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The Human Brain during the First

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Lengths: Atlas of Human Central
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Edition Shirley A. Bayer
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The Human Brain during the
First Trimester 40- to 42-mm
Crown-Rump Lengths
This sixth of 15 short atlases reimagines the classic 5-volume Atlas of Human Central Nervous System Development.
This volume presents serial sections from specimens between 40 mm and 42 mm with detailed annotations, together
with 3D reconstructions. An introduction summarizes human CNS development by using high-resolution photos of
methacrylate-embedded rat embryos at a similar stage of development as the human specimens in this volume. The
accompanying Glossary gives definitions for all the terms used in this volume and all the others in the Atlas.

Features
• Classic anatomical atlas
• Detailed labeling of structures in the developing brain offers updated terminology and the identification of unique
developmental features, such as, germinal matrices of specific neuronal populations and migratory streams of young
neurons
• Appeals to neuroanatomists, developmental biologists, and clinical practitioners
• A valuable reference work on brain development that will be relevant for decades
ATLAS OF
HUMAN CENTRAL NERVOUS SYSTEM DEVELOPMENT
Series

Volume 1: The Human Brain during the First Trimester 3.5- to 4.5-mm Crown-Rump Lengths

Volume 2: The Human Brain during the First Trimester 6.3- to 10.5-mm Crown-Rump Lengths

Volume 3: The Human Brain during the First Trimester 15- to 18-mm Crown-Rump Lengths

Volume 4: The Human Brain during the First Trimester 21- to 23-mm Crown-Rump Lengths

Volume 5: The Human Brain during the First Trimester 31- to 33-mm Crown-Rump Lengths

Volume 6: The Human Brain during the First Trimester 40- to 42-mm Crown-Rump Lengths

Volume 7: The Human Brain during the First Trimester 57- to 60-mm Crown-Rump Lengths

Volume 8: The Human Brain during the Second Trimester 96- to 150-mm Crown-Rump Lengths

Volume 9: The Human Brain during the Second Trimester 160- to 170-mm Crown-Rump Lengths

Volume 10: The Human Brain during the Second Trimester 190- to 210-mm Crown-Rump Lengths

Volume 11: The Human Brain during the Third Trimester 225- to 235-mm Crown-Rump Lengths

Volume 12: The Human Brain during the Third Trimester 260- to 270-mm Crown-Rump Lengths

Volume 13: The Human Brain during the Third Trimester 310- to 350-mm Crown-Rump Lengths

Volume 14: The Spinal Cord during the First Trimester

Volume 15: The Spinal Cord during the Second and Third Trimesters and the Early Postnatal Period
The Human Brain during the
First Trimester 40- to 42-mm
Crown-Rump Lengths
Atlas of Human Central Nervous System
Development, Volume 6

Shirley A. Bayer
Joseph Altman
First edition published 2023
by CRC Press
6000 Broken Sound Parkway NW, Suite 300, Boca Raton, FL 33487-2742

and by CRC Press


4 Park Square, Milton Park, Abingdon, Oxon, OX14 4RN

CRC Press is an imprint of Taylor & Francis Group, LLC

© 2023 Shirley A. Bayer and Joseph Altman

Reasonable efforts have been made to publish reliable data and information, but the author and publisher cannot assume responsibility for the
validity of all materials or the consequences of their use. The authors and publishers have attempted to trace the copyright holders of all material
reproduced in this publication and apologize to copyright holders if permission to publish in this form has not been obtained. If any copyright mate-
rial has not been acknowledged please write and let us know so we may rectify in any future reprint.

Except as permitted under U.S. Copyright Law, no part of this book may be reprinted, reproduced, transmitted, or utilized in any form by any elec-
tronic, mechanical, or other means, now known or hereafter invented, including photocopying, microfilming, and recording, or in any information
storage or retrieval system, without written permission from the publishers.

For permission to photocopy or use material electronically from this work, access www.copyright.com or contact the Copyright Clearance Center,
Inc. (CCC), 222 Rosewood Drive, Danvers, MA 01923, 978-750-8400. For works that are not available on CCC please contact mpkbookspermissions@
tandf.co.uk

Trademark notice: Product or corporate names may be trademarks or registered trademarks and are used only for identification and explanation
without intent to infringe.

Library of Congress Cataloging‑in‑Publication Data

LCCN 2022008216

ISBN: 978-1-032-18334-3 (hbk)


ISBN: 978-1-032-21937-0 (pbk)
ISBN: 978-1-003-27065-2 (ebk)

DOI: 10.1201/9781003270652

Publisher’s note: This book has been prepared from camera-ready copy provided by the authors.

Access the Support Material: www.routledge.com/9781032183343

Typeset in Times Roman by KnowledgeWorks Global Ltd.


CONTENTS

$&.12:/('*0(176 YL
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PART I. INTRODUCTION -------------------------------------------------------------------------------------------1
Organization of the Atlas -----------------------------------------------------------------------------------1
Plate Preparation ---------------------------------------------------------------------------------------------1
Development in Specimens (CR 40 42 mm) ---------------------------------------------------------2
References --------------------------------------------------------------------------------------------------- 13

PART II. 42-mm Crown-Rump Length, M841, Frontal/Horizontal ----------------------------------------- 14


Plates 1–20 A–B ------------------------------------------------------------------------------------------16–55

PART III. 42-mm Crown-Rump Length, C886, Horizontal ---------------------------------------------------- 56


Plates 21–22 A–B ----------------------------------------------------------------------------------------58–61
Plates 23–30 A–D -----------------------------------------------------------------------------------------62–93
Plates 31–38 A–B -------------------------------------------------------------------------------------- 94–109

PART IV. 40-mm Crown-Rump Length, C6658, Sagittal ------------------------------------------------------110


Plates 39–49 A–D ------------------------------------------------------------------------------------- 112-155
+LJK0DJQL¿FDWLRQ3ODWHV–65 A–B-------------------------------------------------------------- 156-187
ACKNOWLEDGMENTS

We thank the late Dr. William DeMyer, pediatric neurologist at Indiana University Medical Center,
for access to his personal library on human CNS development. We also thank the staff of the National
Museum of Health and Medicine that were at the Armed Forces Institute of Pathology, Walter Reed Hos-
pital, Washington, D.C. when we collected data in 1995 and 1996: Dr. Adrianne Noe, Director; Archibald
J. Fobbs, Curator of the Yakovlev Collection; Elizabeth C. Lockett; and William Discher. We are most
grateful to the late Dr. James M. Petras at the Walter Reed Institute of Research who made his darkroom
facilities available so that we could develop all the photomicrographs on location rather than in our labo-
ratory in Indiana. Finally, we thank Chuck Crumly, Neha Bhatt, and Kara Roberts, Michele Dimont, and
Rebecca Condit for expert help during production of the manuscript.
Taylor & Francis
Taylor & Francis Group
http://taylorandfrancis.com
YLLL

