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Published by
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Email: editorial@panstanford.com
Web: www.panstanford.com
British Library Cataloguing-in-Publication Data

A catalogue record for this book is available from the British Library.
VIRAl NAnOpARTIclES: TOOlS FOR MATERIAlS ScIEncE And BIOmEdIcInE

All rights reserved. This book, or parts thereof, may not be reproduced in any form
or by any means, electronic or mechanical, including photocopying, recording or
any information storage and retrieval system now known or to be invented, without
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Printed in Singapore

Contents
Preface vii
Chapter 1 An Introduction to VNPs and Nanotechnology 1
Chapter 2 Overview of the Manifold VNPs Used in

Nanotechnology 13
Chapter 3 Production of VNPs, VLPs, and Chimeras 49
Chapter 4 The Art of Bioconjugation: Functionalization

of VNPs 75
Chapter 5 Encapsulating Materials within VNPs 123
Chapter 6 VNPs as Templates for Materials Synthesis 145
Chapter 7 Playing “Nano-Lego”: VNPs as Building Blocks for the

Construction of Multi-Dimensional Arrays 171
Chapter 8 VNPs as Tools for Nanomedicine 207
Chapter 9 Summary and Outlook 247
Appendix A 249
Appendix B 250
Appendix C 251
Appendix D 255
Appendix E 264
Index 267

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Preface
Viral nanotechnology is a young and emerging discipline. A highly interdis-

ciplinary field, viral nanotechnology inhabits the interface between virol-
ogy, chemistry, and materials science. The field exploits viral nanoparticles
(VNPs) for potential applications in diverse fields that range from electron-
ics and energy to novel materials and medicine.
From a materials science point of view, VNPs are attractive building
blocks for several reasons. In general, VNPs are on the nanometer-
size scale; they are monodisperse with high degree of symmetry and
polyvalency; they can be produced with ease on large scale; they are
exceptionally stable and robust; and they are biocompatible, and in
some cases, orally bioavailable. VNPs are “programmable” units that can
be modified by either genetic modification or chemical bioconjugation
methods.
This book will give a survey of the applications of VNPs in fields
ranging from materials science to biomedicine. An introduction to the
field of viral nanotechnology is given in Chapter 1. Chapter 2 provides an
overview of the many different VNP building blocks currently in use for
viral nanotechnology. Various methods have been established that allow
efficient production of VNPs as well as their non-infectious counterparts,
the virus-like particles (VLPs), as discussed in Chapter 3. A large variety
of bioconjugation methods have been applied and optimized for VNPs
that facilitate chemical modification, fine-tuning, and immobilization of
VNPs; these techniques are described in Chapter 4. Chapter 5 summarizes
strategies to entrap or encapsulate non-natural cargos into VNPs and VLPs.
Mineralization chemistries are discussed in Chapter 6. With the different
chemistries in hand, a range of highly interesting VNP-based materials
has been fabricated with potential applications in sensors, electronics,
and medicine. Toward the development of nanoelectronic devices,
Chapter 7 describes the efforts that have been directed to the generation
of thin-film arrays of VNPs immobilized on solid supports. Last but
not least, Chapter 8 provides an overview of the advances of designing
VNPs for biomedical applications, including their uses as vaccines,
imaging modalities, targeted therapeutics, and gene therapies.
As this young discipline matures, a new era has begun in which
pathogens have become useful material building blocks for next-generation

viii Preface
nanodevices. The field has even greater promise given the enormous variety
of available VNPs with their vast diversity of sizes, structures, chemical
reactivities, and biocompatibilities. This promise drives an inspiring field
with wide-ranging opportunities for discovery.
La Jolla, CA, December 2009 Nicole F. Steinmetz, PhD
Marianne Manchester, PhD

Chapter 1
AN INTRODUcTiON TO VNPs aND

NaNOTEcHNOLOGY
Viruses have long been studied as pathogens, with the goal of understanding
viral infection and disease. More recently viruses have begun to be regarded
as building blocks and tools for nanotechnology. Viruses are exploited as
platforms; that is, they are used as templates or scaffolds for the design of
novel nanomaterials. A wide variety of viral platforms have been studied and
utilized for applications ranging from materials to medicine. This chapter
will provide an introduction to the role of viral nanoparticles (VNPs) in
nanotechnology.
1.1 WHAT IS NANO?
Nanotechnology is a highly interdisciplinary field that brings together

researchers from different scientific backgrounds and has created a novel
common language. It is a collective term for a broad range of novel topics
concerned with matter on the nanometer scale. Nanotechnology sits at the
interface of biology, chemistry, physics, material science, and medicine.
Nanotechnology is found and applied in nearly every scientific area.
Nano is a somewhat fashionable term; in common speech, it is used as
a prefix to denote something that is smaller than usual. As of this writing,
the term “Apple iPod nano” is the first hit on a Google search of the term
“nano.” The word nano is derived from the ancient Greek word for dwarf.
In a scientific context the prefix nano is used to describe “a billionth of
something.” A nanometer is a billionth of a meter (10−9 m = 1 nm), and a
nanosecond is a billionth of a second (10−9 s = 1 ns).
Figure 1.1 illustrates how tiny nano is. An aphid insect (Fig. 1.1,
panel A) is about 1 mm in size. The aphid is about 1,000,000 times bigger
than a nanometer. Human hairs (Fig. 1.1, panel B) have an average diameter
Viral Nanoparticles: Tools for Materials Science and Biomedicine

By Nicole F. Steinmetz and Marianne Manchester
www.panstanford.com

2 An Introduction to VNPS and Nanotechnology
of 100 µm and are thus 10 times smaller than the aphid and still 100,000
times bigger than a nanoparticle. Panel C shows a plant cell, which is about
10 µm in size and thus 10,000 bigger than a nanometer. Particles formed by
the plant virus Cowpea mosaic virus (CPMV; Fig. 1.1, panel D) are about 30
nm in diameter and thus nanoparticles. One would need around 50,000,000
VNPs to fill up the interior of a cell.
A single atom is a fraction of a nanometer in size; molecules, including
biological molecules, are typically nanometers in size and can thus be
regarded as nanoobjects. In recent years a range of biological molecules
have been exploited for nanosciences and nanotechnology. Nucleic acids,
for example, are used as construction materials to generate highly ordered
2D and 3D structures and assemblies such as nanotubes and nanocages.
A main theme in nanotechnology is controlled self-assembly, with the goal
being to generate functional materials with a high degree of precision.
Nanotechnology, then, requires chemical and physical control at the molecular
level. Nucleic acids, proteins, and viruses are essentially naturally occurring
nanomaterials capable of self-assembly with a high degree of precision.
This property, coupled to the relative ease of experimentally controlling
and producing biological nanomaterials, has led to tremendous interest in
their nanotechnology applications. Viruses, and VNPs in particular, possess a
number of traits that make them exceptionally outstanding candidates.
Figure 1.1 Scanning electron micrographs of aphids on leaf (A), human hair (B),
fractured plant cell (C), and Cowpea mosaic virus particles (D). Panel A–C provided by
courtesy of Kim C. Findlay, John Innes Centre, Norwich (UK). Panel D from Steinmetz,
N. F., et al., unpublished.

