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Advances in PET

The Latest in Instrumentation,

Technology, and Clinical Practice
Jun Zhang
Michael V. Knopp
Editors

123
Advances in PET
Jun Zhang • Michael V. Knopp
Editors

Advances in PET
The Latest in Instrumentation,
Technology, and Clinical Practice
Editors
Jun Zhang Michael V. Knopp
Department of Radiology Department of Radiology
The Ohio State University The Ohio State University
Columbus, OH Columbus, OH
USA USA

ISBN 978-3-030-43039-9    ISBN 978-3-030-43040-5 (eBook)

https://doi.org/10.1007/978-3-030-43040-5

© Springer Nature Switzerland AG 2020

This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of
the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation,
broadcasting, reproduction on microfilms or in any other physical way, and transmission or information
storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology
now known or hereafter developed.
The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication
does not imply, even in the absence of a specific statement, that such names are exempt from the relevant
protective laws and regulations and therefore free for general use.
The publisher, the authors and the editors are safe to assume that the advice and information in this book
are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or the
editors give a warranty, expressed or implied, with respect to the material contained herein or for any
errors or omissions that may have been made. The publisher remains neutral with regard to jurisdictional
claims in published maps and institutional affiliations.

This Springer imprint is published by the registered company Springer Nature Switzerland AG
The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland
Preface

First and foremost, I sincerely welcome and greatly appreciate Dr. Suleman Surti
and Dr. Joel S. Karp (Chaps. 1 and 7), Dr. Daniel A. Pryma (Chap. 2), Dr. Dennis
R. Schaart (Chap. 3), Dr. David F.C. Hsu and Dr. Craig S. Levin (Chap. 4), Dr. Jun
Zhang and Dr. Michael V. Knopp (Chap. 5), and Dr. Michael E. Casey and Dr.
Dustin Osborne (Chap. 6) for their coauthorship and contribution to the first edition
of this book.
For decades, researchers have been interested in replacing photomultiplier tubes
with semiconductor photosensors. With the disruptive technology leap of solid state
PET detectors, improvements in PET equipment are happening at a staggering pace
in both PET/CT and PET/MRI. For the first time, clinical PET scanners have
reached 200–400 ps for time-of-flight timing resolution, 3 mm for spatial resolu-
tion, and >20 cps/kBq for sensitivity. Research into innovative PET technologies,
methodologies, and clinical practice is expanding, with the potentials of enabling
better PET image quality, reduced PET dose, and accelerated PET scans.
This book is a guide to new and emerging PET technology, instrumentation, and
its place in clinical practice to discover potential values of solid state PET and pro-
mote the growth of PET imaging. It is primarily addressed to medical physicists,
nuclear medicine physicians, and technologists who wish to learn and acquire
knowledge of solid state digital PET imaging including silicon photomultiplier-
based PET detector technology, system characteristics and performance, as well as
clinical practices. The basic principles of PET physics with elementary concepts are
not dedicatedly described in this book.
We have tried to write the book in a concise and logical manner. It is organized
into three categories: I. the basics of PET imaging, which focuses on the current
status and future direction of PET technology and clinical practice (Chaps. 1 and 2);
II. solid state digital PET instrumentation, technology, and clinical practice, which
focuses on both the fundamentals of silicon photomultipliers for time-of-flight PET
(Chap. 3) and the evaluation of emerging PET instrumentation and technologies
across manufactures (Chaps. 4, 5, and 6); III. future prospective of PET/CT imag-
ing, focusing on the EXPLORER family’s long axial field-of-view total-body PET
instrumentation and technologies (Chap. 7).
We believe that it is a new era for PET imaging. The new-generation solid state
PET instrumentation and technologies have brought exciting times also evolving
challenges to PET imaging. It has created new initiatives and opportunities in

v
vi Preface

low-dose PET, fast PET imaging, better lesion detectability, optimal imaging per-
formance, novel tracer applications, as well as whole-body dynamic scans, which
could revolutionize our understanding and treatment of disease through overall
improved technologies and experiences.
In the end, it is a team’s work. It has been my great pleasure and privilege to work
together with all the coauthoring professionals in writing this book. A sincere appre-
ciation to Dr. Michael V. Knopp for his great help and support to me in providing
the solid state PET platform and fulfilling the book as the coeditor. A great and
special appreciation to Dr. Joel S. Karp and Dr. Suleman Surti for their professional
guidance and meaningful suggestions to me in the book design and write-ups. I am
indebted to many of my peers and colleagues: Dr. Timothy Turkington, Dr. Nathan
C. Hall, Dr. Osama Mawlawi, Dr. David Hintenlang, Dr. Mona Natwa, and more for
their inspiration and support to me. Taking this opportunity, we are very grateful to
Ms. Margaret Moore, Ms. Marie Felina, Ms. Kayalvizhi Eswaran, Mr. Prakash
Marudhu, and other publishing staff of Springer for their various help and coopera-
tion in bringing our labor to fruition.
To my wife, Zhaoxia, and my daughters, Grace and Rachel, with love and
appreciation.

Columbus, OH, USA Jun Zhang

Contents

Part I Basics Science of Positron Emission Tomography

1 Current Status of PET Technology������������������������������������������������������������   3

Suleman Surti and Joel S. Karp
2 Current Status and Future Directions of PET in Clinical Practice�������� 15
Daniel A. Pryma

Part II Solid State Digital PET Instrumentation, Technology

and Clinical Practice

3 Introduction to Silicon Photomultipliers for Time-of-Flight PET���������� 27

Dennis R. Schaart
4 New-Generation Silicon Photomultiplier-­Based
Clinical PET/CT and PET/MR Systems���������������������������������������������������� 41
David F. C. Hsu and Craig S. Levin
5 Solid-State Digital Photon Counting PET/CT������������������������������������������ 53
Jun Zhang and Michael V. Knopp
6 Siemens Biograph Vision 600����������������������������������������������������������������������   71
Michael E. Casey and Dustin R. Osborne

