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Brains as Engines of Association
Brains as Engines
of Association
AN OPERATING PRINCIPLE FOR
NERVOUS SYSTEMS
Dale Purves
1
1
Oxford University Press is a department of the University of Oxford. It furthers
the University’s objective of excellence in research, scholarship, and education
by publishing worldwide. Oxford is a registered trade mark of Oxford University
Press in the UK and certain other countries.
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© Oxford University Press 2019
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a retrieval system, or transmitted, in any form or by any means, without the
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above should be sent to the Rights Department, Oxford University Press, at the
address above.
You must not circulate this work in any other form

and you must impose this same condition on any acquirer.
Library of Congress Cataloging-in-Publication Data

Names: Purves, Dale, author.
Title: Brains as engines of association : an operating principle for nervous systems /
Dale Purves.
Description: New York, NY : Oxford University Press, [2019] |
Includes bibliographical references and index.
Identifiers: LCCN 2018035410 | ISBN 9780190880163
Subjects: LCSH: Brain. | Neurobiology. | Neural circuitry.
Classification: LCC QP356.25 .P86 2019 | DDC 612.8/2—dc23
LC record available at https://lccn.loc.gov/2018035410
9 8 7 6 5 4 3 2 1
Printed by Sheridan Books, Inc., United States of America

CONTENTS
Preface ix
Acknowledgments xiii
PART I } What Nervous Systems Do for Animals

1. Putting the Question in Perspective 3
Introduction 3
Life on Earth 3
Defining Life 5
Energy 6
Evolution 7
Mechanisms 8
Teleology 10
Conclusion 11
Suggested Reading 11
2. Organisms Without Nervous Systems 13

Introduction 13
Bacteria 13
Protists 16
Plants 18
The General Strategy 21
Conclusion 21
3. Organisms With Nervous Systems 23

Introduction 23
Defining Nervous Systems 23
The Emergence of Nervous Systems 26
The Emergence of Central Nervous Systems 28
What Do Brains Add? 28
Conclusion 33
vi { Contents
PART II } Nervous Systems as Engines of Association

4. The Organization of Nervous Systems 37
Introduction 37
Stimuli 37
Preneural Processing 38
Neural Processing 39
Behavioral Output 43
Neural Systems and Subsystems Are Interactive 45
Conclusion 47
5. The Problem 48
Introduction 48
Vision as an Example 49
The Basic Challenge 49
An Answer in General Terms 51
Qualia Determined by Empirical Ordering 54
Perceptual Discrepancies 54
Mechanisms 55
Other Modalities 56
Conclusion 57
6. Neural Associations 59
Introduction 59
Associations Wrought by Evolution 59
Associations Wrought by Lifetime Learning 61
Associations Wrought by Culture 65
Behavioral Categories of Associations 67
Reward 68
Behavioral Responses as Reflexes 69
What Is Associated? 70
Counterarguments 70
Conclusion 71
PART III } Evidence that Neural Systems Operate Empirically

7. Evidence from Lightness and Color 75
Introduction 75
Luminance and Lightness 75
Analyzing the Occurrence of Luminance Patterns 78
Effects of Other Luminance Patterns 80
Contents } vii
Spectral Energy and Color 84

The General Strategy 86
Conclusion 88
8. Evidence from Geometry 89

Introduction 89
Seeing Intervals 89
Seeing Angles 93
Seeing Object Sizes in Two Dimensions 96
Seeing Object Sizes in Three Dimensions 97
Seeing Stereo Depth 100
Conclusion 103
9. Evidence from Motion 104

Introduction 104
Apparent Motion 104
The Perception of Speed 106
Implications for the Perception of Time 110
The Perception of Direction 112
Conclusion 115
10. Evidence from Audition 117

Introduction 117
Sound Signals 117
Sources of Tones 118
Sound Signal Spectra 119
The Problem in Audition 120
An Empirical Approach 122
Evidence from Speech 123
Evidence from Music 127
Implications for Any Sensory System 129
Conclusion 129
PART IV } Alternative Concepts of Neural Function

11. The Major Options 133
Introduction 133
Neural Function as Feature Detection 133
Neural Function as Statistical Inference 136
Neural Function as Predictive Coding 137
viii { Contents
Neural Function as Efficient Coding 138