Where Did It All Begin? A History of VNPs 3
1.2 WHERE DID IT ALL BEGIN? A HISTORY OF VNPs: FROM

PATHOGENS TO BUILDING BLOCKS
The word virus is Latin and means “poison.” Viruses are infectious agents,
and generally pathogens. It was not, however, until the end of the 19th
century that viruses were discovered as infectious agents. The first virus
to be recognized as an infectious agent distinct from bacteria was the plant
pathogen Tobacco mosaic virus (TMV) (Zaitlin, 1898). Today more than
5,000 viruses have been discovered and described, although this likely
represents a fraction of those found in nature. Viruses cause many human
diseases, from the common cold and chicken pox to more serious infections
such as AIDS (acquired immune deficiency syndrome, which is caused by the
Human immunodeficiency virus [HIV]) and SARS (severe acute respiratory
syndrome, which is caused by SARS coronavirus). Virology — the science of
studying viruses — is thus a highly important discipline in regard to human
health.
Viruses infect all forms of life. Generally, animal viruses infect animals,
including humans; plant viruses infect plants; and bacteriophages infect
bacteria. Archaeal viruses are those that infect Archaea. Archaea show
similarities with bacteria as well as with eukaryotes, and although they are
prokaryotes, it has been suggested that they are more closely related to the
eukaryotes (Woese & Fox, 1977).
In their simplest form, viral particles consist of a nucleic acid genome and
a protective protein coat termed the capsid. Some viruses have additional
structural features such as a lipid envelope, or they may consist of separate
head and tail structures (discussed in Chapter 2). In brief, the nucleic acid
genome encodes the genetic information that is needed to produce viral
progeny. In addition to cellular attachment, an important function of the
capsid of non-enveloped viruses is the protection of the nucleic acid genome.
This tends to make non-enveloped viral particles extremely robust. With a few
exceptions, nearly all viruses utilized in nanotechnology are non-enveloped
particles. The envelope for enveloped viruses also plays a role in the initial
stages of the infection process, including binding to surface receptors and
internalization into the host cell.
Viruses have now been studied for more than 100 years, and detailed
knowledge about the structure and function of many viruses has been
gathered. For many years the emphasis has been on the understanding of
viral infection and disease, and it still is. Being able to control or treat viral
infections is an important goal in human medicine (as well as veterinary
medicine and agriculture). Every year novel viruses or virus strains evolve
with the potential to cause disease and death worldwide. For instance, at the
time of writing this book, the Influenza virus strain H1N1 (also referred to as

“swine flu”) has emerged and quickly spread all over the world. The science
of fundamental virology will always play an important role in medicine.
By the 1950s, researchers had begun thinking of viruses as tools in
addition to pathogens. Bacteriophages, for example, played a key role in
the development of molecular biotechnology. Bacteriophage genomes and
components of the protein expression machinery have been widely utilized
as tools for understanding fundamental cellular processes such as nucleic
acid replication, transcription, and translation. Virus genomes are small
and the genetic elements that control expression of the genome are highly
efficient and multifunctional. On the basis of these properties, several viruses
have been exploited as expression systems in biotechnology. Several cloning
vectors are derivatives of bacteriophages, and typical examples include the
Escherichia coli phages l and M13. Various phage-encoded promoters (DNA
sequences that facilitate transcription of DNA into RNA) have been utilized
to regulate gene expression. An overview of tools used for molecular biology
can be found in Molecular Cloning: A Laboratory Manual, by Sambrook and
Russell (2001). The use of viruses as cloning and expression vectors is not
restricted to phages; plant viruses, insect viruses, and mammalian systems
have also been engineered for these purposes. Some of these systems are
discussed in Chapter 3.
Another early application evaluated was bacteriophage therapy, the use
of bacteriophages to combat bacterial infections. With the development of
antibiotics (compounds that kill bacteria), which have proven to be more
efficient and comprehensive compared with bacteriophage therapy, few
efforts were made toward its further development. Other applications include
bacteriophage-mediated microbial control (applied in the food industry) and
phage display technologies that allow screening for biological protein-binding
partners. More recent developments include the use of bacteriophages for
vaccine production and gene delivery approaches. Developments in using
bacteriophages for biotechnological applications were reviewed by Clark and
March (2006) and Marks and Sharp (2000).
In the 1970s many efforts focused on the production of virus-like particles
(VLPs) for use in anti-viral vaccines (reviewed in Garcea & Gissmann, 2004;
Grgacic & Anderson, 2006; Ludwig & Wagner, 2007). A VLP is a particle
consisting of the capsid but lacking the genome. A VLP is the replication-
deficient and thus non-infectious counterpart of a VNP. Chimeric VLPs and
VNPs have also been designed. A chimera is a genetically modified version
of a naturally occurring particle or cell. In vaccine development, chimeras
are used as carriers or platforms for the presentation of antigenic sequences
(sequences that induce an immune response) of other pathogens (reviewed
in Garcea & Gissmann, 2004; Grgacic & Anderson, 2006; Ludwig & Wagner,

Where Did It All Begin? A History of VNPs 5
2007). More details and insights on the use of viruses in vaccine development
are given in Chapter 8.
In the 1980s researchers began exploiting plant viruses as expression
vectors (a DNA-based plasmid that promotes the expression of foreign
genes) to produce pharmaceutical proteins in plants. Advantages of protein
production in plants are the absence of contamination with animal products,
low production costs, and — when using viral expression vectors —
achievement of high expression levels. A range of pharmaceutically relevant
proteins including therapeutic antibodies have been successfully produced
using viral vectors such as TMV, CPMV, and Potato virus X (PVX) (Awram et
al., 2002; Canizares et al., 2005; Johnson et al., 1997; Porta & Lomonossoff,
1998; Scholthof et al., 1996).
Viruses Became VNPs. Beginning about 20 years ago, the focus on
exploiting viruses and their capsids for biotechnology began to shift
toward using them for nanotechnology applications. Douglas and Young
(Montana State University, Bozeman, MT, USA) were the first to consider
the utility of a virus capsid as a nanomaterial (Douglas & Young, 1998).
The virus of interest in their studies was the plant virus Cowpea chlorotic
mottle virus (CCMV). CCMV is a highly dynamic platform with pH- and
metal ion-dependent structural transitions (see Section 2.2.2). Douglas
and Young made use of these capsid dynamics and exchanged the natural
cargo (nucleic acid) with a synthetic material, in this case encapsulating
the organic polymer polyanetholesulfonic acid. Since then many materials
have been encapsulated into CCMV and other VNPs (discussed in detail
in Chapter 5). The system was further engineered to allow not only the
entrapment of materials but also the size-constrained and spatially
controlled synthesis of materials within both the capsid and other protein
cages (discussed in Chapters 5 and 6). A protein cage is a hollow, generally
spherical protein structure that is typically assembled by multiple copies of
protein monomers and thus has similarities to a viral capsid.
At about the same time, the research team led by Mann (University of
Bristol, UK) pioneered a new area using the rod-shaped particles of TMV. The
particles were used as templates for the fabrication of a range of metallized
nanotube structures using mineralization techniques (Shenton et al., 1999).
These techniques have received great attention during recent years. In
particular the contributions of Belcher and colleagues at the Massachusetts
Institute of Technology (MIT, Cambridge, MA, USA) led to the development
of a new technology that allowed for the generation of a large range of
mineralized nanotubes and nanowires for use in batteries and data storage
devices (Lee et al., 2009; Nam et al., 2006; Nam et al., 2008). Mineralization

