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The Biology of Caves and Other
Subterranean Habitats
THE BIOLOGY OF HABITATS SERIES
This attractive series of concise, affordable texts provides an integrated over-
view of the design, physiology, and ecology of the biota in a given habitat, set
in the context of the physical environment. Each book describes practical
aspects of working within the habitat, detailing the sorts of studies which
are possible. Management and conservation issues are also included. The
series is intended for naturalists, students studying biological or environ-
mental science, those beginning independent research, and professional
biologists embarking on research in a new habitat.

The Biology of Rocky Shores Colin Little and J. A. Kitching

The Biology of Polar Habitats G. E. Fogg
The Biology of Lakes and Ponds Christer Brönmark and
Lars-Anders Hansson
The Biology of Streams and Rivers Paul S. Giller and Björn Malmqvist
The Biology of Mangroves Peter J. Hogarth
The Biology of Soft Shores and Estuaries Colin Little
The Biology of the Deep Ocean Peter Herring
The Biology of Lakes and Ponds, 2nd Edition
Christer Brönmark and Lars-Anders Hansson
The Biology of Soil Richard D. Bardgett
The Biology of Freshwater Wetlands Arnold G. van der Valk
The Biology of Peatlands Håkan Rydin and John K. Jeglum
The Biology of Mangroves and Seagrasses, 2nd Edition Peter J. Hogarth
The Biology of African Savannahs Bryan Shorrocks
The Biology of Polar Regions, 2nd Edition David N. Thomas et al.
The Biology of Deserts David Ward
The Biology of Caves and Other Subterranean Habitats David C. Culver
and Tanja Pipan
The Biology of Alpine Habitats Laszlo Nagy and Georg Grabherr
The Biology of Rocky Shores, 2nd Edition Colin Little, Gray A. Williams
and Cynthia D. Trowbridge.
The Biology of Coral Reefs Charles R.C. Sheppard, Simon K. Davy &
Graham M. Pilling
The Biology of
Caves and Other
Subterranean
Habitats
David C. Culver and Tanja Pipan

1
3
Great Clarendon Street, Oxford OX2 6DP
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10 9 8 7 6 5 4 3 2 1
Preface

We are in a golden age of the study of subterranean biology. Twenty-five

years ago, when one of us (DCC) wrote a book on the biology of caves,
it was easy to read and discuss all the non-taxonomic literature on cave
biology written in English. The only book length treatment of cave biol-
ogy at that time in English was the translation from the French of Albert
Vandel’s Biospeleology. Most speleobiologists were not writing in English
and the discipline remained largely a national one. Art Palmer, the author
of a recent introductory text on cave geology, points out that theories
of cave development were developed independently (and in strikingly
parallel ways) three times—first in Serbo-Croatian, next in French, and
finally in English. Speleobiologists as well kept reinventing the wheel—
who knows how many biologists discovered and rediscovered that the
Pleistocene may have driven animals into caves. Twenty-five years ago, for
American speleobiologists, but much less so for European biologists, spe-
leobiology meant the biology of caves. There was scarcely any recognition
or awareness of non-cave subterranean environments among American
speleobiologists.
How times have changed. The scope of speleobiology has expanded to
include those subterranean1 habitats whose inhabitants include blind, de-
pigmented species with compensatory increases in other sensory structures.
The globalization of subterranean biology and collaboration among spe-
leobiologists has been made possible, especially because of Internet and
World Wide Web. The growing and now nearly universal use of English as
the language of scientific communication has opened up new avenues for
cooperation and collaboration. New technology, including the possibility of
sequencing DNA molecules (Porter 2007), the availability of increasingly
sophisticated software for phylogenetic reconstruction, and the possibility

1
We use subterranean in the sense of organisms living in natural spaces. The word
subterranean is also frequently applied to organisms that create their own spaces—
especially mammals such as mole rats, termites, and plant roots. The word hypogean
is sometimes used in the sense we use subterranean, but its use is uncommon, and we
use enough uncommon words as it is. There are many precedents for the way we use
the word, such as the International Society for Subterranean Biology and its journal
Subterranean Biology.
vi PREFACE

of storing and analysing large quantities of spatial information (especially

databases and Geographic Information Systems), has created new potenti-
alities in the analysis of subterranean species and communities. This com-
bined with new conceptual advances, such as vicariance biogeography, the
joint analysis of evolution and development (evo-devo), and ecosystem
models, has led to the current golden age, with an accompanying explo-
sion of published information.
In the past 20 years, several milestone books on subterranean biology
have been published, including Groundwater Ecology (Gibert et al. 1994a),
the three-volume Encyclopaedia Biospeologica (Juberthie and Decu
1994–2001), Subterranean Ecosystems (Wilkens et al. 2000), Encyclopedia
of Caves (Culver and White 2005), and Encyclopedia of Caves and Karst
Science (Gunn 2004). Collectively they have advanced the field of subter-
ranean biology by leaps and bounds, but none of them are introductory
accounts. Hence this book.
We hope that this book is accessible to a wide variety of readers. We have
assumed no training in biology beyond a standard university year-long
course, and we have tried to make the geological and chemical incursions
self-contained. An extensive glossary should help the readers through any
terminological rough spots.
We have organized this book around what seem to us to be the major
research areas and research questions in the field. To provide a context
for these questions, we review the different subterranean environments
(Chapter 1), what the energy sources are for subterranean environ-
ments given that the main energy source in surface environments—
photosynthesis—is missing (Chapter 2), and the main inhabitants of these
underground domains (Chapter 3). The research areas that we focus on
are as follows:
• How are subterranean ecosystems defined and organized, and how in
particular does organic carbon move through the system (Chapter 4)?
• How do species interact and how do these interactions, such as competi-
tion and predation, organize, and constrain subterranean communities
(Chapter 5)?
• How did subterranean organisms evolve the bizarre morphology of
elongated appendages, no pigment, and no eyes (Chapter 6)?
• What is the evolutionary and biogeographic history of subterranean
species? Are they in old, relict lineages (Chapter 7)? How does their
distribution relate to past geologic events?
• What is the pattern of diversity of subterranean faunas over the face of
the earth (Chapter 8)?
We close by “putting the pieces together” and examining some represen-
tative and exemplary subterranean communities (Chapter 9), and how to
conserve and protect them (Chapter 10).
 PREFACE vii

