chemical Basis of Life

elements by the atomic number corresponding to the number of protons,

elements can also be defned by an atomic mass, which is the combined
number of protons and neutrons. In the case of normal hydrogen, both
the atomic number and the atomic mass are equal to one since there
are no neutrons, whereas for helium the atomic number is two and the
atomic mass is four since the helium nucleus also has two neutrons.
The primary elements making up greater than 95% of the mass of living
matter are oxygen, carbon, hydrogen, and nitrogen (TABLE 1.1).

Table 1.1 Common Elements Making Up Living Things

Element Symbol Atomic number Common Percentage in Percentage in Percentage in
atomic mass human body Earth’s crust Universe
Hydrogen H 1 1 9.5 0.14 91
Carbon C 6 12 18.5 0.03 0.02
Nitrogen N 7 14 3.3 Trace 0.04
Oxygen O 8 16 65 47 0.06
Sodium Na 11 23 0.2 2.8 Trace
Phosphorous P 15 31 1.0 0.07 Trace
Chlorine Cl 17 35 0.2 0.01 Trace
Potassium K 19 39 0.4 2.6 Trace
Calcium Ca 20 40 1.5 3.6 Trace
Iron Fe 26 56 Trace 5.0 Trace
Atomic numbers and masses of selected elements and their approximate percentage by weight in the human body, the Earth’s crust, and
the Universe.

Variations in the number of neutrons result in different

forms of an element
Elements can exist in different forms called isotopes that are defned by
a different number of neutrons. All isotopes of a particular element have
the same atomic number but have different atomic masses. For example,
a form of hydrogen with one proton and one neutron in the nucleus is
called deuterium and has an atomic mass of two. The atomic number,
however, is still one. Similarly, tritium is another form of hydrogen
with an atomic mass of three due to one proton and two neutrons. The
various isotopes of an element are chemically the same in regard to
their interactions with other elements. In nature elements are generally
represented by a predominant isotope and a mixture of the other possible
isotopes.
Some isotopes are unstable and tend to break apart or decay. These are
referred to as radioactive isotopes or radioisotopes. Energy is released
when these unstable isotopes break apart and this energy is called
radioactivity. The released energy can be in the form of subatomic
particles or electromagnetic radiation similar to X-rays. For example,
carbon-14 (14C) is an isotope of carbon containing six protons and eight
neutrons. Carbon-14 is not as stable as the more common carbon-12 (12C)
consisting of six protons and six neutrons. To increase stability, one of the
neutrons of carbon-14 converts into a proton by ejecting an electron from
the nucleus. This ejected electron is the radioactivity that is detectable
and can be measured. Since one of the neutrons converts to a proton, the
atomic number is now seven and the atom is now nitrogen-14 (14N), a
common and stable isotope of nitrogen. Radioactivity and radioisotopes
are widely used in biomedical research and medicine (TABLE 1.2) due to

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 ATOMIC STRUCTURE 5

the well-defned properties of radioactive decay and the ease at which

radioactivity can be detected and measured. Various imaging techniques,
such as positron emission tomography (PET) scans (BOX 1.1), can be used
to evaluate the integrity and function of internal organs.

Table 1.2 Examples of Radioactivity in Diagnosis and Treatment of Disease

• Imaging techniques such as X-rays and magnetic resonance imaging (MRI) to evaluate internal organs
• Use of radioactive substances to evaluate physiology such as using radioactive iodine to evaluate thyroid function
• Positron emission tomography (PET) scans to evaluate organ function
• Radiation therapy to treat cancer

Positron Emission Tomography

BOX 1.1
Positron emission tomography, commonly known technology determines the distribution of the
as PET, is used to observe metabolic processes radiotracers in specifc organs or tissues. The most
in the body as a means to diagnose disease. often used radiotracer is a radioactive analog of
Radionuclides are incorporated either into glucose. Glucose is the central metabolite for
compounds normally used by the body, such as cellular energy production (Chapter 3) and tissues
glucose, or into molecules that bind to receptors or or organs using excessive glucose will show up as
other sites of drug action. Such labeled compounds ‘hot spots’ on the PET scan. PET scans are widely
are often called radiotracers. Following injection, used in the diagnosis of cancer and neurological
ingestion, or inhalation of the radiotracer, PET disorders.