Part III Future Prospective of PET/CT Instrumentation

and Technology

7 Future Prospects of PET Instrumentation and Technology�������������������� 95

Suleman Surti and Joel S. Karp

Index�������������������������������������������������������������������������������������������������������������������� 105

vii
Contributors

Michael E. Casey, PhD Siemens Medical Solutions, Knoxville, TN, USA

University of Tennessee Graduate School of Medicine, Knoxville, TN, USA
David F. C. Hsu, PhD Electrical Engineering and Radiology at Stanford University,
Stanford, CA, USA
Joel S. Karp, PhD University of Pennsylvania, Philadelphia, PA, USA
Michael V. Knopp, MD, PhD The Ohio State University, Columbus, OH, USA
Craig S. Levin, PhD Radiology/Physics/Electrical Engineering/Bioengineering at
Stanford University, Stanford, CA, USA
Dustin R. Osborne, PhD, DABSNM Siemens Medical Solutions,
Knoxville, TN, USA
University of Tennessee Graduate School of Medicine, Knoxville, TN, USA
Daniel A. Pryma, MD Perelman School of Medicine at the University of
Pennsylvania, Philadelphia, PA, USA
Dennis R. Schaart, PhD Delft University of Technology, Radiation Science &
Technology Department, Delft, The Netherlands
Suleman Surti, PhD University of Pennsylvania, Philadelphia, PA, USA
Jun Zhang, PhD The Ohio State University, Columbus, OH, USA

ix
Abbreviations

APD Avalanche photodiode

CQIE Center for Quantitative Imaging Excellence
cPET Conventional PET
CS Compton scattering
DCR Dark count rate
DOI Depth of interaction
DPC Digital photon counting
dPET Digital PET
dSiPM Digital silicon photomultiplier
FDG 18F-fluorodeoxyglucose
FOV Field of view
FWHM Full width at half maximum
FWTM Full width at tenth maximum
HD High definition
LOR Line of response
Lu Lutetium
LYSO Lutetium-yttrium oxyorthosilicate
ML-EM Maximum likelihood expectation maximization
MOS Metal oxide semiconductor
NEC Noise equivalent count
NECR Noise equivalent count rate
NEMA National Electrical Manufacturers Association
OS-EM Ordered subsets expectation maximization
PDE Photon detection efficiency
PDPC Philips Digital Photon Counting
PE Photoelectric absorption
PMT Photomultiplier tube
PSF Point spread function
PVE Partial volume effect
QC Quality check
RC Recovery coefficient
SBR Sphere-to-background ratio
SD Standard definition
SiPM Silicon photomultiplier

xi
xii Abbreviations

SNR Signal-to-noise ratio

SPAD Single-photon avalanche diode
SUV Standardized uptake value
TDC Time-to-digital converter
TOF Time of flight
TSV Through-silicon vias
UHD Ultrahigh definition
 Part I
Basics Science of Positron Emission
Tomography
Current Status of PET Technology
1
Suleman Surti and Joel S. Karp

1.1 Introduction

In recent years, PET/CT imaging has played an important clinical role as a molecu-
lar imaging tool for diagnosis and staging of cancer in patients [1]. Used predomi-
nantly with 18F-fluorodeoxyglucose (FDG) as the radiotracer that acts as a glucose
analog, PET/CT has significantly influenced the management of cancer patients [2,
3] and is reimbursed for initial and follow-up imaging of most cancer types [4]. In
addition, PET has also been shown to play an important role in guiding cancer treat-
ment by characterizing the tumor biology as well as monitoring tumor response to
therapy [1, 5]. A more thorough overview of the current status in clinical practice is
given in Chap. 2. Modern time-of-flight (TOF) PET scanners provide sufficient sen-
sitivity and signal-to-noise ratio (SNR) performance so that clinical FDG scans with
excellent diagnostic quality can be completed in 10–15 min using bed translation to
cover a patient from “eyes to thighs.”
Beyond FDG, there are other tracers coming into more widespread use that
have very different imaging characteristics. Some of these tracers have lower pho-
ton flux because of lower dose (68Ga-labelled) and/or low positron emission
branching fraction (89Zr-labelled or 124I-labelled), requiring increased PET system
sensitivity to achieve reliable quantitative images especially for dose calibration.
In addition, the quantitative performance of images from some of these tracers
(e.g., 89Zr-labelled or 124I-labelled) will also require corrections for coincidence
data acquired in the presence of additional single photons. Finally, increased posi-
tron energy will require improvements in the point spread function (PSF) model
used during image reconstruction to better account for the increased range of the
positrons for some of these non-18F-labelled tracers. Hence, while modern PET

S. Surti (*) · J. S. Karp

University of Pennsylvania, Philadelphia, PA, USA
e-mail: surti@pennmedicine.upenn.edu; joelkarp@pennmedicine.upenn.edu

© Springer Nature Switzerland AG 2020 3

J. Zhang, M. V. Knopp (eds.), Advances in PET,
https://doi.org/10.1007/978-3-030-43040-5_1
4 S. Surti and J. S. Karp

scanners may provide excellent quality and quantitative FDG images, moving
PET in new areas requires continued technical improvements and capabilities.

1.2 Current Status of TOF PET/CT Scanners

TOF PET scanners were originally developed in the early 1980s [6–11] when the
primary application was in brain and cardiac imaging using compounds tagged with
short-lived radioisotopes, such as 15O-water, 11C-acetate, and 82Rb. While providing
good timing resolution as well as reduced dead time, the primary limitations of
these systems were lower sensitivity due to the use of interplane septa for 2D imag-
ing and poor spatial resolution arising due to the choice of scintillator and photosen-
sor. With the development of new scintillators in the late 1990s and early 2000s, a
new generation of TOF PET scanners, now all PET/CT, was introduced in the
mid-­2000s. These scanners were optimized for the primary application of detection
and staging of cancer using 18F-FDG. In addition to providing good system timing
resolution and spatial resolution, these scanners overcame the limitations of low
sensitivity by enabling fully 3D imaging (no interplane septa). All of these scanners
use lutetium-based scintillators (LSO and LYSO) and utilize light-sharing detectors
to achieve high spatial resolution (4–5 mm) with photomultiplier tubes (PMTs) of
25–39 mm size. Due to the detector design, the coincidence timing resolution of
these scanners lies within the range of 450–600 ps – very similar to the scanners
developed in the 1980s but with superior spatial resolution and sensitivity. Recent
years have seen an introduction of a new solid-state-based photosensor (silicon pho-
tomultiplier, or SiPM) that provides excellent intrinsic timing performance on par or
better than the conventional PMTs while providing flexibility in detector size that is
not available with PMTs. This has led to the commercial development of a new
generation of “digital” PET scanners using SiPM arrays with reduced or almost no
light sharing in the detector design. These new scanners have much improved sys-
tem coincidence timing resolution (210–390 ps) with similar or improved spatial
resolution through the use of smaller crystals. The benefits of TOF for clinical imag-
ing were well established [12–18], and so it is expected that improved TOF resolu-
tion will increase these benefits, particularly for patients with larger body mass
index (BMI).