With the exception of Chapters 1–3, where we have attempted to provide a

comprehensive geographic and taxonomic review of the basics, we have fo-
cused on a few particularly well-studied cases. Although we have provided
case studies from throughout the world, readers from South America and
Asia will no doubt find a North American and European bias. Of this
we are certainly guilty, but in part this bias is because of longer tradi-
tions of study of subterranean life in Europe and North America. We have
provided an extensive bibliography and hope that interested readers will
pursue the subjects further. When English language articles were avail-
able, we have highlighted them but we also have not hesitated to include
particularly important or unique papers in other languages.
A cautionary word about place names. Many species are limited to a single
cave, well, or underflow of a brook, and, if for no other reason, this makes it
important to accurately give place names. Throughout the book we have iden-
tified the country and state or province in which a site is located. We have,
whenever possible, retained the spelling of the local language. Translation
runs the risk of confusing anyone trying to identify a particular cave or site,
and also runs the risk of repeating the word cave in different languages, as
in Postojnska Jama Cave (Postojna Cave Cave). Postojnska Jama already has
names in three languages (Slovene, Italian, and German) and there is no need
to add a fourth. Maps of sites mentioned in the text are provided.
Even to us, the field of subterranean biology seems especially burdened
with obscure terminology. While there is a temptation to ignore it as much
as possible, it is widespread in the literature and some of it is even useful.
We have defined many terms in the text when we first use them, and have
included an extensive glossary to aid readers.
Besides the fascination of their bizarre morphology (which cannot really
be overrated), there are two main reasons for biologists to be interested in
subterranean faunas. One is numerical. Nearly all rivers and streams have
an underlying alluvial system in which its residents never encounter light.
Approximately 15% of the Earth’s land surface is honeycombed with caves
and springs, part of landscape called karst that is moulded by the forces of
dissolution rather erosion of rock and sediment. In countries such as Cuba
and Slovenia, this is the predominant landform.
But there is a more profound reason for biologists to study subterranean
biology. Subterranean species can serve as model systems for several im-
portant biological questions. As far as we can determine, it was Poulson
and White (1969) who first made this notion explicit but it is implicit in
the writings of many subterranean biologists. This is a recurring theme
throughout this book, and we just list some of the possibilities here:
• Subterranean ecosystems can serve as models of carbon (rather than
nitrogen and phosphorus) limited ecosystems and ones where most
inputs are physically separated from the community itself.
viii PREFACE

• Subterranean communities can serve as a model of species interactions

because the number of species is small enough that all pairwise inter-
actions can be analysed and then combined into a community-wide
synthesis.
• The universal feature of loss of structures (regressive evolution) is espe-
cially obvious in subterranean animals, with a clear basis, that in turn
can allow for detailed studies of adaptation.
• The possibilities of dispersal of subterranean species are highly con-
strained and so the species (and lineages) can serve as models for vicar-
iant biogeography.
• The highly restricted ranges and specialized environmental require-
ments can serve as a model for the protection of rare and endangered
species.
Whatever reasons you have for reading this book, we hope it leads you to
a fascination with subterranean biology, one that lasts a lifetime.
Acknowledgements

The field of subterranean biology is blessed with a strong, cooperative

group of scholars from all over the world, and we could not have written
this book without the help of many of them. We especially thank Janez
Mulec for reading the entire manuscript and making many helpful sugges-
tions. Daniel W. Fong, Horton H. Hobbs III, William R. Jeffery, William K.
Jones, Megan Porter, Peter Trontelj, and Maja Zagmajster all read selected
chapters and helped us avoid many mistakes. Several colleagues provided
unpublished photographs and drawings—Gregor Aljančič, Marie-Jose
Dole-Olivier, Annette Summers Engel, Horton H. Hobbs III, Hannelore
Hoch, William R. Jeffery, Arthur N. Palmer, Borut Peric, Slavko Polak,
Megan Porter, Mitja Prelovšek, Nataša Ravbar, Andreas Wessel, Jill Yager,
and Maja Zagmajster. Colleagues also provided us with preprints and
answered sometimes naive questions—Louis Deharveng, Marie-Jose Dole-
Olivier, Stefan Eberhard, Annette Summers Engel, Daniel W. Fong, Franci
Gabrovšek, Janine Gibert, Benjamin Hutchins, Florian Malard, Georges
Michel, Pedro Oromi, Metka Petrič, Megan Porter, Katie Schneider, Boris
Sket, Peter Trontelj, Rudi Verovnik, and Maja Zagmajster. Jure Hajna and
Franjo Drole of the Karst Research Institute ZRC SAZU devoted many
hours to scanning and producing diagrams. Maja Kranjc, in charge of the
magnificent library at the Karst Research Institute, has constantly helped
even in the face of increasingly panic-stricken requests for books and jour-
nals. Daniel W. Fong, Benjamin Hutchins, Karen Kavanaugh, and Wanda
Young cheerfully handled our many requests for materials from American
University while we were writing the book at the Karst Research Institute
in Slovenia.
We are especially grateful to the Karst Research Institute ZRC SAZU,
especially the head of the institute, Dr. Tadej Slabe and the administra-
tive assistant, Sonja Stamenković, for making the writing go as smoothly
as possible. Tadej Slabe provided time for TP to work, space for DCC to
work, and an appointment to DCC as Associate Researcher. Financial sup-
port was provided by Ad Futura (Javni sklad Republike Slovenije za razvoj
kadrov in štipendije) to DCC during his stay in Slovenia.
x ACKNOWLEDGEMENTS

A project of this magnitude was a burden on both of our families, and we

are especially grateful to our spouses, Gloria Chepko and Miran Pipan, for
providing both understanding and support.
Postojna, Slovenia
March 2008
 Contents

Site Maps and Gazetteer xiv

1 The subterranean domain 1
1.1 Introduction 1
1.2 Caves 4
1.3 Interstitial habitats 16
1.4 Superficial subterranean habitats 19
1.5 Summary 22

2 Sources of energy in subterranean environments 23

2.1 Introduction 23
2.2 Sources of energy 23
2.3 Summary 39

3 Survey of subterranean life 40

3.1 Introduction 40
3.2 Temporary subterranean visitors and residents 40
3.3 Residents of cave entrances 43
3.4 Ecological and evolutionary classifications 45
3.5 Taxonomic review of obligate subterranean species 48
3.6 Subterranean organisms in the laboratory 69
3.7 Collecting stygobionts and troglobionts 71
3.8 Summary 73

4 Ecosystem function 75
4.1 Introduction 75
4.2 Scale and extent of subterranean ecosystems 76
4.3 Stream reaches 78
4.4 Caves 81
4.5 Karst basins 87
4.6 Summary 90
xii CONTENTS

5 Biotic interactions and community structure 91

5.1 Introduction 91
5.2 Species interactions—generalities 91
5.3 Predator–prey interactions—beetles and cricket eggs
in North American caves 93
5.4 Competition and other interactions in Appalachian
cave streams 97
5.5 Competition as a result of eutrophication 101
5.6 Community analysis—generalities 102
5.7 Epikarst communities 103
5.8 Interstitial groundwater aquifer 105
5.9 Overall subterranean community structure in the
Jura Mountains 106
5.10 Summary 108

6 Adaptations to subterranean life 109

6.1 Introduction 109
6.2 History of concepts of adaptation in subterranean
environments 110
6.3 Adaptation in amblyopsid cave fish 113
6.4 Adaptation in the amphipod Gammarus minus 119
6.5 Adaptation of the cave fish Astyanax mexicanus 125
6.6 How long does adaptation to subterranean life take? 129
6.7 Summary 130

7 Colonization and speciation in subterranean

environments 131
7.1 Introduction 131
7.2 Colonization of subterranean environments 133
7.3 What determines success or failure of colonizations? 135
7.4 Allopatric and parapatric speciation 136
7.5 Vicariance and dispersal 142
7.6 Evolutionary and distributional history of A. aquaticus 151
7.7 Summary 153

8 Geography of subterranean biodiversity 155

8.1 Introduction 155
8.2 The struggle to measure subterranean biodiversity 156
8.3 Caves as islands 162
8.4 Global and regional species richness 166
8.5 Summary 177
 CONTENTS xiii

9 Some representative subterranean communities 179

9.1 Introduction 179
9.2 Superficial subterranean habitats 180
9.3 Interstitial habitats 183
9.4 Cave habitats 187
9.5 Summary 194

10 Conservation and protection of subterranean

habitats 195
10.1 Introduction 195
10.2 Rarity 196
10.3 Other biological risk factors 199
10.4 Threats to the subterranean fauna 200
10.5 Site selection 208
10.6 Protection strategies 209
10.7 Preserve design 212
10.8 Summary 214

Glossary 215
References 221
Index 247
Site Maps and Gazetteer

List of sites mentioned in text. The associated number refers to the num-
bers on the maps. Several sites in Bosnia & Herzegovina, France, Slovenia,
and West Virginia (USA) were so close to each other that they are rep-
resented by the same number. All sites can be found on one of the three
maps, except for sites 29 and 51.