Electrons orbit the nucleus at defned distances,

forming shells
The positive charge of the nucleus attracts the orbiting electrons and thus
the electrons are drawn close to the nucleus. However, the negatively
charged electrons repel each other and therefore only a limited number
of electrons can occupy the space close to the nucleus. In the case of
large atoms with many electrons, the electrons need to be distributed at
increasing distances from the nucleus. These increasing distances are
called electron shells and only a fxed number of electrons can occupy
a particular shell. For example, only two electrons can occupy the space
closest to the nucleus and the next shell can accommodate up to eight
electrons. The electrons of an atom fll in the shells closest to the nucleus
frst and then expand outward. For example, the carbon atom with six
electrons has two electrons in the frst shell and four electrons in the
second shell, whereas oxygen with eight electrons has two electrons in
the frst shell and six in the second shell (FIGURE 1.2).

carbon atom oxygen atom

FIGURE 1.2 Electron shells in carbon and
– – oxygen. The electrons in an atom distribute in
6 protons + 8 protons + ‘shells’ defned by the distance from the nucleus. The
– 6 neutrons – 8 neutrons frst shell can only hold two electrons and the next
shell can hold up to eight electrons. Carbon has four
–
– + ++ – – + ++ – electrons in its outer shell and oxygen has six.
+ +
+ +
– –

– – –

+ proton neutron – electron

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 6 CHAPTER 1 Chemical Basis of Life

The electrons orbiting the nucleus can be gained, lost, or

shared with other atoms
The electrons in the outermost shells tend to be rather dynamic and
especially in atoms with a partially flled outermost electron shell. Atoms
with partially flled outer shells have the propensity to gain or lose electrons
in order to fll or empty the outermost shell. Having a completely full
outermost electron shell increases the stability of the atom. For example,
atoms in which the outer shell is completely full in its normal state do
not react with other atoms and are called inert. Conversely, atoms with
partially full outer shells are more likely to interact with other atoms by
forming chemical bonds.

INTERACTIONS BETWEEN ATOMS

Atoms can combine with other atoms to form molecules. For example,
water – defned chemically as H2O – is composed of two hydrogen atoms
(H) and one oxygen atom (O). The interaction between atoms to form
molecules involves gaining, losing, or sharing electrons in the outermost
shell. Losing, gaining, or sharing electrons between two or more
atoms results in the formation of chemical bonds. Chemical bonds
are attractive forces that hold the atoms close together. Each element
has a characteristic set of possible chemical bonds that can be formed,
which depends largely on the confguration of its electrons in the outer
shell. These chemical bonds can be either weak forces or strong forces
(TABLE 1.3).

Table 1.3 Types of Bonds between Atoms or Molecules

Bond type Nature of the bond/interaction
Strong forces Covalent A sharing of electrons between atoms
Weak forces Hydrogen Interactions between a hydrogen atom involved in a polar covalent bond and another atom involved in a polar covalent bond
Ionic Interactions between positive and negatively charged atoms
Hydrophobic Due to the exclusion of hydrophobic molecules by water

Atoms share electrons in a covalent bond

An atom with a partially full outer electron shell can become more stable
by sharing electrons with another atom and thereby form a covalent
bond. For example, hydrogen has a single electron in a shell designed for
two electrons. By combining with another hydrogen atom, the two atoms
can share the electrons (FIGURE 1.3) to form molecular hydrogen (H2). In
this case, each electron contributed by each hydrogen atom orbits both
nuclei and, thus, each atom now appears to have a full electron shell and
the hydrogen molecule is more stable than the two individual hydrogen
atoms. Therefore, the natural state of hydrogen is predominantly two
hydrogen atoms bound by a covalent bond and free hydrogen atoms are
relatively rare. The covalent bond is depicted as a single line between the
atoms (H—H).
Two oxygen atoms also interact to form molecular oxygen (O2) in
a similar manner (see Figure 1.3). However, in this case each oxygen
atom has six electrons in its outer shell and needs two electrons to
complete the shell. Therefore, two electrons are shared to fll the shell
and this results in a double covalent bond that is denoted by a double
line between the atoms (O=O). And if three electrons are shared by two

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 INTERACTIONS BETWEEN ATOMS 7

single covalent bond formation

–
–

+ + + + +

–
–

hydrogen (H) hydrogen (H) molecular hydrogen (H2)

double covalent bond formation

– –
– – – –
– – –
–
– –
– – + ++ – – + – – + ++ – – + ++ + ++
+ + + +
+ + + – – +
– –
– –
– – – –
– –

oxygen (O) oxygen (O) molecular oxygen (O2)

(double bond is formed)