and metal deposition techniques as well as their potential applications are

discussed in Chapters 6 and 7.
A third direction began a few years later. In 2002, the first study was
reported in which bioconjugation chemistries had been applied to a VNP.
The research teams led by Johnson and Finn (The Scripps Research Institute,
La Jolla, CA, USA) showed, in a proof-of-concept study, that small chemical
modifiers such as organic dyes and nanogold particles could be covalently
attached to the surface of CPMV. Attachment and display was achieved
with atomic precision (Wang et al., 2002). Since then, various chemistries
ranging from standard techniques utilizing commercially available reagents
to complex and advanced reactions have been developed (discussed in
Chapter 4). The establishment of a wide variety of bioconjugation protocols
for VNPs was an important development and can be regarded as fundamental
to viral nanotechnology. Functional molecules such as therapeutic or imaging
molecules for drug delivery and imaging applications, for example, can be
covalently attached and displayed on the VNPs, and this has opened the door
for developing “smart” devices for medical applications.
The field of viral nanotechnology is still a young discipline that is rapidly
evolving. A broad range of VNP platforms have been exploited or show
promise in applications ranging from the development of battery electrodes
to medical imaging and drug delivery.
1.3 WHY VNPs? MATERIALS PROPERTIES OF VNPs
When a virologist looks at a virus, he or she might see a pathogen, an infectious

agent that is causing a disease. What does a chemist or a materials scientist
see in VNPs? Working at the interface of chemistry and medicine, we see
tiny building blocks, platforms that can be tuned with functionalities. A VNP
can be regarded as a platform that is used as a template or scaffold for the
generation of functional materials. The regular surface properties of VNPs
allow one to covalently attach functional molecules (termed functionalizing).
VNPs are programmable and tunable, as they can be functionalized with
a broad range of molecules used for manifold applications. The covalent
modification can also lead to a tuning of the materials properties; for
example, the charge properties can be altered by attaching neutral groups
to charged surface groups on the viral capsid.
VNPs occur in two basic shapes, icosahedral and rod (Fig. 1.3). An
icosahedron is a polyhedron with 20 triangular faces; the icosahedral
symmetry description is explained in detail in Chapter 2. More complex
structures such as head–tail bacteriophages, enveloped viruses, and even
spindle- and bottle-shaped particles can also be found. For example, the
particles of Acidianus bottle-shaped virus (ABV) indeed look like a bottle

Why VNPS? Materials Properties of VNPs 7
(Fig. 1.2). ABV is an archaeal virus of the family Ampulliviridae with particles
about 230 nm long and 4–75 nm wide (Prangishvili et al., 2006).
Figure 1.2 Transmission electron micrograph of Acidianus bottle-shaped virus.

The scale bar represents 100 nm. Reproduced with permission from Prangishvili D.,
Forterre P., and Garrett R. A (2006) Viruses of the Archaea: a unifying view, Nat. Rev.
Microbiol., 4(11), 837–848.
Icosahedral particles range in their size from 18 nm to 500 nm. Rod-

shaped or filamentous VNPs can reach up to 2 µm in length. The structure of
many viruses has been solved to atomic or near-atomic resolution, and many
of these structures and accompanying structural information can be found
at the Virus Particle ExploreR database (VIPER; at www.viperdb.scripps.
edu). Structures of VNPs currently under investigation and use for potential
applications in nanotechnology are shown in Fig. 1.3. We will discuss each
of these systems throughout the book and highlight the characteristics and
materials properties of each VNP in Chapter 2.
A whole library of VNPs has become available, offering a large variety of
building blocks with varying structural and chemical properties. Each VNP
can be selected for its most suitable applications. For example, the reversible
permeability of CCMV has led to its use as a constrained chemical reaction
vessel (Douglas et al., 2002; Douglas & Young, 1998, 1999). Filamentous
or rod-shaped VNPs such as the bacteriophage M13 and TMV have been
metal-coated and used as nanowires and nanotubes in the fabrication of
lithium–ion battery electrodes and data storage devices (Lee et al., 2009;
Nam et al., 2006, 2008; Tseng et al., 2006). The development of VNPs for
nanotechnology applications has fueled the search for novel VNPs possessing

Icosahedral plant viruses An insect virus
BMV CCMV CPMV HCRSV RCNMV TYMV FHV
Icosahedral bacteriophages
MS2 Qb
HK97 P22 T7
Icosahedral mammalian viruses
CPV
Ad
Rod-shaped plant virus and a filamentous phage
m13
tmv
Figure 1.3 A snapshot of the viral nanoparticles (VNPs) that are currently exploited
and developed for materials science and medicine. The list is continuously growing;
these are the VNPs in use as of May 2009. Icosahedral plant viruses: Brome mosaic
virus (BMV), Cowpea chlorotic mottle virus (CCMV), Cowpea mosaic virus (CPMV),
Hibiscus chlorotic ringspot virus (HCRSV), Red clover necrotic mottle virus (RCNMV),
Turnip yellow mosaic virus (TYMV). Icosahedral insect virus: Flock house virus (FHV).
Icosahedral bacteriophages: HK97, P22, T7, MS2, and Qβ. Note P22 and T7 are head-
tail phages. The tail is shown for P22, not for T7. Icosahedral mammalian viruses:
Adenovirus and Canine parvovirus (CPV). Rod-shaped and filamentous viruses: Tobacco
mosaic virus (TMV), a plant virus, and M13, a bacteriophage. Images of the following

A General Introduction to Virus Structure and Replication 21
Figure 2.6 The structure of the viral capsid and the icosahedral asymmetric unit of
Cowpea mosaic virus (CPMV). The capsid is comprised of two viral proteins, the S and
L subunits, which form three β-sandwich domains in the icosahedral asymmetric
unit. These three domains occupy comparable positions shown in a T = 3 quasi-
equivalent surface lattice. These are designated A (domains forming pentamers)
and B and C (domains forming hexamers). Numerals following the letter indicate
positions related by the icosahedral symmetry to A, B, and C. The S subunit forms
the domain occupying the A positions in an icosahedral lattice and these are shown
in blue. The L subunit is formed by two domains, C and B5, shown in green and red,
respectively. (A). A space-filling drawing of the CPMV capsid. All atoms are shown
as spheres of 1.8 Å in diameter. (B). A schematic presentation of the CPMV capsid.
The S subunit occupies A positions around the fivefold axis; the two domains of
L subunit occupy the B and C positions. The quaternary interactions at A/B5 and
C/C2 interfaces are pseudo-equivalent. Selected icosahedral symmetry axes are
also shown. (C). A ribbon diagram of the three β-barrel domains that comprise the
icosahedral asymmetric unit. N-termini of the S and L subunits are in the interior,
and C-termini are in the exterior. All three domains are variants of canonical jelly-
roll β sandwiches. (D). A schematic diagram of the asymmetric unit with icosahedral
symmetry axes. The A domains surround the fivefold axes, and the B and C domains
alternate around the threefold axis. Reproduced with permission from Lin, T.,
Chen, Z., Usha, R., Stauffacher, C. V., Dai, J. B., Schmidt, T., and Johnson, J. E. (1999)
The refined crystal structure of Cowpea mosaic virus at 2.8 Å resolution, Virology,
265(1), 20–34.