Abisso di Trebiciano, Italy 1

Alpena Cave, West Virginia, USA 2
Ayyalon Cave, Israel 3
Baradla/Domica, Slovakia/Hungary 4
Bayliss Cave, Queensland, Australia 5
Bellissens, France 65
Blue Lake Rhino Cave, Oregon, USA 6
Bracken Cave, Texas, USA 7
Carlsbad Caverns, New Mexico, USA 8
Lechuguilla Cave, New Mexico, USA 8
Cave Spring Cave, Arkansas, USA 9
Cesspool Cave, Virginia, USA 10
Col des Marrous, France 65
Columbia River basalt, Washington, USA 11
Cueva de Villa Luz, Mexico 12
Devil’s Hole, Nevada, USA 13
Dillion Cave, Indiana, USA 14
Dorvan-Cleyzieu, France 15
Edwards Aquifer, Texas, USA 16
Flathead River, Montana, USA 17
Greenbrier Valley, West Virginia, USA 18
Grotta di Frasassi, Italy 19
Grotte de Sainte-Catherine, France 20
Gua Salukkan, Sulawesi, Indonesia 21
HaLong Bay, Vietnam 22
Hellhole, West Virginia, USA 23
Hidden River Cave, Kentucky, USA 24
Inner Space Caverns, Texas, USA 25
Jameos del Agua, Tenerife, Canary Islands 26
Kartchner Caverns, Arizona, USA 27
Kavakuna Matali System, Papua New Guinea 28
 SITE MAPS AND GAZETTEER xv

Kazumura Cave, Hawaii, USA 29

Križna jama, Slovenia 30
Lachein Creek, France 20
Lobau Wetlands, Austria 31
Logan Cave, Arkansas 9
Logarček, Slovenia 32
Lower Kane Cave, Wyoming, USA 33
Lower Potomac, District of Columbia, USA 34
Lubang Nasib Bagus, Sarawak, Malaysia 35
Mammoth Cave, Kentucky, USA 36
McClean’s Cave, California, USA 37
Grotte de Moulis, France 20
Old Mill Cave, Virginia, USA 38
Organ Cave, West Virginia, USA 39
Otter Hole Cave, Wales, United Kingdom 40
Paka, Slovenia 41
Peştera Movile, Romania 42
Peştera Urşilor, Romania 43
Pivka River, Slovenia 46
Pless Cave, Indiana, USA 44
Popovo Polje, Bosnia & Herzegovina 45
Postojna-Planina Cave System, Slovenia 46
Rhône River at Lyon, France 47
Robber Baron Cave, Texas, USA 48
Robe River, Western Australia, Australia 49
San Marcos Spring, Texas, USA 50
São Mateus Cave, Goiás, Brazil 51
Sarang and Subis Karst, Borneo, Malaysia 52
Scott Hollow Cave, West Virginia, USA 39
Segeberger Höhle, Germany 53
Shelta Cave, Alabama, USA 54
Shihua Cave, China 55
Sierra de El Abra, Mexico 56
Silver Spring, Florida, USA 57
Sotano de las Golandrinas, Mexico 60
South Platte River, Colorado, USA 61
Šipun, Croatia 58
Škocjanske jame, Slovenia 59
Tantabiddi Well, Western Australia, Australia 62
Thompson Cedar Cave, Virginia, USA 63
Thornhill Cave, Kentucky, USA 64
Tour Laffont, France 65
Trebišnjica River System, Bosnia & Herzegovina 45
Triadou well, France 66
 SITE MAPS AND GAZETTEER xvi

Tular, Slovenia 67
Vjetrenica, Bosnia & Herzegovina 45
Voronja, Abkhazia 68
Walsingham Cave, Bermuda 69
Ward’s Cove, Virginia, USA 70
Young-Fugate Cave, Virginia, USA 71
Zink Cave, Indiana, USA 72
 11 17

6 33

34
61 72 2
37
13 24 38 36 71 70
9
27 54
8 69
7 25 57

56
60

12

53
40

4
43
47 1 41
66 31 68
19 58 42
65

3
26
 55

22

52

21 28

49 5

62
 1 The subterranean domain

1.1 Introduction
Beneath the surface of the earth are many spaces and cavities. These spaces
can be very large—some cave chambers such as the Sarawak Chamber,
with an area of over 21,000,000 m3 in Lubang Nasib Bagus (Good Luck
Cave) in Sarawak, Malaysia (Waltham 2004), can easily accommodate the
world’s largest aircraft. They can also be very small, such as the spaces
between grains of sand on a beach. These spaces can be air-fi lled, water-
fi lled, or even filled with petroleum. All of these spaces share one very
important physical property—the complete absence of sunlight. This is
a darkness that is darker than any darkness humans normally encoun-
ter, a darkness to which our eyes cannot acclimate no matter how long
one waits. There are some habitats that are dark and yet have some light.
The ocean abyss is nearly without light but many organisms of the abyss,
such as the well-known angler fish, produce their own light with the help
of microbes. In addition, the heat of deep sea vents is high enough that
light is emitted (Van Dover 2000). In subterranean habitats, with very rare
exceptions, this does not happen. The most notable exception is that of
glow-worms (actually fungus gnat larvae) in a few caves in Australia and
New Zealand. But even in these special cases, organisms cannot use light
to find their way about, to find food, to find mates, and so on.
Taken together, the water-fi lled and air-fi lled cavities are quite com-
mon, perhaps more common than surface habitats. Over 94% of the
world’s unfrozen freshwater is stored underground, compared with only
3.6% found in lakes and reservoirs, with the rest in soil, rivers, and the
atmosphere (Heath 1982). Heath estimates that there are 521,000 km3
of subsurface spaces and cavities in the soils and bedrock of the United
States, and most of these contain water. Whitman et al. (1998) indicate
that between 6% and 40% of the total prokaryotic (organisms with no
nuclear membrane such as bacteria) biomass on the planet may be in the
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Ulcerative endocarditis, infectious osteomyelitis, pulmonary
gangrene, general pyæmia, and, as is claimed by a few authors,
typhoid fever, are often accompanied by multiple abscesses in the
brain-substance. Usually the foci are small, as may be readily
inferred from the fact that they are of embolic origin, the emboli being
usually so small as to lodge in very small vessels, and that the
fatality of the primary disease is so great as to cut short life before
the abscess can reach larger dimensions. For the same reasons the
symptoms they produce are rarely distinctive. In chronic lung
affections accompanied by putrescence in bronchiectatic or other
cavities cerebral abscesses are not uncommon occurrences. Under
these circumstances, although we must assume an embolic origin,
the abscess is rarely multiple, and the symptoms are as marked as
in the ordinary varieties. Thus a patient suffering from chronic
phthisis, with or without prodromal malaise or somnolence,
experiences formications and pain in his right leg; he then notices a
slight halt in walking; twitches appear in the affected extremity; it
becomes distinctly paretic. The arm then becomes affected in like
manner; the pupils become unequal; a severe chill occurs, followed
by delirium, convulsions, coma, and death.