FIGURE 1.3 Covalent bonds. In molecular hydrogen one electron from each atom is shared to form a single covalent bond. The two electrons are shared
equally between the two nuclei and therefore the frst electron shell is flled for both nuclei. In molecular oxygen two electrons from each atom are shared to form
a double covalent bond. By sharing two electrons there are now eight apparent electrons in the outermost electron shell instead of six, and thus the shell is flled.

atoms a triple bond is formed as in the case of nitrogen gas (N≡N) which
has fve electrons in the outer shell.
In the cases of molecular hydrogen (H2), molecular nitrogen (N2), and
molecular oxygen (O2), both nuclei are identical and the electrons
spend an equal amount of time with each nucleus. This equal sharing of ˜+ +
electrons results in symmetrical molecules that are electrically neutral. – –
However, not all covalent bonds exhibit such equality in electron sharing – –
–
between atoms. In covalent bonds between different atoms, the nucleus
with the most protons has a greater positive charge and therefore attracts + ++ ˜–
+
the electrons more strongly. In other words, the electrons spend more +
time over the more positively charged nucleus. This results in a molecule –
that is asymmetric in terms of the distribution of electrons and the – –
– –
molecule exhibits a polarity. For example, water is formed between an
oxygen atom and two hydrogen atoms (FIGURE 1.4). The electrons spend ˜+ +
more time over the oxygen atom, resulting in a slight negative pole over
the oxygen atom and slight positive poles over the hydrogen atoms.

Chemical reactions involve the breaking of

FIGURE 1.4 Polarity of water. Water is formed
covalent bonds via covalent bonds between two hydrogen (H) atoms
Covalent bonds are signifcantly stronger than the weak forces between and one oxygen atom (O). In each of the two bonds
electrons are shared between hydrogen and oxygen.
atoms and molecules. However, not all covalent bonds are of equal
Since the oxygen nucleus has more protons and a
strength since some are more stable than others. For example, the bond greater positive charge than the hydrogen nucleus
forming molecular hydrogen (H2) is extremely stable and diffcult to break. the electrons spend more time orbiting the oxygen
Other covalent bonds are not as strong and can more easily be broken. nucleus. This unequal sharing of the electrons
results in a slight negative charge (designated by δ−)
Energy is released during chemical reactions in which weaker bonds are near the oxygen atom and slight positive charges
broken and more stable bonds are formed. For example, fuels such as (designated by δ+) near the hydrogen atoms.

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 8 CHAPTER 1 Chemical Basis of Life

gasoline and molecular oxygen (O2) are composed of relatively unstable

O
bonds. Burning fuels in the presence of oxygen is a chemical reaction that
H H
˜– results in the formation of carbon dioxide (CO2) and water (H2O) – both of
˜+
˜– which contain highly stable bonds. This results in a release of energy in
O ˜+ O H
the form of heat. A similar process occurs in living organisms during the
H H ˜– H H
+ metabolism of carbohydrates to generate energy (Chapter 3). In this case
˜+ ˜– ˜ O
H the energy is captured and used to maintain the living organism.
H
O ˜ + ˜– O
H H
H Hydrogen bonds play important roles in biology
Another type of chemical bond between atoms and molecules is the
hydrogen bond. This type of bond is weaker than a covalent bond, but
FIGURE 1.5 Hydrogen bonds between water nevertheless, it plays an important role in the structure and properties of
molecules. The partial positive charge (δ+) of
hydrogen (H) interacts with the partial negative
water as well as being important in the structure and function of many
charge (δ−) of oxygen (O) as depicted by the dashed biological molecules. The basis of the hydrogen bond is the slight polarity
lines. This placement of hydrogen between two of the covalent bond formed between a hydrogen atom and atoms such
atoms with partial negative charges is called a as oxygen and nitrogen. The hydrogen atom involved in such a polar
hydrogen bond. covalent bond has a slight positive charge and is attracted to another atom
with a slight negative charge. For example, oxygen has a slight negative
charge when it forms a covalent bond with hydrogen. Therefore, water
molecules interact with each other via hydrogen bonds (FIGURE 1.5).