1.3 Hardware Design

1.3.1 Scintillator

As mentioned earlier, lutetium-based scintillators are currently being used in all

modern commercial whole-body TOF PET scanners. This choice is driven by the
combination of high stopping power, high light output, and fast decay time of these
scintillators which leads to high system sensitivity as well as very good energy and
spatial and timing resolutions – all necessary characteristics for a modern fully 3D
1 Current Status of PET Technology 5

TOF PET system [19–21]. The two scintillators used most commonly are closely
related: cerium-doped lutetium oxy-orthosilicate (Lu2SiO5(Ce) or LSO(Ce)) and
lutetium-yttrium oxy-orthosilicate (Lu1.8Y0.2SiO5(Ce) or LYSO(Ce)). In the early to
mid-1990s, LSO(Ce), usually referred to as LSO, was first developed and intro-
duced as a PET scintillator [22] as a replacement for BGO. At the time BGO was
the primary scintillator being used in commercial PET but had limitations due to its
poor light output and long decay time – less than ideal properties for fully 3D PET
in comparison with NaI(Tl), which was also in use commercially [19]. While LSO
was first used in a small animal PET scanner [23] and subsequently incorporated
into a brain [24, 25] and a whole-body PET scanner [26], it was later recognized that
it also had very good timing resolution that could be used in the development of
TOF PET scanners [20, 21]. Following a similar trajectory, LYSO was first utilized
in a dedicated small animal PET scanner [27] and subsequently used in the produc-
tion of a new generation of commercial TOF PET/CT system [28]. While the cur-
rent version of the Lu-based scintillators all uses varying levels of Ce doping, there
have been efforts to change the dopant in order to achieve improved performance.
For instance, a co-doped version of LSO using calcium has been developed with
increased light output and shorter decay time than the LSO(Ce) scintillator [29],
leading to further improvements in timing resolution [30]. Similarly, calcium and
magnesium co-doped versions of LYSO have also been reported to produce higher
light output than cerium-doped LYSO [31], implying improved timing performance.
Thus, there is potential for improved lutetium-based scintillators that could be a
direct replacement for the current versions of commercially used LSO and LYSO.
While BGO is relatively inexpensive and was a preferred PET scintillator prior
to the development of LSO, the slow time scale of the luminescence process made
it impossible to use it for TOF PET. However, it has been noted that the passage of
charged electrons produced within BGO by the annihilation photons leads to the
emission of Cherenkov light that can be detected by the SiPM devices (high quan-
tum efficiency) [32]. The time scale of Cherenkov emission is very fast leading to a
very fast signal and potential for fast timing resolution that is appropriate for TOF
PET [33]. Several studies have been performed recently [34, 35] with best results
indicating a coincidence timing resolution as good as 330 ps (FWHM) can be
achieved with a 20-mm-long crystal [34]. However, the light output from Cherenkov
emission is very low; thus, it is still necessary to utilize the (slower) scintillation
light to determine both energy and spatial localization of the gamma interaction,
and it remains to be seen how practical it is to use Cherenkov timing for TOF PET
imaging with BGO.

1.3.2 Photosensor

Since the development of early PET scanners, PMTs have been the photosensor of
choice for all clinical systems. The high gain and consequently high signal-to-noise
ratio of the PMT signal lead to very good energy resolution, and plano-concave
photocathodes combined with careful dynode design have made it possible to
6 S. Surti and J. S. Karp

achieve very fast timing with cost-effective PMTs in sizes suitable (25–39 mm
diameter) to combine with multi-crystal PET detector arrays. The development of
new super and ultra bialkali, plano-concave photocathodes with increased quantum
efficiency (QE) and improved dynode structure to improve signal rise time has led
to further improvements in the timing resolution achieved with PMTs. Single-­
channel PMTs have been the standard in PET systems where a light-sharing method
is used to achieve spatial resolution significantly better than the PMT size. In addi-
tion to new photocathode materials, new fabrication methods have led to the devel-
opment of fast multi-anode PMTs where a single photocathode is shared by several
small anodes in a single PMT package. A fairly common multi-anode PMT design
has a 5 × 5 cm2 cross section with an 8 × 8 or 16 × 16 array of anodes for readout
(Fig. 1.1). These PMTs provide additional flexibility in developing fast PET detec-
tors with minimal light sharing to achieve high detector spatial resolution; however,
their complex design makes them considerably more expensive than the single-­
channel PMT.
The last 15–20 years has seen the introduction of a new solid-state photosensor
(SiPM) that is compact and fast, has high gain and low noise, and is insensitive to
magnetic fields [36–40]. Although this technology was initially expensive, the cost
has decreased as the devices become more widely used. Details about SiPM tech-
nology are discussed later in Chaps. 3–6.
Since SiPM devices can be fabricated in small sizes, as opposed to PMTs, they
provide great flexibility in developing high-resolution PET detectors. They also
operate at low bias voltage (few tens of volts) as opposed to PMTs (about thousand
volts), and being solid-state technology can be made nonmagnetic that is necessary
for PET/MR scanners that incorporate a PET detector ring inside the magnet bore

Fig. 1.1 From left to right, pictures of 25 mm diameter (R9800), 38 mm diameter (R9420), and
51 mm diameter (R7724) single-channel PMTs and a 64-channel H8500 MAPMT. All PMTs are
manufactured by Hamamatsu. The H8500 is 5 × 5 cm2 in size and the individual anodes are about
6.25 × 6.35 mm2. (Images courtesy of Hamamatsu)
1 Current Status of PET Technology 7

to enable simultaneous PET and MR scanning. Additional work to further improve

the performance of SiPMs is ongoing, ranging from increase in photon detection
efficiency (PDE) due to increased microcell density and quantum efficiency, as well
as the development of 3D digital SiPMs (wafer-level integration of the SiPM and
readout electronics). While the current version of digital SiPM technology [41, 42]
has increased the flexibility and performance to some degree, there are some limita-
tions in this design. For instance, a single time-to-digital converter (TDC) is used to
obtain timing information for one or more SiPM channels. However, there is a tim-
ing skew attached to the signal from each microcell based on its location within the
SiPM channel that leads to a degradation of the timing performance of the SiPM
channel due to the use of a common TDC. One aspect of 3D digital SiPM develop-
ment work is to fabricate a dedicated TDC for each microcell within a SiPM chan-
nel that will likely lead to a device with significantly improved intrinsic timing
performance. Early SiPM devices were a few millimeters in size, but recent fabrica-
tion techniques have led to the development of larger arrays of these devices that are
suitable for use in modern PET scanners by coupling to comparably sized scintilla-
tor arrays. Early SiPM arrays were fabricated using discrete SiPM devices con-
nected via bond wires on a common printed circuit board that led to relatively large
dead areas between each SiPM channel and hence a lower device PDE. More recent
developments have led to the production of SiPM arrays using through-silicon vias
(TSV) technology to interconnect the individual SiPM channels and significantly
reduce the dead area due to classical wire bonding. An ideal solution will be to fab-
ricate the entire SiPM array on common wafer to produce what are called mono-
lithic arrays. However, a disadvantage of this method is that fabrication errors could
lead to large variations in performance of individual channels on the device leading
to suboptimal performance – a limitation that is not present in the discrete arrays
where each channel can be chosen to provide a uniform performance.