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10, more highly magnified pairing of gametes of different origin; 11, 12,
zygote developing into "A" form. (After Schaudinn.)
Several genera have tests of siliceous or chitinous plates, formed in

the cytoplasm in the neighbourhood of the nucleus, and connected
by chitinous cement. Among these Quadrula (Fig. 10, A) is Lobose,
with square plates, Euglypha (Fig. 8, p. 29), and Paulinella[69] are
Filose, with hexagonal plates. In the latter they are in five longitudinal
rows, with a pentagonal oral plate, perforated by the oval pylome. In
other genera again, such as Cyphoderia (Filosa), the plates are
merely chitinous. Again, the shell may be encrusted with sand-grains
derived directly from without, or from ingested particles, as shown in
Centropyxis, Difflugia (Fig. 10, D), Heleopera, and Campascus when
supplied with powdered glass instead of sand. The cement in
Difflugia is a sort of organic mortar, infiltrated with ferric oxide (more
probably ferric hydrate). In Lecqueureusia spiralis (formerly united
with Difflugia) the test is formed of minute sausage-shaped granules,
in which may be identified the partly dissolved valves of Diatoms
taken as food; it is spirally twisted at the apex, as if it had enlarged
after its first formation, a very rare occurrence in this group. The
most frequent mode of fission in the testaceous Rhizopods (Figs. 8,
10) is what Schaudinn aptly terms "bud-fission," where half the
protoplasm protrudes and accumulates at the mouth of the shell, and
remains till a test has formed for it, while the other half retains the
test of the original animal. The materials for the shell, whether sand-
granules or plates, pass from the depths of the original shell
outwards into the naked cell, and through its cytoplasm to the
surface, where they become connected by cementing matter into a
continuous test. The nucleus now divides into two, one of which
passes into the external animal; after this the two daughter-cells
separate, the one with the old shell, the other, larger, with the new
one.
Fig. 10.—Test-bearing Rhizopods. A, Quadrula symmetrica: B, Hyalosphenia
lata; C, Arcella vulgaris; D, Difflugia pyriformis. (From Lang's Comparative
Anatomy.)
If two individuals of the shelled species undergo bud-fission in close

proximity, the offspring may partially coalesce, so that a monstrous
shell is produced having two pylomes.
Reproduction by fission has been clearly made out in most members

of the group; some of the multinucleate species often abstrict a
portion, sometimes at several points simultaneously, so that fission
here passes into budding[70] (Fig. 9, 2, 6).
Brood-division, either by resolution in the multinucleate species, or

preceded by multiple nuclear division in the habitually 1-nucleate,
though presumably a necessary incident in the life-history of every
species, has only been seen, or at least thoroughly worked out, in a
few cases, where it is usually preceded by encystment, and mostly
by the extrusion into the cyst of any undigested matter.[71]
In Trichosphaerium (Fig. 9) the cycle described by Schaudinn is very

complex, and may be divided into two phases, which we may term
the A and the B subcycles. The members of the A cycle are
distinguished by the gelatinous investment being armed with radial
spicules, which are absent from the B form. The close of the A cycle
is marked by the large multinucleate body resolving itself into
amoeboid zoospores (3), which escape from the gelatinous test, and
develop into the large multinucleate adults of the B form. These, like
the A form, may reproduce by fission or budding. At the term of
growth, however, they retract their pseudopodia, expel the excreta,
and multiply their nuclei by mitosis (7). Then the body is resolved
into minute 2-flagellate microzoospores (8), which are exogamous
gametes, i.e. they will only pair with similar zoospores from another
cyst. The zygote (9-11) resulting from this conjugation is a minute
amoeboid; its nucleus divides repeatedly, a gelatinous test is formed
within which the spicules appear, and so the A form is reconstituted.
In many of the test-bearing forms, whether Lobose or Filose,
plastogamic unions occur, and the two nuclei may remain distinct,
leading to plurinucleate monsters in their offspring by fission, or they
may fuse and form a giant nucleus, a process which has here no
relation to normal syngamy, as it is not associated with any marked
change in the alternation of feeding and fission, etc. In
Trichosphaerium also plastogamic unions between small individuals
have for their only result the increase of size, enabling the produce
to deal with larger prey. Temporary encystment in a "hypnocyst" is
not infrequent in both naked and shelled species, and enables them
to tide over drought and other unfavourable conditions.
Schaudinn has discovered and worked out true syngamic processes,

some bisexual, some exogamous, in several other Rhizopods. In
Chlamydophrys stercorea the pairing-cells are equal, and are formed
by the aggregation of the chromidia into minute nuclei around which
the greater part of the cytoplasm aggregates, while the old nucleus
(with a little cytoplasm) is lost. These brood-cells are 2-flagellate
pairing-cells, which are exogamous: the zygote is a brown cyst; if
this be swallowed by a mammal, the original Chlamydophrys
appears in its faeces.[72]
Centropyxis aculeata, a species very common in mud or moss, allied

to Difflugia, also forms a brood by aggregation around nuclei derived
from chromidia. The brood-cells are amoeboid, and secrete
hemispherical shells like those of Arcella; some first divide into four
smaller ones, before secreting the shell. Pairing takes place between
the large and the small forms; and the zygote encysts. Weeks or
months afterwards the cyst opens and its contents creep out as a
minute Centropyxis. Finally, Amoeba coli produces its zygote in a
way recalling that of Actinosphaerium (pp. 73-75, Fig. 21): the cell
encysts; its nucleus divides, and each daughter divides again into
two, which fuse reciprocally. Thus the cyst contains two zygote
nuclei. After a time each of these divides twice, so that the mature
cyst contains eight nuclei. Probably when swallowed by another
animal they liberate a brood of eight young amoebae. Thus in
different members of this group we have exogamy, both equal and
bisexual, and endogamy.
Most of the Rhizopoda live among filamentous Algae in pools,

ponds, and in shallow seas, etc.; some are "sapropelic" or mud-
dwellers (many species of Amoeba, Pelomyxa, Difflugia, etc.), others
frequent the roots of mosses. Amoeba coli is often found as a
harmless denizen of the large intestine of man. Amoeba histolytica,
lately distinguished therefrom by Schaudinn, is the cause of tropical
dysentery. It multiplies enormously in the gut, and is found extending
into the tissues, and making its way into the abscesses that so
frequently supervene in the liver and other organs. Chlamydophrys
stercorea is found in the faeces of several mammals. The best
monograph of this group is that of Penard.[73]
2. Foraminifera[74]
Sarcodina with no central capsule or distinction of ectosarc; the
pseudopodia fine, branching freely, and fusing where they meet to
form protoplasmic networks, or the outermost in the pelagic forms
radiating, but without a central or axial filament: sometimes
dimorphic, reproducing by fission and by rhizopod or flagellate germs
in the few cases thoroughly investigated: all marine (with the
exception of some of the Allogromidiaceae), and usually provided
with a test of carbonate of lime ("vitreous" calcite, or "porcellanous"
aragonite?), or of cemented particles of sand ("arenaceous"); test-
wall continuous, or with the walls perforated by minute pores or
interstices for the protrusion of pseudopodia.
The classification of Carpenter (into Vitreous or Perforate,