DIAGNOSIS.—There is little difficulty in recognizing the existence of a

cerebral abscess in which well-marked focal and constitutional
symptoms coincide, or where a distinct abscess-producing cause,
such as an ear trouble, a head injury, or a putrid bronchiectasis,
coexists.59 But there are a number of cases, varying from the latent
form to forms with obscure general symptoms, whose recognition is
impossible or at best a matter of conjecture. Such cases may be
readily confounded with certain tumors. The existence of febrile
symptoms, although not excluding tumor, as some tumors are
accompanied by such, is greatly in favor of abscess. On the other
hand, choked disc, which is rare with abscess and found only with
the very largest, is in favor of tumor.
59 Yet a leading and careful authority was misled into making the diagnosis of
abscess in a case of ear disease complicated by a cerebral tumor.
PROGNOSIS AND TREATMENT.—The majority of cerebral abscesses
must, from the nature of the case, be regarded as not influenceable
by medicinal measures or surgical treatment. The miliary and other
abscesses due to general septic causes or to mycotic invasion,
being in the nature of the case but features of intrinsically dangerous
or fatal primary diseases, do not call for special measures. It is
different with those due to local trouble about the head and to
surgical causes. Remarkable advances have been made in the
operative treatment of cerebral abscesses, chiefly owing to the
increasing accuracy of the localization of the affected areas through
the disturbance of their function, and to the perfection of surgical
methods. A number of cases by Gussenbauer, Wernicke, and others
have shown that some abscesses may be accurately located during
life by the focal symptoms produced by their presence. Wernicke's
observation of a large abscess in the occipital lobe showed two
facts. First, it permitted the study of the effect of large abscesses on
the cerebral movements, as it was found after trephining that the
pulsation movement of the brain was lost and the dura tensely
bulging, thus indicating a high degree of cerebral pressure.
Secondly, the operation showed that an abscess can be emptied of
its contents, under moderate aseptic precautions, without provoking
contiguous inflammatory reaction or infecting the meninges.
Notwithstanding these favorable local conditions, the patient died.
Gussenbauer60 was more fortunate. He surmised from the fluctuation
of some symptoms and the predominance of others that his patient
had an abscess in the frontal lobe. The suspicion was verified: an
abscess of the size of an apple was found, opened, and emptied of
its contents. The patient recovered without any immediate untoward
symptom.61
60 Prager medizinische Wochenschrift, 1885, Nos. 1, 2, and 3.

61 Epileptic and focal spasms subsequently developed, which shows that a new
inflammatory or other destructive process may have set in in the vicinity of the
emptied sac.
The uncertainties of localization in some districts of the brain are so
great that a number of attempts to repeat the explorations and
aspiration of Wernicke and Gussenbauer have failed. In one case
recently operated on in New York City the aspirating-needle was run
into the brain-substance in several different directions without
striking the pus. It is a question under such circumstances whether
the chances of an abscess becoming latent, minimal though they be
in cases with pronounced signs, are not to be preferred to those
which an uncertain operation can give. The superficial encephalitic
foci offer far better opportunities for surgical triumphs. Here not only
the symptoms are much more constant, and point more unerringly to
the site of the morbid spot, but there are often other signs, such as
the evidences of impaction of a foreign body, local tenderness on
percussion, or bone disease, which aid in determining the proper
spot for the application of the trephine. Several operations where
traumatic encephalitis existed with or without leptomeningitis of the
convexity, followed by complete recovery, were performed by
Macewen.62
62 The Lancet, 1885, vol. i. p. 881.

The medicinal treatment of abscess of the brain is limited to

derivative methods, whose aim is the relief of pressure—an aim
whose fulfilment is more frequently illusory than otherwise.

Benefit has been claimed from the energetic use of mercury, chiefly
in the form of calomel, by older writers; and recently Handfield Jones
has endorsed its administration, attributing to it a remission in a case
in which it was employed. It must be remembered, however, that
remissions occur spontaneously in this disease, and that the
purgative action of calomel may act well for the time being in an
affection so apt to be associated with hyperæmia and increased
cerebral pressure as is an abscess of the brain.63 In the nature of the
case, even this latter momentarily beneficial effect is at the best
temporary.
63 Brain, October, 1884, p. 398.
The prophylaxis of cerebral abscess can be carried out only in cases
due to cranial and aural affections. The importance of treating all
scalp and cranial injuries under aseptic64 precautions is recognized
by all surgeons. It is generally admitted that the trephining of a bone
suspected to be the site of an ostitic or necrotic process involves
fewer risks than the allowing it to remain. Similar principles govern
the treatment of the inflammatory involvement of the mastoid cells
often complicating otitis media. If trephined at all, these should be
trephined at the earliest moment. It was a belief among the older
aurists that the sudden cessation of an aural discharge was of evil
augury, and that cerebral complications were more apt to follow
under such circumstances than when the ear discharged freely. Von
Tröltsch, Politzer, Gruber, and Toynbee have opposed the exclusive
application of the old dogma. In so far as the older ear-surgeons
regarded a profuse aural discharge as an encouraging sign, in this
respect they were of course wrong. But their observation of the
frequent concurrence of cerebral sequelæ with suppression of
discharge is, I think, borne out by a large number of cases. It does
not apply, however, to the suppression of discharge by the rational
employment of aseptic injections.
64 This term is used in its widest sense here.

Spinal Hyperæmia.

The spinal cord is found to vary considerably in color in different

individuals. To some extent this difference is influenced by the
position occupied by the body after death, but not as markedly as in
the case of the brain. Thus it will be paler in a body which has been
kept in the prone than in one which has been kept in the supine
position.65 It is usually found more injected in persons who have died
of febrile affection than in those who have died of exhausting
diseases; notably is this the case with typhus fever and with subjects
who die with congestive malarial chill. But the most intense
congestion, where the gray substance instead of having the normal
rosy hue appears like a blood-soaked sponge, and the white
substance instead of the tint to which it owes its name has a pink
shade, is found in subjects dying in convulsive disorders complicated
by asphyxia, such as epileptic status, tetanus, and certain toxic
disorders.
65 I have not seen it noticed anywhere, but it is a fact readily demonstrable in any
autopsy made in a well-preserved body that when a short segment of cord is
observed at the moment of section the section surface appears pale; but if it be again
examined after a few moments, it will show a reddish tinge, marked as a faint injection
would be; this tinge distinctly deepens under the eye. I have observed this in
specimens which were not held in the hand, so that the influence of pressure can be
excluded. Nor do I believe that the elasticity of the tissues is such as to account for
the phenomenon.