1.3.3 Detector Design

The properties of the chosen scintillator and photosensor define the best intrinsic
performance that can be achieved by the PET detector. In particular, the crystal size
determines the detector spatial resolution (cross section) and sensitivity (thickness)
as well as light output that in turn affects the detector energy and timing resolution.
Standard commercial PET detectors have used Lu-based crystals that are 4–5 mm
wide and 18–25 mm thick. Until the recent advent of SiPM, PET detectors were
comprised of rectangular crystal arrays coupled to large PMTs (25–39 mm in diam-
eter) via a light-sharing technique such as that utilized in block detectors [43], quad-
rant sharing block detectors [44], or the pixelated Anger-logic detectors [45] (see
examples in Fig. 1.2). These types of detector designs are still in use in modern
PET/CT systems from all major manufacturers [46–49]. While all of these systems
provide very good overall performance, their TOF performance (coincidence timing
resolution in the range of 400–550 ps) is limited due to the practical choices made
in the detector design: light loss and transit time dispersion of scintillation photons
8 S. Surti and J. S. Karp

Fig. 1.2 Picture of a single detector module from a Philips Gemini TF scanner using a pixelated
Anger-logic detector design (left) and a Siemens Biograph mCT scanner using a block detector.
The Gemini TF (and newer Ingenuity TF) detector consists of a 23 × 44 array of 4 × 4 × 22 mm3
LYSO crystals coupled to a hexagonal array of 39-mm-diameter single-channel PMTs. The open
crystal area visible along the lower edge of the detector module would be read out with another row
of PMTs that are not shown here which will straddle this and the neighboring module to form a
continuous detector ring in the scanner. The mCT detector consists of a 13 × 13 array of
4 × 4 × 20 mm3 LSO crystals coupled to a 2 × 2 square array of 25-mm-diameter single-channel
PMTs. Compared to the Gemini TF detector, each mCT detector is completely independent of
neighboring detector modules. (Pictures courtesy of Philips Healthcare and Siemens Healthineers)

Amplifiers

Position encoding
Gain compensation

H10966A-100
(8 × 8 ancode)

L0.95GSO array
(15 × 15, 3 × 3 × 20 mm3)

Fig. 1.3 Pictures of two multi-anode PMT-based PET detectors used in a dedicated breast imag-
ing PET/tomosynthesis system under development at the University of Pennsylvania and a research
whole-body PET scanner. The detector on the left uses a 32 × 32 array of 1.5 × 1.5 × 15 mm3 LYSO
crystals, while the one on the right uses a 15 × 15 array of 3 × 3 × 20 mm3 LGSO crystals. (The
figures are reprinted with permission from [51] (left) and [52] (right))

as they undergo multiple reflections within a long, narrow crystal [20] and the
degrading effects due to the light-sharing mechanism of these PMT-based detectors
[50]. Some of these limitations can be mitigated with the use of multi-anode PMTs
(MAPMT) that require minimal or no light sharing due to small size of the indi-
vidual anodes (Fig. 1.3) [51]. As shown in Fig. 1.3, one can also appreciate the
20 D. A. Pryma

While PET/MR appears to have niches with potentially important clinical utility, it
is likely that it will remain a much smaller market segment compared to PET/CT.