Neural Function as Computation 140
Conclusion 143
12. Summing Up 145

Introduction 145
Obstacles and a Way Around Them 145
Empirical Ranking 146
Winning Games 147
Biological and Artificial Intelligence 148
Consequences for Neuroscience 150
The Status of Reasoning 152
Novel Situations 153
Choice 154
Culture 155
Frequency of Stimuli 155
Conclusion 156
Glossary 159
Bibliography 169
Index 187
PREFACE
This book concerns a question that has bothered me—and no doubt many others—
for a long time: What does the human brain and the rest of the nervous system
actually do?
Based on subjective experience, the question seems gratuitous: Isn’t the brain the
ultimate organizer of our behavior and the seat of the “I” when we think and act?
The term “behavior,” however, includes an enormous range of responses to internal
and external stimuli, and what “organizing” this welter of reactions means is not at
all clear. Likewise, what the “I” might be in this conception of neural function has
been debated for centuries without resolution.
If the goal of biology is to understand how living things succeed in the world
and the mechanisms they use to do so, then this sort of answer only papers over
our ignorance about the function of the nervous system compared to other organs.
By middle school, kids have learned that the respiratory system oxygenates the
blood, that the digestive system infuses blood with nutrients, that the cardiovas-
cular system conveys oxygen and nutrients to body tissues, and so on. An oper-
ating principle of one sort or another has been documented for every major organ
system save one: the brain and the rest of nervous system.1 Although textbooks and
an overwhelming original literature testify to the wealth of neurobiological infor-
mation gleaned over the past century or more, a comparable statement about the
nervous system will not be found.
This deficiency has led some investigators to imagine that the human brain is so
specialized that understanding it will require approaches that have not been needed
to fathom the operating principles of other organ systems. The implication is that
conventional anatomical and physiological thinking is not up to understanding
the human brain, sometimes described as “the most complex object in the known
Universe.”2 Some of the disciplines envisioned as coming to the rescue are com-
puter science, mathematics, statistics, network theory, game theory, graph theory,
cybernetics, systems engineering, cognitive psychology, physics, quantum physics,
philosophy, and “big data” mining. The big data approach (called “connectomics”)
rests on the idea that, if all else fails, documenting the connectivity of the estimated
1
The term “brain” is often used as if the rest of the nervous system doesn’t count for much. In fact,
there are about as many nerve cells in the spinal cord and peripheral nervous system as there are in the
cerebrum, cerebellum, and brainstem (the usual definition of the brain).
2
This truism was recently asserted by Christof Koch, chief scientific officer of the Allen Institute
for Brain Science (June 14, 2013, on NPR’s “Science Friday” with Ira Flatow). ix
x { Preface
86 billion neurons in the human brain should provide the answers sought. Some
thinkers have gone so far as to suggest that understanding the human brain may
require scientific principles not yet conceived or that the function of the brain is so
abstruse that its operation may remain a mystery forever.3 And since interest in the
brain runs as deep in the context of human health as it does in basic science, billions
of dollars are being poured into neurogenomics, psychotropic drug discovery, and
other large-scale efforts aimed at understanding the brain for medical purposes.
In comparison, the thesis here is simple: the operating principle of the brain
and the rest of the nervous system is to make and update neuronal connections
(associations) between inputs (stimuli) and outputs (behaviors) on a wholly empir-
ical basis. At first blush, this statement must seem pedestrian. Putting associations
front and center in thinking about the brain is certainly not new. Both Plato4 and
Aristotle5 emphasized the importance of associations in mental life, as have many
subsequent philosophers (George Berkeley, Thomas Hobbes, John Locke, David
Hume, and Immanuel Kant, to name a few). In psychology, association has also
loomed large in the context of classical and operant conditioning for more than a
century. And, since the discovery of synapses in the late nineteenth century, neu-
ronal associations based on modulating synaptic connections (“neural plasticity”)