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1

INTRODUCTION

ORGANIZATION OF THE ATLAS in the horizontal plane, and C6658 is cut in the sagittal
This is the sixth book in the Atlas of Human Central plane. The three section planes in specimens of the same
Nervous System Development series, 2nd Edition. It deals age give a more complete perspective of the structure of
with human brain development in three normal specimens the brain at this time. As in the previous volumes of the
during the middle first trimester with crown-rump (CR) Atlas, each specimen is presented in serial grayscale pho-
lengths from 40- to 42-mm and estimated gestation weeks tographs of its Nissl-stained sections showing the brain
(GW) from 9.5 to 9.6 (Loughna et al., 2009). These spec- and surrounding tissues (Parts II–IV). The photographs
imens were analyzed in Volume 4 of the 1st Edition (Bay- are shown from anterior to posterior (frontal/horizontal
er and Altman, 2006). One specimen (M841) is from the specimen), dorsal to ventral (horizontal specimen), and
Minot Collection.1 The other two (C886 and C6658) are medial to lateral (sagittal specimen). The dorsal part of
from the Carnegie Collection.2 Both collections are now each frontal/horizontal section is toward the top of the
in the National Museum of Health and Medicine, which page, the ventral part at the bottom, and the midline is in
used to be located at the Armed Forces Institute of Pathol- the vertical center. In the horizontal specimen, the left side
ogy (AFIP) in Walter Reed Hospital in Washington, D.C. of the section is anterior, right side, posterior, and the mid-
When the AFIP closed, the National Museum moved to line is in the horizontal center. In the sagittal specimen,
Silver Springs, MD; this collection is still available for re- the left side of each section is anterior, right side posterior,
search. M841 is cut in the frontal/horizontal plane, C886 top side dorsal, and bottom side ventral.

1. The Minot Collection (designated by an M prefix in the specimen PLATE PREPARATION


number) is the work of Dr. Charles Sedgwick Minot (1852–1914), an
embryologist at Harvard University. Throughout his career, Minot col- All sections of a given specimen were photographed at
lected about 1900 embryos from a variety of species. The 100 human the same magnification. Sections throughout the entire
embryos in the group were probably acquired between 1900 and 1910. specimen were photographed in serial order with Kodak
From our examination of these specimens and their similar appearance,
we assume that they are preserved in the same way, although we could technical pan black-and-white negative film (#TP442).
not find any records describing fixation procedures. The slides contain The film was developed for 6 to 7 minutes in dilution F of
information on section numbers, section thickness (6 µm or 10 µm), and Kodak HC‑110 developer, stop bath for 30 seconds, Ko-
stain (aluminum cochineal).
dak fixer for 5 minutes, Kodak hypo‑clearing agent for 1
2. The Carnegie Collection (designated by a C prefix in the specimen minute, running water rinse for 10 minutes, and a brief
number) started in the Department of Embryology of the Carnegie rinse in Kodak photo‑flo before drying. Negatives were
Institution of Washington. It was led by Franklin P. Mall (1862–1917),
George L. Streeter (1873–1948), and George W. Corner (1889–1981). scanned as color positives at 2700 dots per inch (dpi) with
These specimens were collected during a span of 40 to 50 years and a Nikon Coolscan‑1000 35-mm negative film scanner at-
were histologically prepared with a variety of fixatives, embedding tached to a Macintosh PowerMac G3 computer which had
media, cutting planes, and histological stains. Early analyses of speci-
mens were published in the early 1900s in Contributions to Embryology, a plug‑in driver built into Adobe Photoshop. Images were
The Carnegie Institute of Washington (now archived in the Smithsonian converted to 300 dpi using the non‑resampling method for
Libraries). O’Rahilly and Müller (1987, 1994) have given overviews of image size. Using the powerful features of Adobe photo-
some first trimester specimens in this collection.
2
shop, contrast was enhanced, uneven staining was correct SUHVHQW WRVKRZWKH¿QHGHWDLOVRILPPDWXUHVWUXF
ed, and areas of uneven exposure were slightly darkened tures in the medulla, pons, cerebellum, superior colliculus,
or lightened. FHUHEUDOFRUWH[DQGROIDFWRU\EXOE Figs. 1–8 EHFDXVHWKH
preservation of human specimens does not show detail.
The photos chosen for annotation in Parts II–IV are
presented as companion plates. The ORZPDJQL¿FDWLRQ The structures we are showing in the rat embryos are
plates RI WKH IURQWDOKRUL]RQWDO DQG KRUL]RQWDO VSHFLPHQV some of the immature features in the brain at this time.
are designated as A and B on one set of facing pages. Part In the medulla, young neurons in the posterior extramu
ARQWKHOHIWVKRZVWKHIXOOFRQWUDVWSKRWRZKLOHPart B UDOPLJUDWRU\VWUHDPDUHVWLOOFURVVLQJWKHPLGOLQH Fig. 1 
RQWKHULJKWVKRZVDORZFRQWUDVWFRS\ZLWKDQQRWDWLRQV although in fewer numbers than shown in the specimens
Plates of the sagittal specimen are designated as A through DQDO\]HGLQ9ROXPH %D\HUDQG$OWPDQLQSUHVV 0DQ\
D on two sets of facing pages. Part A on the left page RI WKHVH QHXURQV DULVLQJ LQ WKH SUHFHUHEHOODU 1(3 LQ WKH
VKRZVWKHIXOOFRQWUDVWSKRWRJUDSKRIWKHEUDLQLQWKHVNXOO GRUVDOPHGXOOD KDYHDOUHDG\VHWWOHGLQWKHODWHUDOUHWLFXODU
without labels. Part BRQWKHULJKWSDJHVKRZVORZFRQ DQGH[WHUQDOFXQHDWHQXFOHL%XWWKHSUHFHUHEHOODU1(3LV
trast copies of the same photograph with superimposed VWLOOWKHPRVWDFWLYHJHUPLQDO]RQHLQWKHPHGXOOD Fig. 2 
outlines of brain parts and labels of major brain ventricles and is now generating neurons that will migrate to the pons
and structures. Part C on the second left page shows a and settle in the pontine nuclei. The pons does not show
IXOOFRQWUDVW SKRWR RI D VOLJKWO\ ODUJHU EUDLQ ³GLVVHFWHG´ settling neurons in the pontine nuclei, but they are on their
from its peripheral structures, except cranial sensory struc ZD\LQWKHDQWHULRUH[WUDPXUDOPLJUDWRU\VWUHDPDGH¿QLWH
tures have been preserved. Part D on the second right VWUXFWXUHRQWKHDQWHULRUVXUIDFH Fig. 3 
SDJHLVDORZFRQWUDVWFRS\RIWKHSKRWRLQ&ZLWKPRUH
detailed labeling in the brain. Several KLJKPDJQL¿FDWLRQ 7KH FHUHEHOOXP Fig. 4  LV JUHDWO\ H[SDQGLQJ  0DQ\
plates feature enlarged views of the brain to show tissue or GHHSQXFOHDUQHXURQVDUHOHDYLQJVXSHU¿FLDOSDUWVWRVHWWOH
ganization. This allows users to see the entire section and beneath the growing cortex. The cortical area is always
WKHQFRQVXOWWKHGHWDLOHGPDUNXSLQWKHORZFRQWUDVWFRS\ limited to the part just beneath the mitotically active exter
on the facing page, leaving little doubt about what is being QDOJHUPLQDOOD\HU HJO ZKLFKLVRQHSURQJRIDJHUPLQDO
LGHQWL¿HG7KHODEHOVWKHPVHOYHVDUHQRWDEEUHYLDWHGVR trigone. Purkinje cells are accumulating beneath the egl
WKHUH LV QR ORRNXS RQ D OLVW  'LIIHUHQW IRQWV DUH XVHG WR DQGVRPH¿EHUVDUHDOUHDG\LQDSULPRUGLDOPROHFXODUOD\
label different classes of structures: the ventricular system er. The cerebellar NEP itself is still mitotically active and
is labeled in CAPITALS, the neuroepithelium and other LV PDNLQJ SURJHQLWRUV RI *ROJL FHOOV WKDW ZLOO HYHQWXDOO\
germinal zones in Helvetica bold, transient structures in disperse throughout the granule cell layer.
Times bold italicDQGSHUPDQHQWVWUXFWXUHVLQ7LPHV5R
man or Times bold. Using Adobe Illustrator, labels were Another immature feature is in the tectum of the mid
superimposed and lines were drawn around structural de EUDLQ:HLOOXVWUDWHWKHHYROYLQJOD\HULQJLQWKHVXSHULRU
WDLOVRQWKHORZFRQWUDVWLPDJHV3ODWHVZHUHSODFHGLQWRD FROOLFXOXV Fig. 5  ZKHUH OHQVOLNH EXQGOHV RI D[RQV DUH
ERRNOD\RXWXVLQJ$GREH,Q'HVLJQ)LQDOO\KLJKUHVROX surrounded by migrating neurons in a honeycomb ma
WLRQSRUWDEOHGRFXPHQW¿OHV SGI ZHUHXSORDGHGWR&5& trix. It is our hypothesis that young neurons intermingle
3UHVV7D\ORU )UDQFLVZHEVLWHV DQGKDYHD³KDQGVKDNH´ZLWKWKHD[RQVWKDWZLOOLQQHUYDWH
these neurons in the mature brain.
DEVELOPMENT IN SPECIMENS
(CR 40–42-mm) 7KH HYROYLQJ OD\HULQJ RI WKH QHRFRUWH[ Fig. 6  LQ WKH
The specimens in this volume are equivalent to rat em telencephalon shows a thick cortical plate throughout its
EU\RVRQHPEU\RQLFGD\ ( EDVHGRQRXUPRUSKRORJL HQWLUHH[WHQW&DMDO5HW]LXVQHXURQVDUHSURPLQHQWLQOD\HU
FDOPDWFKLQJ(UDWVKDYHDVLPLODUDSSHDUDQFHWRKXPDQ I, while subplate neurons are delaminating from the lower
VSHFLPHQVIURPWRPPFURZQUXPSOHQJWKV2XU SDUWVRIWKHFRUWLFDOSODWHDQGDUHLQWHUPLQJOLQJZLWK¿EHUV
timetables of neurogenesis use +WK\PLGLQHGDWLQJPHWK in the glial channels at the base of the plate. A wide swath
RGV %D\HUDQG$OWPDQSUHVHQW%D\HU of migrating neurons lies beneath the subplate—the strati
HWDO WRGHWHUPLQHQHXURQDOSRSXODWLRQVWKDW ¿HGWUDQVLWLRQDO¿HOG 67) ZKHUHS\UDPLGDOFHOOVWKDWZLOO
DUHEHLQJJHQHUDWHGLQ(UDWVZHDVVXPHWKDWLVFRPSD reside in various layers of the mature neocortex sojourn to
UDEOHWRQHXURJHQHVLVLQWRPPKXPDQVSHFLPHQV intermingle with incoming axons from the thalamus and
%D\HUHWDO%D\HUDQG$OWPDQ Table other parts of the brain. Indeed, the STF is an important
1 lists populations being generated throughout the neurax staging area for the maturation of cortical circuitry.
LVWKHEUDLQVWHPDQGPLGEUDLQWHFWXP Table 1A WKHGL
HQFHSKDORQ Table 1B WKHSDOOLGXPVWULDWXPDP\JGDOD The hippocampus is another immature telencephalic
DQGVHSWXP Table 1C DQGWKHFHUHEUDOFRUWH[KLSSRFDP VWUXFWXUH Fig. 7 ZKHUHDIHZQHXURQVDUHVHWWOLQJLQWKH
SXVDQGROIDFWRU\VWUXFWXUHV Table 1D):HXVHSKRWRV &$¿HOGVRI$PPRQ¶VKRUQ0RVWOLNHO\WKHVHZLOOUHVLGH
RI PHWKDFU\ODWHHPEHGGHG UDW HPEU\RV RQ ( %D\HU LQ¿HOG&$7KHWKLFNKLSSRFDPSDO1(3KDVDQRWFKHG
3
Table 1A: Neurogenesis by Region Table 1C: Neurogenesis by Region
REGION and CROWN RUMP LENGTH REGION and CROWN RUMP LENGTH
NEURAL POPULATION NEURAL POPULATION 40-42 mm
40-42 mm
PRECEREBELLAR NUCLEI PALLIDUM AND STRIATUM

