Invertebrate Histology -  - E-Book

Invertebrate Histology E-Book

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The first comprehensive reference to Invertebrate Histology Invertebrate Histology is a groundbreaking text that offers a comprehensive review of histology in invertebrates. Designed for use by anyone studying, diagnosing, or researching invertebrates, the book covers all major taxonomic groups with details of the histologic features, with color photographs and drawings that clearly demonstrate gross anatomy and histology. The authors, who are each experts in the histology of their respective taxa, bring together the most recent information on the topic into a single, complete volume. An accessible resource, each chapter focuses on a single taxonomic group with salient gross and histologic features that are clearly described in the text and augmented with color photographs and greyscale line drawings. The histologic images are from mostly hematoxylin and eosin stained microscopic slides showing various organ systems at high and low magnification. In addition, each chapter provides helpful tips for invertebrate dissection and information on how to process invertebrates for histology. This important book: * Presents detailed information on histology of all major groups of invertebrates * Offers a user-friendly text that is organized by taxonomic group for easy reference * Features high-quality color photographs and drawings, with slides showing histology and gross photographs to demonstrate anatomy * Provides details on invertebrate dissection and processing invertebrates for histology Written for veterinary pathologists, biologists, zoologists, students, and other scientists studying these species, Invertebrate Histology offers the most updated information on the topic written by over 20 experts in the field.

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Table of Contents

Cover

Title Page

Copyright Page

List of Contributors

Foreword

1 Echinodermata

1.1 Introduction

1.2 Gross Anatomy

1.3 Histology

References

2 Porifera

2.1 Introduction

2.2 Gross Anatomy

2.3 Histology

2.4 Organ Systems

Acknowledgments

Abbreviations for Figures

References

3 Cnidaria

3.1 Introduction

3.2 Gross Anatomy

3.3 Histology

3.4 Conclusion

Acknowledgments

Disclaimer

Appendix 3.1 Specimen Relaxation and Common Fixative Formulations

Appendix 3.2 Basic Histology Protocol for Processing Scleractinian Corals (refer to Price and Peters (2018) for more detailed techniques)

References

4 Mollusca

4.1 Introduction

4.2 Gross Anatomy

4.3 Histology

4.4 Histology Processing Techniques (Table 4.1)

References

5 Mollusca

5.1 Introduction

5.2 Gross Anatomy

5.3 Histology (Table 5.1)

Disclaimer

References

6 Mollusca

6.1 Introduction

6.2 Gross Anatomy

6.3 Histology (Table 6.1)

References

7 Annelida

7.1 Introduction

7.2 Gross Anatomy

7.3 Histology

References

8 Arthropoda

8.1 Introduction

8.2 Gross Anatomy

8.3 Histology (Table 8.1)

Acknowledgments

Disclaimer

References

9 Arthropoda

9.1 Introduction

9.2 Gross Anatomy

9.3 Histology

Disclaimer

References

10 Arthropoda

10.1 Introduction

10.2 Gross Anatomy

10.3 Histology

Disclaimer

References

11 Arthropoda

11.1 Overview

11.2 Gross Anatomy of Adults

11.3 Histology

Acknowledgments

References

12 Arthropoda

12.1 Introduction

12.2 Gross Anatomy

12.3 Histology

Disclaimer

References

Index

End User License Agreement

List of Tables

Chapter 1

Table 1.1 Organs for histologic evaluation in Echinodermata.

Table 1.2 Cuticular layers in echinoderms (Holland).

Chapter 2

Table 2.1 Organs for histologic evaluation in Porifera.

Chapter 3

Table 3.1 Tissues, structures, and cells for histologic evaluation in cnidari...

Chapter 4

Table 4.1 Organs for histologic evaluation in Gastropoda.

Chapter 5

Table 5.1 Organs for histologic evaluation in Cephalopoda.

Chapter 6

Table 6.1 Organs for histologic evaluation in Bivalvia.

Chapter 7

Table 7.1 Organs for histologic evaluation in Annelida.

Chapter 8

Table 8.1 Organs for histologic evaluation in Arachnida.

Chapter 9

Table 9.1 Organs for histologic evaluation in Merostomata.

Chapter 10

Table 10.1 Organs for histologic evaluation in Merostomata.

Chapter 11

Table 11.1 Organs for histologic evaluation in decapods.

Chapter 12

Table 12.1 Organs for histologic evaluation in Insecta.

List of Illustrations

Chapter 1

Figure 1.1 Representative image of the aboral (a) and oral (b) surface of a ...

Figure 1.2 Representative image of the aboral (a) and oral (b) surface of a ...

Figure 1.3 Image of a white sea urchin (

Tripneustes ventricosus

) demonstrati...

Figure 1.4 Representative image of the ventral (a) and lateral (b) aspects o...

Figure 1.5 Gross necropsy image (a) of a bat star (

Patiria miniata

) open at ...

Figure 1.6 Gross necropsy images of urchin open at necropsy. Images include ...

Figure 1.7 Gross necropsy images of a California giant sea cucumber opened a...

Figure 1.8 Low‐magnification image of the histology of the body wall of an (...

Figure 1.9 Histology of the epidermis of a sunflower sea star (

Pycnopodia he

...

Figure 1.10 Low‐magnification image of the histology of a sunflower sea star...

Figure 1.11 Higher magnification image of the histology of an ochre sea star...

Figure 1.12 Histology of the base of a white sea urchin spine at the ball an...

Figure 1.13 Histology of white sea urchin appendages including pedicellaria ...

Figure 1.14 Histology of the madreporite (a) and stone canal (b) in a mottle...

Figure 1.15 Histology of the water vascular (radial) canal in a white urchin...

Figure 1.16 Histology of a tube foot in a mottled star. 25×, HE. C, connecti...

Figure 1.17 Histology of the cardiac (a) stomach in a mottled star (100×, HE...

Figure 1.18 Histology of the pyloric cecae of a mottled star. 25×, HE.

Figure 1.19 Histology of the large intestine of a white urchin. 200×, HE.

Figure 1.20 Low‐magnification histology of anatomy of Aristotle's lantern in...

Figure 1.21 Axial gland in a white urchin. 400×, HE.

Figure 1.22 Tiedmann's body in a mottled star. 100×, HE.

Figure 1.23 Histology of gills (papulae) in a white urchin showing epidermal...

Figure 1.24 Histology of the ventral nerve cord (N) in a mottled star. 100×,...

Figure 1.25 Histology of the ovary (a) in a Caribbean thorny star, and testi...

Figure 1.26 Nutritive support cells in the gonad of a sand dollar. 100×, HE....

Chapter 2

Figure 2.1 Gross anatomy. (a) Scheme of sponge organization; black arrows: w...

Figure 2.2 The ectosome, dermal membrane, and cortex. (a) Semithin section o...

Figure 2.3 Cuticle, exopinacoderm, and pores. (a) Histologic section of

Aply

...

Figure 2.4 Cells of the internal environment. (a) Lophocyte of

Chondrilla

sp...

Figure 2.5 Mesohyl cellular strands of

Aplysina cavernicola

. (a) General vie...

Figure 2.6 Types of aquiferous systems in sponges. (a) Asconoid aquiferous s...

Figure 2.7 Inhalant aquiferous system. (a,b) Ostia and inhalant canals lined...

Figure 2.8 Choanocyte chambers. (a) Syconoid choanocyte chambers in

Sycettus

...

Figure 2.9 Skeleton. (a) Inorganic skeleton (SiO

2

) of

Haliclona

sp. with reg...

Figure 2.10 Reproduction, female. (a) Oviparous gonochoric sponge

Aplysina c

...