In attempting to apply the fact that varying degrees of vascular

injection are found on post-mortem examinations of the spinal cord
to the elucidation of certain clinical phenomena, we encounter the
same difficulties and sources of error that confronted us in the study
of nutritive brain disorders. The majority of writers have therefore
contented themselves with making a careful clinical study of the
mostly subjective signs of disorders which, once designated as
spinal anæmia and hyperæmia, are now classified under the non-
committal titles of spinal irritation and exhaustion (neurasthenia), as
in this volume. A number of these disorders, like the so-called
anæmic paralysis of Bouchut, Leroy d'Etiolles, Beroliet, Baimer, and
Brandis, would to-day be considered as hysterical or reflex; and a
few of the instances cited by their contemporaries as cases of spinal
hyperæmia have been since demonstrated to resemble the initial
phases of organic diseases of the cord.

The causes of active spinal hyperæmia are either direct, as when the
spinal centres are overtasked by muscular strain either through over-
exertion or through toxic convulsions, surprised by violent shocks,
such as concussion accidents, or collateral, as when a physiological
discharge (menstruation) or a pathological one (hemorrhoidal flux) is
suddenly checked. A few cases are reported where carbonic-oxide-
gas poisoning provoked spinal hyperæmia. But, like the alleged
cases of spinal hyperæmia after continued and exanthematic fevers,
they were probably cases of incipient or established myelitis.
Hammond claims that surface chilling exerts the same congesting
influence on the cord which he claims for the brain; but no definite
observations have been made in this direction.

Passive spinal hyperæmia has been attributed to obstructive cardiac

and chronic pulmonary affections. In such cases, as with most
causes acting on the circulation of blood in the nerve-centres, the
coexisting cerebral congestion usually masks the spinal. It is a
question how far the intense hyperæmia of the cord found in some
cases of tetanus, strychnia-poisoning, and the condition called
hydrophobia66 is primary and an indication of neural hyperexcitability,
and how far it is secondary to the asphyxia attending the last phases
of these convulsive states. The weight of opinion is in favor of an
acceptation of the latter as the chief or only factor.
66 In a case of hydrophobia clinically as well marked as has been recorded, which I
had the privilege of examining through the courtesy of Kretschmar, both brain and
cord were found remarkably anæmic.

Over-exertion and sexual excesses are frequently followed by a

sensation of fulness and tension in the sacral and lumbar regions,
which may be relieved by lying prone, while it is aggravated while
lying supine.67 From the location of this pain it is evident that it is not
due to congestion of the cord or its membranes, but to fulness of the
vertebral and spinal veins of the lower segment of the vertebral
column. A similar sensation, which may be relieved by the same
change in position or by a hemorrhoidal flux, is complained of by
patients suffering from portal obstruction. The veritable symptoms of
hyperæmia manifest themselves in the parts which receive their
nervous supply from the affected districts. The reflexes are usually
more active; paræsthesias of different kinds, such as formication,
tingling, and creeping sensations, are common; and there is more or
less motor weakness, the limbs feeling heavy and sometimes being
the seat of an acute pain. As a rule, these symptoms are limited to
the lower half of the body.
67 Although this fact has been questioned, I have no doubt whatever that it is true,
from a large number of observations. In many subjects suffering from the results of
excessive venery or masturbation, an intolerable, sometimes pulsating, feeling in the
lumbo-sacral region is only relieved by raising the lower end of the trunk with the back
up. This condition is influenced by a change of residence to a district having a
different level above the sea, and consequently a different barometric condition.

There is very little question when these symptoms exist for any
length of time, and become aggravated, that more subtle nutritive
changes than are covered by the single term hyperæmia become
responsible for them. In a pure hyperæmia the position-test of
Brown-Séquard, which shows relief when the patient is upright or
prone and aggravation when he is supine, particularly if the
gravitation of blood to the cord be facilitated by raising the head and
extremities, ought to yield constant results. But in some cases,
particularly those of long-standing, the very opposite is noted: the
patient's symptoms are aggravated by standing or sitting up, and
relieved by lying down. Here there is probably exhaustion or
malnutrition of the nerve-elements, rendering them abnormally
sensitive to exertion. This view is supported by the fact that
molecular disturbances, such as those which probably accompany
simple concussion, predispose the patient to the development of the
symptoms of spinal hyperæmia, and aggravate them if established
previous to such accident.

Hammond,68 who in his chapter on Spinal Congestion and Anæmia

follows rather the older authors, such as Ollivier, than the newer and
either more cautious or more sceptical writers on the subject,
describes the symptoms of congestion as comprising belt
sensations, paraplegia, erections of the penis, muscular twitches,
loss of expulsive power, and incontinence of the bladder, paralysis of
the abdominal muscles, paralysis of the anal sphincter, loss or
abolition of reflex excitability, diminution of electro-muscular
contractility, and occasionally hyperæsthesia and shooting pains. It is
not doubtful for a moment that if such a case were to occur in
hospital experience it would be regarded as one of organic disease,
and not incipient, but well-established organic disease of the cord.
Those making the diagnosis would have their opinion strengthened
if, as Hammond states, the process took place with great rapidity
and had a tendency to extend itself and eventually involve the whole
cord, or if, as Brown-Séquard is by him cited as stating, bed-sores
occurred in addition. Although Hammond describes certain
anatomical changes, such as increased development of blood-
vessels and distension and injection of them, I am unable to find any
cases recorded as spinal congestion during life, and carefully
examined with a due regard to sources of error after death, in which
such changes were found. It is true that after strychnine- and
cocaine-poisoning an intense hyperæmia69 of the cord is found. In
mammals it is of a far more pronounced character than in reptiles,
and usually more marked in proportion to the existing asphyxia. That
the characteristic toxic effects of these drugs is not to be sought for
in their direct or indirect congesting influence is shown by the fact
that exsanguinated frogs can be made to undergo strychnine tetanus
when their blood is replaced by a saline solution according to the
method of Salkowski. Little support, therefore, could be derived from
a pretended analogy between toxic and pathological hyperæmias,
even if the phenomena of both were similar; which is not the case.
68 Diseases of the Nervous System, 7th ed., p. 392.

69 It is true that in animals which are so organized that the congestion cannot be
attributed to asphyxia, as I showed (Hammond Prize Essay of the American
Neurological Association, 1878) in some experiments on strychnine, arterial
congestion and small foci of hemorrhage were found in the upper cervical cord of
frogs who had been kept in continuous strychnine tetanus for over seventy days.

