Parasitology

Parasitology
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Parasitology Highly detailed textbook on parasites and parasite relationships The fully revised edition of Parasitology: An Integrated Approach holds true to its engaging and easy-to-read approach. It comprehensively covers the complex and dynamic interaction between the parasite and its host ranging from invertebrates to vertebrates. Following an integrated approach, the authors explain how the study of parasites requires an understanding of biological concepts such as growth and reproduction, molecular biology, biochemistry, immunology, and pathology. In this second edition, they further address parasites and parasite relationships in the grand scheme of global changes and their impact. This textbook also reviews the often-neglected positive aspects of parasite infections and how humans have used parasites for their own advantage. Parasitology: An Integrated Approach, 2nd edition includes supplementary learning resources such as self-assessment quizzes, practical exercises, and an extensive collection of photographs. Now includes parasite life cycles in colour Strong focus on parasite interactions with other pathogens such as bacteria and viruses Discusses major advancements in the field of parasite diagnostics Additional image material and learning resources (quizzes, practical exercises) provided online A valuable and comprehensive learning resource for undergraduate students in the biological, biomedical and veterinary sciences and in medicine. It is also of interest to postgraduates and professionals with an interest including but not limited to parasitology, animal welfare, ecology, and medical microbiology.

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Alan Gunn. Parasitology

Table of Contents

List of Tables

List of Illustrations

Guide

Pages

Parasitology. An Integrated Approach

Preface

About the Companion Website

1 Animal Associations and the Importance of Parasites. CONTENTS

1.1 Introduction

1.2 Animal Associations

1.2.1 Symbiosis

1.2.1.1 Symbionts

1.2.1.2 The Importance of Symbionts to Blood‐feeding Organisms

1.2.2 Commensalism

1.2.3 Phoresis

1.2.4 Mutualism

1.2.5 Parasitism

From Welcome Guest to Villain: The Derivation of the Term ‘Parasite’