Figure 2.11 Follicle. (a) Beginning of follicle development in

Oscarella nic

...

Figure 2.12 Tissue modification during sexual reproduction in

Halisarca duja

...

Chapter 3

Figure 3.1 Cnidarian body forms: polyp and medusa.

Figure 3.2 Gross anatomy of a scleractinian polyp from an imperforate coral....

Figure 3.3 Overview of key structures in a polyp from the scleractinian cora...

Figure 3.4 Gross anatomy of an octocoral (gorgonian polyp).

Figure 3.5 Subgross photomicrograph of a sea pen (order Pennatulacea). The e...

Figure 3.6 Undecalcified

Acropora cervicornis

section prepared using resin e...

Figure 3.7

Acropora cervicornis

tissue section with aragonite skeleton disso...

Figure 3.8 Diagram showing the surface body wall of a scleractinian polyp, s...

Figure 3.9 Diagram of the key features of a scleractinian polyp mesentery. F...

Figure 3.10 Body wall from a scleractinian coral,

Orbicella annularis

. The s...

Figure 3.11 Body wall from a scleractinian coral,

Orbicella cavernosa

. The m...

Figure 3.12 Body wall from a branching perforate scleractinian coral,

Acropo

...

Figure 3.13 Body wall from an aggregating anemone (

Anthopleura

sp.) showing ...

Figure 3.14 Cross‐section of a moon jelly (

Aurelia aurita

). The epithelial l...

Figure 3.15 Body wall of the gorgonian,

Antillogorgia americana

, sectioned a...

Figure 3.16 Body wall of a sea pen (order Pennatulacea). The epidermis (E) i...

Figure 3.17 High magnification of the surface of a sea fan,

Gorgonia ventali

...

Figure 3.18 High magnification of the surface of another gorgonian,

Plexaure

...

Figure 3.19 Epidermal mucocytes secreting through apical pores (

arrows

) in

O

...

Figure 3.20 Diagram of a cnidocyte (

left

) ejecting venom from the tip of the...

Figure 3.21 Exumbrella from a moon jelly (

Aurelia auria

). Embedded within th...

Figure 3.22 Transmission electron microscopy of nematocysts from a spotted j...

Figure 3.23 The tightly wound tropocollagen‐rich tubules of spirocysts are e...

Figure 3.24 Ciliated surface body wall of the burrowing sea anemone,

Ceriant

...

Figure 3.25 Epidermal pigment cells (

arrows

), possibly containing fluorescen...

Figure 3.26 Melanin‐like pigment‐containing granular amoebocytes (aka amoebo...

Figure 3.27 Squamous calicoblasts (

arrowhead

) in the calicodermis deep in th...

Figure 3.28 Calicoblasts (

arrows

) containing eosinophilic granules in the ca...

Figure 3.29 Corticocytes (

arrow

) secrete the proteinaceous axis and desmocyt...

Figure 3.30 Low magnification of a fixed, decalcified sample from the sea fa...

Figure 3.31 Section through the deep basal body wall of the scleractinian co...

Figure 3.32 Vacuole‐bound symbiotic dinoflagellates in the gastrodermis of

O

...

Figure 3.33 Transmission electron microscopy of zooxanthellae from a spotted...

Figure 3.34 Large slightly basophilic mucocytes (

arrows

) line the gastroderm...

Figure 3.35 Mesenteries of

Montastraea cavernosa

. The thickness of mesenteri...

Figure 3.36 Mesenteries of aggregating anemone (

Anthopleura

sp.). The mesent...

Figure 3.37 Mesenterial filament of a sea pen (order Pennatulacea). The term...

Figure 3.38 Mesenterial filaments of aggregating anemone (

Anthopleura

sp.). ...

Figure 3.39 Cnidoglandular band and mesentery sections in

Pseudodiploria

sp....

Figure 3.40 Higher magnification of Figure 3.39. Nematocysts in cnidoglandul...

Figure 3.41 Higher magnification of Figure 3.39. Gastrodermal granular pigme...

Figure 3.42 Pigmented areas on mesenterial filaments of

O. faveolata

. Gross ...

Figure 3.43 Large gastrodermal granular gland cells found in the mesentery o...

Figure 3.44 Granular cells in the calcifying hydrozoan fire coral,

Millepora

Figure 3.45 Mesoglea (

asterisk

) in coral

Orbicella cavernosa

. Epidermis is o...

Figure 3.46 Mesoglea in gorgonian

Gorgonia ventalina

showing spaces left by ...

Figure 3.47 Mesogleal pleats (P) in

Fungia

sp. 600×. HE.

Figure 3.48 Mesogleal pleats (

arrowheads

) in

Orbicella cavernosa

have thick ...

Figure 3.49 Arrays of myonemes (longitudinal section) (MN) from epitheliomus...

Figure 3.50 Myonemes (

arrowheads

; cross‐section), basal extensions of epithe...

Figure 3.51 Contracted bands of myonemes in cross‐section from a burrowing a...

Figure 3.52 Surface body wall of

Manicina areolata

, showing the epidermis (E...

Figure 3.53 Mesoglea of a moon jelly (

Aurelia aurita

). Amoebocytes are scatt...

Figure 3.54 Mesoglea of a fleshy scleractinian coral,

Colpophyllia natans

, s...

Figure 3.55 Mesoglea of a sea fan (

Gorgonia ventalina

), containing multiple ...

Figure 3.56 Mesenteries in an aggregating anemone (

Anthopleura

sp.). Spermar...

Figure 3.57 Spermatozoa in

Acropora

sp. 600×. HE.

Figure 3.58 Spermatozoa in an aggregating anemone (

Anthopleura

sp.). 200×. H...

Figure 3.59 Multiple stages of developing oocytes from the jellyfish

Aurelia

...

Figure 3.60 Oocytes in

Acropora

sp. 200×. HE.

Figure 3.61 Oocytes (

asterisks

) in an unknown species of gorgonian. The tiss...

Figure 3.62 Oocytes (

arrowheads

) developing within the medusa stage in the f...

Figure 3.63

Hydra

spp. exist as solitary polyps and do not have a medusoid f...

Figure 3.64 A myxozoan plasmodium located in the gill arch of the fish

Lepom

...

Figure 3.65 Polar capsules of myxozoan spores found in gill arch of

Lepomis

...

Figure 3.66 Fragments of coral specimens enrobed in agar.

Figure 3.67 Decalcification of coral fragment enrobed in agar. (a) Remove ex...

Chapter 4

Figure 4.1 Shell of

Lobatus gigas

. The apex (

arrow

) of the shell was formed ...

Figure 4.2 (a)

Helix pomatia

in anesthetized state. The head (

arrowhead

) and...

Figure 4.3 Two female

Lobatus

gigas

, removed from shell; (a) mantle cavity u...

Figure 4.4 Dissected

Helix

in water cover displaying the internal organs. AG...

Figure 4.5 Comparison of integument in (a) mantel of

Littorina ziczac

, 400×,...

Figure 4.6 Hypobranchial gland of

Cittarium pica

. The gland is composed of c...

Figure 4.7 (a) Connective tissue of the body of

Lobatus gigas

. Fibrous conne...

Figure 4.8 Ventral (interior) mantel of

Lobatus

gigas

. Epidermal cells are c...

Figure 4.9 Dorsal (exterior) mantel of

Lobatus

gigas

. Epidermal cells are le...

Figure 4.10 Mantle edge of

Lobatus

gigas

. Arrows designate the mantle edge g...

Figure 4.11 Ink gland of mantle edge of

Aplysia dactylomela

. A large flask‐s...

Figure 4.12 Overview of the mantle collar with zones of

Helix

. E, epithelium...