Porcellanous or Imperforate, and Arenaceous), according to the
structure of the shell, had proved too artificial to be used by Brady in
the great Monograph of the Foraminifera collected by the
"Challenger" Expedition,[75] and has been modified by him and
others since then. We reproduce Lister's account of Brady's
classification.[76] We must, however, warn the tyro that its
characterisations are not definitions (a feature of all other recent
systems), for rigid definitions are impossible: here as in the case, for
instance, of many Natural Orders of Plants, transitional forms making
the establishment of absolute boundaries out of the question. In the
following classification we do not think it, therefore, necessary to
complete the characterisations by noting the extremes of variation
within the orders:—
1. Allogromidiaceae: simple forms, often fresh-water and similar to

Rhizopoda; test 0, or chitinous, gelatinous, or formed of cemented particles,
whether secreted platelets or ingested granules. Biomyxa, Leidy =
Gymnophrys, Cienk.; Diaphorodon, Archer; Allogromia, Rhumbl. (= Gromia,
auctt.[77] nec Duj.) (Fig. 14, 1); Lieberkühnia, Cl. and Lachm. (Fig. 12);
Microgromia, R. Hertw. (Fig. 11); Pamphagus, Bailey.
2. Astrorhizidaceae: test arenaceous, often large, never truly chambered, or if
so, asymmetrical. Astrorhiza, Sandahl; Haliphysema, Bowerb.; Saccammina,
M. Sars (Fig. 13, 1); Loftusia, Brady.
3. Lituolidaceae: test arenaceous, often symmetrical or regularly spiral,
isomorphous with calcareous forms: the chambers when old often
"labyrinthine" by the ingrowth of wall-material. Lituola, Lam.; Reophax, Montf.;
Ammodiscus, Reuss; Trochammina, Parker and Jeffreys.
4. Miliolidaceae: test porcellanous, imperforate, spirally coiled or cyclic, often
chambered except in Cornuspira: simple in Squamulina. Cornuspira, Max
Sch.; Peneroplis, Montf.; Miliolina, Lam. (incl. Biloculina (Fig. 15), Triloculina,
Quinqueloculina (Figs. 14, 4; 15, B), Spiroloculina (Fig. 13, 5) of d'Orb.);
Alveolina, d'Orb.; Hauerina, d'Orb.; Calcituba, Roboz; Orbitolites, Lam.;
Orbiculina, Lam.; Alveolina, Park. and Jeffr.; Nubecularia, Def.; Squamulina,
Max Sch. (Fig. 14, 3).
5. Textulariaceae: test calcareous, hyaline, perforated; chambers increasing
in size in two alternating rows, or three, or passing into a spiral. Textularia,
Def.; Bulimina, d'Orb.; Cassidulina, d'Orb.
6. Cheilostomellaceae: test vitreous, delicate, finely perforated, chambered,
isomorphic with the spiral forms of the Miliolidaceae. Cheilostomella, Reuss.
7. Lagenaceae: Test vitreous, very finely perforate, chambers with a distinct
pylome projecting (ectosolenial), or turned in (entosolenial), often succeeding
to form a necklace-like shell. Lagena, Walker and Boys (Fig. 13, 2);
Nodosaria, Lam. (Fig. 13, 3); Cristellaria, Lam.; Frondicularia, Def. (Fig. 13, 4);
Polymorphina, Lam.; Ramulina, Wright.
8. Globigerinidae: test vitreous, perforate; chambers few, dilated, and
arranged in a flat or conical spiral, usually with a crescentic pylome to the
last. Globigerina, d'Orb. (Figs. 13, 6; 16, 2); Hastigerina, Wyv. Thoms.;
Orbulina, d'Orb. (Fig. 16, 1).
9. Rotaliaceae; test vitreous, perforate, usually a conical spiral (like a snail),
chambers often subdivided into chamberlets, and with a proper wall, and
intermediate skeleton traversed by canals. Rotalia, Lam. (Fig. 14, 2);
Planorbulina, d'Orb. (Fig. 13, 9); Polytrema, Risso; Spirillina, Ehr. (non-
septate); Patellina, Will.; Discorbina, P. and J. (Fig. 13, 7).
10. Nummulitaceae: test usually a complex spiral, the turns completely
investing their predecessors: wall finely tubular, often with a proper wall and
intermediate skeleton. Fusulina, Fisch.; Polystomella, Lam.; Nummulites,
d'Orb. (Fig. 13, 11); Orbitoides, d'Orb.
The Allogromidiaceae are a well-marked and distinct order, on the

whole resembling the Rhizopoda Filosa, and are often found with
them in fresh water, while all other Foraminifera are marine. The type
genus, Allogromia (Fig. 14, 1), has an oval chitinous shell.
Microgromia socialis (Fig. 11) is often found in aggregates, the
pseudopodia of neighbours fusing where they meet into a common
network. This is due to the fact that one of the two daughter-cells at
each fission, that does not retain the parent shell, remains in
connexion with its sister that does: sometimes, however, it retracts
its pseudopodia, except two which become flagella, wherewith it can
swim off. The test of Pamphagus is a mere pellicle. In Lieberkühnia
(Fig. 12) it is hardly that; though the body does not give off the fine
pseudopodia directly, but emits a thick process or "stylopodium"[78]
comparable to the protoplasm protruded through the pylome of its
better protected allies; and from this, which often stretches back
parallel to the elongated body, the reticulum of pseudopodia is
emitted. Diaphorodon has a shell recalling that of Difflugia (Fig. 10,
D, p. 55), formed of sandy fragments, but with interstices between
them through which as well as through the two pylomes the
pseudopodia pass. In all of these the shell is formed as in the
Rhizopods once for all, and does not grow afterwards; and the fresh-
water forms, which are the majority, have one or more contractile
vacuoles; in Allogromia they are very numerous, scattered on the
expanded protoplasmic network.
Fig. 11.—Microgromia socialis. A, entire colony; B, single zooid; C, zooid which

has undergone binary fission, with one of the daughter-cells creeping out of
the shell; D, flagellula. c.vac, Contractile vacuole; nu, nucleus; sh, shell.
(From Parker and Haswell, after Hertwig and Lesser.)
Fig. 12.—Lieberkühnia, a fresh-water Rhizopod, from the egg-shaped shell of
which branched pseudopodial filaments protrude. (From Verworn.)
The remaining marine families may all be treated of generally, before

noting their special characters. Their marine habitat is variable, but in
most cases restricted. A few extend up the brackish water of
estuaries: a large number are found between tide-marks, or on the
so-called littoral shelf extending to deep water; they are for the most
part adherent to seaweeds, or lie among sand or on the mud. Other
forms, again, are pelagic, such as Globigerina (Figs. 13, 6, 16, 17)
and its allies, and float as part of the plankton, having the surface of
their shells extended by delicate spines, their pseudopodia long and
radiating, and the outer part of their cytoplasm richly vacuolated
("alveolate"), and probably containing a liquid lighter than sea water,
as in the Radiolaria. Even these, after their death and the decay of
the protoplasm, must sink to the bottom (losing the fine spines by
solution as they fall); and they accumulate there, to form a light oozy
mud, the "Globigerina-ooze" of geographers, at depths where the
carbonic acid under pressure is not adequate to dissolve the more
solid calcareous matter. Grey Chalk is such an ooze, consolidated by
the lapse of time and the pressure of superincumbent layers. Some
Foraminifera live on the sea bottom even at the greatest depths, and
of course their shell is not composed of calcareous matter.
Foraminifera may be obtained for examination by carefully washing
sand or mud, collected on the beach at different levels between tide-
marks, or from dredgings, or by carefully searching the surface of
seaweeds, or by washing their roots, or, again, by the surface or
deep-sea tow-net. The sand used to weight sponges for sale is the
ready source of a large number of forms, and may be obtained for
the asking from the sponge-dealers to whom it is a useless waste
product. If this sand is dried in an oven, and then poured into water,
the empty shells, filled with air, will float to the surface, and may be
sorted by fine silk or wire gauze.
From the resemblance of the shells of many of them to the Nautilus