Pontine nuclei Olfactory tubercle (small neurons)

SUPERIOR COLLICULUS Caudate and putamen


Nucleus accumbens
stratum album
Islands of Calleja
stratum lemnisci
AMYGDALA
stratum griseum intermediate
Central nucleus
stratum griseum superficial
Intercalated masses
INFERIOR COLLICULUS Amygdalo-hippocampal area
Anterior intermediate
SEPTUM
Posterior intermediate
Lateral nucleus
Anteromedial
Posteromedial

Table 1D: Neurogenesis by Region


Table 1A. Neural populations in the precerebellar nuclei, mesencephalic
REGION and CROWN RUMP LENGTH
tegmentum, superior colliculus, and inferior colliculus that are being NEURAL POPULATION 40-42 mm
generated in rats on Embryonic day (E) 17 (comparable to humans at CR
NEOCORTEX and LIMBIC CORTEX
31-33-mm) . Green dots indicate the amount of neurogenesis occurring:
one dot=<15%; two dots=15-90%. This same dot notation is used for Layer V
all of the remaining parts (B-E) of Table 1. Layers IV-II
OLFACTORY CORTEX
Layer II (anterior)
Layer II (posterior)
Table 1B: Neurogenesis by Region
REGION and CROWN RUMP LENGTH Layers III-IV (anterior)
NEURAL POPULATION 40-42 mm HIPPOCAMPAL REGION
PREOPTIC AREA/ Entorhinal cortex Layer III
HYPOTHALAMUS
Medial preoptic nucleus Subiculum (deep)

Sexually dimorphic nucleus Subiculum (superficial)

Periventricular preoptic nucleus Ammon’s Horn CA1

Median preoptic nucleus Ammon’s Horn CA3


Arcuate nucleus Ammon’s Horn CA4
Tuberommillary nucleus OLFACTORY BULB

Medial mammillary n. (ventral) Internal tufted cells (main bulb)