Figure 4.13 Histologic structure of the mantle collar of

Helix

. (a) Mantle c...

Figure 4.14 Undecalcified thin ground section of the shell of

Helix

. Ch, con...

Figure 4.15 Pedal glands of

Helix

. (a) Overview of pedal protein gland and s...

Figure 4.16 Columellar muscle of

Triplofusus papillosus

. Myocytes are arrang...

Figure 4.17 Radular apparatus of

Cerithium

sp. Within the buccal cavity (B),...

Figure 4.18 Posterior radular apparatus of

Nerita peloronta

. Odontoblasts ar...

Figure 4.19 Pharynx and radula of

Helix

. (a) Overview of the cross‐section o...

Figure 4.20 Esophagus of

Cerithium

sp. The esophagus has a very thin wall co...

Figure 4.21 Longitudinal section of the esophagus of

Helix

. BB, brush border...

Figure 4.22 Crop and salivary gland in

Helix

. (a) Longitudinal section of th...

Figure 4.23 Stomach of

Cerithium

sp. In areas, the gastric mucosa is covered...

Figure 4.24 Style sac of

Lobatus

gigas

. A distinct fold in the wall is a ref...

Figure 4.25 Intestine of

Cittarium pica

. Prominences reflect typhlosoles (

ar

...

Figure 4.26 Anus of

Lobatus

gigas

. The mucosa (M) is lined by simple to pseu...

Figure 4.27 Midgut of an active

Helix

. (a) Overview of the midgut embedded i...

Figure 4.28 Rectum of

Helix

. (a) Overview of cross‐section of the rectum. 50...

Figure 4.29 Digestive gland of

Cittarium pica

. Acini (

asterisks

) are lined b...

Figure 4.30 Digestive gland of

Helix

. (a) Overview of the digestive gland an...

Figure 4.31 Ceras of

Limenandra confusa

. Cerata are extensions of the mantle...

Figure 4.32 Kidney of

Tegula excavata

. Hemolymphatic spaces (H) separate tub...

Figure 4.33 Histology of renal sac of active and inactive

Helix

. (a) Histolo...

Figure 4.34 Renal sac and ureters in

Helix

. (a) Overview of primary ureter a...

Figure 4.35 Nephridial gland of

Lobatus

gigas

. Tubular folds are lined by si...

Figure 4.36 Heart of

Lobatus

gigas

. The myocardium consists of trabeculae (

a

...

Figure 4.37 Overview of heart in pericardium and striated heart muscle of

He

...

Figure 4.38 Histologic structure of the heart of

Helix

. (a) Ventricular wall...

Figure 4.39 Pericardium of

Cittarium

pica

. The pericardium consists of a mem...

Figure 4.40 Different areas of the pericardium of

Helix

. ELM, epithelial lay...

Figure 4.41 Hemolymphatic vessels in

Helix

. (a) Artery above subpedal gland....

Figure 4.42 Hemocyte‐producing organ in

Helix

. (a) Overview of hemocyte‐prod...

Figure 4.43 Type I hemocytes in the hemocyte‐producing organ and in the lung...

Figure 4.44 Type II hemocytes of

Helix

. (a) Type II hemocytes in the lung an...

Figure 4.45 Gill of

Tegula excavata

. The gill is bipectinate, bearing lamell...

Figure 4.46 Lung of

Helix

. (a) Overview of cross‐section of the lung. 30×. H...

Figure 4.47 Nerves of

Helix

. (a) Large nerve from subesophageal ganglion of

Figure 4.48 Ganglion and nerve of

Lobatus

gigas

. Nerves (N) are composed of ...

Figure 4.49 Ganglion of

Aplysia californica

. Large neurons predominate. 100×...

Figure 4.50 Cerebral ganglia of

Helix

. (a) Lobes and cell types and neural s...

Figure 4.51 Overview of ommatophore with digitate ganglion and collar cells ...

Figure 4.52 Male gonad of

Cerithium

sp. The gonad marginates digestive gland...

Figure 4.53 Female gonad of

Tegula ecavata

. Vitellogenic oocytes (

asterisks

)...

Figure 4.54 Ovotestis of

Helix

. (a) Overview of the gonad. 15×. HE. (b) Basi...

Figure 4.55 Spermatogenetic stages of

Helix

. EC, epithelial (nutritive) cell...

Figure 4.56 Oogenic stages of

Helix

. FC, follicular cell; FL, follicular cel...

Figure 4.57 Gonoducts and dart sac of

Helix

. (a) Histologic structure of the...

Figure 4.58 Nidamental gland of

Lobatus

gigas

. Mucopolysaccharide‐secreting ...

Figure 4.59 Prostate of

Lobatus

gigas

. Tubules are lined by simple columnar ...

Figure 4.60 Comparison of eyes. (a)

Corallophilia

sp. 400×. HE. (b)

Littorin

...

Figure 4.61 Eye and tentacular ganglion complex of

Helix

. (a) Overview with ...

Figure 4.62 Comparison of osphradia. (a)

Nerita peloronta

. The well‐develope...

Chapter 5

Figure 5.1 Color variation and patterns of

Hapalochlaena

spp. chromatophores...

Figure 5.2 External anatomy of cephalopods. (a) Nautilus. c, cirri; e, eye; ...

Figure 5.3 Decapodiforme, cuttlefish (

Sepia officinalis

) photograph showing ...

Figure 5.4 Giant Pacific octopus (

Enteroctopus dofleini

) with sessile, radia...

Figure 5.5 Cuttlefish (

Sepia

sp.) showing the mantle (M), eye (E), arms (A),...

Figure 5.6 Giant Pacific octopus (

Enteroctopus dofleini

) beak

in situ

(backg...

Figure 5.7 Giant Pacific octopus (

Enteroctopus dofleini

), the mantle of whic...

Figure 5.8 Excretory organs of a giant Pacific octopus (

Enteroctopus doflein

...

Figure 5.9 Giant Pacific octopus (

Enteroctopus dofleini

)

circulatory system....

Figure 5.10 Formalin‐fixed cuttlefish (

Sepia officinalis

), the mantle of whi...

Figure 5.11 (a) Necropsy of a common cuttlefish (

Sepia officianalis

). Portio...

Figure 5.12 Giant Pacific octopus (

Enteroctopus dofleini

). (a) Hectocotylus ...

Figure 5.13 Mantle incision of a Giant Pacific octopus (

Enteroctopus doflein

...

Figure 5.14 (a) Cuttlefish (

Sepia officinalis

). The cuttlebone attached to m...

Figure 5.15 Cuttlefish (

Sepia officinalis

) mantle. Columnar epithelial cells...

Figure 5.16 Cuttlefish (

Sepia officinalis

) skin showing melanophores and iri...

Figure 5.17 Arm of giant Pacific octopus (

Enteroctopus dofleini

). Note that ...

Figure 5.18 Section through a sucker on the arm of a cuttlefish (

Sepia offic

...

Figure 5.19 Section through the buccal mass of a giant Pacific octopus (

Ente

...

Figure 5.20 Radula bearing chitinous teeth in a chambered nautilus (

Nautilus

...

Figure 5.21 Types of salivary gland from giant Pacific octopus (

Enteroctopus

...

Figure 5.22 Esophagus of a northern flamboyant cuttlefish (

Metasepia pfeffer

...

Figure 5.23 Cecum of a giant Pacific octopus (

Enteroctopus dofleini

). The mu...

Figure 5.24 Intestine of a giant Pacific octopus (

Enteroctopus dofleini

). Th...

Figure 5.25 Digestive gland of a giant Pacific octopus (

Enteroctopus doflein

...