they were at first described as minute Cephalopods, or Cuttlefish, by
d'Orbigny,[79] and their true nature was only elucidated in the last
century by the labours of Williamson, Carpenter, Dujardin, and Max
Schultze. At first they possess only one nucleus, but in the adult
stage may become plurinucleate without dividing, and this is
especially the case in the "microsphaeric" states exhibited by many
of those with a complex shell; the nucleus is apt to give off fragments
(chromidia) which lie scattered in the cytoplasm. At first, too, in all
cases, the shell has but a single chamber, a state that persists
through life in some. When the number of chambers increases, their
number has no relation to that of the nuclei, which remains much
smaller till brood-formation sets in.
The shell-substance, if calcareous, has one of the two types,

porcellanous or vitreous, that we have already mentioned, but
Polytrema, a form of very irregular shape, though freely perforated,
is of a lovely pink colour. In the calcareous shells sandy particles
may be intercalated, forming a transition to the Arenacea. In these
the cement has an organic base associated with calcareous or
ferruginous matter; in some, however, the cement is a phosphate of
iron. The porcellanous shells are often deep brown by transmitted
light.
Fig. 13.—Shells of Foraminifera. In 3, 4, and 5, a shows the surface view, and b
a section; 8a is a diagram of a coiled cell without supplemental skeleton; 8b
of a similar form with supplemental skeleton (s.sk); and 10 of a form with
overlapping whorls; in 11a half the shell is shown in horizontal section; b is
a vertical section; a, aperture of the shell; 1-15, successive chambers, 1
being always the oldest or initial chamber. (From Parker and Haswell, after
other authors.)
Despite the apparent uniformity of the protoplasmic body in this

group, the shell is infinitely varied in form. As Carpenter writes, in
reference to the Arenacea, "There is nothing more wonderful in
nature than the building up of these elaborate and symmetrical
structures by mere jelly-specks, presenting no traces whatever of
that definite organisation which we are accustomed to regard as
necessary to the manifestations of conscious life.... The tests (shells)
they construct when highly magnified bear comparison with the most
skilful masonry of man. From the same sandy bottom one species
picks up the coarsest quartz grains, unites them together with a
ferruginous cement, and thus constructs a flask-shaped test, having
a short neck and a single large orifice; another picks up the finer
grains and puts them together with the same cement into perfectly
spherical tests of the most extraordinary finish, perforated with
numerous small pores disposed at pretty regular intervals. Another
species selects the minutest sand grains and the terminal portions of
sponge-spicules, and works them up together—apparently with no
cement at all, but by the mere laying of the spicules—into perfect
white spheres like homoeopathic globules, each showing a single-
fissured orifice. And another, which makes a straight, many-
chambered test, the conical mouth of each chamber projecting into
the cavity of the next, while forming the walls of its chambers of
ordinary sand grains rather loosely held together, shapes the conical
mouths of the chambers by firmly cementing together the quartz
grains which border it." The structure of the shell is indeed variable.
The pylome may be single or represented by a row of holes
(Peneroplis, Orbitolites), or, again, there may be several pylomes
(Calcituba); and, again, there are in addition numerous scattered
pores for the protrusion of pseudopodia elsewhere than from the
stylopodium, in the whole of the "Vitrea" and in many "Arenacea";
and, as we shall see, this may exercise a marked influence on the
structure of the shell.
In some cases the shell is simple, and in Cornuspira and Spirillina

increases so as to have the form of a flat coiled tube. In Calcituba
the shell branches irregularly in a dichotomous way, and the older
parts break away as the seaweed on which they grow is eaten away,
and fall to the bottom, while the younger branches go on growing
and branching. The fallen pieces, if they light on living weed, attach
themselves thereto and repeat the original growth; if not, the
protoplasm crawls out and finds a fresh weed and forms a new tube.
In the "Polythalamia" new chambers are formed by the excess of the
protoplasm emerging and surrounding itself with a shell, organically
united with the existing chamber or chambers, and in a space-
relation which follows definite laws characteristic of the species or of
its stage of growth, so as to give rise to circular, spiral, or irregular
complexes (see Fig. 13).
Fig. 14.—Various forms of Foraminifera. In 4, Miliola, a, shows the living animal;
b, the same killed and stained; a, aperture of shell; f, food particles; nu,
nucleus; sh, shell. (From Parker and Haswell, after other authors.)
In most cases the part of the previously existing chamber next the
pylome serves as the hinder part of the new chamber, and the old
pylome becomes the pore of communication. But in some of the
"Perforata" each new chamber forms a complete wall of its own
("proper wall," Fig. 13, 8b), and the space between the two adjacent
walls is filled with an intermediate layer traversed by canals
communicating with the cavities of the chambers ("intermediate
skeleton"), while an external layer of the same character may form a
continuous covering. The shell of the Perforata may be adorned with
pittings or fine spines, which serve to increase the surface of support
in such floating forms as Globigerina, Hastigerina, and the like (Fig.
17). In the "Imperforata" the outer layer is often ornamented with
regular patterns of pits, prominences, etc., which are probably
formed by a thin reflected external layer of protoplasm. In some of
the "Arenacea" a "labyrinthine" complex of laminae is formed.
A very remarkable point which has led to great confusion in the study
of the Foraminifera, is the fact that the shell on which we base our
characters of classification, may vary very much, even within the
same individual. Thus in the genus Orbitolites the first few chambers
of the shell have the character of a Milioline, in Orbiculina of a
Peneroplis. The arrangements of the Milioline shell, known as
Triloculine, Quinqueloculine, and Biloculine respectively, may
succeed one another in the same shell (Figs. 14 4, 15). A shell may
begin as a spiral and end by a straight continuation: again, the
spherical Orbulina (Fig. 16 1) is formed as an investment to a shell
indistinguishable from Globigerina, which is ultimately absorbed. In
some cases, as Rhumbler has pointed out, the more recent and
higher development shows itself in the first formed chambers, while
the later, younger chambers remain at a lowlier stage, as in the case
of the spiral passing into a straight succession; but the other cases
we have cited show that this is not always the case. In Lagena (Fig.
13 2) the pylome is produced into a short tube, which may protrude
from the shell or be turned into it, so that for the latter form the genus
Entosolenia was founded. Shells identical in minute sculpture are,
however, found with either form of neck, and, moreover, the
polythalamial shells (Nodosaria, Fig. 13 3), formed of a nearly
straight succession of Lagena-like chambers, may have these
chambers with their communications on either type. Rhumbler goes
so far as to suggest that all so-called Lagena shells are either the
first formed chamber of a Nodosaria which has not yet become
polythalamian by the formation of younger ones, or are produced by
the separation of an adult Nodosaria into separate chambers.
Fig. 15.—A, Megalospheric; B, microspheric shell of Biloculina. c, The initial
chamber. The microspheric form begins on the Quinqueloculina type. (From
Calkins' Protozoa.)
Many of the chambered species show a remarkable dimorphism, first