1.2.5.1 Intra‐specific Parasites

1.2.6 Parasitoids

Parasitoid: Virus Interactions

1.2.7 The Concept of Harm

1.3 Parasite Hosts

Parasites of Parasites

1.4 Zoonotic Infections

1.5 The Co‐evolution of Parasites and Their Hosts

1.5.1 The Red Queen’s Race Hypothesis

1.5.2 Parasites in the Fossil Record

1.5.3 Parasites and the Evolution of Sexual Reproduction

1.6 Parasitism as a ‘Lifestyle’: Advantages and Limitations

1.7 The Economic Cost of Parasitic Diseases

1.7.1 DALYs: Disability‐Adjusted Life Years

1.8 Why Parasitic Diseases Remain a Problem

2 Taxonomy. CONTENTS

2.1 Introduction

2.2 Viruses: A Special (Unresolved) Case

2.3 Taxonomic Hierarchy

2.3.1 The Binomen System

2.4 Kingdom Protista

2.5 Kingdom Animalia

2.5.1 Parazoa

2.5.2 Eumetazoa

3 Parasitic Protozoa Part A: Phyla Rhizopoda, Metamonada, Apicomplexa. CONTENTS

3.1 Introduction

3.2 Phylum Rhizopoda

3.2.1 Entamoeba histolytica

Entamoebae and Amoebic Dysentery

3.2.2 Entamoeba dispar

3.2.3 Entamoeba moshkovskii

3.2.4 Entamoeba gingivalis

3.2.5 Naegleria fowleri

3.2.6 Balamuthia mandrillaris

3.2.7 Genus Acanthamoeba

3.3 Phylum Metamonada

3.3.1 Order Diplomonadida

3.3.1.1 Genus Giardia

3.3.1.1.1 Giardia duodenalis

3.3.2 Order Trichomonadida

3.3.2.1 Histomonas meleagradis

3.3.2.2 Trichomonas vaginalis

3.3.2.3 Trichomonas tenax

3.3.2.4 Trichomonas gallinae

3.3.2.5 Tritrichomonas foetus

3.3.2.6 Pentatrichomonas hominis

3.4 Phylum Apicomplexa

Plastids in Parasites

3.4.1 Genus Plasmodium

How Malaria Has Influenced the Course of History

3.4.1.1 Plasmodium Life Cycle

3.4.1.2 Plasmodium falciparum

3.4.1.3 Plasmodium vivax

3.4.1.4 Plasmodium ovale

3.4.1.5 Plasmodium malariae

3.4.1.6 Plasmodium knowlesi

3.4.2 Genus Theileria

3.4.2.1 Theileria Life Cycle

3.4.2.2 Theileria parva

3.4.3 Genus Babesia

3.4.3.1 Babesia Life Cycle

3.4.3.2 Babesia bigemina

3.5 Subclass Coccidiasina

3.5.1 Genus Eimeria

3.5.1.1 Eimeria tenella

3.5.2 Genus Isospora

3.5.3 Genus Cystoisospora

3.5.3.1 Cystoisospora (Isospora) belli

3.5.4 Genus Cyclospora

3.5.4.1 Cyclospora cayetanensis

3.5.5 Genus Sarcocystis

Human Sarcocystis Infections

3.5.6 Genus Toxoplasma, Toxoplasma gondii

Population structure of Toxoplasma gondii

3.5.7 Genus Neospora

3.5.7.1 Neospora caninum

3.5.8 Genus Cryptosporidium

4 Parasitic Protozoa Part B: Phylum Kinetoplastida; Parasitic Algae and Fungi. CONTENTS

4.1 Introduction

4.2 Phylum Kinetoplastida

4.2.1 Genus Leishmania

4.2.1.1 Leishmania Life Cycle

How Harming the Vector Facilitates Transmission

How Leishmania Establishes Within Mammalian Phagocytes

4.2.1.2 Visceral Leishmaniasis

4.2.1.3 Post Kala‐Azar Dermal Leishmaniasis

4.2.1.4 Cutaneous Leishmaniasis

4.2.2 Genus Trypanosoma

4.2.2.1 Trypanosoma brucei

Genomic Regulation in Trypanosomes

How Trypanosomes Alter Tsetse Fly Physiology to Facilitate Transmission

4.2.2.2 Trypanosoma congolense

4.2.2.3 Trypanosoma evansi

4.2.2.4 Trypanosoma equinum

4.2.2.5 Trypanosoma equiperdum

4.2.2.6 Trypanosoma cruzi

4.3 Phylum Chlorophyta

4.3.1 Genus Prototheca

4.4 Kingdom Fungi

4.4.1 Phylum Microspora

5 Platyhelminth and Acanthocephalan Parasites. CONTENTS

5.1 Introduction

5.2 Phylum Platyhelminthes

5.2.1 Trematoda

5.2.1.1 Family Fasciolidae

5.2.1.1.1 Fasciola hepatica/Fasciola gigantica

Environmental DNA (eDNA) and Parasite/Intermediate Host Monitoring

5.2.1.1.2 Fasciolopsis buski

5.2.1.2 Family Dicrocoeliidae

5.2.1.2.1 Dicrocoelium dendriticum

5.2.1.3 Family Paramphistomatidae

5.2.1.3.1 Genus Calicophoron: Calicophoron daubneyi

5.2.1.4 Family Opisthorchiformes

5.2.1.4.1 Clonorchis sinensis

5.2.1.4.2 Opisthorchis viverrini

5.2.1.4.3 Opisthorchis felineus

5.2.1.5 Family Paragonomidae

5.2.1.5.1 Paragonimus westermani

5.2.1.6 Family Cathaemasiidae: Genus Ribeiroia

5.2.1.7 Family Schistosomatidae: Schistosoma mansoni, Schistosoma japonicum, Schistosoma haematobium

5.3 Class Cestoda

5.3.1 Order Pseudophyllidea/Diphyllobothridea

5.3.1.1 Genus Diphyllobothrium

5.3.1.1.1 Diphyllobothrium latum

5.3.2 Order Cyclophyllidea

5.3.2.1 Family Taeniidae

5.3.2.1.1 Taenia solium

5.3.2.1.2 Taenia saginata

5.3.2.1.3 Taenia hydatigena

5.3.2.1.4 Taenia multiceps

5.3.2.2 Genus Echinococcus

5.3.2.2.1 Echinococcus granulosus

5.3.2.2.2 Echinococcus multilocularis

Has Successful Anti‐Rabies Control Resulted in an Increased Risk of Echinococcus multilocularis Infection in Europe?