Figure 5.26 Atrophic digestive gland of a senescent giant Pacific octopus (

E

...

Figure 5.27 Ink sac complex of a northern flamboyant cuttlefish (

Metasepia p

...

Figure 5.28 Renal appendages of giant Pacific octopus (

Enteroctopus dofleini

Figure 5.29 Renal appendage of a giant Pacific octopus (

Enteroctopus doflein

...

Figure 5.30 Pancreatic appendages of a northern flamboyant cuttlefish (

Metas

...

Figure 5.31 Systemic heart (i.e., cardiac muscle) of a northern flamboyant c...

Figure 5.32 Branchial heart (BH) and branchial appendage (BA) of a giant Pac...

Figure 5.33 Branchial appendage of a giant Pacific octopus (

Enteroctopus dof

...

Figure 5.34 Aorta of a northern flamboyant cuttlefish (

Metasepia pfefferi

). ...

Figure 5.35 Hemolymph from a giant Pacific octopus (

Enteroctopus dofleini

) w...

Figure 5.36 Body wall of a common octopus (

Octopus vulgaris

) with interstiti...

Figure 5.37 White body of a common cuttlefish (

Sepia officinalis

). The white...

Figure 5.38 Branchial gland and gills of a giant Pacific octopus (

Enteroctop

...

Figure 5.39 Gills of a chambered nautilus (

Nautilus pompilius

) (a) and a gia...

Figure 5.40 Branchial gland of a northern flamboyant cuttlefish (

Metasepia p

...

Figure 5.41 Brain of an unspecified cephalopod revealing the brain surroundi...

Figure 5.42 Brain of a giant Pacific octopus (

Enteroctopus dofleini

). The op...

Figure 5.43 Central nervous system of a northern flamboyant cuttlefish (

Meta

...

Figure 5.44 Peripheral nerve of a chambered nautilus (

Nautilus pompilius

). P...

Figure 5.45 Saccular ovary of a common octopus (

Octopus vulgaris

) on wet mou...

Figure 5.46 Mature follicle of a giant Pacific octopus (

Enteroctopus doflein

...

Figure 5.47 Oviduct of a giant Pacific octopus (

Enteroctopus dofleini

) has d...

Figure 5.48 Oviductal gland of a giant Pacific octopus (

Enteroctopus doflein

...

Figure 5.49 Paired nidamental gland in a northern flamboyant cuttlefish (

Met

...

Figure 5.50 Male gonad in a northern flamboyant cuttlefish (

Metasepia pfeffe

...

Figure 5.51 Vas deferens of a two‐spot octopus (

Octopus bimaculoides

) is lin...

Figure 5.52 Male gonad in a northern flamboyant cuttlefish (

Metasepia pfeffe

...

Figure 5.53 Spermatophoric glands I and II of a two‐spot octopus (

Octopus bi

...

Figure 5.54 Spermatophoric glands II with glandular parenchyma with basal nu...

Figure 5.55 Needham's sac of a two‐spot octopus (

Octopus bimaculoides

) conta...

Figure 5.56 Anterior portion of the eye of a common cuttlefish (

Sepia offici

...

Figure 5.57 Anterior portion of the eye of a common cuttlefish (

Sepia offici

...

Figure 5.58 Retina of a common cuttlefish (

Sepia officinalis

). The distal la...

Chapter 6

Figure 6.1 D‐shaped eastern oyster (

Crassostrea virginica

) larvae.

Figure 6.2 Early Prodissoconch II (umbo) stage of the eastern oyster. a, sto...

Figure 6.3 Pediveligers of hard clams (

Mercenaria mercenaria

). Arrows indica...

Figure 6.4 Diagram of a pediveliger of the European flat oyster (

Ostrea edul

...

Figure 6.5 A hard clam with labeled directions. a, dorsal; b, posterior; c, ...

Figure 6.6 Fixed, shucked hard clam (left demibranchs and a portion of the v...

Figure 6.7 Diagram of an Atlantic surf clam (

Spicula solidissima

) with left ...

Figure 6.8 Diagram illustrating the direction of water flow from the mantle ...

Figure 6.9 Intact eastern oyster. a, anterior (ligament); b, dorsal.

Figure 6.10 Shucked eastern oyster. a, digestive gland; b, pericardial sac w...

Figure 6.11 Diagram of an oyster showing the opening of the gill's water tub...

Figure 6.12 Shucked Atlantic sea scallop (

Placopecten magellanicus

), fixed i...

Figure 6.13 Diagram of a pink scallop (

Chlamys hastata

) with left valve and ...

Figure 6.14 Pallial mantle (membranous mantle) of the central zone. a, shell...

Figure 6.15 Mantle edge of an eastern oyster showing three lobes. a, periost...

Figure 6.16 Mantle edges of a hard clam. a, lobe 1 (outer lobe); b, periostr...

Figure 6.17 Mantle edge of a hard clam. a, periostracal fold; b, periostracu...

Figure 6.18 Sagittal section through a juvenile hard clam. a, gastric shield...

Figure 6.19 Section of a hard clam. a, style in the style sac; b, intestinal...

Figure 6.20 Section of a hard clam showing the typhlosole in the descending ...

Figure 6.21 Diagram of the digestive system of an eastern oyster. an, anus; ...

Figure 6.22 Section of a hard clam. a, rectum surrounded by the ventricle of...

Figure 6.23 Section of a hard clam. a, type I cells of the digestive gland d...

Figure 6.24 Section of the digestive gland from an Atlantic sea scallop (

Pla

...

Figure 6.25 Section through the body wall of a giant clam with numerous zoox...

Figure 6.26 Section of a demibranch from a hard clam. a, water tubule; b, in...

Figure 6.27 Section of a plica from a hard clam. a, ostia; b, collagenous ro...

Figure 6.28 Section of a demibranch from a hard clam. Note the food groove (

Figure 6.29 Section showing hemocytes in the connective tissues of a hard cl...

Figure 6.30 Section of an eastern oyster showing an artery surrounded by ves...

Figure 6.31 Section of a juvenile hard clam. a, lumen of the auricular (gree...

Figure 6.32 Section of a hard clam showing kidney tubules surrounded by sinu...

Figure 6.33 Section of a hard clam's gonadal tissue. a, developing oocyte; b...

Figure 6.34 Section of a hard clam's pedal ganglia.

Figure 6.35 Diagram of a scallop mantle eye.

Figure 6.36 Mantle eyes of a bay scallop (

Argopecten irradians

).

Figure 6.37 Section of a hard clam's foot. a, sinusoids of the foot; b, muco...

Figure 6.38 Section of the foot of a juvenile hard clam. a, byssal gland; b,...

Chapter 7

Figure 7.1

Lumbricus terrestris

in anesthetized state. A, anus; CL, clitellu...

Figure 7.2 (a) One dorsal cut is enough to reveal the internal organs of the...

Figure 7.3 Dissected

Lumbricus

in water cover displaying the internal organs...

Figure 7.4 Structure of body wall layers in different regions of

Lumbricus

. ...

Figure 7.5 Morphology of epidermal cell types of

Lumbricus

after formalin an...

Figure 7.6 Setae and setal sac of

Lumbricus

. (a) Setal arrangement: closely ...

Figure 7.7 Organization of clitellar epidermis of

Lumbricus

. (a) Morphologic...

Figure 7.8 Foregut with chromophil cells in parasagittal section (Bouin fixa...

Figure 7.9 Histology of oral cavity. Oral cavity and its wall (Bouin fixatio...

Figure 7.10 Histology of pharynx. (a) Pharyngeal wall and chromophil cells (...