THALAMUS/EPITHALAMUS External tufted cells (main bulb)
Paraventricular ANTERIOR OLFACTORY NUCLEUS
Paratenial Pars externa
Medial habenula AON proper

curve that extends downward to connect with the choroid These cells appear to enter the bulb from the lateral side,
plexus. Part of that germinal zone is probably a glioepi- where their trailing axons accumulate in the lateral olfacto-
thelium that will generate oligodendroglia in the fimbria, ry tract. Once inside, migrating cells fan out and peel off
but most of it is the source of the dentate migration. Many in curved arrays to settle in the primordial mitral cell layer
cells are migrating from the NEP in the “dentate notch” throughout the entire extent of the main olfactory bulb.
and accumulate beneath Ammon’s horn. This NEP is the
source of a dispersed germinal matrix that will produce All terms used in the tables, figures, and plates identify
granule cells in the dentate gyrus. A clump of cells is al- maturing and unique developmental structures that are of-
ready visible on E18. The dentate migration is visible in a ten unfamiliar or have long been forgotten from courses in
rudimentary form in the specimens analyzed in Volume 4 medical and graduate school. To refresh memories, please
of this series (Bayer and Altman, in press). consult the extensive definitions of these terms in the Glos-
sary (Bayer, in press) that accompanies the Atlas.
The main developmental event in the olfactory structures
is the mitral cell migration into the olfactory bulb (Fig. 8).
4
THE LOWER MEDULLA IN AN E18 RAT EMBRYO
A
Reticular formation
TECTUM

Midline raphe
TEGMENTUM

PONS
Inferior olive
CEREBELLUM
Posterior intramural
Posterior extramural Ascending
tracts from the
Migratory streams spinal cord
MEDULLA
A
0.5 mm
B

0.1 mm
Posterior extramural migratory
stream crosses midline

Figure 1. Frontal/horizontal section of the posterior brain in an E18 rat embryo at a similar stage as human specimens in this volume. The two posterior migratory streams (arrows) of precere-
bellar nuclear neurons are visible. The waning intramural stream takes a trajectory through the medullary parenchyma and heads for the inferior olive. The extramural stream runs beneath the pia
meninx on the outer edge of the medulla. Neurons cross the midline beneath the inferior olive and will settle on the opposite side in the lateral reticular and external cuneate precerebellar nuclei.
(3µ methacrylate section, toluidine blue stain). Source: braindevelopmentmaps.org (E16 coronal archive)
Figure 2. Frontal/horizontal section of the elaborate precerebellar
THE PRECEREBELLAR NEUROEPITHELIUM neuroepithelium (red outline) in an E18 rat embryo at a similar
stage as human specimens in this volume. This is the ger-
IN AN E18 RAT EMBRYO minal source of the pontine reticular and pontine pre-
cerebellar nuclei that are currently being generated.
Young neurons migrate out of the neuroepithelium
(arrows) to form the anterior extramural
migratory stream that travels beneath the
pia to the pons. (3µ methacrylate sec-
tion, toluidine blue stain). Source:
braindevelopmentmaps.org (E18
coronal archive)

Anterior extramural
migratorystream

Choroid plexus

Young neurons migrate


forward toward the pons

0.1 mm

5
6

THE ANTERIOR PONTINE SURFACE


IN AN E18 RAT EMBRYO
migratorystream
amural
or extr
nteri A

Figure 3. The anterior extramural migratory stream (red


outline) is a lens-shaped structure on the anterior pontine
surface at the base of the mesencephalic flexure. The neu-
rons are migrating perpendicular to the section plane, so
they are rounded rather than spindle-shaped. These cells
will settle in the core of the large pontine precerebellar
nuclei at the base of the pons in the mature brain. (3µ
methacrylate section, toluidine blue stain). Source: brain-
developmentmaps.org (E17 coronal archive)

0.1 mm
7

THE CEREBELLUM
IN AN E18 RAT EMBRYO Fiber bundles in the cerebellum
may be the primordial inferior
(deep) and superior (superficial)
A peduncles.
A

Locus coeruleus
Superior peduncle?

Migrating
deep nuclear
neurons

Inferior peduncle?

Migrating
Purkinje
neurons 1 External
germinal
layer (egl)

Note that inferior peduncle fibers


are surrounded by migrating
Cerebellar neuroepithelium 2
Germinal
neurons, allowing for interactions.
Choroid plexus 3 trigone
0.5 mm
Purkinje neurons settle beneath the egl.
B Primordial molecular layer

Figure 4. The cerebellum in a frontal/horizontal section


of a rat embryo on E18. A, view of the entire cerebellum
showing the external germinal layer (egl), the cerebellar
Mitotic cells in neuroepithelium, and the choroid plexus as the three
the egl “prongs” of the germinal trigone. The trigone wraps
around the back and sides of the cerebellum. The egl is a
secondary matrix that will produce the basket, stellate, and
granule cells in the cerebellar cortex much later. Deep
nuclear neurons are migrating downward, while Purkinje
neurons go upward (arrows) to settle beneath the egl.
B, detail of the primordial cortex showing Purkinje neu-
rons settling beneath the mitotically active egl. A prim-
itive molecular layer is a new feature. (3µ methacrylate
section, toluidine blue stain). Source: braindevelopment-
Migrating
Purkinje maps.org (E17 coronal archive)
neurons

0.1 mm
8
THE SUPERIOR COLLICULUS
IN AN E18 RAT EMBRYO
HAS A HONEYCOMB
MATRIX

Stratum I
Visual processing layers

Strata II-IV

Layers II-VII contain migrating young neurons (arrows).


Stratum V
and
intermediate
magnocellular
layer
Sensorimotor processing layers

Strata VI-VII

Neuroepithelium/
ependyma
0.1 mm

Figure 5. Sagittal section of the E18 rat brain (inset) showing a high magnification detail of the superior colliculus. The layers are still poorly defined,
but superficial (I-IV) and deep layers (VI-VII) are tentatively identified by the honeycomb matrix (red outlines of fiber bundles) in layer V and the
intermediate magnocellular layer. We postulate that the honeycomb matrix is a region where incoming axons interact with young superior colliculus
neurons, a staging area for connections in the mature brain. (3µ methacrylate section, toluidine blue stain) Source: braindevelopmentmaps.org (E18
sagittal archive)
9

THE NEOCORTEX
IN AN E18 RAT EMBRYO

Cajal-Retzius neurons Predominantly


cut transversely Layer VI neurons
reside in the cortical plate

Layer I

Cortical
plate

Subplate
among
glial
channels
are migrating perpendicular to the section plane

Upper STF
(former Layer VI
Many cells in the STF and the SVZ

sojourn zone)

Lower STF
(Layer V
sojourn zone)

Subventricular
zone (SVZ) and
migrating cells
cut transversely

Neuroepithelium
mainly generating
neurons in Layers
IV-II and the dis-
persed germinal
cells in the SVZ

0.1 mm
STF - stratified transitional field
Figure 6. Sagittal section of the E18 rat brain (inset) showing high-magnification detail of the neocortex. In the frontal/horizontal section plane, many
neurons are migrating parallel and appear as long, spindle-shaped cells. Here, all cells appear to be migrating radially, but that is because the spin-
dle-shaped cells are being cut transversely (white-outlined arrow). This cortex is in the middle of the ventrolateral (more mature) to dorsomedial (less
mature) region. The predominant occupants of the cortical plate are layer VI neurons; subplate neurons have mostly delaminated from the cortical
plate and are mingling with the glial channels beneath it. Layer V neurons sojourn deep in the STF, but many are migrating upward toward the cortical
plate, where they will stack up above the layer VI neurons. Layers IV-II neurons are being generated in the primary cortical neuroepithelium and in
the subventricular zone. Eventually, they will sojourn in the deepest part of the STF. The STF is the area where the initial layout of cortical circuitry is
being established with other parts of the brain. (3µ methacrylate section, toluidine blue stain)
Source: braindevelopmentmaps.org (E18 sagittal archive)
10
THE HIPPOCAMPUS IN AN E18 RAT EMBRYO
Sojourning pyramidal cells