Ionic bonds are formed between atoms of

opposite charges
Atoms can also gain or lose electrons to stabilize the outermost electron
shell. The loss of an electron results in a positively charged atom, whereas
the gain of an electron results in a negatively charged atom. Charged
atoms or molecules are called ions. Positively charged ions are called
cations and negatively charged ions are called anions. Atoms that have
an almost empty outer shell tend to lose those electrons and become
cations. Conversely, atoms with an almost full outer shell tend to gain
electrons and become anions. Therefore, atoms with a nearly empty
outer electron shell tend to donate electron(s) to atoms with a nearly
full outer electron shell. For example, the formation of sodium chloride
(NaCl) is due to sodium donating an electron to chlorine (FIGURE 1.6).
This donor/acceptor relationship results in sodium becoming positively
charged (Na+) and chlorine becoming a negatively charged anion called
chloride (Cl−). These two oppositely charged ions now exhibit a mutual
attraction and this electrical attraction is an ionic bond.
+ –
transfer of
– electron – – – –
– – – – – – – –
– – – – – – – –

– – – – – – – – – – –
+11 +17 +11 +17
– – – – – – – – – – – –

– – – – – – – –
– – – –
Na Cl Na+ Cl–
sodium atom chlorine atom sodium ion chlorine ion
sodium chloride (NaCl)

FIGURE 1.6 Ionic bond formation in sodium chloride. The sodium atom (Na) donates its lone electron in the outermost shell to
chlorine (Cl) which accepts the electron to fll its outermost shell. This results in sodium having 10 electrons and a net positive charge
of +1 and chlorine having 18 electrons and a net negative charge of −1. The resulting sodium cation (Na+) and chloride anion (Cl−) are
attracted to each other due to their opposite charges to form an ionic bond.

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 INTERACTIONS BETWEEN ATOMS 9

Electrostatic interactions between oppositely charged ions can result in

the formation of ionic crystals which are generally called salts. The atoms
of ionic crystals are held together in an orderly array of alternating anions
and cations. For example, sodium chloride, common dietary salt, forms a
cubic array (FIGURE 1.7). In the crystal form these ionic interactions are
quite strong and diffcult to break. However, in an aqueous environment
ionic bonds are easily broken. The polar water molecules interact with Na+ Cl–
the salt ions to break the ionic bonds and dissolve the salt in water.

Many important biological processes involve the gain and FIGURE 1.7 Crystal structure of sodium
chloride. Anions and cations can interact to form
loss of electrons ionic crystals in which the anions and cations
alternate to form a lattice.
Oxidation is the loss of electrons and reduction is the gain of electrons.
These oxidation and reduction reactions occur together in pairs with one
substance being oxidized by donating electrons to another substance. The
substance receiving the electrons is reduced. In the example of sodium
chloride, sodium is oxidized and chlorine is reduced to chloride. These
combined reactions are often called redox reactions as a combination
of reduction and oxidation. Many important biological processes involve
redox reactions. For example, the aerobic metabolism of glucose to
produce energy is a series of redox reactions in which glucose is ulti-
mately oxidized to carbon dioxide (CO2) and oxygen is reduced to water
(Chapter 3).
Substances with the ability to cause other substances to lose electrons
are called oxidizing agents. Oxygen is the quintessential oxidizing agent,
hence the term oxidation is used to describe the loss of electrons in
general. In contrast, reducing agents are substances that cause other
substances to gain electrons.

Inappropriate oxidation causes molecular damage

Although redox reactions are a normal part of cellular metabolism,
oxidation potentially presents problems for organisms. For example, a
possible by-product of aerobic metabolism is the generation of reactive FURTHER
REDUCTION REDUCTION
oxygen species (ROS). The aberrant generation of ROS represents a O2 O– H2O
situation in which the reduction of oxygen was not immediately coupled (detoxifcation)
molecular ROS water
with an oxidation reaction. Due to their excess electrons, ROS are highly oxygen
unstable and readily react with a lot of other molecules. This high reactivity ABERRANT
of ROS causes cellular damage by inappropriately oxidizing important REACTIONS
cellular molecules (FIGURE 1.8). Just as rust is viewed as an undesirable
oxidation of iron, inappropriate oxidation of biological molecules causes oxidative
cellular damage and disease. damage

Oxidative damage is involved in many diseases. Thus, cells have

mechanisms to remove and detoxify ROS and minimize the undesirable FIGURE 1.8 Reactive oxygen species. Reactive
consequences of redox reactions. In addition, some chemicals called oxygen species (ROS) are formed by the reduction of
antioxidants function as strong reducing agents and can remove ROS by molecular oxygen. ROS can cause oxidative damage
reducing them to less reactive compounds. For this reason, antioxidants by oxidizing other molecules and destroying their
normal function. ROS can be further reduced to
are generally considered to be healthy supplements to our diet (BOX 1.2).
non-toxic forms such as water to prevent aberrant
It should also be noted, though, that ROS are also exploited by the oxidation.
immune system to kill pathogens (Chapter 9).

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