Figure 7.11 Morphology and position of esophagus and calciferous glands in s...

Figure 7.12 Esophagus and calciferous gland of segment X. (a) Esophagus and ...

Figure 7.13 Esophagus and calciferous glands of segment XI. (a) Esophagus wi...

Figure 7.14 Esophagus and calciferous glands of segment XII. (a) Esophagus a...

Figure 7.15 Histology of the crop (formalin fixation), 15× and 350× (inset)....

Figure 7.16 Wall of the gizzard (formalin fixation), 40× and 600× (inset). B...

Figure 7.17 Histology of the midgut wall. (a) Midgut and typhlosole in cross...

Figure 7.18 Histology of the hindgut. Formalin fixation, 40× and 550× (inset...

Figure 7.19 Microstructure of the ciliated funnel/nephrostome (drawings afte...

Figure 7.20 Histology of the nephridial duct. (a) Overview of metanephridium...

Figure 7.21 Histomorphology of blood vessels. (a) Ventral blood vessel; 600×...

Figure 7.22 Coelomocytes. (a) Morphology of coelomocytes after Bouin fixatio...

Figure 7.23

Monocystis

infection and encapsulation. (a)

Monocystis

oocytes a...

Figure 7.24 Histology of the cerebral and subesophageal ganglia. (a) Longitu...

Figure 7.25 Some histologic features of the ventral nerve cord. (a) Overview...

Figure 7.26 Overview of male genital organs in segments IX–XI (longitudinal ...

Figure 7.27 Testis and spermatogenic phases. (a) Testis in the lobe of semin...

Figure 7.28 Sperm reservoir of seminal vesicle and vas deferens. (a) Overvie...

Figure 7.29 Seminal receptacle/spermatheca. (a) Seminal receptacle/spermathe...

Chapter 8

Figure 8.1 Gross anatomy. Dorsal (

left

) and ventral (

right

) view of therapho...

Figure 8.2 Gross anatomy. Dorsal (

left

) and ventral (

right

) view of scorpion...

Figure 8.3 Mesolense image of unidentified Pseudoscorpion from Peru.

Figure 8.4 Gross anatomy. Dorsal (

left

) and ventral (

right

) view of ascarid ...

Figure 8.5 Dissection of a theraphosid spider (

Therophosa stirmi

). The proso...

Figure 8.6 Dissection of a theraphosid spider (

Therophosa stirmi

) using a di...

Figure 8.7 Dissection of a scorpion (

Parabuthus transvaalicus

). The prosomal...

Figure 8.8 Transverse section of wolf spider cuticle showing a simple epider...

Figure 8.9 Transverse section of cuticle of an unspecified arachnid species....

Figure 8.10 Prosoma of a birdeater tarantula (

Theraphosa blondi

). The endost...

Figure 8.11 Endosternite of a birdeater tarantula (

Theraphosa blondi

). Cells...

Figure 8.12 Cheliceral muscle in an unspecified wolf spider species. Longitu...

Figure 8.13 Skeletal muscle of a birdeater tarantula (

Theraphosa blondi

). Sk...

Figure 8.14 Oral cavity setae of a zebra tarantula (

Aphonopelma seemanni

). S...

Figure 8.15 Esophagus of Asian jungle scorpion (

Heterometrus

sp.). The esoph...

Figure 8.16 Sucking stomach of an Arizona blond tarantula (

Aphonopelma chalc

...

Figure 8.17 Midgut tube of a birdeater tarantula (

Theraphosa blondi

). The mi...

Figure 8.18 Longitudinal section of an unspecified arachnid limb demonstrati...

Figure 8.19 Midgut diverticula of birdeater tarantula (

Theraphosa blondi

). T...

Figure 8.20 Stercoral sac of a birdeater tarantula (

Theraphosa blondi

). The ...

Figure 8.21 Stercoral sac of a birdeater tarantula (

Theraphosa blondi

). The ...

Figure 8.22 Stercoral sac of a birdeater tarantula (

Theraphosa blondi

). The ...

Figure 8.23 Nephrocytes of a vinegaroon (order Thelyphonida). Nephrocytes ar...

Figure 8.24 Malphighian tubule of a zebra tarantula (

Aphonopelma seemanni

). ...

Figure 8.25 Coxae (CX) of two walking legs of a Trinidad chevron tarantula (

Figure 8.26 Coxal gland of a birdeater tarantula (

Theraphosa blondi

). The co...

Figure 8.27 Coxal gland of a birdeater tarantula (

Theraphosa blondi

). The co...

Figure 8.28 Coxa of a Trinidad chevron tarantula (

Psalmopoeus cambridgei

). T...

Figure 8.29 Heart of a vinegaroon (order Thelyphonida). Thin ligaments (

arro

...

Figure 8.30 Melanin and hemocytes in the midgut diverticula of a zebra taran...

Figure 8.31 Book lungs of an unspecified arachnid. The book lungs (BL) are i...

Figure 8.32 Book lungs of a vinegaroon (order Thelyphonida). Thin chitinous ...

Figure 8.33 Limb of an unspecified arachnid. The tracheoles occur singly (

ar

...

Figure 8.34 Brain of Asian jungle scorpion (

Heterometrus

sp.). The supra‐ (a...

Figure 8.35 Spermatheca of a zebra tarantula (

Aphonopelma seemanni

). An inva...

Figure 8.36 Ovary of a zebra tarantula (

Aphonopelma seemanni

). Ovarian folli...

Figure 8.37 Uterus and fetuses of Asian jungle scorpion (

Heterometrus

sp.). ...

Figure 8.38 Testes of a birdeater tarantula (

Theraphosa blondi

). Multiple lo...

Figure 8.39 Cross‐section of an opisthosoma of a birdeater tarantula (

Therap

...

Figure 8.40 Longitudinal section of an eye of an orb‐weaver spider (family A...

Figure 8.41 Basal segment of fang within an unspecified arachnid species. 20...

Figure 8.42 Venom gland of a birdeater tarantula (

Theraphosa blondi

). Subjac...

Figure 8.43 Silk glands of a Trinidad chevron tarantula (

Psalmopoeus cambrid

...

Chapter 9

Figure 9.1 External photograph of the dorsum of a horseshoe crab,

Limulus po

...

Figure 9.2 External photograph of the ventrum of

L. polyphemus

. Walking and ...

Figure 9.3 External photograph of the ventrum of

L. polyphemus

. A gill operc...

Figure 9.4 Illustrated sagittal section of

L. polyphemus

. am, arthrodial mem...

Figure 9.5 Illustrated ventral dissection approach of

L. polyphemus

. The ven...

Figure 9.6 Body wall of

L. polyphemus

. The cuticle is organized into a poorl...

Figure 9.7 Arthrodial membrane of

L. polyphemus

. The cuticle is focally thin...

Figure 9.8 Epidermis of

L. polyphemus

. The epidermis is a simple layer of ce...

Figure 9.9 Cartilage of

L. polyphemus

. Numerous lacunae are surrounded by a ...

Figure 9.10 Hepatopancreas of

L. polyphemus

. The majority of the organ is co...

Figure 9.11 Collecting tubules of

L. polyphemus

. Light and dark cells are ar...

Figure 9.12 Interstitial cells of

L. polyphemus

. Large, polygonal, interstit...

Figure 9.13 Ventriculus of

L. polyphemus

. The mucosa is overlain by a thick ...

Figure 9.14 Ventriculus of

L. polyphemus

. Higher magnification of the ventri...

Figure 9.15 Intestine of

L. polyphemus

. A simple mucosa is subtended by rese...

Figure 9.16 Coxal gland of

L. polyphemus

. Zones of the coxal gland are pictu...