2.3 Future Directions

Because of the high sensitivity and spatial resolution of PET compared with other
functional imaging modalities, there are innumerable ongoing trials testing PET
radiopharmaceuticals. These range from elemental salts such as 124I-sodium iodide
to radiolabeled intact antibodies or nanoparticles. The majority of agents in later
phase clinical trials are small molecules or peptides, most of which are labeled with
the relatively short-lived isotopes 18F and 68Ga, though some are labeled with longer-­
lived isotopes such as 64Cu (T½ 12.7 hours), 89Zr (T½ 3.3 days), or 124I (T½ 4.2 days).
Most agents under development seek to detect or characterize some specific
aspect of disease such as receptor status or antigen expression for one of two rea-
sons. Specific cancer detection agents have been developed for diseases that have
low FDG uptake. Alternatively, many agents have been developed more for disease
characterization and to evaluate the patient for a specific targeted therapy than for
detection. This is a reasonable approach given the high sensitivity but lower speci-
ficity of FDG PET/CT. However, because FDG PET/CT is imperfect with malig-
nant diseases that are poorly FDG avid and many benign processes that result in
false positives, general cancer detection radiopharmaceuticals have been developed
as potential FDG replacements. Finally, while oncologic imaging is the primary
current use of FDG PET/CT and the focus of much research, there is also a great
deal of work on PET radiotracers for non-oncologic diseases.
Fibroblast activation protein (FAP) is a protein associated with the fibroblasts
that are overexpressed in the vast majority of epithelial cancer stroma. Thus, it can
be possible to detect the tumor stroma rather than the tumor itself using a target that
should be more cancer-specific than FDG. Therefore, among potential targets for a
general cancer imaging radiotracer, FAP agents appear to be the most promising
[45, 46]. However, it is important to note that they are still very early in develop-
ment and also that the bar to supersede FDG PET/CT is quite high since FDG PET/
CT has excellent diagnostic accuracy despite its limitations.
Among agents developed for detection of poorly FDG avid diseases, radiophar-
maceuticals to detect prostate-specific membrane antigen (PSMA) are certainly the
closest to clinical use and most widely investigated, with two agents nearing likely
FDA approval [47–50]. These agents appear to have much higher sensitivity for
detection of disease than other available imaging techniques while preserving very
high specificity. They are also paired with PSMA-targeted therapeutics and so will
likely be used for both diagnostic and theranostic purposes.
Outside of oncology, there is a great deal of ongoing research on neurologic and
cardiac imaging. For example, given the ongoing opioid crisis, there is a great deal
of effort to use PET to help understand and, hopefully, develop treatments for addic-
tion [51, 52]. Additionally, development of tracers specific for the neurodegenera-
tion seen in Parkinson disease has been an ongoing effort [53, 54]. In cardiology,
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Cyathophyllidae, 394
Cyathophyllum, 394
Cycads, spermatozoa of, 38
Cyclidium, 137
Cyclocnemaria, 397
Cyclomyaria, 325
Cyclops, host of Choanophrya, 159;
of Rhyncheta and other Suctoria, 159 f., 162;
of Vorticellidæ, 158
Cycloseridae, 404
Cydippidea, 417
Cydippiform stage of Lobata and Cestoidea, 414
Cydonium milleri, 222
Cymbonectes, 306
Cymbonectinae, 306
Cyphoderia, 52
Cyrtoidea, 79
Cyst (a closed membrane distinct from the cytoplasm around a
resting-cell or apocyte), 37, 39;
cellulose-, 37;
chitinous, 37;
growth of vegetal cell in, 37;
of Protozoa present in dust, 47;
of Centropyxis aculeata, 57;
of Chlamydophrys stercorea, 57;
of Amoeba coli, 57;
of Actinophrys sol, 72;
of Actinosphaerium, 73 f.;
of Flagellata, 109, 117 f.;
brood-, of Paramoeba eilhardii, 116 n.;
of Bodo saltans, 117;
of Opalina, 123 f.;
of Volvocaceae, 128;
of Dinoflagellates, 131;
of Pyrocystis, 131, 132;
of Ciliata, 147;
of Colpoda cucullus, 147, 153;
 temporary (hypnocyst) of Rhizopoda, 57;
of Proteomyxa, 88;
of Myxomycetes, 91;
-wall, of Acystosporidae, 104 f.
Cystiactis, 382
Cystid—see Cystoidea
Cystiphyllidae, 394
Cystoflagellata, 110, 132 f.
Cystoidea, 580, 597 f.
Cytogamy, 33 f.
Cytoplasm, 6;
of ovum of Sea-urchin, 7;
granular, nutritive, of muscle cell, 19;
in cell-division by mitosis, 26 f.;
during syngamy, 34

Dactylometra, 311, 323;

D. lactea, 312
Dactylopores, 257
Dactylozooids, 264;
of Hydractinia, 264;
of Millepora, 259;
of Siphonophora, 299
Dale, on chemiotaxy, 22
Dallinger, W. H., and Drysdale, C., on Protozoa, 44, 45;
on organisms of putrefaction, 44, 116 f.;
on life-histories of Flagellates, 116 f.
Dallingeria, 111, 112, 119;
anchoring flagella of, 114;
D. drysdali, gametes of, 116 n.
Dalyell, 317 n., 375
Dangeard, on brood-division in active Chlamydomonadidae, 115;
on Flagellata, 119 n.
Dantec, Le, on protoplasmic movements, 16 n.;
on peptic digestion in Protozoa, 16
Darwin, 328 n., 360, 391
Darwinella, 221