The introduction of subaqueous caissons for workmen engaged in

the building of bridges, in which those employed labor under
abnormal atmospheric pressure, has led to the development of a
previously-unknown cerebro-spinal affection known as the caisson
disease, and in which, it is generally supposed, either congestion or
hemorrhage of the spinal cord occurs in consequence of sudden
changes of vascular pressure resulting from sudden diminution of
the barometric pressure. Clinically, this affection has been studied in
England, France, and above all in America in connection with the
building of the East River Bridge and the one over the Mississippi at
St. Louis.70 Experiments by Hoppe-Seyler, Bert, and I. Rosenthal
have shown that a sudden diminution of pressure leads to
hemorrhages in various tissues, and, according to the two first-
named, a development of gas occurs in the vascular and other fluids
of the body. A number of peculiar symptoms which do not specially
interest us here occur in conjunction with the so-called caisson
disease: these are—pain in the ear, with or without otitis sicca;
peculiar pains in the joints, which occur on leaving the caisson, and
are probably due to hyperæmia of the joint-surfaces and sudden
increase of the intra-articular fluid; and retardation of the pulse-rate.
In some cases cerebral hyperæmia is added, the patients tottering
about as if drunk. The spinal symptoms consist of a paraplegiform
affection. The paralysis is usually sudden; in some cases the patient
a few minutes after stepping from the air-chamber falls down
perfectly helpless as far as the lower half of the body is concerned.
The expulsive power of the bladder is usually weakened, and there
is anæsthesia to all forms of sensation in the affected limbs, as well
as diminished electro-cutaneous sensibility. The patient often
complains of a strange feeling, as if the lower half of his body were a
foreign substance. With this the electro-muscular reactions are
normal. In the majority of cases these symptoms disappear entirely
in from three to ten days, but occasionally they remain longer;
imperfect recovery of motion and sensation occurs, or, as happened
in a few cases, one of which was carefully examined during life by
Lehwess and after death by Leyden, death occurs as in myelitis. In
the only case where an autopsy and careful microscopical
examination were made under these circumstances71 peculiar
fissures were found in the substance of the spinal cord, surrounded
by areas of reactive myelitis and filled with granule-cells. The
absence of any pigmentary relics of a hemorrhage induced Leyden
to assume that the lacunæ were not of hemorrhagic origin. He
inclines to the view that they were due to the escape of gas from the
blood-plasma, and consequent multilocular inflation of the tissue. If
his observation be confirmed, it constitutes a strong objection to the
hyperæmia theory of the caisson disease. There is neither
permanent hyperæmia nor congestive or hemorrhagic myelitis
developed, as far as the limited material thus far studied permits a
conclusion.
70 Clark, St. Louis Med. and Surg. Journ., cited from Hammond, loc. cit.

71 E. Leyden, Archiv für Psychiatrie, ix. p. 316.

Pure spinal hyperæmia rarely presents itself for treatment. The form
due to over-exertion is recovered from by rest in a very short time;
that due to suppressed discharges, by the re-establishment of the
latter or by the application of leeches to the lumbo-sacral and iliac
region. Ergotin is recommended by Hammond in very large doses. It
is a question whether this drug may not exert a bad effect in
protracted cases where its use has to be continued for a long time.72
In using it, it is well to bear in mind that imperfect nutrition of nerve-
elements is perfectly compatible with an increased blood-amount.
72 A young physician, who for a long period took ergotin in twelve-grain doses for the
relief of symptoms regarded as congestive, acquired a tolerance of the drug such as I
have not seen recorded anywhere, and in addition presents some obscure signs of
cerebellar disease and initial optic-nerve atrophy.

Strychnia has been given with benefit in the caisson disease—

another evidence, as this drug is theoretically contraindicated in true
hyperæmia, that this disease is not, as Hammond and the majority of
authors with him regard it, essentially a congestive affection. The
treatment of those numerous cases in which signs of venous fulness
accompany spinal exhaustion and irritation is detailed in the articles
dealing with those affections.

Spinal Anæmia.
Anæmia of the cord-substance proper, like hyperæmia, is practically
inseparable from the corresponding condition of the membranes.
The influence of a reduced blood-amount on the functional activity of
the spinal cord is more susceptible of exact demonstration than the
corresponding nutritive disturbance of the brain. As the functions of
this segment of the nervous axis are far simpler than those of the
higher organ, there is more unanimity among observers as to the
interpretation of their disordered states. In Stenon's experiment, and
the more elaborate modifications made by those who have followed
his method, it is found that interference with the supply of arterial
blood to the spinal cord is followed by abolition of the function of the
gray matter; if the supply be still further diminished, the functions of
the white tracts become eliminated; next the peripheral nerves, and
ultimately the muscles themselves, lose their normal excitability. On
the re-establishment of the circulation these various parts regain
their functional capacity in the inverse order of its suspension—the
muscles first, next the nerves, then the white substance, and last the
gray substance of the cord. The initial symptoms of some cases of
myelitis from refrigeration correspond more nearly to such a result of
artificial anæmia of the cord than they do to anything that is
customarily regarded as hyperæmia.73
73 I have seen distinct pallor of the spinal meninges on dipping the posterior
extremities of a dog, whose cord had been exposed, into water. It is to be remarked,
however, that other observers, notably Hammond, have either obtained different
results or interpreted the consequences of refrigeration differently.

No one has gone farther than Hammond in erecting a theoretical

anatomical framework which elaborately provides for the
accommodation of various symptoms of spinal anæmia. He
describes anæmia of the posterior columns, and sharply
discriminates between it and anæmia of the antero-lateral columns.
It is a question whether the conducting tracts of these columns are
seriously affected in their functions by anæmia as long as the
centres of innervation are well nourished. Undoubtedly, it is the gray
substance of the cord which is most vulnerable to the influence of
disturbed circulation and nutrition, as Stenon's experiment has
shown; and a glance at the distribution of the blood-vessels will show
that a partial anæmia or hyperæmia, limited to special cornua in any
considerable length of the cord, is an exceedingly improbable
occurrence. With regard to isolated anæmia of the white columns, it
is to be admitted that the posterior are most vulnerable to
malnutrition. But it is doubtful whether this vulnerability is so great as
to allow of an exclusively posterior anæmia, or whether a protracted
anæmia of this kind could exist for years as a purely symptomatic—
or, as some designate it, functional—disorder.

Hammond candidly states that in specifically locating the lesions in

these affections he is aware that post-mortem examinations are
wanting to support them, and admits that what he calls anæmia of
special parts of the cord is the spinal irritation of most authors, and in
part the reflex paraplegia of others.

The most clearly-established form of cerebral anæmia is the one

which is indicated by the ischæmic paraplegia of Jaccoud and the
paraplegia following profuse hemorrhages. The former is produced
by all causes which, by obstructing the flow of arterial blood in the
abdominal or thoracic aorta, cut off the proper blood-supply to the
cord, which the latter receives through the intercostal and upper
lumbar arteries. Aneurism, compression by tumors, and embolism of
the aorta produce this result. The consequence is paraplegia
corresponding in all features of its development to the phenomena
observed in Stenon's experiment. These features, already detailed,
suffice to show that it is not the anæmia of the peripheral nerves and
muscles that is chiefly responsible for the paraplegia, but the
insufficient irrigation of the gray and white substance of the cord
itself. The same is true of the paraplegia following hemorrhage which
has been noted after uterine, renal, and enteric hemorrhages. Both
affections are exceedingly rare.

The influence of general anæmia on the functions of the spinal cord

is not susceptible of accurate study. The cerebral enlargement of the
nervous axis is so much more unfavorably situated than the cord that
it suffers first and most when general anæmia is present. The
consequence is that the signs of cerebral anæmia mask those of
spinal anæmia. It is supposed, however, by many authorities that the
effect of anæmia on the cord may be regarded as an auxiliary factor
in the production of hysterical and neurasthenic symptoms.