Hippocampal NEP

M
igr
Dentate NEP ati
in dentate notch ng
py
ra
mi
da
lc
ell
s

A3?
P

dC
ial GE

el
Fi
Fimbr

m
rdiu
i mo
pr
te
nta
De

Choroid plexus 0.1 mm


Figure 7. The hippocampus in a sagittal section of an E18 rat embryo (inset). The thick hippocampal neuroepithelium (NEP) is adjacent to the dentate NEP; the fimbrial glioepithelium (GEP) thins and spreads
out among fibers that may be afferents from the septum. The hippocampal NEP has a pyramidal cell sojourn zone basally, and many cells are migrating from there (straight arrows) into primordial field CA3.
There is a stream of cells (the dentate migration, curved arrow) heading into the dentate primordium (red outline). (3µ methacrylate section, toluidine blue stain) Source: braindevelopmentmaps.org
(E18 sagittal archive)
11

THE OLFACTORY BULB


AMYGDALA CE
RE IN AN E18 RAT EMBRYO

N
BE

LO
OLFACT0RY LL

EP L
HA
BULB UM

NC A
LE B A S
SUBTHALAMUS
AND
HYPOTHALAMUS PONS
TE

Olfactory nerve bundles

Olfactory
NEP

In
ter
na
lp
lex
ifo
Se rm
ttl lay
ing er
Ex mi
ter tra
na l/tu
lp
Ol lex fte
fac ifo dc
tor rm e
yn lay lls
erv er
ex
Trajectory of
migrating neurons

Lateral olfactorytract
Lateral olfactory tract
Figure 8. A horizontal section of the rat brain (inset) on E18
showing detail of the olfactory bulb. The mitral and tufted cell
migrations come in from the lateral basal telencephalon and
enter the internal plexiform layer to peel off and settle in the 0.25 mm
primordial mitral/tufted cell layer (curved arrow with branches).
(3µ methacrylate section, toluidine blue stain) Source: brainde-
velopmentmaps.org (E17 coronal archive)
Taylor & Francis
Taylor & Francis Group
http://taylorandfrancis.com


REFERENCES

%D\HU6$$OWPDQ-  Development of the Cerebel- %D\HU6$$OWPDQ- LQSUHVV The Human Brain during
lar System in Relation to Its Evolution, Structure, and the First Trimester 21- to 23-mm Crown Rump
Function&5&3UHVV%RFD5DWRQ)/ Lengths. Atlas of Human Central Nervous System
Development9ROXPH7D\ORU )UDQFLV&5&3UHVV
%D\HU6$ SUHVHQW www.braindevelopmentmaps.
org/DERUDWRU\RI'HYHORSPHQWDO1HXURELRORJ\ %D\HU6$$OWPDQ- LQSUHVV Glossary to Accompany
2FDOD)/ 7KLVZHEVLWHLVDQLPDJHGDWDEDVHRI Atlas of Human Central Nervous System Development,
PHWKDFU\ODWHHPEHGGHGQRUPDOUDWHPEU\RVDQGSDUDI 7D\ORU )UDQFLV&5&3UHVV
¿QHPEHGGHGUDWHPEU\RVH[SRVHGWR+7K\PLGLQH
%D\HU6$$OWPDQ-5XVVR5-=KDQJ;  7LPH
%D\HU6$ LQSUHVV Glossary to Accompany Atlas of tables of neurogenesis in the human brain based on
Human Central Nervous System Development. Taylor experimentally determined patterns in the rat. Neuro-
)UDQFLV&5&3UHVV toxicology 14

%D\HU6$$OWPDQ-  Neocortical Development, %D\HU6$$OWPDQ-5XVVR5-=KDQJ;  (PEU\


5DYHQ3UHVV1HZ<RUN ology. In: Pediatric Neuropathology6HUJH'XFNHWW
(G:LOOLDPVDQG:LONLQVSS
%D\HU6$$OWPDQ-  'HYHORSPHQW6RPHSULQ
ciples of neurogenesis, neuronal migration and +RFKVWHWWHU)  Beiträge zur Entwicklungsgeschichte
neural circuit formation. In: The Rat Nervous System, des menschlichen Gehirns9RO/HLS]LJXQG:LHQ
QG(GLWLRQ*HRUJH3D[LQRV(G$FDGHPLF3UHVV 'HXWLFNH
2UODQGR)ORULGDSS
/RXJKQD3&LWW\/(YDQV7&KXGOHLJK7  )HWDO
%D\HU6$$OWPDQ- SUHVHQW www.neurondevelop VL]HDQGGDWLQJ&KDUWVUHFRPPHQGHGIRUFOLQLFDO
ment.org 7KLVZHEVLWHKDVGRZQORDGDEOHSGI¿OHVRI obstetric practice, Ultrasound
RXUVFLHQWL¿FSDSHUVRQUDWEUDLQGHYHORSPHQWJURXSHG
E\VXEMHFW 2¶5DKLOO\50OOHU)  Developmental Stages in
Human Embryos, Carnegie Institution of Washington,
%D\HU6$$OWPDQ-  Atlas of Human Central 3XEOLFDWLRQ
Nervous System Development, Volume 4: The Human
Brain during the Late First Trimester.&5&3UHVV 2¶5DKLOO\50OOHU)  The Embryonic Human
Brain:LOH\/LVV1HZ<RUN
14

PART II: M841


CR 42 mm (GW 10.6)
Frontal/Horizontal

This specimen is number 841 in the Minot Collection, The diencephalic neuroepithelia have thinned out
referred to here as M841, with a crown rump length (CR) of because nearly all neuronal populations are either near
42 mm, and estimated to be at gestational week (GW) 10.6. their last days of generation or have already been gener-
7KH IHWXV ZDV HPEHGGHG LQ SDUDI¿Q FXW LQ  —P WKLFN ated. Nuclear borders are indistinct throughout the dien-
sections, and stained with borax carmine and Lyon’s blue. cephalon because most neurons are migrating, blurring the
No information is available on date of collection (some- boundaries.
WLPHEHWZHHQDQG DQGWKH¿[DWLYHXVHG6LQFH
there is no photograph of this brain before it was embedded The midbrain tegmentum, pons, and medulla are lined by
and cut, a specimen from Hochstetter (1919) that is compa- a thin neuroepithelium being transformed to an ependyma
UDEOHWR0KDVEHHQPRGL¿HGWRVKRZWKHDSSUR[LPDWH because the majority of neurons have been produced. Neu-
section plane and external features of the brain at GW10.6 rons throughout this part of the brain are still migrating,
(Fig. 9). Like most of the specimens in this volume, the but many have settled, making some nuclear groups more
sections are not cut exactly in one plane; M841 is midway GH¿QLWH7KHUHLVDSURPLQHQWJHUPLQDO]RQHLQWKHPHV-
between frontal and horizontal. Photographs of 22 sections encephalic tectum, where many neuronal populations are
(Plates 1-22 DUHLOOXVWUDWHGDWORZPDJQL¿FDWLRQVKRZLQJ KDYLQJWKHLU¿QDOGD\VRIJHQHUDWLRQ6RPHOD\HULQJLQWKH
H[FHOOHQWWLVVXHGHWDLO8QIRUWXQDWHO\D¿QHJUDQXODUSUH- tectum is beginning to differentiate.
FLSLWDWHLVYLVLEOHDWKLJKPDJQL¿FDWLRQ
The thick precerebellar neuroepithelium is an exception
In the cerebral cortex, the neuroepithelium is prom- in the medulla because it is still generating neurons. The
inent and there is probably a small subventricular zone anterior extramural and posterior extramural migratory
present (not visible because of the inability to see detail at streams are subpial accumulations in the medulla and pons.
KLJKPDJQL¿FDWLRQ 7KHUHLVDVWUDWL¿HGWUDQVLWLRQDO¿HOG
67) ZLWKOD\HUVZHGHVLJQDWHDV67)67)DQG67) 7KHFHUHEHOOXPKDVDGH¿QLWHEXWWKLQQLQJQHXURHSLWKH-
The ventroateral-to-dorsomedial maturation gradient in the lium at the ventricular surface. Probably most of the Pur-
FRUWLFDOSODWHDQG67)LVHYLGHQW6WUHDPVRIQHXURQVDQG kinje neurons are either sojourning in a thick dense layer
JOLDDSSHDUWROHDYH67)DQGHQWHUWKHODWHUDOPLJUDWRU\ outside the neuroepithelium, while others are migrating
stream. A massive neuroepithelium/subventricular zone upward to form a primordial layer beneath the external ger-
overlies the amygdala, nucleus accumbens, and striatum minal layer (egl) that partially covers the cerebellum. The
(caudate and putamen) where neurons (and glia) are being cortex is just beginning to form beneath the parts coverec
generated. The olfactory evagination is at the junction of by the egl.
the basal telencephalon and the cerebral cortex.
15