have been widely accepted as the basis of learning and memory. The question at
hand is not whether neuronal associations are important in neural function: they
clearly are. The goal in what follows is to explain why the empirical determination
of neuronal connections is inevitable, how useful associations are forged in a phys-
ical world we can’t apprehend, and what the consequences are for understanding
the behavioral responses elicited by neural inputs.
The core of the argument is that biological sensing systems don’t have the ability
to measure the physical characteristics of the objective world in which behaviors
must be carried out. Given this quandary, the gist of what follows is that we and
every other organism—whether it has a nervous system or not—depend entirely on
associations between stimuli and behavior made empirically on the basis of species
and lifetime experience. The evolution of nervous systems simply gives members
of the Animal Kingdom a wider reach in doing the same things that any biological
agent must do to survive and reproduce. If making empirical associations over evo-
lutionary and individual time is indeed the operating principle of nervous systems,
then no special assistance from other disciplines, no novel scientific principle, and
3
The linguist Noam Chomsky has been the most notable advocate of this position. See “Science,
Mind, and Limits of Understanding.” The Science and Faith Foundation (STOQ), The Vatican,
January 2014.
4
Plato (~370 BCE) Phaedrus, trans. by A. Nehamas and P. Woodruff. From Plato: Complete Works,
ed. by John M. Cooper. Indianapolis, IN: Hackett.
5
Aristotle (1906) De Memoria et Reminiscentia. G. R. T. Ross, ed. Cambridge: Cambridge
University Press.
Preface } xi
no investment in schemes for reinventing the brain in silico or knowing the brain’s
every synaptic connection are needed.6
Since buying into this idea entails giving up the assumption that brains compute
representations of the world—or that they compute anything at all—the argument
will be a hard sell. As in any scientific endeavor, the test of a theory is whether it
can make sense of phenomena that have been around for a long time but never
explained. Oddly, for modern neuroscience, the most salient phenomena that need
to be explained are perceptions, one of the end products of nervous systems like
ours. Perceptions—what we actually see, hear, feel, taste, smell, etc.—are strange
indeed, and, at this point in the trajectory of neuroscience, understanding percep-
tual phenomenology may be the best way to get at how brains operate. Whatever
one’s concept of neural function may be, it must be able to explain what we end up
perceiving.
The book is divided into four roughly equal parts. The first part (“What Nervous
Systems Do for Animals”) is intended to set the stage for understanding the emer-
gence of neural systems as a particularly effective way to achieve what all organisms
must accomplish: survival and reproduction. The second part (“Neural Systems as
Engines of Association”) lays out the general argument that biological sensing sys-
tems face a daunting problem: they cannot measure the parameters of the world we
live in as physical instruments do. As a result, nervous systems must make and up-
date associations (synaptic connections) on the basis of empirical success or failure
over both evolutionary and individual time. The third part (“Evidence that Neural
Systems Operate Empirically”) reviews evidence accumulated over the past 20 years
that supports this interpretation in vision and audition, the sensory systems that
have been most studied from this or any other perspective. Finally, the fourth part
(“Alternative Concepts of Neural Function”) considers the pros and cons of other
interpretations of how brains operate.
Although I have written the book for a broad audience, some readers may find
one section or another superficial, while others may find some parts too detailed.
I hope not too many will be put off by my failure to always find the middle ground.
Obviously, I think that understanding the operating principle of nervous systems is
worth thinking hard about, whether or not one agrees with my take on this puzzle.
Dale Purves
Durham, North Carolina
2018
6
This bald statement does not mean that information from other domains is irrelevant. Quite
the contrary. For example, an unexpected source of support for a wholly empirical theory of nervous
system function has come from recent advances in artificial intelligence (see Chapters 11 and 12).
ACKNOWLEDGMENTS
I am deeply grateful to a host of excellent collaborators over the past 20 years,