Dasygorgiidae, 333 (= Chrysogorgiidae, 355)
Davenport on protoplasmic movements, 16 n., 19 n.
Dawydoff, 423
Dead men's fingers (= Alcyonium digitatum, 349)
Death, 11;
by diffluence, granular disintegration or solution, 14 f.;
by desolution, 15;
necessary, of colonial cells of Volvox, 128;
in Volvox and in Metazoa, compared, 130
Deep-sea deposits (Foraminifera), 70
Degen, on functions of contractile vacuole, 15 n.
Degeneration, senile, among Ciliata, 148
Deglutition in Podophrya trold, 159
—see also Ingestion of food
Deiopea, 419
Deiopeidae, 419
Delage, on protoplasm, 3 n.;
on syngamy, 34 n.;
on motion of flagella, 114 n.;
on Sponges, 168, 174, 226;
and Hérouard, on Protozoa, 46
Delap, M. J., 311 n.
Deltoid plate of Blastoidea, 599
Demospongiae, 195, 209 f.
Dendoryx, 224
Dendrite, 444
Dendrobrachia, 409
Dendrobrachiidae, 409
Dendroceratina, 209, 220, 221
Dendrochirota, 568, 569, 572, 577, 578
Dendrocometes, 159, 160, 161 f.
Dendrograptidae, 281
Dendrograptus, 281
Dendrophyllia, 404
Dendy, 188 n., 192, 274, 275
Depastrella, 321
Depastridae, 320, 321
Depastrum cyathiforme, 321
Depressor muscles of compasses of Echinus esculentus, 527
Dercitus bucklandi, 221
Dermal, gill—see Papula;
membrane, 170
Dermalia, 201
Dermasterias, 471
Desma (the megasclere which forms the characteristic skeletal
network of the Lithistida, an irregular branched spicule), 215,
224
Desmacella, 224
Desmacidon, 222
Desmophyes, 307
Desmophyinae, 307
Desmothoraca, 71
Desolution of protoplasm, 11 f.
Deutomerite, 98
Development, of Sponges, 226;
of Scyphozoa, 316;
of Alcyonaria, 341;
of Zoantharia, 373;
of Echinodermata, 601 f.
Dextrin, 15
Diadematidae, 531, 532, 538 f., 558
Diadematoid type of ambulacral plate, 531, 539
Dialytinae, 192
Diancistra (a spicule resembling a stout sigma, but the inner margin
of both hook and shaft thins out to a knife edge and is
notched), 222
Diaphorodon, 59;
shell of, 60
Diaseris, 404;
asexual reproduction, 388
Diatomaceae, skeleton of, 84;
symbiotic with Collosphaera, 86
Diatomin, 86;
(?) in coloured Flagellates, 115 n.
Dichoporita, 598, 599
Dicoryne, 268, 270
Dictyoceratina, 220
Dictyocha, 110
Dictyochida (= Silicoflagellata, 110), 79;
in Phaeocystina, 86 f.
Dictyocystis, 137;
test of, 152
Dictyonalia, 201
Dictyonema, 281
Dictyonina, 202
Dictyostelium, 90
Dicyclica, 594
Dicymba, 308
Dicystidae, 97
Didinium, 137;
trichocysts of, 143
Didymium, 90;
D. difforme, 92
Didymograptus, 282
Diffluence, 14 f.
Difflugia, 52;
D. pyriformis, 55;
test of, 55
Digestion, 9, 15 f.;
of reserves in brood-formation, 33;
in Carchesium, 147;
in Starfish, 440
Digestive system—see Alimentary Canal
Dileptus, 137, 152 n.
Dill, on Chlamydomonas, etc., 119 n.
Dimorpha, 70, 73, 75 n., 112
Dimorphism of chambered Foraminifera, 67 f.
Dinamoeba, 51;
test of, 53
Dinenympha, 111, 115;
undulating membranes of, 123
Dinobryon, 110, 112
Dinoflagellata, 110, 113, 130, 131, 132;
plastids of, 40;
nutrition of, 113
Dinoflagellate condition of young Noctiluca, 134
Diphyes, 303, 307
Diphyidae, 306
Diphyopsinae, 307
Diplacanthid, 457
Dipleurula, definition, 605;
forms of, 605-608
Diplocyathus, 277
Diplodal, 210
Diplodemia, 223
Diploëpora, 346
Diplograptus, 281, 282
Diplomita, 111
Diplopore, 597, 599
Diploporita, 598, 599
Diprionidae, 282
Directives, 367
Disc, of Vorticellidae, 155, 158;
of Ophiothrix fragilis, 484
Discalia, 309
Discohexaster, 200
Discoidea, 77
Discoidea, 558
Discomedusae (= Ephyropsidae, 322 + Atollidae, 322 + Discophora,
323)
Discomorpha, 137
Discooctaster, 200
Discophora, 310, 316, 323 f.
Discorbina, 59, 63;
reproduction of, 69
Discosomatidae, 383
Diseases, produced by Coccidiidae, 102;
by Acystosporidae, 103 f.;
 by Flagellates, 119 f.;
by Trypanosomes, 119 f.;
Protozoic organisms of, 43 f.
Dissepiments, 385, 387
Dissogony, 419
Distichopora, 284, 286
Distomatidae, 110
Distribution of Protozoa, 47;
of Sponges, in space, 239 f.;
in time, 241
Disyringa dissimilis, 209, 214, 215
Diverticulum—see Caecum
Division, binary, 10;
reduction-, 75 n.
Dixon and Hartog on pepsin in Pelomyxa, 16
Dobie, 167
Doederlein, 193 n.
Doflein, 46;
on parasitic and morbitic Protozoa, 94 n.;
on syngamy of Cystoflagellates, 135
Dohrn, on carnivorous habits of Sphaerechinus, 516
Dolichosporidia (= Sarcosporidiaceae), 98, 108
Doramasia, 306;
D. picta, 303
Dorataspis, 78;
skeleton, 80
Dorocidaris—see Cidaris
Dorsal elastic ligament of Antedon rosacea, 587
Dorso-central plate of Echinarachnius parma, 543
Dourine, disease of horses and dogs, 119
Drepanidium (= Lankesterella), 97, 102
Dreyer, on genera and species of Radiolaria, 87 f.;
on skeleton of Radiolaria, 82 n.
Dropsy, ascitic, associated with Leydenia, 91
Drysdale and Dallinger, on organisms of putrefaction, 44 f., 116 f.
Dual force of dividing cell, 26 f.
Duboscq, Léger and, sexual process in Sarcocystis tenella, 108 n.
Duerden, 261, 369 n., 371, 373, 374, 389, 397 n., 400 n., 403, 405,
406
Dujardin, on sarcode (= protoplasm), 3;
on Protozoa, 45;
on true nature of Foraminifera, 62 f.;
on Sponges, 167
Dust, containing cysts of Protozoa, 47;
of Flagellata, 118
Dysentery, in Swiss cattle, caused by Coccidium, 102;
tropical, caused by Amoeba coli, 57
Dysteria, 137, 153;
oral apparatus, 145;
shell, 141