Figure 9.7KHODWHUDOYLHZRIWKHEUDLQDQGXSSHUFHUYLFDOVSLQDOFRUGIURPDVSHFLPHQZLWKDFURZQUXPSOHQJWKRIPP PRGLÀHGIURP)LJXUH


7DEOH9,,+RFKVWHWWHU VHUYHVWRVKRZWKHDSSUR[LPDWHORFDWLRQVDQGFXWWLQJDQJOHVRIWKHLOOXVWUDWHGVHFWLRQVRI0LQWKHIROORZLQJSDJHV7KH
VPDOOLQVHWLGHQWLÀHVWKHPDMRUVWUXFWXUDOIHDWXUHV7KHOLQHLQWKHFHUHEHOOXPDQGGRUVDOHGJHVRIWKHSRQVDQGPHGXOODLVWKHFXWHGJHRIWKHPHGXOODU\
YHOXP
16
17
18

20
21
22
23
24
25
26
27
28

30
31
32
33
34
35
36
PLATE 11A
CR 42 mm,
GW 10.6, M841
Frontal/Horizontal
Section 235

LAYERS OF THE CORTICAL


STRATIFIED TRANSITIONAL
FIELD (STF)
STF1 Superficial fibrous layer,
with an early
developmental stage (t1)
when many cells are
migrating through it,
followed by a late stage
(t2) with sparse cells.
Endures as the
subcortical white matter.

STF4 Complex middle layer


where sojourning and
migrating cortical
neurons grow
corticofugal axons and
intermingle with
corticopetal axons.

STF5 Deep cellular layer that is


prominent during the first
trimester, the first sojourn
zone to appear outside
the germinal matrix.
Another random document with
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The Project Gutenberg eBook of Infiltration
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and most other parts of the world at no cost and with almost no
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you are located before using this eBook.

Title: Infiltration

Author: Algis Budrys

Release date: November 4, 2023 [eBook #72026]

Language: English

Original publication: New York, NY: Royal Publications, Inc, 1958

Credits: Greg Weeks, Mary Meehan and the Online Distributed


Proofreading Team at http://www.pgdp.net

*** START OF THE PROJECT GUTENBERG EBOOK


INFILTRATION ***
INFILTRATION

By ALGIS BUDRYS

If werewolves exist, they don't necessarily


conform to all the superstitions people have.
They may even know fear....

[Transcriber's Note: This etext was produced from


Infinity October 1958.
Extensive research did not uncover any evidence that
the U.S. copyright on this publication was renewed.]
I
Sunset. They're coming for me, tonight, he knew as he woke.
Sunset. Not really—if he were to get dressed now, and go out on the
street, the red globe would still be hanging over the cliffs of New
Jersey. But the shadow of the building next door had fallen over his
apartment windows, and he sleepily pushed a cigarette between his
numb lips and swung his feet over the side of the bed, fumbling with a
match as he walked over to the small radio on the windowsill and
turned it on. There was a double-header between the Giants and
Cincinnati—the first game was probably in its last inning.
Sunset—odd, how the conditioning worked. Was it conditioning? Or
were the old wives' tales not so absurd, after all? But he could go out
in the sunlight—had done it many times. His tan proved it. He
touched silver and cold iron countless times each day, crossed
running water—and he'd gone to church every Sunday, until he was
twelve. No, there was a core of truth under the fantastically complex
shell of nonsense, but the old limitations were not part of it. He
shrugged. Neither were most of the powers.
Still, he liked to sleep in the daytime. His schedule seemed to gain an
hour at night, lose one in the morning, until, almost unnoticeably, it
had slipped around the clock.
He went into the bathroom while the worn tubes in the radio warmed
up slowly, and washed his face, brushed his teeth, shaved. He
combed his hair, then paused thoughtfully. Wouldn't do any harm. No
full moon in here, either, he thought, looking up at the circular
fluorescent tube in the ceiling, but he noticed no impediment as he
coalesced, dropped to all fours, and ran his pelt against the curry-
combs he had screwed to the bathroom door. He did a thorough job,
enjoying it, and, after he had realigned, walked out of the bathroom in
time to hear the Giants making their final, fruitless out of the first
game. Five-Zero, Cincinnati, and he grimaced in disgust. Four shut-
outs in the last five games.
He laughed at himself, then, for actually being annoyed. Still and all, it
wasn't the first time a man became emotionally involved in a mirage.
Was it a mirage? True, there weren't really any such things as the
San Francisco Giants—but a man could certainly be expected to
forget that, occasionally, if he were part of the same illusion at least
half the time. And certainly, such stuff as dreams are made of is solid
enough when you are yourself a dream.
He went out in the kitchen and started coffee, then came back and
sat down next to the radio, hardly listening to the recap of the game.