especially Beau Lotto, for his remarkable imagination and art, and Bill Wojtach,
for holding my feet to the fire on a multitude of issues based on his grasp of both
neuroscience and philosophy. Special thanks also to Catherine Howe and Zhiyong
Yang for insights that I could never have come up. Finally, I am indebted to Henry
Greenside for advice about physics, to Larry Inderbitzen for his perspective as a
psychiatrist, to Shannon Ravenel and Gordon Smilnak for carefully reading and
correcting the manuscript and to Jan Troutt for her fine work on the art.
xiii
PART I }
What Nervous Systems

Do for Animals
1}
Putting the Question in Perspective
Introduction
Basic to the question of whether the brain and the rest of the human nervous
system have a simple operating principle are some central facts about biology and
its relation to neuroscience. What nervous systems do is best appreciated in the con-
text of what all organisms must accomplish in order to survive and prosper, with or
without neural assistance.
By definition, biology is the study of life. Whereas physics and chemistry concern
laws and principles that govern the behavior of everything in the cosmos, biology is
concerned with the subset of entities that live, reproduce, and die. The needed back-
ground for considering nervous systems is thus a definition of life, some knowledge
about the origins of life on Earth, and what all organisms must do to get along in
their niches. Although my understanding of these issues is no more than that of any
other student who pays a modicum of attention to the broader sweep of scientific
progress, this chapter considers some points of consensus. The aim is to situate the
quest for a principle of neural function in the context of biology writ large.
Life on Earth
Contrary to the belief of four of every ten citizens in the United States today, life
on Earth has been around for a lot longer than the roughly 6,000 years stipulated
in the Bible.1 Based on geological observations, cosmic microwave radiation, radi-
oactive decay, and extrapolation back in time from the rate of cosmic expansion,
current estimates are that we are living in a Universe about 13–14 billion years after
1
A Gallup poll in 2014 showed that 42% of US citizens believe that God created humans less than
10,000 years ago, a devastating comment on our educational system and the politics that poor educa-
tion leads to. The citizens of other developed countries do a lot better. 3
4 { What Nervous Systems Do for Animals
FIGURE 1.1 Sedimentary rocks in Western Australia formed by mats of bacteria that lived several
billion years ago.
Photograph taken by Paul Harrison (Reading, UK). Stromatolithes dans la réserve naturelle marine de
Hamelin.
its strange origin. Our solar system, however, is a good deal younger. The Sun is
an average yellow dwarf star in early middle age, having formed, together with the
planets, by condensation from a gaseous cloud of matter about 4.6 billion years
ago. The Sun has a remaining lifetime of about 5 billion years before it turns into a
red giant star, engulfs the nearest planets (including Earth), and then cools gradu-
ally to become a white dwarf and ultimately a black dwarf.2 The Earth is thus about
4.5 billion years old, with something like 5 billion years to go.
As soon as the initially molten surface of the Earth cooled to 100° centigrade,
liquid water and organic molecules already present would have made the emergence of
life possible, and it didn’t take long for that to happen. Surface cooling to the boiling
point of water is estimated to have taken only a few hundred million years, meaning
that the conditions for life on Earth were present by about 4 billion years ago.
The oldest evidence for life in the fossil record is in rocks that formed about 3–4
billion years ago (Figure 1.1).3 These organisms were much like today’s cyanobacteria
(previously called blue-green “algae”), a phylum of photosynthetic bacteria that
remains prevalent.4 Like plants that produce oxygen as “waste,” these organisms
began to contribute oxygen to an atmosphere that was initially oxygen-free.
2
Since dwarf stars cool very slowly, their lifetimes are enormously long. Thus, the ultimate fate of
the Sun is tied to the as yet unknown fate of the Universe.
3
The recent discovery in Canada of tubular fossils that may have been made by bacteria could ex-
tend the age of life on Earth to as much as 4.2 billion years.
4
Six kingdoms of life are generally recognized today: eubacteria (like cyanobacteria and the bac-
teria in our gut), archaebacteria (found today in deep-sea thermal vents and other extreme niches),
protists (single-cell organisms with nuclei), fungi, plants, and animals. The last four are called eukaryotes
Putting the Question in Perspective } 5
In terms of the Earth’s history, then, the emergence of life was very early. Of
course, life on Earth is unlikely to have been first off the mark or in any way special.
Since there are an estimated 100–400 billion stars in our Milky Way Galaxy at var-
ious stages of their lifetimes—many with “rocky” planets like ours at a “habitable”
distance from their stars—and about 100 billion other galaxies, the implication is
clear. The Universe must be teeming with life.
Defining Life
Attempts to define life on Earth have a long and tortured history, beginning in
most Western accounts with the opinions of Aristotle on the subject.5 Although
local skirmishes over the definition of life continue, most of the major battles have
long since been settled. A philosophical battle fought in previous centuries con-
cerned whether life depends on some special quality that transcends the known laws
of physics and chemistry. The idea that it does is called “vitalism,” and its major
proponents were René Descartes in the seventeenth century, Friedrich Nietzsche in
the eighteenth century, and Henri Bergson the late nineteenth and early twentieth
centuries. Much like Aristotle before them, the metaphysical thinking of these
philosophers was based on a “life force” that permeated animate objects, directing
material entities toward a goal, which, for Nietzsche, was a “moral” good. Bergson
coined the phrase “élan vital” and saw this force as a necessary complement to
the “blindness” of Darwinian natural selection. In scientific circles, this idea was
laid low by Hermann von Helmholtz’s demonstration in the 1840s that living
tissues (muscle) use energy and produce heat, consistent with the first law of ther-
modynamics and the physical behavior of everything else in the material world.
Nonetheless, the attempt to draw some spiritual or religious significance from life
and its trajectory on Earth continued to influence the thinking of many notable
biologists. Examples include embryologist Hans Driesch, population geneticist
Sewall Wright, geneticist Julian Huxley, and paleo-biologist Pierre Teilhard de
Chardin. Most biologists today, however, are at best agnostic on the issue of a goal,
moral or otherwise, or indeed any goal at all. The consensus has long been that life
depends on chemistry and physics, not an overarching teleology or properties not
explainable by the laws that govern all material things.
This widely held opinion notwithstanding, simple definitions of life on a mate-
rial basis are elusive. Dictionary definitions such as “The quality that distinguishes
animate from inanimate objects” are uninformative, and more incisive definitions
because their cells have distinct nuclei, whereas the bacterial kingdoms, which do not have nuclei, are
called prokaryotes.
5
A good historical account is Dyson F (1999). The Origins of Life. New York: Cambridge
University Press.
entail a long list of attributes. The chemical and physical properties of life include
organic composition; metabolism; and the generation of metabolic waste products
such as methane, carbon dioxide, or, in the case of photosynthetic organisms, ox-
ygen. Since these are the simplest signatures of life on Earth, they are the properties
that biologists interested in extraterrestrial life are now looking for on Mars, other
planets of our solar system or their moons, and planets orbiting other stars. More
complex entries on the list are replication, the imposition of local organization
(loss of entropy in thermodynamic terms), growth by cell division, and evolu-
tion. However, the characteristic of living things most germane to the operational
principle of nervous systems is actively seeking out sources of energy.
Energy
Energy is needed to do any kind of work, whether in terms of ordinary conversation