5.3.2.3 Family Anoplocephalidae

5.3.2.3.1 Anoplocephala perfoliata

5.3.2.3.2 Moniezia expansa and Moniezia benedeni

5.4 Phylum Acanthocephala

Homosexual Interactions in the Acanthocephalan Moniliformis dubius

6 Nematode Parasites. CONTENTS

6.1 Introduction

6.2 Class Enoplea. 6.2.1 Genus Trichuris

6.2.2 Genus Trichinella

6.2.2.1 Trichinella spiralis

Invasion of Muscle Cells by Trichinella spiralis

6.3 Class Rhabdita. 6.3.1 Genus Strongyloides

6.3.1.1 Strongyloides stercoralis

6.3.2 Genus Ancylostoma

Arrested Development (Hypobiosis) in Hookworm Larvae

6.3.3 Genus Necator

6.3.4 Genus Ascaris

6.3.5 Genus Enterobius: Enterobius vermicularis

6.3.6 Genus Toxocara: Toxocara canis

6.3.7 Genus Anisakis

6.3.8 Family Onchocercidae

6.3.8.1 Genus Onchocerca

6.3.8.2 Genus Wuchereria, Wuchereria bancrofti

6.3.8.3 Genus Brugia

6.3.8.4 Genus Loa, Loa loa

6.3.9 Family Dracunculidae: Dracunculus medinensis

7 Arthropod Parasites. CONTENTS

7.1 Introduction

7.2 Phylum Chelicerata

7.2.1 Family Demodicidae

7.2.2 Family Sarcoptidae

7.2.2.1 Genus Sarcoptes

7.2.2.2 Genus Notoedres, Notoedres cati

7.2.2.3 Genus Knemidocoptes

7.2.3 Family Psoroptidae

7.2.3.1 Genus Psoroptes

7.2.4 Suborder Ixodida

7.2.4.1 Family Argasidae

7.2.4.1.1 Argas persicus

7.2.4.2 Family Ixodidae

7.3 Phylum Crustacea

7.3.1 Subclass Copepoda

7.3.2 Infraclass Cirripedia

7.3.3 Subclass Branchiura

7.3.4 Subclass Pentastomida

7.4 Subphylum Hexapoda

7.4.1 Order Phthiraptera (Lice)

7.4.2 Order Siphonaptera (Fleas)

Tungiasis

7.4.3 Order Diptera (True Flies)

7.4.3.1 Suborder Nematocera

7.4.3.2 Suborder Brachycera

Wound Myiasis

7.4.3.2.1 Family Calliphoridae

Blowfly Strike in Sheep

7.4.3.2.1.1 Genus Chrysomya

7.4.3.2.1.2 Genus Cochliomyia

7.4.3.2.1.3 Genus Auchmeromyia

7.4.3.2.1.4 Genus Cordylobia

7.4.3.2.2 Family Sarcophagidae

7.4.3.2.3 Family Oestridae

7.4.3.2.3.1 Subfamily Oestrinae

7.4.3.2.3.2 Subfamily Gasterophilinae

7.4.3.2.3.3 Subfamily Hypodermatinae

7.4.3.2.3.4 Subfamily Cuterebrinae

7.4.3.2.4 Family Streblidae

7.4.3.2.5 Family Nycteribiidae

8 Parasite Transmission. CONTENTS

8.1 Introduction

8.2 Contaminative Transmission

Geohelminths and Geophagy

Contaminative Transmission of Cyclospora spp

8.3 Transmission Associated with Reproduction. 8.3.1 Sexual (Venereal) Transmission

8.3.2 Transmission within Gametes

8.3.3 Congenital Transmission

8.4 Autoinfection

8.5 Nosocomial Transmission

8.6 Active Parasite Transmission

8.7 Hosts and Vectors. 8.7.1 Paratenic Hosts

8.7.2 Intermediate Hosts

8.7.3 Vectors

The Role of Symbionts in the Life of Tsetse Flies and Their Transmission of Trypanosome Parasites