Odd, how it had all started. Being suddenly marooned on this planet,
forced to survive, somehow, through the long years while waiting for
rescue. How many years had it been, now? Some five hundred
thousand, in the subjective reference for this particular universe. He
knew the formula for conversion into objective time—it all worked out
to the equivalent of about six months—but that wasn't what mattered,
as long as they'd all had to survive in this universe.
Sleep—suspended animation, if you wanted to call it that—had been
the only answer. And they couldn't do that, directly. They'd had to
resort to chrysalids.
He smiled to himself, got up, and turned down the fire under the pot
until the coffee was percolating softly.
The original plan had snowballed, somewhat.
Resolving chrysalids was one thing—making them eternal was
another, and unnecessary. It was far simpler to arrange the chrysalids
so they'd be able to reproduce themselves. And, of course, in order to
survive, and take care of itself, a chrysalis had to have some
independent intelligence.
And, so it worked. The chrysalis housed a sleeper, operating
unawares and completely independent of him—or her—until the
chrysalis wore out. Then the sleeper was passed on to a new
chrysalis, with neither of the chrysalids involved—nor, for that matter,
the sleeper—conscious of the transfer. So it would continue, through
the weary, subjective years; generation upon generation of
chrysalids, until, finally, the paramathematical path drifted back to
touch this universe, and the sleepers could wake, and continue their
journey.
And if the human race chose to speculate on its origins in the
meantime, well, that was part of the snowball.
He got up again, and turned off the flame under the coffeepot. Now, if
I were a sorcerer—as defined by Cotton Mather's ilk, of course—, he
thought, I should be able to (a) turn the fire off without getting up, or,
(b) generate the flame without the use of Con Edison's gas, or, (c), if I
had any self-respect at all, conjure hot coffee out of thin air. His lips
twisted with nausea as he thought that nine out of ten people would
expect him to be drinking blood, as a matter of course.
He sighed with some bitterness, but more of resignation. Well, that
was just another part of the snowball.
Because the chrysalids had done a magnificent job in all three of its
subdivisions. They had kept the sleepers safe—and reproduced, and
used their intelligence to survive. They had survived in spite of
pestilence, famine, and flood—by learning enough to wipe out the
first two, and control the third. It would seem that progress was not a
special quality to be specially desired. Most of the chrysalids were
consumed by a fierce longing for the Good Old Days, as a matter of
fact. It was merely the inescapable accretion to sheer survival.
And so came civilization. With civilization: recreation. In short, the
San Francisco Giants, and—He reached over, suddenly irritated at
the raspy-voiced and slightly frantic recapitulation of the lost
ballgame, and changed the station. And Beethoven.
He relaxed, smiling slightly at himself once again, and let the music
sing to him. Chrysalids, eh? Well, they certainly weren't his kind of
life, free to swing from star to star, riding the great flux of Creation
from universe to universe. But whence Beethoven? Whence
Rembrandt, Da Vinci, and Will Shakespeare, hunched over a mug of
ale and dashing off genius on demand, with half an eye on the
serving wench?
He shook his head. What would happen to this people, when the
sleepers woke?
The snowball. Ah, yes, the snowball. That was a good part of it—and
he and his kind were another.
If we had known, he thought. If we had known how it would be...?
But, they hadn't known. It had been just a petty argument, at first.
Nobody knew, now, who had started it. But there were two well-
defined sides, now, and he was an Insurgent, for some reason. The
winning side gives the names that stick. They were Watchers—an
honorable name, a name to conjure up trust, and duty, and loyalty.
And he was an Insurgent. Well, let it stand. Accept the heritage of
dishonor and hatred. Somewhere, sometime, a gage was flung, and
he was heir to the challenge.
The chrysalids solved the problem of survival, of course. But the
problem of rescue had remained. For rescue, in the sense of help
from an outside agency, would be disastrous. When the path shifted
back, they had to learn of it themselves, and go on of their own
accord—or go into slavery. For there is one currency that outlives
document and token. Personal obligation. And, if they were so
unlucky as to have an actual rescuer, the obligation would be high—
prohibitively so.
The solution had seemed simple, at first. In each generation of
chrysalids, there would be one aware individual—one Watcher, to
keep guard, and to waken the rest should the path drift back in the
lifetime of his chrysalis. Then, when that particular chrysalis wore out,
the Watcher would be free to return to sleep, while another took his
place.
His mouth twisted to one side as he took a sip of coffee.
A simple, workable plan—until someone had asked, "Well and good.
Excellent. And what if this high-minded Watcher realizes that we,
asleep, are all in his power? What if he makes some agreement with
a rescuer, or, worse still, decides to become our rescuer when the
path drifts back? What's to prevent him, eh? No," that long-forgotten,
wary individual had said, "I think we'd best set some watchers to
watch the Watcher."
Quis custodiet?
What had it been like? He had no way of knowing, for he had no
memory of his exact identity. That would come only with Awakening.
He had only a knowledge of his heritage. For all he knew, it had been
he who raised the fatal doubt—or, had been the first delegated
Watcher. He shrugged. It made no difference. He was an Insurgent
now.
But he could imagine the voiceless babel among their millions—the
argument, the cold suspicion, the pettiness. Perhaps he was passing
scornful judgment on himself, he realized. What of it? He'd earned it.
So, finally, two groups. One content to be trustful. And the other a
fitful, restless clan, awakening sporadically, trusting to chance alone,
which, by its laws, would insure that many of them were awake when
the path drifted back. The Insurgents.
So, as well, two basic kinds of chrysalids. The human kind, and the
others. Wolves, bears, tigers. Bats, seals—every kind of living thing,
except the human. The Insurgent kind.
And so the struggle began. It was a natural outgrowth of the
fundamental conflict. Which side had tried to over-power the first
chrysalis? Who first enslaved another man? he thought, and half-
snarled.
That, too, was unimportant now. For the seed had been planted. The
thought was there. Those who are awake can place those who sleep
under obligation. Control the chrysalids, and you control the sleepers
within. But chrysalids endure for one generation, and then the
sleepers pass on.
What then? Simplicity. Group your chrysalids. Segregate them. Set
up pens for them, mark them off, and do it so the walls and fences
endure through long years.
This is my country. All men are brothers, but stay on your side of the
line, brother.
Sorry, brother—you've got a funny shape to your nose. You just go
live in that nice, walled-off part of my city, huh, brother?
Be a good fellow, brother. Just move to the back of the bus, or I'll
lynch you, brother.
And the chrysali die, the sleepers transfer—into another chrysalis in
the same pen. SPQR. Vive, Napoleon! Sieg Heil!