or physics. But this statement only says what energy does, not what it is. Physicists—
perhaps most famously Richard Feynman in his Messenger Lectures at Cornell in
19646—readily admit that energy remains a concept that is not fully understood.
Although energy comes in many forms (e.g., kinetic, electromagnetic, chemical, me-
chanical, gravitational, and nuclear), it resists being reduced to a “thing.”
If there is a common denominator other than conservation,7 it is that energy is
needed for events to occur.8 Without energy, nothing happens. An event can thus be
defined as the transfer of energy from one material entity to another at a particular
moment in time in a particular place.
It follows that organisms must acquire energy in some form to enable the events
that allow them to survive long enough to reproduce. And they (including us) must
compete with other organisms to get their fair share of energy in whatever ecolog-
ical niche they occupy.
Although inanimate objects are passively influenced by energy and change ac-
cordingly (think geological change), living things pursue energy sources to obtain
nutrients needed for metabolism, growth, reproduction, successful progeny, and,
in our own species, surrogates for these basic needs in the form of money, political
power, the admiration of others, and so on. Thus, seeking energy is arguably the key
property that differentiates living from non-living things.
6
Available on the Internet thanks to Bill Gates who bought the rights. In addition to being one of
the great physicists in the twentieth century, Feynman was indisputably its most charismatic, if some-
times opaque, teacher. See Feynman R (2001). The Character of Physical Law (Messenger Lectures,
1964). Cambridge, MA: MIT Press.
7
See The Feynman Lectures on Physics, volume 1, c hapter 4, for an extended discussion of energy.
8
One caveat is that, under special circumstances, momentum can be transferred without a transfer
in energy and can thus make things happen as well. Another is that all bets are off in the quantum
domain.
Evolution
The process by which organisms have come to obtain energy in the widely varied
niches they occupy on Earth is evolution. In an article written for school teachers
in 1973, the geneticist Theodosius Dobzhansky stated that “Nothing in biology
makes sense save in the light of evolution.” This cogent generalization is worth
keeping in mind when examining the functions of any organism or organ system.
Since evolution—from cyanobacteria to us—gave rise to nervous systems some-
where along the way, it is important to clarify the basic features of this process.
Without the changes over time wrought by evolution there would be no nervous
systems to worry about.
In 2009, the sesquicentennial of Charles Darwin’s 1859 masterpiece On the
Origin of Species by Means of Natural Selection brought forth a wealth of books
and articles on the many ways evolution by natural selection has influenced modern
biology and human intellection. The opinion of many scientists that The Origin of
Species is the most important book ever written seems on target, at least from the
perspective of biology. The gist of Darwin’s ideas taught worldwide (with some
egregious exceptions) is that organismal variation on Earth can be fully accounted
for by natural selection.9
Foremost among other contributors to the evidence for evolution at about the
same time was the field biologist Alfred Russel Wallace, whose collecting expeditions
in the Amazon and Malaysia were in many ways similar to Darwin’s forays in South
America as the “naturalist” on the 5-year voyage of the HMS Beagle.10 Wallace
published an account of his expeditions in 1853, and he had read Darwin’s account
of his trip in Voyage of the Beagle, published in 1839. But although both men had
been thinking about how the variety of species they witnessed came to be, neither
was explicit about an underlying mechanism.
In 1854, Wallace embarked on an 8-year expedition to the Malay archipelago
that included Singapore, Java, Borneo, and New Guinea, a full account of which he
eventually published in 1876 (The Geographical Distribution of Animals). In 1858,
however, Wallace wrote a nine-page essay titled “On the Tendency of Varieties
to Depart Indefinitely from the Original,” in which he laid out many of the same
ideas that Darwin had been thinking about for 20 years but had not yet written up.
9
As many commentators have pointed out, the concept of evolution did not arise de novo with
Darwin, who was arguably only an eloquent midwife for ideas that had become prominent if less well
formed than his own. By the mid-nineteenth century, educated Europeans were already primed by
Darwin’s predecessors in their thinking and writing: Jean-Baptiste Lamarck’s 1809 treatise on evolution;
Charles Lyell’s geological evidence for the antiquity of the Earth in Principles of Geology, published
in 1830; and Erasmus Darwin’s theory of evolution had already tilled the soil. What differentiated
Darwin’s thinking from these forerunners was a mechanism: natural selection.
10
The primary purpose of the voyage that began in 1831 (the ship’s second) was mapping the South
American coast. On the ship’s first voyage, Robert Fitzroy, the captain featured in many accounts, took
over command of the Beagle after the original captain killed himself in 1828.
Wallace mailed the paper to Darwin from abroad for comment, and it was the re-
ceipt of Wallace’s essay that prompted Darwin to quickly compose a précis of his
own work. To ensure that both men got credit, Darwin (whose son was fatally ill
at the time) arranged to have his friends Charles Lyell and Joseph Hooker present
Wallace’s essay along with his own theory to the Linnaean Society in 1858.11
Darwin was puzzled, however, by aspects of the evidence he could not explain.
Among these was the problem of adaptation, which, then as now, presents a con-
tentious issue. Adaptation refers to the general idea that evolution is based on “fit-
ness,” enshrined in Herbert Spencer’s phrase “survival of the fittest.” In this view,
all changes wrought by natural selection should be adaptive, meaning better suited
to reproductive success when compared with other members of a cohort lacking
the novel trait in question. But it was obvious to Darwin that there were flagrant
exceptions: the tail of the male peacock and the elaborate nest crafted by male
bower birds are the usual examples drawn from many others. Darwin argued that
such patently nonadaptive phenotypic features and behaviors12 provided an advan-
tage in attracting mates, thus fitting an adaptational perspective by virtue of what
he termed “sexual selection.” Sexual selection refers to attraction prior to mating
as another factor that promotes reproductive success. The idea was reinvigorated
in the 1930s by the quantitative work of the British statistician Ronald Fisher and
remains a robust field of evolutionary biology today.
Mechanisms
With respect to a heritable basis for evolution by natural selection, for Darwin and
Wallace, genetics lay in the future. Although Darwin and Gregor Mendel were
contemporaries, Mendel did not begin his work on pea plants until 1856, and it
remained unknown in England until nearly the turn of the century.13,14 As a result,
Darwin and Wallace were uninformed about the fundamentals of genetics that are
now taken for granted in “neo-Darwinian” theory.
11
Darwin’s rush to publication was made on the basis of Thomas Henry Huxley’s urging. Although
Wallace has gotten less credit over the years than he deserved, Darwin was the better writer, clearer
thinker, and far better connected socially and scientifically with friends like Huxley, Lyell, and Hooker
to promote his ideas. Wallace also made the mistake of exempting human culture and morality from
his evolutionary schema, while Darwin went on to publish The Descent of Man in 1871, arguing that
humans are just another species in the biological mix. Nonetheless, Darwin seems to have been extraor-
dinarily fair-minded in sharing credit, much more so than most scientists seeking priority on far less
important issues.
12
Phenotypes are often thought of as comprising gross anatomical features. They actually include
all an organism’s properties, from molecular mechanisms to mentation. Whether some subtle trait is
adaptive is rarely a simple story.
13
Bowler PJ (2003). Evolution: The History of an Idea. Berkeley: University of California Press.
14
Conversely, Mendel is said to have read On the Origin of Species in German translation in 1863.
In addition to the rapid growth of “classical genetics” in the early twentieth