Figure 9.17 Coxal gland of

L. polyphemus

. Zones 2–3 of the coxal gland show ...

Figure 9.18 Heart of

L. polyphemus

. The heart has an outer epicardium and an...

Figure 9.19 Heart of

L. polyphemus

. Higher magnification of the heart shows ...

Figure 9.20 Artery and nerve of

L. polyphemus

. Nerve is contained in an arte...

Figure 9.21 Vascular sinus of

L. polyphemus

. Hemolymph is composed of eosino...

Figure 9.22 Inflammatory coagulum of

L. polyphemus

. A central coagulum (

aste

...

Figure 9.23 Book gills of

L. polyphemus

. The bases of the gills have large m...

Figure 9.24 Book gills of

L. polyphemus

. The gills are composed of two paral...

Figure 9.25 Corpora pedunculata of

L. polyphemus

. Numerous bands of Kenyon c...

Figure 9.26 Brain of

L. polyphemus

. Large neurons occur in clusters and are ...

Figure 9.27 Hepatopancreas and ovary of

L. polyphemus

. Ovarian tissue is adm...

Figure 9.28 Ovary of

L. polyphemus

. Large oocytes, surrounded by a thin memb...

Figure 9.29 Testis of

L. polyphemus

. Multiple seminiferous tubules are pictu...

Figure 9.30 Compound eye of

L. polyphemus

. The cuticle is bowed outward and ...

Figure 9.31 Compound eye of

L. polyphemus

. Four ommatidia are pictured, each...

Figure 9.32 Sensilla of

L. polyphemus

. The cuticle is focally expanded by se...

Chapter 10

Figure 10.1 Schematic of a transverse section of a diplopod with the segment...

Figure 10.2 Schematic of a longitudinal section of a myriapod. Br, brain; FG...

Figure 10.3 Transverse histologic section of a giant African millipede (

Arch

...

Figure 10.4 Detail of the layers of the cuticle of a giant African millipede...

Figure 10.5 Body wall of a giant desert centipede (

Scolopendra heros

). The e...

Figure 10.6 The arthrodial membrane between two tergites in a giant African ...

Figure 10.7 The parietal fat body in a giant African millipede (

Archispirost

...

Figure 10.8 Longitudinal and cross‐sections of skeletal muscle fibers (SM) i...

Figure 10.9 Head of a giant African millipede (

Archispirostreptus gigas

) sho...

Figure 10.10 Venom gland of giant desert centipede (

Scolopendra heros

). The ...

Figure 10.11 Longitudinal section through the foregut (esophagus) of a giant...

Figure 10.12 Cross‐section of the midgut of a giant African millipede (

Archi

...

Figure 10.13 Cross‐section of the hindgut of a giant African millipede (

Arch

...

Figure 10.14 The perivisceral fat body (PvF) of a giant African millipede (

A

...

Figure 10.15 Cross‐sections of Malpighian tubules in a giant African millipe...

Figure 10.16 Heart of a giant African millipede (

Archispirostreptus gigas

). ...

Figure 10.17 Hemolymphatic vessel of a giant African millipede (

Archispirost

...

Figure 10.18 Spiracles of a giant African millipede (

Archispirostreptus giga

...

Figure 10.19 Longitudinal section through the cerebral ganglion (brain) of a...

Figure 10.20 The ventral nerve cord of a giant African millipede (

Archispiro

...

Figure 10.21 Testes of a giant desert centipede (

Scolopendra heros

). The tes...

Figure 10.22 Ovary and oviduct of a giant African millipede (

Archispirostrep

...

Figure 10.23 Ovary of a giant African millipede (

Archispirostreptus gigas

). ...

Figure 10.24 Corneal lenses of a giant African millipede (

Archispirostreptus

...

Figure 10.25 Eye of a giant desert centipede (

Scolopendra heros

). The cornea...

Chapter 11

Figure 11.1 Dorsal view of

H. americanus

. (a) Telson; (b) uropods; (c) somit...

Figure 11.2 Ventral view of the mouth parts of

H. americanus

. Maxillipeds we...

Figure 11.3 Second right pereiopod of

H. americanus

. (a) Epipodite; (b) gill...

Figure 11.4 Ventral surface of a male

H. americanus

. (a) Gonadopore; (b) mod...

Figure 11.5 Ventral surface of a female

H. americanus

. (a) Gonadopore; (b) m...

Figure 11.6 Sagittal cut through the seminal receptacle. (a) Receptacle pouc...

Figure 11.7 Lateral view of trichobranchiate gills of a

H. americanus

with b...

Figure 11.8 Line drawings of three types of decopod crustacean gills.(a)...

Figure 11.9 Ventral surface of

Hemigrapsus sanguineus

(Japanese shore crab)....

Figure 11.10 Ventral surface of hard carapace showing chromatophores (

arrows

Figure 11.11 Dissection of male

H. americanus

. (a) Cardiac stomach; (b) pylo...

Figure 11.12 Dissection of a female

H. americanus

. (a) Hepatopancreas; (b) o...

Figure 11.13 Dorsally opened stomach of

H. americanus

. (a) Uncalcified gastr...

Figure 11.14 Dissection of a female

H. americanus

. Dorsally opened stomach. ...

Figure 11.15 Line drawing of the dissected neural cord of

H. americanus

....

Figure 11.16 Junction of the hard and soft carapace of

H. americanus

. (a) Ar...

Figure 11.17 Histology of the carapace of a lobster in C

4

/D

o

phase of develo...

Figure 11.18 Histology of a tegmental gland of

H. americanus

. (a) Pyramidal ...

Figure 11.19 Histology of the stomach of a lobster. (a) Esophagus; (b) cardi...

Figure 11.20 Histology of the plyloric stomach of a crayfish. (a) Dorsal cha...

Figure 11.21 Histology of the hepatopancreas of a crayfish. Blind‐ended dige...

Figure 11.22 Histology of the midgut of

H. americanus

. (a) Circular muscle; ...

Figure 11.23 Histology of the hindgut of

H. americanus

. (a) Chitin lining th...

Figure 11.24 Histology of the blind end of a hepatopancreatic tubule. E–E ce...

Figure 11.25 Histology of a proximal part of a hepatopancreatic tubule of a ...

Figure 11.26 Histology of the heart of a crayfish. (a) Adventitia; (b) stria...

Figure 11.27 Histology of a medium‐sized artery. (a) Internal intima (basal ...

Figure 11.28 Hemocyte types adjacent to the internal intima of the dorsal po...

Figure 11.29 Encapsulation (decapod granuloma). (a) Central melanization (up...

Figure 11.30 Histology of a focus of hematopoietic tissue surrounding a smal...

Figure 11.31 Histology of the kidney (green gland) of a crayfish. (a) Coelom...

Figure 11.32 Line drawing of the hemolymph circulation in an outer filament ...

Figure 11.33 Histology of a cross‐section of an outer gill filament of a cra...

Figure 11.34 Line drawing of the filaments of

Janus novaehollandiae

(rock lo...

Figure 11.35 Histology of a cross‐section of a middle gill filament of

H. am

...

Figure 11.36 Line drawing of a phyllobranchiate gill lamellae of a crab (

Car

...

Figure 11.37 Photomicrograph of a thoracic ganglia. (a) Ganglia; (b) nerve; ...

Figure 11.38 Histology of a large nerve. (a) Small arteriole; (b) neural she...

Figure 11.39 Histology of the ovary of a crayfish. (a) Connective tissue cap...

Figure 11.40 Histology of the seminal receptacle of

H. americanus

. (a) Chiti...

Figure 11.41 Testis of

H. americanus

. (a) Spermatogonia; (b) spermatids; (c)...