Earthworm, Monocystis parasitic in, 95

Echinanthidae, 549
Echinarachnius, 548, 549;
E. parma, 542 f., 543, 544, 545, 547;
shape, 542;
sphaeridia, 545;
internal skeleton, 545;
habits, 546;
alimentary canal, 546;
Aristotle's lantern, 546;
tube-feet, 547
Echinaster, 439, 462
Echinasteridae, 455, 458, 462
Echinating, 217
Echinidae, 539, 558
Echininae, 539
Echinocardium, 549;
E. cordatum, 549 f., 551, 552;
habitat, 549;
shape, 550;
spines, 550;
sphaeridia, 551;
alimentary canal, 551;
 tube-feet, 551;
habits, 552;
stone-canal, 552;
E. flavescens, 555;
E. pennatifidum, 555
Echinoconidae, 558
Echinocyamus, 548, 549;
E. pusillus, 549
Echinocystites, 557
Echinodermata, 425 f.
Echinoid stage in the development of a Holothuroid, 615
Echinoidea, 431, 503 f.;
compared with Holothuroidea, 560;
with Blastoidea, 580;
mesenchyme of larva, 604;
development of, 607, 608, 609, 613, 614;
phylogeny of, 622
Echinolampas, 554
Echinometra, 542
Echinomuricea, 356
Echinoneus, 553, 553
Echinonidae, 553
Echinopluteus, 607, 608;
metamorphosis of, 613, 614
Echinosphaerites, 598;
E. aurantium, 598
Echinothuriidae, 530, 531, 532, 535, 558, 560
Echinus, 533, 539;
E. esculentus, 504 f. 505, 507, 511-515;
locality, 504;
spines, 506;
pedicellariae, 506;
corona, 511;
periproct, 513;
peristome, 513;
alimentary canal, 516;
water-vascular system, 516 f.;
 nervous system, 518 f.;
sphaeridia, 524;
perihaemal spaces, 524 f.;
genital system, 528;
blood-system, 529;
larva, 507;
E. acutus, 540;
pedicellariae, 509;
E. alexandri, pedicellaria, 510;
E. elegans, 539, 540;
pedicellariae, 510;
E. microtuberculatus, 540;
E. miliaris, 540, 542, 549;
E. norvegicus, 539, 540
Economic uses of Foraminifera, 69 f.
Ectocoele, 367
Ectoderm, 246
Ectoplasm (= ectosarc), 6, 46 f., 50;
of Amoeba, 5;
of Rhizopoda, 51 f.;
of Heliozoa, 71 f.;
of Radiolaria, 79 f. (see also Extracapsular protoplasm);
regeneration of, in Radiolaria, 35;
of Collozoum inerme, 76;
of Gregarines, 96 f.;
of Ciliata, 141 f.;
of Stylonychia, 140;
of Suctoria, 159;
of Trachelius ovum, 153;
of Vorticella, 156
Ectopleura, 268
Ectosarc—see Ectoplasm
Ectosome, 170
Ectyoninae, 217
Edrioasteroidea, 580, 596
Edwardsia, 328, 366, 368, 376;
E. allmani, 377;
 E. beautempsii, 376, 377;
E. carnea, 377;
E. goodsiri, 377;
E. tecta, 377;
E. timida, 376, 377
Edwardsia stage of Zoantharia, 367
Edwardsiidae, 377
Edwardsiidea, 367, 371, 375, 395
Egg, fertilised, 31;
of Metazoa, 32 f.;
of bird, 32;
various meanings of, 34;
of affected Silkworm moths transmitters of pébrine, 107
Ehrenberg, on Protozoa, 45 f.;
on skeletons of Radiolaria, 87 f.;
on Ciliata, 146;
on Suctoria, 162
Eimer and Fickert, on classification of Foraminifera, 58 n.
Elasipoda, 569, 571, 577, 578
Electric, currents, stimulus of, 19, 22;
shock, action on Amoeba, etc., 7
Eleutheria, the medusa of Clavatella, 265
Eleutheroblastea, 253
Eleutheroplea, 279
Eleutherozoa, 430, 560, 577, 579, 583;
development of, 602 f.;
larva of, 605;
phylogeny of, 621, 622
Elevator muscles of compasses of Echinus esculentus, 527
Ellipsactinia, 283
Ellis, 167
Embryonic type of development, 601
Encystment, 37, 39;
of animal cells, 37;
of vegetal cells, 37, 39;
growth during, 37;
of zygote, general in Protista, 34;
 of Rhizopoda, 57;
temporary, of Rhizopoda, 57 (see also Hypnocyst);
of Heliozoa, 72 f.;
of Actinophrys, 72;
of Actinosphaerium, 73 f.;
of Proteomyxa, 88, 89
of Myxomycete zoospores, 90 f.;
of Sporozoa, 96 f.;
of zygote of Sporozoa, 95 f.;
of Gregarines, 95 f., 98;
of Lankesteria, 95;
of Monocystis, 96;
of Coccidiidae, 97 f.;
of Coccidium, 100, 101;
of archespore or pansporoblast of Myxosporidiaceae, 107;
of Flagellates, 115, 117 f.;
of zygote of Bodo saltans, 117;
of Opalina, 123 f.;
of oosperm of Volvocaceae, 127 f., 129 f.;
of Dinoflagellates, 131;
of Ciliata, 147;
of Colpoda cucullus, 147, 153
Endocyclica, 529, 530 f., 556, 559
Endoderm, 246
Endogamy, in Amoeba coli, 57;
in Actinosphaerium, 73 f., 75 (diagram);
in Stephanosphaera, 128
Endogenous budding in Suctoria, 160 f., 162
Endoparasitic Suctoria, 86, 160 f.
Endoplasm (= endosarc, q.v.), 6
—see also Intracapsular protoplasm (Radiolaria)
Endoral, cilia, 139;
undulating membrane, 139
Endosarc (= endoplasm), 6;
of Gregarines, 95 f.;
branching, of Noctiluca, 110, 133;
of Loxodes and Trachelius, 144, 153;
 of Ciliata, 143 f.;
of Stylonychia, 140;
of Suctoria, 161
Endosphaera, 159, 161
Energy, changes of, in living organism, 8, 13;
sources of, 13 f.
Entocnemaria, 394
Entocoele, 367
Entosolenia, 66
Entz, Geza, on Choanoflagellates, 121 n.;
on structure of Vorticella, 157 n.
Eocene Foraminifera, 70
Eolis (= Aeolis), 248
Eophiura, 501
Eozoon, 70 n.
Epaulettes, 315
Epenthesis, 281
Ephelota, 159, 160;
E. bütschliana, cytological study of, 162
Ephydatia, 217, 225;
E. fluviatilis, structure, etc., 174 f., 176, 177, 178, 179
Ephyra, 317
Ephyropsidae, 322
Epiactis (usually placed in the order Zoanthidea, 404);
E. marsupialis, 379;
E. prolifera, 379
Epibulia, 308
Epidemic, of pébrine among Silkworms, 107;
among Fish, due to Costia necatrix, 119;
to Myxosporidiaceae, 107;
to Icthyophtheirius, 152
—see also Diseases, Fever
Epigonactis fecunda, 379
Epimerite of Gregarines, 97, 98 f.
Epineural canal, of Ophiothrix fragilis, 481;
of Echinus esculentus, 515
Epiphysis, of jaw, of Echinus esculentus, 526;
 of jaws of Diadematidae, 531;
absent in Cidaridae and Arbaciidae, 531
Epiphytic Protozoa, 48
Epiplasm (= cytoplasm of a brood-mother-cell remaining over
unused in brood-formation), 96
Epistrelophyllum, 403
Epistylis, 138, 158;
E. umbellaria, nematocysts of, 249
Epitheca, 386
Epizoanthus, 406;
on Hyalonema, 204;
E. glacialis, infested by Gregarines, 99;
E. incrustatus, 406;
E. paguriphilus, 406;
E. stellaris, 406
Epizoic, Protozoa, 48;
Ciliata, 158;
Suctoria, 158, 162
—see also Symbiosis
Equatorial plate (= the collective chromosomes at the equator of the
spindle in mitosis), 25, 27
Equiangular, 185
Errina, 284, 286;
E. glabra, 286;
E. ramosa, 286
Ersaea, 306;
E. picta, 303
Esperella, 225, 231
Esperiopsis, 225
Euaster (a true aster in which the actines proceed from a centre,
contrasting with the streptaster), 184
Eucalyptocrinus, 596
Eucharidae, 420
Eucharis, 420;
E. multicornis, 416, 418 f., 420
Enchlora, 417
Eucladia, 502
Euclypeastroidea, 549
Eucopidae, 277, 280
Eudendrium, 269, 270
Eudiocrinus, 594
Eudorina, 111, 128 f.
Eudoxia, 306;
E. eschscholtzii, 303
Euglena, 110;
barotaxy of, 20;
nutrition of, 113;
E. viridis, 124
Euglenaceae, 110, 124;
pellicle of, 113
Euglenoid motion, 124;
of Sporozoa, 50
Euglypha, 52;
 in fission, 29;
test, 29, 54
Eunicea, 356;
spicules, 335, 336
Eunicella, 356;
spicules, 335, 336;
E. cavolini, 356
Eupagurus prideauxii, 378, 381;
E. bernhardus, 378
Eupatagus, 553
Euphyllia, 401
Euplectella, 204;
E. aspergillum (Venus's Flower-Basket, 197);
E. imperialis, 206;
E. suberea, 202, 204, 205, 221
Euplexaura, 356
Euplokamis, 418
Euplotes, 138
Eupsammiidae, 402, 404
Eurhamphaea, 419
Eurhamphaeidae, 419
Euryalidae, 501
Eurypylous, 210
Euspongia, 221
Eutreptia, 110;
E. viridis, 124
Evacuation of faeces by mouth in Noctiluca, 133
Evans, 179 n., 217
Excretion, 13 f.;
in Sponges, 172;
in Asterias rubens, 437;
in Echinus esculentus, 527, 528;
in Antedon rosacea, 587
Excretory, granules, 6;
of Ciliata, 144;
pore of contractile vacuole of Flagellates, 110;
of Trachelius ovum, 153
Exogametes of Trichosphaerium, 54
Exogamy, 34 n.;
in Rhizopoda, 56 f.;
in Foraminifera, 68 f.
Expansion of Amoeboid cell, 16 f.
Exsert, septa, 398, 399
Extracapsular protoplasm, of Phaeodaria, 76;
of Radiolaria, 79 f. (see also Ectoplasm)
Eye of Asterias, 445 f., 446;
of Echinoidea, 512
Eye-spot of coloured Protista, 21, 125 f.