noted by Schlumberger, and finally elucidated by J. J. Lister and
Schaudinn. It reveals itself in the size of the initial chamber;
accordingly, the two forms may be distinguished as "microspheric"
and "megalospheric" respectively (Fig. 15), the latter being much the
commoner. The microspheric form has always a plurality of nuclei,
the megalospheric a single one, except at the approach of
reproduction. Chromidial masses are, however, present in both
forms. The life-history has been fully worked out in Polystomella by
Schaudinn, and in great part in Polystomella, Orbitolites, etc., by
Lister; and the same scheme appears to be general in the class, at
least where the dimorphism noted occurs. The microspheric form
gives birth only to the megalospheric, but the latter may reproduce
megalospheric broods, or give rise to swarmers, which by their
(exogamous) conjugation produce the microspheric young. The
microspheric forms early become multinucleate, and have also
numerous chromidia detached from the nuclei, which they ultimately
replace. These collect in the outer part of the shell and aggregate
into new nuclei, around which the cytoplasm concentrates, to
separate into as many amoeboid young "pseudopodiospores" as
there are nuclei. These escape from the shell or are liberated by its
disintegration, and invest themselves with a shell to form the initial
large central chamber or megalosphere.
Fig. 16.—1, Orbulina universa. Highly magnified. 2, Globigerina bulloides. Highly
magnified. (From Wyville Thomson, after d'Orbigny.)
In the ordinary life of the megalospheric form the greater part of the
chromatic matter is aggregated into a nucleus, some still remaining
diffused. At the end of growth the nucleus itself disintegrates, and
the chromidia concentrate into a number of small vesicular nuclei,
each of which appropriates to itself a small surrounding zone of thick
plasm and then divides by mitosis twice; and the 4-nucleate cells so
formed are resolved into as many 1-nucleate, 2-flagellate swarmers,
which conjugate only exogamously.[80] The fusion of their nuclei
takes place after some delay: ultimately the zygote nucleus divides
into two, a shell is formed, and we have the microsphere, which is
thus pluri-nucleate ab initio. As we have seen, the nuclei of the
microsphere are ultimately replaced by chromidia, and the whole
plasmic body divides into pseudopodiospores, which grow into the
megalospheric form.
Fig. 17.—Shell of Globigerina bulloides, from tow-net, showing investment of

spines. (From Wyville Thomson.)
In the Perforate genera, Patellina and Discorbina, plastogamy
precedes brood formation, the cytoplasms of the 2-5 pairing
individuals contracting a close union; and then the nuclei proceed to
break up without fusion, while the cytoplasm aggregates around the
young nuclei to form amoebulae, which acquire a shell and separate.
In both cases it is the forms with a single nucleus, corresponding to
megalospheric forms that so pair, and the brood-formation is, mutatis
mutandis, the same as in these forms. Similar individuals may
reproduce in the same way, in both genera, without this plastogamic
pairing, which is therefore, though probably advantageous, not
essential. If pseudopodiospores form their shells while near one
another, they may coalesce to form monsters, as often happens in
Orbitolites.[81]
The direct economic uses of the Foraminifera are perhaps greater

than those of any other group of Protozoa. The Chalk is composed
largely of Textularia and allied forms, mixed with the skeletons of
Coccolithophoridae (pp. 113-114), known as Coccoliths, etc. The
Calcaire Grossier of Paris, used as a building stone, is mainly
composed of the shells of Miliolines of Eocene age; the Nummulites
of the same age of the Mediterranean basin are the chief constituent
of the stone of which the Pyramids of Egypt are built. Our own Oolitic
limestones are composed of concretions around a central nucleus,
which is often found to be a minute Foraminiferous shell.
The palaeontology of the individual genera is treated of in

Chapman's and Lister's recent works. They range from the Lower
Cambrian characterised by perforated hyaline genera, such as
Lagena, to the present day. Gigantic arenaceous forms, such as
Loftusia, are among the Tertiary representatives; but the limestones
formed principally of their shells commence at the Carboniferous.
The so-called Greensands contain greenish granules of "glauconite,"
containing a ferrous silicate, deposited as a cast in the chambers of
Foraminifera, and often left exposed by the solution of the
calcareous shell itself. Such granules occur in deep-sea deposits of
the present day.[82]
3. Heliozoa
Sarcodina with radiate non-anastomosing pseudopodia of granular
protoplasm, each with a stiff axial rod passing into the body plasma;
no central capsule, nor clear ectoplasm; skeleton when present
siliceous; nucleus single or multiple; contractile vacuole (or vacuoles)
in fresh-water species, superficial and prominent at the surface in
diastole; reproduction by fission or budding in the active condition, or
by brood-formation in a cyst, giving rise to resting spores;
conjugation isogamous in the only two species fully studied; habitat
floating or among weeds, mostly fresh water.
1. Naked or with an investment only when encysted.

Aphrothoraca.—Actinolophus F.E. Sch.; Myxastrum Haeck.;
Gymnosphaera Sassaki; Dimorpha (Fig. 37, 5, p. 112) Gruber;
Actinomonas Kent; Actinophrys Ehrb.; Actinosphaerium St.; Camptonema
Schaud; Nuclearia Cienk.
2. Invested with a gelatinous layer, sometimes traversed by a firmer elastic

network.
Chlamydophora.—Heterophrys Arch.; Mastigophrys Frenzel;
Acanthocystis, Carter.
3. Ectoplasm with distinct siliceous spicules.

Chalarothoraca.—Raphidiophrys Arch.
4. Skeleton a continuous, fenestrated shell, sometimes stalked.

Desmothoraca.—Myriophrys Penard; Clathrulina Cienk.; Orbulinella Entz.
This class were at first regarded and described as fresh-water

Radiolaria, but the differences were too great to escape the greatest
living specialist in this latter group, Ernst Haeckel, who in 1866
created the Heliozoa for their reception. We owe our knowledge of it
mainly to the labours of Cienkowsky, the late William Archer, F. E.
Schulze, R. Hertwig, Lesser, and latterly to Schaudinn, who has
monographed it for the "Tierreich" (1896); and Penard has published
a more recent account.
Fig. 18.—Actinophrys sol. a, Axial filament of pseudopod; c.v, contractile

vacuole; n, nucleus. (From Lang's Comparative Anatomy, after Grenacher.)
Actinophrys sol Ehrb. (Fig. 18) is a good and common type. It owes
its name to its resemblance to a conventional drawing of the sun,
with a spherical body and numerous close-set diverging rays. The
cytoplasm shows a more coarsely vacuolated outer layer, sometimes
called the ectosarc, and a denser internal layer the endosarc. In the
centre of the figure is the large nucleus, to which the continuations of
the rays may be seen to converge; the pseudopodia contain each a
stiffish axial filament,[83] which is covered by the fine granular plasm,
showing currents of the granules. The axial filament disappears
when the pseudopodia are retracted or bent, and is regenerated
afterwards. This bending occurs when a living prey touches and
adheres to a ray, all its neighbours bending in like the tentacles of a
Sundew. The prey is carried down to the surface of the ectoplasm,
and sinks into it with a little water, to form a nutritive vacuole. Fission
is the commonest mode of reproduction, and temporary plastogamic
unions are not uncommon. Arising from these true conjugations
occur, two and two, as described by Schaudinn. A gelatinous cyst
wall forms about the two which are scarcely more than in contact
with their rays withdrawn. Then in each the nucleus divides into two,
one of which passes to the surface, and is lost (as a "polar body"),
while the other approaches the corresponding nucleus of the mate,
and unites with it, while at the same time the cytoplasms fuse. Within
the gelatinous cyst the zygote so formed divides to produce two
sister resting spores, from each of which, after a few days, a young
Actinophrys escapes, as may take place indeed after encystment of
an ordinary form without conjugation.
Fig. 19.—Actinosphaerium eichornii. A, entire animal with two contractile