century, another milestone on the path to understanding the mechanisms under-
lying evolution was theoretical physicist Erwin Schrödinger’s 1944 book What Is
Life? In a volume of less than 100 pages, Schrödinger speculated that evolution’s
essential feature must be chemically mediated inheritance conveyed by a code
embodied in an aperiodic crystal. He thus anticipated the discovery of the struc-
ture of DNA in the early 1950s and the molecular biology that followed. The basis
of natural selection, he argued, is “jump-like” heritable changes (mutations) in
these molecules, which provided a “working ground” that natural selection then
plowed. The more specific concept of genes and their ability to mutate is usually
attributed to German physicist Max Delbruck, a major figure in the unfolding mo-
lecular genetic revolution.15 Delbruck’s establishment of a summer course on bac-
terial viruses16 at Cold Spring Harbor Laboratories on Long Island in 1945 brought
together many of the founders of molecular biology, Salvador Luria and Alfred
Hershey preeminent among them.17
This is not to say that molecular biology holds all the answers to questions about
the mechanisms that natural selection relies on to propel evolution. For instance,
it is difficult to explain in genetic terms the evolution of altruism, a phenomenon
best exemplified by social insects. One doesn’t have to know much about ants, bees,
wasps, and termites to recognize the puzzle: many species of these insect families
live in colonies with a single queen and a host of sterile workers whose reproduc-
tive fitness, in Darwinian terms, is nil. As a result, most members of these species,
among the most prevalent on the planet, are not “fit” as individuals, at least not in
conventional terms. Darwin, whose initial interest in biology came from collecting
beetles, was well aware of this issue, which he regarded as another “special diffi-
culty” for his theory. This particular enigma was resolved by the theory of kin selec-
tion proposed by theoretical biologist William Hamilton in 1964. The theory points
out that giving up one’s reproductive success can nonetheless be adaptive for the
species if the altruist’s genes are more effectively passed on by relatives as a result
of self-sacrificing behavior.
An ongoing battle about the mechanism of Darwinian evolution concerns
the “target” of natural selection. For the Oxford zoologist and writer Richard
Dawkins—who, along with John (“J. B. S.”) Haldane, Peter Medawar, and Steven
Jay Gould, is one of the very best popularizers of modern biology—the target is
the gene itself, the idea being that organisms are simply vehicles for increasing the
15
As a dedicated reductionist, Delbruck rejected the idea of quantum indeterminacy and seemed at
ease with his unsettling conclusion that we are statistically determined machines.
16
Because they depend on the molecular machinery of a host cell to survive and reproduce, viruses
are not included among the kingdoms of living things. The viruses studied by Delbruck and colleagues
are called bacteriophage viruses.
17
Delbruck, Hershey, and Luria were awarded the Nobel Prize in Physiology or Medicine in 1969
for their work on molecular genetics.
prevalence of particular genes (thus Dawkins’s phrase “the selfish gene”).18,19 In