8.8 Host Factors. 8.8.1 Host Identification

8.8.2 The Influence of Host Behaviour on Parasite Transmission

Baby Care and Strongyloides fuelleborni kellyi Infection

8.8.3 Religion and Parasite Transmission

8.8.4 War and Parasite Transmission

8.8.5 Parasites Influencing Host Behaviour

The Effect of Toxoplasma gondii on Human Behaviour

8.9 Co‐Transmission and Interactions Between Pathogens

8.10 Environmental Factors. 8.10.1 Natural Environmental Variables

8.10.2 Pollution

Sewage Effluent, Toxoplasma gondii and Marine Sentinel Species

8.10.3 Climate Change

A Perfect Storm: Did Global Warming Contribute to Disease in African Lions?

9 Immune Reactions to Parasitic Infections. CONTENTS

9.1 Introduction

9.2 Invertebrate Immunity

9.3 Vertebrate Immunity

9.3.1 Innate Immunity

Innate Immunity to Trypanosome Infection

9.3.2 Adaptive Immunity

9.3.3 Cell‐Mediated Immunity

9.4 Innate Immunity to Parasites. 9.4.1 Physical Factors

9.4.2 Chemical and Microbial Factors

9.4.3 Acute Inflammatory Response

9.4.4 Cell‐Mediated Immune Reactions

9.5 Adaptive Immune Reactions to Parasites

9.6 Microbiomes and Host Immune Reactions to Parasites

9.7 Avoiding the Host Immune Response

Hide in a ‘safe house’