Figure 11.42 Vas deferens containing a developing spermatophore. (a) Spermat...

Figure 11.43 Line drawing of decapod ommatidia.(a) Cornea; (b) corneagen...

Figure 11.44 Histology of a sagittal section through the eye of a crayfish i...

Figure 11.45 Line drawing of the rhabdom of the compound eye of crayfish (

Pr

...

Figure 11.46 Histology of a sagittal section through the eye and eyestalk of...

Figure 11.47 Histology of the polygonal cells of the mandibular organ (arrow...

Chapter 12

Figure 12.1 Sagittal section of a honeybee (

Apis mellifera

). Scale bar, 1.0 ...

Figures 12.2–12.4 Transverse sections of the head of a honeybee (

Apis mellif

...

Figure 12.5 Body wall of a gypsy moth caterpillar (

Lymantria dispar

). The bo...

Figure 12.6 Sagittal section of a gypsy moth caterpillar (

Lymantria dispar

)....

Figure 12.7 Body wall of a gypsy moth caterpillar (

Lymantria dispar

). Higher...

Figure 12.8 Body wall of a gypsy moth caterpillar (

Lymantria dispar

) with ep...

Figure 12.9 Skeletal muscle of a honeybee (

Apis mellifera

). The skeletal mus...

Figure 12.10 Forewing of a beetle (unspecified species). The wing is compose...

Figure 12.11 Transverse section of the head of a honeybee (

Apis mellifera

) a...

Figure 12.12 Hypopharyngeal glands in a worker honeybee (

Apis mellifera

). 40...

Figure 12.13 Junction of the crop (C), proventriculus (P), and ventriculus (...

Figure 12.14 Proventriculus of a honeybee (

Apis mellifera

). The cuticular li...

Figure 12.15 Foregut of a gypsy moth caterpillar (

Lymantria dispar

). The lum...

Figure 12.16 Midgut (ventriculus) of a gypsy moth caterpillar (

Lymantria dis

...

Figure 12.17 Ventricular mucosa of a honeybee (

Apis mellifera

). EC, epitheli...

Figure 12.18 Hindgut of a gypsy moth caterpillar (

Lymantria dispar

). The lum...

Figure 12.19 Fat body of a gypsy moth caterpillar (

Lymantria dispar

). The fa...

Figure 12.20 Malpighian tubules of a honeybee (

Apis mellifera

). The tubules ...

Figure 12.21 Heart of a honeybee (

Apis mellifera

). The heart is a tubular or...

Figure 12.22 Tracheae (T) in the superficial aspect of the hemoceolom of a g...

Figure 12.23 Tracheoles in a gypsy moth caterpillar (

Lymantria dispar

). Trac...

Figure 12.24 Transverse section of a ventral nerve cord of a gypsy moth cate...

Figure 12.25 Longitudinal section of a ventral nerve cord of a gypsy moth ca...

Figure 12.26 Section through the head of a carpenter ant (

Camponotus

sp.). T...

Figure 12.27 Longitudinal section of the abdomen of a mated honeybee queen. ...

Figure 12.28 Photomicrograph of a coronal section of a newly emerged honeybe...

Figure 12.29 Transverse section of the head of a honeybee (

Apis mellifera

). ...

Figure 12.30 Section through an ocellus in a honeybee (

Apis mellifera

). The ...

Figure 12.31 Section through a compound eye in a honeybee (

Apis mellifera

). ...

Figure 12.32 Sensilla of a honeybee (

Apis mellifera

). 2×. HE. Inset, 30×.

Figure 12.33 Venom sac of a honeybee (

Apis mellifera

). The venom sac is line...

Guide

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Invertebrate Histology

 

Edited by

Elise E.B. LaDouceur, DVM, DACVP

Chief, Extramural Projects and Research

Joint Pathology Center

Silver Spring, MD, USA

 

 

 

 

 

 

 

 

 

 

 

 

This edition first published 2021© 2021 John Wiley & Sons, Inc.

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Library of Congress Cataloging-in-Publication Data

Names: LaDouceur, Elise E., editor.Title: Invertebrate histology / edited by Elise E LaDouceur.Description: Hoboken, NJ : Wiley-Blackwell, 2021. | Includes bibliographical references and index.Identifiers: LCCN 2020024338 (print) | LCCN 2020024339 (ebook) | ISBN 9781119507659 (cloth) | ISBN 9781119507666 (adobe pdf) | ISBN 9781119507604 (epub)Subjects: MESH: Invertebrates–anatomy & histologyClassification: LCC QL363 (print) | LCC QL363 (ebook) | NLM QL 363 | DDC 592–dc23LC record available at https://lccn.loc.gov/2020024338LC ebook record available at https://lccn.loc.gov/2020024339

Cover Design: WileyCover Image: © (cephalopod) Francesco Martini, (histology of an eye) Damien Laudier

Thank you to my mentors who nurtured my passion for invertebrates, especially Michael Garner, Kevin Keel, and Patricia Pesavento, as well as the entire veterinary anatomic pathology department at University of California, Davis.

This book is dedicated to my husband and go-to consultant for all things pathology and life, Andrew Cartoceti.

List of Contributors

Ilze K. BerzinsOne Water, One Health, LLC, Golden Valley, MN, USA

Alvin C. CamusUniversity of Georgia College of Veterinary Medicine, Athens, GA, USA

John E. CooperWildlife Health Services, UK

Michelle M. DennisCenter for Conservation Medicine and Ecosystem Health Department of Biomedical Sciences Ross University School of Veterinary Medicine Basseterre, St Kitts and Nevis Department of Biomedical and Diagnostic Services University of Tennessee College of Veterinary Medicine Knoxville, TN, USA

Jennifer A. Dill-OkuboFlorida Department of Agriculture and Consumer Services, Kissimmee, FL, USA

Alexander EreskovskyInstitut Méditerranéen de Biodiversité et d’Ecologie Marine et Continentale (IMBE), Aix Marseille University, CNRS, IRD, Avignon University, Marseille, France Department of Embryology, Faculty of Biology, Saint-Petersburg State University, Saint-Petersburg, Russia Koltzov Institute of Developmental Biology, Russian Academy of Sciences, Moscow, Russia

Michael M. GarnerNorthwest ZooPath, Monroe, WA, USA

Benjamin KennedyVeterinary Invertebrate Society, Venomtech Ltd, Discovery Park, Sandwich, Kent, UK

György KriskaInstitute of Biology Eötvös Loránd University MTA Centre for Ecological Research, Danube Research Institute, Budapest, Hungary

Elise E.B. LaDouceurJoint Pathology Center, Silver Spring, MD, USA

Damien LaudierLaudier Histology, New York, NY, USA

Andrey LavrovDepartment of Embryology, Faculty of Biology, Saint-Petersburg State University, Saint-Petersburg Pertsov White Sea Biological Station, Biological Faculty, Lomonosov Moscow State University, Moscow, Russia

Péter LőwDepartment of Anatomy, Cell and Developmental Biology Eötvös Loránd University Budapest, Hungary

Kinga MolnárDepartment of Anatomy, Cell and Developmental Biology, Eötvös Loránd University, Budapest, Hungary

Alisa L. NewtonWildlife Conservation Society, Bronx, NY, USA Disney’s Animals, Science and Environment Orlando, FL, USA

Esther C. PetersEnvironmental Science and Policy, George Mason University, Fairfax, VA, USA

Katie J. RoordaJohns Hopkins University, Baltimore, MD, USA

Elemir SimkoWestern College of Veterinary Medicine, University of Saskatchewan, Saskatoon, Canada

Roxanna SmolowitzAquatic Diagnostic Laboratory Roger Williams University Bristol, RI, USA

Steven A. TrimVenomtech Ltd Discovery Park Sandwich, Kent, UK

Sarah C. WoodWestern College of Veterinary Medicine, University of Saskatchewan, Saskatoon, Canada

Roy P.E. YanongTropical Aquaculture Laboratory Fisheries and Aquatic Sciences Program School of Forest Resources and Conservation Institute of Food and Agricultural Sciences University of Florida, Ruskin, FL, USA

Foreword

Veterinary medicine is a dynamic profession that began over 250 years ago to heal and protect working and warring equids along with livestock for food and other human‐use products. The profession has come a long way since the 1700s, most notably in the breadth of species embraced, and the information that exists and is being explored related to this taxonomic diversity. Increasing human population growth, commerce, technology, and animal welfare are all contributing to this expansion. Our profession is more diverse than ever, and a growing part of that diversity is the inclusion of over 97% of the animal kingdom: the invertebrates.