this view, phenotypes, including the “tools” many organisms create (the cocoon
of a butterfly, a spider’s web, the stick a monkey uses to extract termites from a
rotting log, human instruments), are all ways of giving particular genes an advan-
tage. Despite the ingenuity of this idea, most evolutionary biologists still hew to the
Darwinian idea that natural selection can act on any phenotypic feature.
Lest anyone take away from this overview the idea that genetics is now well
understood and free from fundamental debates, the articles collected in the book
Genetic Explanations: Sense and Nonsense20 are instructive. Among the points made
by some of today’s leading geneticists is that no one knows quite how to define a
gene, that the links between genotype and phenotype remain deeply uncertain, and
that efforts to pin human diseases on specific genotypes is, in most instances, a sus-
pect goal. Nonetheless, evolution and the mechanisms that underlie it are how some
organisms eventually came to have nervous systems.
Teleology
Perhaps the most heated question over the decades has been whether the evolution
of life by natural selection has a “purpose,” or even a clear direction. The most pas-
sionate biologist weighing in on this issue was Stephen Jay Gould who, like most
biologists today, answered in the negative. It is obvious that the course of evolution
over the past several billion years exemplifies increasing organismal complexity,
and, in this limited sense, the evolution of living things has a “direction.” But the
tendency toward complexity is presumably the result of organisms occupying ever
more niches that demand specialization.21 Gould went on to argue that, despite
this apparent direction, the evolution of increasing organismal complexity is blind
with respect to any particular phenotype, the increasing complexity of organisms
that occupy different niches notwithstanding. That is, if evolution on Earth were
to run all over again from day one—or, as Gould put it, if the tape of life could be
rewound and restarted—the species extant today might be quite different.22
This idea bears on another argument put forward in the 1970s by Gould and
his colleague, Niles Eldredge. They suggested that evolution is not a continuum of
small changes, as Darwin supposed, but is actually characterized by long periods
18
The idea of the gene as the target of natural selection stems from papers written in the 1960s by
W. D. Hamilton and G. C. Williams.
19
The picture is further complicated by increasing evidence today for naturally occurring hybrids,
where two “species” interbreed.
20
Krimsky S, Gruber J (eds) (2013). Genetic Explanations: Sense and Nonsense. Cambridge,
MA: Harvard University Press.
21
In our case, nearly the entire nonaqueous surface of the planet at this point in history.
22
Explained in Gould SJ (1989). Wonderful Life: The Burgess Shale and the Nature of History.
New York: Norton.
of stasis punctuated by rapid change, an idea for which they found empirical evi-
dence in the fossil record of invertebrates. Although basically a refinement of neo-
Darwinian theory (i.e., the rate of evolutionary change is clearly variable, and the
data are always confounded by gaps in fossil record), the idea that evolution pro-
ceeds as a “punctuated equilibrium” continues to engender persistent (and often
acrimonious) debate.23
It seems likely that this and other ongoing arguments will soon enough be in-
formed by the discovery of life on other planets or their moons in our solar system
and beyond.
Conclusion
The issues discussed here supply a patently superficial account of the history of life
on Earth. They nonetheless provide a foundation for asking what brains—or, more
properly, what nervous systems—add to this general account of biology.
Suggested Reading
Bowler PJ (2003). Evolution: The History of an Idea. Berkeley: University of California Press.
Darwin C (1859). On the Origin of Species by Means of Natural Selection, or the Preservation
of Favoured Races in the Struggle for Life (1st edition). London: John Murray.
Darwin C (1868). The Variation of Animals and Plants Under Domestication. London: John
Murray.
Darwin C (1871). The Descent of Man, and Selection in Relation to Sex (1st edition).
London: John Murray.
Darwin C (1872). The Origin of Species by Means of Natural Selection, or the Preservation
of Favoured Races in the Struggle for Life (6th edition). London: John Murray.
Darwin C, Wallace AR (1858). On the Tendency of Species to form Varieties; and on the
Perpetuation of Varieties and Species by Natural Means of Selection. J Proc Linnaean
Soc London Zool 3:46–50.
Dawkins R (1976). The Selfish Gene. Oxford: Oxford University Press.
Dawkins R (1986). The Blind Watchmaker. New York: WW Norton.
Dyson F (1999). The Origins of Life. New York: Cambridge University Press.
Feynman R (2001). The Character of Physical Law (Messenger Lectures, 1964). Cambridge,
MA: MIT Press.
Gould SJ (1977). Ever Since Darwin. New York: WW Norton.
23
A recent finding that undermines the idea of punctuated equilibrium as a general rule is that the
bacterium Escherichia coli goes on making small improvements over the course of 60,000 generations in
the same environment. Thus, what appears to be a period of stasis may be only a slowing of continual
change (Lenski R et al. [2015]. Sustained fitness gains and variability in fitness trajectories in the long-
term evolution experiment with Escherichia coli. Proc Roy Soc B 282:2015–2292).
Gould SJ (1989). Wonderful Life: The Burgess Shale and the Nature of History. New York:
W.W. Norton.
Gould SJ (2002). The Structure of Evolutionary Theory. Cambridge, MA: Belknap Press.
Krimsky S, Gruber J (eds.) (2013). Genetic Explanations: Sense and Nonsense. Cambridge,
MA: Harvard University Press.
Rosenblueth A, Wiener N, Bigelow J (1943). Behavior, purpose and teleology. Phil Sci
10:18–24.
Simon HA (1962). The architecture of complexity. Proc Am Phil Soc 106:467–482.
Tasker E (2017). The Planet Factory: Exoplanets and the Search for a Second Earth. New
York: Bloomsbury Sigma.
Tattersall I (2012). Masters of the Planet: The Search for Human Origins. New York: St.
Martin’s-Griffin.
Tyson, N (2017). Astrophysics for People in a Hurry. New York: WW Norton.
Wallace AR (1869). The Malay Archipelago. New York: Harper.
Wallace AR (1870). Contributions to the Theory of Natural Selection (2nd edition).
New York: Macmillan.
Wallace AR (1876). The Geographical Distribution of Animals. New York: Harper.
2}
Organisms Without Nervous Systems
Introduction
According to current estimates, the kingdoms of life on Earth comprise at least