vacuoles (c.vac); B, a portion much magnified, showing alveolate
cytoplasm, pseudopodia with axial rods, non-nucleate cortex (cort), multiple
nuclei (nu) of endoplasm (med), and food-vacuole (chr). (From Parker and
Haswell.)
The axial rods of the pseudopodia may pass either to the

circumference of the nucleus or to a central granule, corresponding,
it would appear, to a centrosome or blepharoplast; or again, in the
plurinucleate marine genus Camptonema, each rod abuts on a
separate cap on the outer side of each nucleus. The nucleus is
single in all but the genera Actinosphaerium, Myxastrum,
Camptonema, and Gymnosphaera. The movements of this group
are very slow, and are not well understood. A slow rolling over on the
points of the rays has been noted, and in Camptonema they move
very decidedly to effect locomotion, the whole body also moving
Amoeba-fashion; but of the distinct movements of the species when
floating no explanation can be given. The richly vacuolate ectoplasm
undoubtedly helps to sustain the cell, and the extended rays must
subserve the same purpose by so widely extending the surface.
Dimorpha (Fig. 37, 5, p. 112) has the power of swimming by
protruding a pair of long flagella from the neighbourhood of the
eccentric nucleus; and Myriophrys has an investment of long
flagelliform cilia. Actinomonas has a stalk and a single flagellum in
addition to the pseudopodia; these genera form a transition to the
Flagellata.
Several species habitually contain green bodies, which multiply by

bipartition, and are probably Zoochlorellae, Chlamydomonadidae of
the same nature as we shall find in certain Ciliata (pp. 154, 158) in
fresh-water Sponges (see p. 175), in Hydra viridis (p. 256), and the
marine Turbellarian Convoluta (Vol. II. p. 43).
Reproduction by fission is not rare, and in some cases

(Acanthocystis) the cell becomes multinuclear, and buds off 1-
nucleate cells. In such cases the buds at first lack a centrosome, and
a new one is formed first in the nucleus, and passes out into the
cytoplasm. These buds become 2-flagellate before settling down. In
Clathrulina the formation of 2-flagellate zoospores has long been
known (Fig. 20, 3). In Actinosphaerium (Figs. 19, 21), a large
species, differing from Actinophrys only in the presence of numerous
nuclei in its endoplasm, a peculiar process, which we have
characterised as endogamy, results in the formation of resting
spores. The animal retracts its rays and encysts; and the number of
nuclei is much reduced by their mutual fusion, or by the solution of
many of them, or by a combination of the two processes. The body
then breaks up into cells with a single nucleus, and each of these
surrounds itself with a wall to form a cyst of the second order.

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Viral Nanoparticles Tools for Materials

Science and Biomedicine 1st Edition

Composite Materials: Applications in Engineering,

Advances in Neutron Optics Fundamentals and

Tools For Chemical Product Design From Consumer

CRC materials science and engineering handbook Fourth

Materials engineering science processing and design

Silver nanoparticles for antibacterial devices :

A Matlab Primer for Technical Programming for Materials

Vaccine Technologies for Veterinary Viral Diseases

Viral.indd 1 3/7/11 3:13:12 PM

Copyright © 2011 by Pan Stanford Publishing Pte. Ltd.

Viral_Nanoparticles_TP.indd 1 3/1/11 1:38:12 PM

Copyright © 2011 by Pan Stanford Publishing Pte. Ltd.

British Library Cataloguing-in-Publication Data

VIRAl NAnOpARTIclES: TOOlS FOR MATERIAlS ScIEncE And BIOmEdIcInE

ISBN 978-981-4267-45-8 (Hardcover)

Copyright © 2011 by Pan Stanford Publishing Pte. Ltd.

Chapter 1 An Introduction to VNPs and Nanotechnology 1

Chapter 2 Overview of the Manifold VNPs Used in

Chapter 3 Production of VNPs, VLPs, and Chimeras 49

Chapter 4 The Art of Bioconjugation: Functionalization

Chapter 5 Encapsulating Materials within VNPs 123

Chapter 6 VNPs as Templates for Materials Synthesis 145

Chapter 7 Playing “Nano-Lego”: VNPs as Building Blocks for the

Chapter 8 VNPs as Tools for Nanomedicine 207

Chapter 9 Summary and Outlook 247

Copyright © 2011 by Pan Stanford Publishing Pte. Ltd.

Copyright © 2011 by Pan Stanford Publishing Pte. Ltd.

Viral nanotechnology is a young and emerging discipline. A highly interdis-

Copyright © 2011 by Pan Stanford Publishing Pte. Ltd.

Copyright © 2011 by Pan Stanford Publishing Pte. Ltd.

AN INTRODUcTiON TO VNPs aND

1.1 WHAT IS NANO?

Nanotechnology is a highly interdisciplinary field that brings together

Viral Nanoparticles: Tools for Materials Science and Biomedicine

Copyright © 2011 by Pan Stanford Publishing Pte. Ltd.

Copyright © 2011 by Pan Stanford Publishing Pte. Ltd.

1.2 WHERE DID IT ALL BEGIN? A HISTORY OF VNPs: FROM

Copyright © 2011 by Pan Stanford Publishing Pte. Ltd.

Copyright © 2011 by Pan Stanford Publishing Pte. Ltd.

Copyright © 2011 by Pan Stanford Publishing Pte. Ltd.

and metal deposition techniques as well as their potential applications are

1.3 WHY VNPs? MATERIALS PROPERTIES OF VNPs

When a virologist looks at a virus, he or she might see a pathogen, an infectious

Copyright © 2011 by Pan Stanford Publishing Pte. Ltd.

Figure 1.2 Transmission electron micrograph of Acidianus bottle-shaped virus.

Icosahedral particles range in their size from 18 nm to 500 nm. Rod-

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Icosahedral plant viruses An insect virus

BMV CCMV CPMV HCRSV RCNMV TYMV FHV

Rod-shaped plant virus and a filamentous phage

Copyright © 2011 by Pan Stanford Publishing Pte. Ltd.

Copyright © 2011 by Pan Stanford Publishing Pte. Ltd.

Several genera have tests of siliceous or chitinous plates, formed in

If two individuals of the shelled species undergo bud-fission in close

Reproduction by fission has been clearly made out in most members

Brood-division, either by resolution in the multinucleate species, or

In Trichosphaerium (Fig. 9) the cycle described by Schaudinn is very

Schaudinn has discovered and worked out true syngamic processes,

Centropyxis aculeata, a species very common in mud or moss, allied

Most of the Rhizopoda live among filamentous Algae in pools,

The classification of Carpenter (into Vitreous or Perforate,