Dr LaDouceur and her internationally recognized contributors have assembled an organized, easy to navigate, comprehensive, and richly illustrated work focused on the microanatomy and histology of the invertebrates. It is certainly the only book of its kind on the market and one that is long overdue. The text is richly illustrated with beautiful images, drawings, and micrographs, detailing the normal gross and microscopic anatomy of the species covered. Chapters also describe how to properly and efficiently process invertebrate tissues for histology. This is critically important as standard vertebrate tissue‐processing methods frequently do not apply to invertebrates. Anatomic features like chitinous shells, glass spicules, calcium carbonate skeletons, and mesoglea, to name a few, may require specialized fixatives, processing, and staining techniques.

One of the biggest challenges for a clinician or pathologist is being able to recognize and become familiar with what is normal about an animal. This challenge is especially pertinent when dealing with nondomestic species. There is no greater or more diverse animal classification than the invertebrates, estimated to include over 1.3 million described species (and it's likely that the global total could be 10 times this number), representing at least 40 phyla. The editor and authors have wisely focused on the taxa that are the most economically important and/or in need of conservation, protection, and veterinary support. This includes species commonly displayed in zoos and aquaria, taxa that are utilized in the laboratory for research, and animals that are kept as pets.

This detailed and thorough text is a windfall for our profession and anyone working on the health and welfare of these animals. Pathologists, veterinary clinicians, histology technicians, invertebrate zoologists, and students studying in these areas will all find this book highly useful and important for their work. The timing for this book could not be better. I'm sure you, the reader, will agree with me, and find this one of the most important references on your bookshelf, in your laboratory, or digitally on your computer.

Gregory A. Lewbart

Raleigh, NC, USA

1 Echinodermata

Alisa L. Newton1,2 and Michelle M. Dennis3,4

1 Wildlife Conservation Society, Bronx, NY, USA

2 Disney’s Animals, Science and Environment, Orlando, FL, USA

3 Center for Conservation Medicine and Ecosystem Health, Department of Biomedical Sciences, Ross University School of Veterinary Medicine, Basseterre, St Kitts and Nevis

4 Department of Biomedical and Diagnostic Sciences, University of Tennessee College of Veterinary Medicine, Knoxville, TN, USA

1.1 Introduction

Phylum Echinodermata consists of three subphyla (Asterozoa, Echinozoa, and Crinozoa) and five main classes. Subphylum Asterozoa contains two extant classes: Asteroidea (sea stars, sea daisies) and Ophiuroidea (brittle and basket stars). Echinozoa contains two extant classes: Echinoidea (sea urchins, sand dollars) and Holothuroidea (sea cucumbers). Subphylum Crinozoa contains only one extant class: Crinoidea (feather stars, sea lilies). There are 7000 living species of echinoderms (Mulcrone 2005). All are marine and almost exclusively benthic. Some subphyla are mobile (Asterozoa, Echinozoa) and others are sessile (Crinozoa), though some sea lilies have been documented to swim significant distances. Echinoderms do not appear to have near relatives among other invertebrate phyla.

Most members of Echinodermata are dioecious and undergo sexual reproduction, with a few species reproducing asexually. Holothuroids are gonochoric (Leake 1975). Asexual reproduction through fragmentation may occur in some Asteroidea and Holothuroidea due to trauma or predation. The diet varies widely by class, with Asterozoa being carnivorous, Echinozoa and Crinozoa being vegetarian browsers and filter feeders, and Holothuroidea being detritivores. Significant conservation concerns and anthropogenic stressors include commercial fisheries, which impact diet availability, particularly clams, mussels, and oysters, and the pet trade through individual animal collection and the collection of coral and live rock causing habitat loss. Environmental concerns include habitat destruction and direct animal impacts due to ocean acidification. Population declines due to disease such as the Caribbean Diadema antillarum mortality event in 1983–1984 (Carpenter 1990; Lessios 2016) and “wasting disease” events across multiple species of asteroid (Hewson et al. 2014; Menge et al. 2016) have more recently received significant focus. Certain Asteroidea are keystone species in their ecosystems, critical for controlling prey populations and diversity. Echinoidea and Holothuroidea are of paramount importance to marine ecosystems because of respective roles in counteracting macroalgal competition with corals, and recycling nutrients from decaying organic matter.

1.2 Gross Anatomy

Uniting features of all echinoderms include radial symmetry (pentamerous symmetry), a tricoelomate body cavity, and a body wall composed of calcite endoskeletal plates (dermal ossicles) connected by “mutable collagenous tissue.” Most internal features, including the alimentary system, reproductive system, nervous system, respiratory system, and a unique water vascular system, share similar basic plans between the subphyla. The basic echinoderm body plan has 10 divisions: five radii (rays or arms) which alternate with five interradii (interrays). Typically, there is an oral surface with a central mouth and an aboral surface that contains the anus. Despite these commonalities, morphology does vary widely and thus representative examples of each subphylum are discussed separately.

The asteroid (sea star) body plan consists of a central disc with typically five but in some species (sun stars) up to 40 or more individual rays. Rays are broad based and arise from the lateral margins of the disc. They taper distally and each ray terminates in one or more tentacle‐like sensory tube feet and a red eyespot. The aboral surface is dorsal and contains the anus at the center of the central disc, which may not be grossly apparent. The madreporite, bearing openings of the water vascular system, is on one side of the disc near the interradius of the first and second rays (Figure 1.1a). The oral surface is ventral and in contact with the substrate. Originating at the mouth and extending the length of each ray is a prominent groove, the ambulacrum (ambulacral groove). Two to four rows of tube feet (podia) lie within the ambulacral groove (Figure 1.1b). The margins are lined by moveable spines that can close over the top of the groove. Ophiurids (brittle and basket stars) demonstrate similar morphology. They typically have five rays, but these are distinctly offset from a round to pentagonal central disc. The rays are typically very long, slender, and very flexible. In basket stars the rays are highly branched. The disc has a proportionally smaller diameter compared to most sea stars. Ophiurid rays lack an ambulacral groove and the tube feet lack distal suckers as they are not typically used for movement.

Figure 1.1 Representative image of the aboral (a) and oral (b) surface of a chocolate chip sea star (Protoreaster nodosus) demonstrating pentamerous symmetry. Labels include (A) radius, (B) interradius, (C) mouth, (D) ambulacral groove, (E) anus, and (F) madreporite.

Figure 1.2 Representative image of the aboral (a) and oral (b) surface of a purple urchin (Arabacia punctulata) demonstrating pentamerous symmetry. Labels include (A) ambulacral plates, (B) interambulacral plates, (C) mouth, (D) anus, and (E) madreporite.