How far the spinal cord is liable to suffer from arterial spasm is as yet
a matter of conjecture. It is supposable that just as a powerful
psychical impression provokes a sudden spasm of the cerebral
arteries, so a peripheral irritation may provoke a spasm of the spinal
arteries. In this way the reflex paralyses, motor and vaso-motor, are
explained by many writers.

The subject of reflex palsy has been so much confused by improper

cataloguing—if it can be so called—that some of the best authorities
have become sceptical as to its occurrence. Among the chief
sources of error has been the attributing to irritation of the genital
organs various convulsive, psychical, and paralytic disorders.
Adherence of the prepuce and its excessive length were charged
with being responsible for idiocy, imbecility, epilepsy, and every form
of paraplegia and panplegia. It was further claimed that instances of
complete cure of each of these affections had followed the removal
of the exuberant or adherent prepuce. I cannot find a single instance
recorded where such a cure was effected in any of our large medical
centres, so as to prove convincing to critical colleagues. On the
contrary, L. C. Gray74 has shown that various surgical procedures
have been needlessly resorted to on this erroneous theory in cases
of organic diseases of the spinal cord. I have seen two unfortunate
children suffering from the worst forms of anterior poliomyelitis, one
afflicted with pseudo-hypertrophic paralysis, and several
hydrocephalous and microcephalous idiots, whose prepuces had
been sacrificed to the theory alluded to—it is needless to add without
any result, good or bad.
74 Reflex Irritation from Genital Irritation. In this paper written communications from all
or nearly all neurologists in the United States—certainly including all those of national
fame and large experience—are cited, in which they testify to never having seen a
 case of this character cured by operations on the penis (Annals of Anatomy and
Surgery, Jan. and Feb., 1882.)

The possibility of a reflex paralysis occurring from genital irritation in

the male cannot be denied; among the lower animals a ligature
around the spermatic cords sometimes produces paraparesis, and
paraplegia is a common complication of renal and vesical troubles in
others. But analogous observations in man are rare, and becoming
rarer with our increasing acumen in diagnosis. In females peculiar
reflex disturbances are found associated with uterine and ovarian
derangements. In one case of retroflexion, with possible dislocation
of the ovary, referred to me by H. J. Boldt, there is a remarkable
vaso-motor paralysis of the right arm during each menstrual period:
this member becomes greatly enlarged, of a purplish-blue color, and
cold. Equally remarkable are the reflex disturbances resulting from
the presence of worms in the intestinal canal. Every form of spinal
and cerebral paralysis, even aphasia, has been observed in
connection with helminthiasis. Such disorders yield as rapidly as
they are developed to the exhibition of vermifuges.

Special interest has been aroused by the discovery laid down in the
joint treatise of J. W. Mitchell, Morehouse, and Keen of reflex
paralysis following injuries, observed in the War of the Rebellion. The
cases cited by them appear singular on first sight. The paralysis is
often observed in parts of the body which are not only remote from
the seat of injury, but have no direct connection, physiologically or
otherwise, with it. The hand may be injured and the opposite leg
paralyzed.

Since Mitchell, Morehouse, and Keen first announced the existence

of this peculiar form of reflex paralysis a careful search has been
made by military surgeons engaged in other campaigns for like
results. Notably was this done in the Franco-Prussian War. A number
of confirmatory instances have been collected, some of which rival in
singularity those related by the discoverers of the affection. In one
case a unilateral paralysis agitans followed a punctured wound of the
opposite shoulder, and in another reflex aphasia followed a gunshot
wound of the lumbar region.75 A discrimination is to be made
between such cases where the paralysis, anæsthesia, or neuralgia is
an immediate result of the injury, and those where they follow after
weeks or months. In the latter instance we have not true reflex
disorders to deal with, an ascending neuritis having been found in
the few cases which could be carefully followed up.76
75 Sanitäts Bericht über die deutschen Heere im Krieg gegen Frankreich, 1870-71,
vol. vii.—abstracted in Neurologisches Centralblatt, 1886, p. 207.

76 In a case of Mollenhauer's, vesical paralysis and paresis of the right leg occurred
six years ago (1880) in a veteran of our civil war who had a gunshot wound of the
right hand, with signs, which are still present, of occasional exacerbation of brachial
neuritis. Prodromal signs of paresis were noticed at intervals since his return from the
campaign. The bladder trouble and paresis are now apparently stationary. Such a
case can be accounted for only on the assumption of an organic cord-change
secondary to a neuritis.

The theory that the reflex paralysis from utero-ovarian, intestinal, and
surgical affections, when acutely produced, is due to central
anæmia, is as acceptable as any other would be in the absence of
decisive observations.

Spinal anæmia will but rarely present itself as a subject for special
and separate treatment. When not associated with an intrinsically
grave condition, such as aortic obstruction, dysentery, fatal
hemorrhage, or typhoid fever, it is an exceedingly benign affection,
rapidly yielding to tonic and restorative measures combined with
rest.

Embolism, Thrombosis, Hemorrhage, and Abscess of the Spinal

Cord.

Although the spinal cord is a segment of the same central organ as

the brain, nourished in a similar way, and subject to the same
physiological and pathological laws, lesions of the vascular
apparatus, which play so important a part in brain pathology, play a
comparatively insignificant one in that of the spinal cord. Embolic,
thrombic, and primary hemorrhagic lesions of the cord are so rare
that their possible existence has even been denied. A primary
thrombosis of the cord has not yet been satisfactorily demonstrated
to occur independently of syphilitic lesions; and when it occurs the
ensuing tissue-changes, as described by Heubner, Julliard, and
Greiff, are usually in the background as compared to the
gummatous, sclerotic, or meningitic changes which coexist. The
clinical as well as the anatomical picture is accordingly either one of
a myelitis or meningitis, as the cases of Charcot-Gombault, Heubner,
McDowell, Wilks, Wagner, Zambaco, Homolle, Winge, Moxon,
Schultze, Westphal, Julliard, and Greiff show. (See Myelitis and
Spinal Meningitis.)

With regard to the occurrence of hemorrhage into the substance of

the spinal cord (hæmato-myelia), it is so rare an occurrence that I
can recall but a single case in which I entertained the diagnosis of
this lesion; and in that very case I am unable to declare that it was
not a hemorrhagic myelitis. Aneurismal changes of the spinal
arteries are comparatively of rare occurrence, and as other
predisposing causes to primary vascular rupture are rare in the cord,
the probability of its occurrence is very much diminished. Hebold,77
in a young girl who had developed severe cerebro-spinal symptoms
during a period of nine months following an erysipelatous disorder,
found the upper dorsal cord, on section, dotted with numerous
reddish and round points. These points corresponded to aneurismal
dilatations of the vessels. As there were other inflammatory and
vascular lesions in the same subject, the author referred their
causation to a general constitutional vice, the result either of the
phlegmonous or of a tuberculous disorder.
77 Archiv für Psychiatrie, xvi. 3. Rupture of miliary and other aneurisms in the
meninges has been reported by Astley Cooper, Traube, and others. It is remarkable
that such cases are more and more rarely recorded from year to year in inverse ratio
to the accuracy of our spinal autopsies. I have never found a miliary aneurism below
 the uppermost cervical level of the cord. On the other hand, I have found extensive
spinal hemorrhage in cases where the vessels of the cord proper were fairly healthy.