Some of the time it was the Watchers. Some of the time it was the
Insurgents. And some of the time, of course, the chrysalids evolved
their own leaders, and imitated. For, once the thing had begun, it
could not be stopped. The organization was always more powerful
than the scattered handsful. So, the only protection against
organization was organization.
But it was not organization in itself that was the worst of it. It was the
fact that the only way to control the other side's penned chrysalids
was to break down a wall in the pen, or to build a larger pen including
many of the smaller ones.
And, again, it was too late, now, to decide who had been at fault.
Who first invented War?
The way to survive war is to wage more decisive war. The chrysalids
had to survive. They learned. They ... progressed? ... by so doing.
They progressed from bows to ballistas to bombs. From arbalests to
aircraft to A-bombs. Phosphorus. Chlorine. HE. Fragmentation.
Napalm. Dust, and bacteriological warfare. Thermopylae, Crecy, the
Battle of Britain, Korea, Indo-China, Indonesia.
And try to believe as you sit here, Insurgent, that none of this is real,
that it is all a phase, acted out by dolls of your own creation in a sham
battle that is really only a bad dream in the unfamiliar bed of a lodging
for the night!
Chrysalids they might be, Insurgent, he lashed himself, but it was the
greed and suspicion of all your kind—Insurgent and Watcher alike—
that set this juggernaut to rolling!
He took another sip of coffee, and almost gagged as he realized it
had grown cold. He got up and walked into the kitchen with the cup in
his hand. He threw the rest of the coffee in the sink, washed out the
cup, and turned on the burner under the coffeepot.
One more thing—one more development, born of suspicion.
For the original one-Watcher plan had been abandoned, of course.
And here, again, there was no telling whose blame it was. Quis
custodiet ipsos custodes? Who will watch the Watchers? There had
been many Watchers to a generation—how many, no one knew. They
balanced each other off, and they checked the random number of
Insurgents who awoke in each generation. So, more Insurgents
awoke to check the Watchers—and, more.
In spite of what the Transylvanians believed, a wolf is no match for a
man, except under special conditions. A tiger can pull a man down—
but cannot fire back at the hunters. A seal is prey to the Eskimo.
So, "werewolves." Child of fear, of Watcher propaganda, and of one-
tenth fact. The animals were Insurgent chrysalids, right enough. But,
for an awake Insurgent to compete with a Watcher, the Insurgent, too,
had to be a man—or something like it.
The coffee had warmed up. He poured himself a fresh cup, and
added cream and sugar absently. The refrigerator was empty. He
reached in and turned it off. No more need for that, after tonight.
So, that was the power the Insurgents had. The only power, and the
Watchers had it, as well. They could resolve their chrysalids into any
form they chose—realign. A wolf could become a man—without hair
on his palm, and with garlic on his breath, if he so chose. A man—a
Watcher, of course—could become a wolf.
Thus, the final development. Espionage and counter-espionage.
Infiltration. Spying, if you chose.
The Insurgent smiled bitterly, and drained the cup. And propaganda,
of course. Subtle—most of it indirect, a good deal of it developed by
the chrysalids themselves, but propaganda, nevertheless. Kill the evil
ones—kill the eaters of dead flesh, the drinkers of blood. They are the
servants of the Evil One.
He almost retched.
But, you could hardly blame them. It was a war, and, in a war, you
play all your cards, even if some of them were forced into your hand.
Yes, and I've played genuine werewolf on occasion, when I had to.
He started to wash the coffeepot and the cup—then, threw both into
the garbage can. He walked back to the radio and dialed it away from
Eroica and back to baseball. The Giants were losing, Three-Zero, in
the third inning.
The house phone buzzed. He went to it slowly, picked it calmly off the
hook.
"Yes, Artie?"
"Mister Disbrough, there's a couple of guys coming up to see you. I'm
not supposed to tell you about it, but.... Well, I figured ..." the
doorman said.
"All right. Thanks, Artie," he answered quietly. He almost hung up,
then thought of something. "Artie?"
"Yes, Mister Disbrough?"
"There'll be a couple of fifths of Dewar's in my cupboard. I won't be
back for a while. You and Pete are welcome to them. And thanks
again."
He hung up and began to dress, realigning his chrysalis to give him
the appearance of clothing. The doorbell rang, and he went to open it
for the two men from the FBI.

II
What difference did it make, what particular pen he represented?
Rather, since the sober-faced men knew very well which pen it was,
why should it be so necessary to them for him to confirm what they
already knew without a shadow of a doubt?
"Now, then, Mister Disbrough," one of the FBI men said, leaning his
hands on the edge of the table at which the Insurgent was sitting, "we
know who sent you."
Good. Why bother me, then?
"We know where you got your passport, we know who met you at the
dock, we know your contacts. We have photographs of everyone
you've met and talked to, we have tapes of every telephone call
you've made or received. We also know that you are the top man in
your organization here."
And? They were chrysalids, every one of them. Perhaps there was no
Watcher behind them—perhaps. But he'd been picked up a little too
quickly. The net had folded itself around him too soon. No—there had
to be a Watcher. He wished they'd stop this talking and bring him out.
"Now, I'd simply like to point out to you that this is an airtight case. No
lawyer in the world will be able to break it down. You'll retain counsel,
of course. But, I'd simply like to point out to you that there'll be no
point to any denial you may make to us. We know what you've been
doing. I'd suggest you save your defense for the trial."
He looked up at him and smiled ruefully. "If you've got a list of
charges," he said, "I'll be glad to confess to all of them—provided, of
course, that it is a complete list."
I'm sure it doesn't list me as a werewolf, he thought. I wonder what
the sentence would be—death by firing squad equipped with silver
bullets?
But, then, he wasn't going to confess to that, anyway.
"Um!" The FBI man looked suspicious. Obviously, he'd expected
nothing of the kind.
"No strings," the Insurgent reassured him. "The job's over, and it's
time to punch the clock."
Which was just about the way it was. But he wanted that Watcher. If
he was in the office at all, he'd almost have to come out to witness
the confession. After all, the Insurgent was supposed to be a pretty
big fish.
The FBI man went into a cubicle office set off to one side. When he
came out, carrying a sheaf of paper, the Watcher was with him.
The Insurgent felt the hackles standing up on the back of his neck,
and something rumbled inaudibly at the base of his throat. He knew.
He could tell. He could smell Watcher every step of the way, from the
day he had docked until now, when the scent—half there, half the
pure intuition of instinct—rose up before him in an over-powering
wave.
Then he saw the look of distaste crawl across the Watcher's face,
and he barked a laugh that drew curious looks from the men in the
office. Hello, brother.
He saw the bulge of the hip holster on the Watcher's belt, and
laughed again. So, we play the game, he thought. We add up scores,
and, in the end, the side with the most points wins. Forget that there
should be no sides, that every point, no matter for whom scored, is a
mark of shame and disgrace.
He wondered, briefly, whether the Watcher was of his kind by choice,
or whether it was simply something that had happened, as it was with
him. Probably. Two separate heritages had met, represented by
identical individuals who happened to have awakened in dissimilar
chrysalids.
Will we remember? he wondered. When we awaken, will we
remember this? How we battled, blinded, in the shadows of our own
casting? Or was there more mercy in Creation than they, themselves,
had shown to the chrysalids? He had three brothers among the
sleepers. When they woke, would they embrace, not remembering
that each had killed the other countless times? Or forgetting that they
had stood together, on some battlefield? Would all the old comrades,
all the bitter enemies, be wiped from memory? He hoped so. With
every segment of his being, he hoped so, for there was no peace,
through eternity, if it was otherwise.
He stood up, lightly, tensing the muscles in his calves. The FBI men,
suddenly alert, began to move for him, but he'd maneuvered things
so that none of them were close enough to him.
The Watcher went pale.
"Shall I coalesce, brother?" the Insurgent asked, the words rumbling
out of his throat, a grin of derision baring his teeth.
"No!" The Watcher was completely frightened. Words could be
explained away, particularly if they sounded like nonsense to the
other men in the room. But a werewolf, fanging the throat of a
Watcher who would have to fight back with his spectacular
weapons.... Nothing in the world could keep the rumors from
spreading. The chrysalids might even learn, finally and irrevocably,
the origin of their species.
"Your obligation, brother," the Insurgent half-laughed, and kept
stalking toward the Watcher. Perhaps he is my brother.
And if he is...?
No difference. The shadows are thick and very dark. One of the other
men shot him in the side, but he sprang for the Watcher, carefully
human, to hold the Watcher to his debt, and the Watcher shot him
three times in the chest, once in the throat, and once in the stomach.
The shape of a cross? Did he believe it himself? Was it true? A plus
sign, cancelling a negative force? Who knew? Shadow, shadow, all is
darkness.
He fell to his knees, coughing, in victory. Score one more for the
Insurgents, and a Watcher, at that!
"Thank you, brother," the Insurgent murmured, and fell into the long
sleep with a grateful sigh.
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