Stay quiet and do not draw attention to yourself

Camouflage

Keep changing your disguise

Put out false information

Neutralise any threats

Exhaust the enemy

9.8 Immunity to Malaria

9.8.1 Plasmodium: Anopheles Interactions

9.8.2 Plasmodium: Human Interactions

9.8.2.1 Innate Immune Mechanisms Against Malaria

9.8.2.2 Antibodies Against Malaria

9.8.2.3 Why Humans Do Not Develop Protective Immunity Against Malaria

9.9 Schistosoma spp. and Hepatitis C Virus Interactions

9.10 HIV‐AIDS and Parasitic Infections

9.10.1 Parasites and the Transmission of HIV

9.10.2 Parasite‐HIV Co‐Infections

9.10.2.1 Leishmania‐HIV Co‐Infections

9.10.2.1.1 The Increasing Problem of HIV‐ Leishmania Co‐Infections

9.10.2.2 Malaria‐HIV Co‐Infections

9.10.2.3 Toxoplasma‐HIV Co‐Infections

9.10.2.4 Microsporidia‐HIV Co‐Infections

10 Pathology Part A: Factors Influencing Pathogenesis, How Parasites Cause Pathology, Types of Pathology. CONTENTS

10.1 Introduction

10.2 Factors Influencing Pathogenesis

10.2.1 Host Factors

10.2.2 Parasite Factors

10.3 Mechanisms By Which Parasites Cause Pathology

10.3.1 Direct Damage

10.3.2 Indirect Damage

10.4 Types of Pathology. 10.4.1 Abortion and Obstetric Pathology

10.4.2 Abscesses and Ulcers

10.4.3 Anaemia

10.4.4 Anorexia

10.4.5 Apoptosis

10.4.6 Autoimmunity

10.4.7 Calcification

10.4.8 Cancer

10.4.9 Castration

10.4.10 Delusional Parasitosis

10.4.11 Diarrhoea

10.4.12 Elephantiasis

10.4.13 Fever

10.4.14 Fibrosis

10.4.15 Granulation

10.4.16 Hyperplasia

10.4.17 Hypertrophy

10.4.18 Hypoplasia and Hypotrophy

10.4.19 Inflammation

10.4.20 Jaundice

10.4.21 Metaplasia

10.4.22 Pressure Atrophy

10.4.23 Psychological Disturbance

11 Pathology Part B: Damage to Specific Organs; Co‐Infections and Pathogenesis. CONTENTS

11.1 Introduction

11.2 Damage to Specific Organs. 11.2.1 Bladder

11.2.2 Brain and Nervous System

11.2.3 Gastrointestinal Tract

11.2.4 Gall Bladder and Bile Ducts

11.2.5 Genitalia

11.2.6 Kidney

11.2.7 Liver

11.2.8 Lungs

11.2.9 Skin

It Started with a Leech Bite

11.2.10 Spleen

11.3 Co‐Infections and Pathogenesis

12 The Useful Parasite. CONTENTS

12.1 Introduction: The Goodness of Parasites?

12.2 The Importance of Parasites for the Maintenance of a Healthy Immune System

12.2.1 Type 1 Diabetes Mellitus

12.2.2 Type 2 Diabetes

12.2.3 Irritable Bowel Syndrome

12.2.4 Inflammatory Bowel Disease

12.3 The Use of Parasites to Treat Medical Conditions

12.3.1 Helminth Therapy

12.3.1.1 Helminth Therapy in Practice

12.3.2 Maggot Therapy

12.3.2.1 Maggot Therapy in Practice

12.3.2.2 How Maggot Therapy Works

12.3.3 Leech Therapy

12.3.3.1 Leech Therapy in Practice

12.3.4 Malaria Therapy (Malariotherapy)

12.4 Parasites as Sources of Novel Pharmaceutically Active Substances

12.5 Parasites as Biological Control Agents

12.5.1 Life Cycle of the Entomopathogenic Nematodes Heterorhabditis and Steinernema

Rhabditid Nematode–Bacteria Relationships

12.6 Parasites as Forensic Indicators

The Use of Parasitoid Wasps to Determine the Minimum Time Since Death

13 The Identification of Protozoan and Helminth Parasites. CONTENTS

13.1 Laboratory Testing for Parasitic Infections: Introduction

13.2 Importance of Correct Identification

13.3 Properties of an Ideal Diagnostic Test

13.4 Isolation of Parasites

Giardia String Test (Entero‐Test)