It is claimed that suppression of the menses, over-exertion, lifting

heavy weights, and concussion are causes of spinal hemorrhage.
The same causes are also mentioned for acute hemorrhagic
myelitis; and it is a question whether the supposed hemorrhage is an
initial lesion or secondary to congestive or anæmic softening.78
78 I have never found vascular ruptures, although carefully searching for them, in the
spinal cord of persons dying instantly after falls from a great height, or, as in one case
which I was fortunately able to secure the cord of, where the subject had been
violently thrown down. Where hemorrhages have been found under these
circumstances they were, as far as I am able to learn from the cases recorded,
meningeal.

The symptoms attributed to spinal hemorrhage are the same, taking

the same locality of the cord, as those of a very rapidly-developed
transverse myelitis. It is unnecessary to enumerate these here in
anticipation of the next section. They are described as being much
more sudden. This suddenness is the only diagnostic aid on which
we can rely.79 The fate of the patient is said by Erb to be decided
within a few days. If he survive the immediate consequences of the
hemorrhage, he is apt to recover, as to life, altogether, with such
permanent atrophies, paralyses, and anæsthesias as are entailed by
the destruction of the tracts and gray substance involved in the
hemorrhage. The treatment recommended for this condition consists
of rest, either in the lateral or prone position, local depletion and
derivation to the intestinal canal, as well as the internal use of
ergotin. The local application of ice, which is also advised, is
probably based on illusory views.80 After the immediate danger is
past the case is to be treated as one of myelitis—a very safe
recommendation in view of the probability that it was a case of
myelitis from the beginning.
79 And even this sign is unavailable as a distinguishing feature in supposed
hemorrhage from concussion, as sudden paraplegias of motion and sensation are
 found in some cases of railway spine, and, although a number of cases terminating
fatally have been examined, there was not always hemorrhage even in the meninges.

80 Until authorities shall have agreed as to what effect the exposure of the bodily
periphery to certain temperatures has on the circulation of the cord, it would be
premature to make any special recommendations as to the temperature at which they
should be kept. I am inclined to believe that while, as is universally accepted, a
general cooling of the bodily surface tends to increase vascular fulness in the cord, as
in all other internal organs, a partial cooling, as of the feet, produces local anæmia at
the level of origin of the nerves supplying the cooled part. Certainly, the bilateral
neural effects of unilateral cooling are in favor of this view.

The descriptions given of the hemorrhagic foci as observed after

death strengthen the view that they were in the majority of cases of
myelitic origin. Usually, they are stated to extend up and down the
cord in the direction of least resistance—that is, in the gray
substance—resembling an ordinary apoplectic clot. But in their
neighborhood there was usually considerable softening, and, to
judge by the descriptions given, this softening differed in no wise
from that which is the characteristic feature of acute myelitis;81 and
often the transition from a peripheral zone of white softening, through
an intermediate zone of red softening, to a central compact clot, is so
gradual as to leave it unquestionable that the softening pre-existed,
and that a vessel had broken down in the midst of the myelitic
detritus. Many ancient foci of myelitis betray the hemorrhagic
complication of their initial period by the presence of pigmented
residue of the absorbed clot.
81 In the latest treatise on nervous diseases published in our language (Ross, loc. cit.,
vol. ii. p. 325) the insufficient foundation on which a whole chapter has been built up is
illustrated by the admission that the usual evidence of acute central myelitis may be
observed far beyond the limits of the hemorrhagic infiltration. If a large area of
softening in the brain were found to contain a central or peripheral clot of blood, and
histologically resembling a typical embolic or thrombic softening, no one would be in
doubt as to which of the two was the primary lesion.

Embolisms and embolic softenings of that part of the spinal cord

which is supplied by the small spinal vessels are so rarely observed
in the dead-house that our knowledge of their possible occurrence
and character is almost entirely the result of experimental
observations or based on analogy. The situation of these vessels,
the angle at which their supply-tubes are given off from the aorta, all
act as protectors of the cord against what is one of the chief dangers
to which the brain is exposed. No definite symptoms have been
attributed to the few doubtful cases of simple embolic occlusion of
the spinal arteries found accidentally in human subjects. Even those
emboli which, when once let loose in the circulation, are found
distributed in nearly every organ of the body, those derived from
ulcerative endocarditis and those due to the invasion of micrococci,
are comparatively rare in the cord. Leyden found multiple capillary
emboli in the spinal cord from the former cause. Small grayish white
foci in a similar distribution were found to be due to an invasion of
cocco-bacteria from a decubitus by Rovigli.82 In this latter case an
increase of pain and muscular spasm in the history of the case of
sclerosis which was thus complicated was attributed to the parasitic
affection.
82 Rivista sperimentale di Freniatria, 1884, x. p. 227.

Just as simple and infectious embolic lesions are frequent in the

brain and rare in the cord, so purulent inflammation or abscess is an
exceptional occurrence in spinal as compared with cerebral
pathology, and probably for the same reasons.

Simple Acute Myelitis.

SYNONYMS.—Spontane (primäre) acute Rückenmarkserweichung,

Softening of the spinal cord, Ramollissement blanc de la moelle,
Myélite aiguë.

As Leyden, whose treatise83 may be regarded as the foundation of

our knowledge on this subject, correctly avers, it is to Abercrombie
and Ollivier that we owe the determination of the existence of that
acute structural disease of the spinal cord, now termed myelitis, as
an affection independent of meningeal changes. The anatomical
descriptions given by these older writers may be accepted at the
present day as models of accurate observation by the naked eye.
Their statement that in acute myelitis the substance of the spinal
cord is softened and changed into a puriform, yellowish, diffluent
mass; that while the disorganization is sometimes more marked in
the posterior, at others in the anterior, and occasionally in the lateral
half, it is most pronounced in the axis of the cord, because the
central gray substance is the favorite starting-point of the morbid
process,—requires no modification to-day. Considerable doubt
existed in the minds of the contemporaries of Ollivier and
Abercrombie as to whether this change was the result of a true
inflammation; and one of the clearest thinkers of the day, Recamier,84
regarded myelitic softening as a lesion peculiar to the nervous
apparatus, and different from ordinary inflammation. I believe that
the most profound investigators of the present day have not been
able to rid themselves of a similar doubt. The discovery of Gluge's
so-called inflammatory corpuscles, which was regarded as settling
the question, only served to confuse the student by the confidence
with which it was urged that they were infallible criteria of the
inflammatory process. Under the non-committal designation of
granule-cells these bodies still flourish in the annals of cerebro-spinal
pathology. As we shall see, a number of products of real disease, of
artifice, and of cadaverous change have passed and do pass muster
under this name. The first substantial progress in our knowledge of
the minute processes underlying inflammation of the spinal cord was
made by Frommann and Mannkopf, but it applied altogether to the
chronic inflammatory or cirrhotic affections of the cord. The
difference between acute and chronic myelitis is greater than is the
difference between acute and chronic inflammation in any other
organ; and it must be admitted that if Recamier is to be regarded as
having erred in asserting that acute myelitis is not a true
inflammation, he is justified in so far as he asserted many features of
the process to be altogether peculiar to the organ affected. Leyden
himself attempted to throw light on the subject by provoking myelitis
experimentally in dogs. He injected Fowler's solution into the spinal