Fascicularia, 348
Fasciole, of Echinocardium, 550, 555;
of Spatangoidea, 553;
of Spatangus, 553;
of Eupatagus, 553;
of Spatangidae, 555
Fats, fatty acids, 15;
in Flagellates, 110, 115;
formation of, 36
Fauré-Fremiet, on attachment of Peritrichaceae, 141 n.
Faurot, 368
Favia, 373, 401
Favosites, 344
Favositidae, 344
Feather-star, 581
Feeding, of Noctiluca, 133, 144;
of Peritrichaceae, 145
—see also Food
Feeler, of Holothuria nigra, 561 f., 566;
of Holothuroidea, 568;
of Dendrochirota, 568, 572;
of Synaptida, 568, 575;
of Molpadiida, 568, 575
Female gamete, 33;
of Pandorina, 128 f.;
 of Acystosporidae, 104 f.;
of Peritrichaceae, 151, 157
—see also Megagamete, Oosphere
Ferment, required for germination, brood-formation, etc., 32 f.
—see also Zymase
Fermentation, organisms of, 43 f.
Fertilisation, 33 f.;
"chemical," 32 n.
Fertilised egg, 31
—see also Oosperm, Zygote
Fertilising tube of Chlamydomonas, 125
Fever, intermittent, malarial, 103 f.;
relapsing, 121;
remittent, 105;
Texas-, Tick, 120;
Trypanosomic, 119 f.
Fewkes, 268 n.
Fibularidae, 549
Fibularites, 559
Fickert, Eimer and, on classification of Foraminifera, 58 n.
Ficulina, 219, 224, 230;
F. ficus, 219
Filoplasmodieae, 90 f.
Filopodia, 47 n.
Filosa, 29, 50, 52 f.;
resemblance to Allogromidiaceae, 59
Finger, 580;
of Cystoidea, 597;
of Blastoidea, 599, 600
Firestone of Delitzet contains fossil Peridinium, 132
Fischer, on fixing reagents, 11;
on structure of flagellum, 114
Fish, rheotaxy of, 21;
epidemics of, due to Myxosporidiaceae, 107;
to Costia necatrix, 119;
to Ichthyophtheirius, 152
Fission, 10, 23 f.;
 equal, 10;
Spencerian, 23;
multiple, 30 f. (see also Brood-division);
of Heliozoa, 72 f.;
of Radiolaria, 84 f.;
radial, in Volvocaceae, 110;
transverse, in Craspedomonadidae, 115 n.;
longitudinal and transverse, of Bodo saltans, 117;
of Opalina, 123;
of Euglenaceae, 124;
of Eutreptia viridis, 124;
of Noctiluca, 133;
of Ciliata, 147 f.;
of Stentor polymorphus, 156;
of Vorticellidae, 157 f.
—see also Bud-fission
Fissiparantes, 387, 400
Fixing protoplasm, 15
Flabellum, 375, 386, 398;
protandry of, 370
Flagella, flagellum, 17 f., 47;
of Protozoa, 47;
formed by altered pseudopodia in Microgromia, 60;
of Heliozoa, 73;
of sperms of Coccidiidae, 102;
of Acystosporidae, 105;
of Flagellata, 109, 114 f.;
of Trichonymphidae, 114;
Delage on mechanism of, 114 n.;
of Bodo saltans, 117;
of Trypanosoma, 121;
of Euglenaceae, 124 f.;
of Maupasia, 124;
of Eutreptia viridis, 124;
of Sphaerella, 126;
of Dinoflagellata, 130, 131;
of Peridinium, 131;
 of Polykrikos, 132;
of Noctiluca, 132, 133 f.
—see also Sarcoflagellum
Flagellar pit, in Flagellates, 110, 124 f.
Flagellata, 17 f., 40, 48 f., 50, 109 f.;
barotaxy of, 20;
galvanotaxy of, 22;
chemiotaxy of, 23;
nutrition of, 40, 113;
of putrefying liquids, 44, 116 f.;
studied by botanists, 45;
as internal parasites, 48, 119 f.;
relations with Acystosporidae, 106;
shell of, 113;
stalk of, 113;
life-history of, 116 f.;
literature of, 119;
saprophytic, 119 f.
Flagellate stage, of Sarcodina, 56 f., 60, 109;
of Heliozoa, 74;
of Radiolaria, 85 f.
—see also Flagellula
Flagellated chamber, 170
Flagellula, 31;
of Proteomyxa, 88, 89;
of Myxomycetes, 91, 92;
of Didymium, 92
—see also Zoospores
Flagellum—see Flagella
Fleming, 168 n.
Flexible Corals, 326
Flint, 219, 241
Floricome, 203
Floscelle, of Echinocardium cordatum, 551;
of Cassidulidae, 554
Flowering plants, male cells of, 38
Flowers of tan (= Fuligo varians), 92 f.;