13.5 Identification from Gross Morphology

13.5.1 Morphological Identification of Entamoeba spp

13.5.2 Morphological Identification of Plasmodium spp. and Babesia spp

13.5.3 Morphological Identification of Taenia spp. Tapeworms

13.5.4 Morphological Identification of Filarial Nematode Infections

13.6 Biochemical Techniques for Identification

13.7 Immunological Techniques for Identification

13.8 Molecular Techniques for Identification

How Diagnostic Techniques Can Influence Epidemiological Studies

13.9 Diagnostic Testing of Parasitic Infections Outside the Laboratory: Introduction

13.9.1 Immunochromatographic (Lateral Flow) Tests

13.9.1.1 Detection of Parasites in Blood with LFDs. 13.9.1.1.1 Detection of Plasmodium spp. with LFDs

13.9.1.1.2 Detection of Leishmania spp. with LFDs

13.9.1.2 Detection of Parasites in Faeces with LFDs. 13.9.1.2.1 Detection of Faecal Protozoa with LFDs

13.9.1.2.2 Detection of Helminths in Faeces with LFDs

13.9.2 Point‐of‐Care Nucleic Acid Amplification Tests (NAATs)

13.9.2.1 Detection of Trichomonas vaginalis Using POCT NAATs

13.9.2.2 Detection of Plasmodium spp. Using POCT NAATs

13.9.2.3 Invertebrate Vector and Intermediate Host Monitoring Using POCT NAATs

14 Parasite Treatment. CONTENTS

14.1 Introduction

14.2 The Ideal Antiparasitic Drug

14.3 Pharmaceutical Drugs

14.4 DNA/RNA Technology

Antisense DNA and RNA

14.5 Molecular Chaperones (Heat Shock Proteins)

14.6 Nanotechnology

Gold Nanoparticles for the Diagnosis and Treatment of Parasitic Infections

14.7 Quantum Dots

14.8 Natural Remedies

Artemisinin and the Treatment of Malaria

14.9 Homeopathy

15 Parasite Vaccines. CONTENTS

15.1 Introduction

15.2 The Design and Use of Vaccines

15.3 Herd Immunity

15.4 Factors Limiting the Production of Commercial Antiparasitic Vaccines

15.5 Properties of an Ideal Vaccine

15.6 Types of Vaccine

15.6.1 Live Attenuated Vaccines

15.6.2 Inactivated Vaccines

15.6.3 Subunit/Recombinant Vaccines

15.6.4 Peptide/Polypeptide Vaccines

15.6.5 Carbohydrate Vaccines

15.6.6 Toxoid (Anti‐toxin) Vaccines

15.6.7 Virus‐Like Particles Vaccines

15.6.8 DNA/RNA Vaccines

15.7 Identification of Antigens for Use in Anti‐parasite Vaccines

15.8 Vaccine Delivery

15.9 Vaccines Against Malaria

15.10 Nanobodies (Single Domain Antibodies)

15.11 Problems with Vaccination Strategies

16 Parasite Control. CONTENTS

16.1 Introduction

16.2 Eradication, Elimination, and Control

16.3 Education

16.4 Environmental Modification and Cultural Control

Organic Livestock Farming and Parasitic Diseases

16.5 Remote Sensing and GIS Technology

16.6 Whether to Treat the Individual or the Population

16.7 Piggy‐Backing Control Programmes

16.8 Disruptions to Control Programmes

16.9 Role of Governments, Foundations, and Aid Organisations

References

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Second Edition

Alan Gunn

.....

Evolution can be defined as a change in gene frequency between generations, but for this to occur three criteria need to be met. First, there must be genetic variation within the population. If the population is genetically homogeneous, then variation can only occur sporadically through random mutation. The second criterion is that the variation must be heritable: if the variation cannot be passed on to offspring, then it will be lost regardless of the benefits it imparts. The third and final criterion is that the variation must influence the probability of leaving reproductively viable offspring. If the variation is beneficial, then the organism possessing it will leave more offspring; however, unless these are reproductively viable, the variation would be quickly lost from the gene pool. Parasites live in close association with their hosts and the two organisms will co‐evolve. The nature of the host: parasite relationship may therefore change with time. For example, provided the three criteria are met, the host will evolve resistance/susceptibility factors depending upon the pressure exerted by the parasite. Although ever greater resistance to infection may appear to be ‘ideal’, this is unlikely to arise if the energetic cost impacts on the ability to leave viable offspring. At the same time, the parasite will evolve virulence/avirulence factors that promote its own survival.

It is often stated that long‐standing parasite: host relationships are less pathogenic than those that have established more recently. This is based on the reasoning that if the parasite kills its host, then it will effectively ‘commit suicide’ because it will have destroyed its food supply. Consequently, over time, it is to be expected that the parasite will become less harmful to its host – that is, it becomes less virulent. However, this assumption is questionable because a pathogen’s virulence often reflects its reproductive success. For example, let us consider two hypothetical strains, A and B, of the same nematode species that lives in the gut of sheep. Strain A is highly virulent and causes the death of the sheep whilst strain B is relatively benign and seldom causes any mortality. At first glance, one might expect that strain B would leave more offspring because its host lives for longer. However, if virulence was linked to the nematode’s reproductive output and the eggs were released at a time when they were likely to infect new hosts, then strain A would bequeath more of its genes to subsequent generations. Consequently, the proportion of strain A in the nematode population would increase with time and there would be constant selection for increasing virulence. The sheep and the parasites may eventually be driven to extinction by these changes, but individual animals (and humans) are almost always driven by their own immediate self‐interest rather than hypothetical future prospects.

.....

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