8.7 million eukaryotic species (protists, fungi, plants, and animals) and many
more prokaryotic species (eubacteria and archaebacteria).1 The fuzziness of these
numbers is understandable: new species are being discovered daily and extant spe-
cies are going extinct, as they always have. Moreover, the concept of a species is not
straightforward. The textbook definition of a biological species is a reproductively
isolated population of organisms that breed. While these criteria are a fine way to
differentiate dogs from cats, they break down for bacteria and other organisms that
multiply by cell division and for organisms that exchange genetic material,2 as well
as for plant and animal hybrids. Whatever the number may actually be, the over-
whelming majority of past and present species don’t have nervous systems. Thus,
before asking what function or functions nervous systems add to animal biology, an
obvious question is how organisms without them get along so well.
Bacteria
As mentioned in Chapter 1, the two bacterial kingdoms (eubacteria and archae-

bacteria) are large, enormously successful forms of life and were the first to appear
on Earth. Like all organisms, bacteria seek out energy sources to make biolog-
ical events happen, the most important of these being survival and reproduction.
Depending on the species, the energy sought can be in the form of light, heat, or
1
Organismal groupings and numbers are in constant revision, and the delineations here are only
general. For more detail see Whittaker (1969), May (1997), and Mora et al. (2011) in the list of suggested
readings at the end of the chapter.
2
The definition of species among bacteria is especially complicated because many varieties transfer
genetic material in a process called conjugation. 13
chemical reactions. By using the rules of plant and animal metabolism in some-
times remarkable ways, bacteria have come to occupy most of the ecological niches
that the Earth provides, including some very forbidding ones.3 Chemolithotrophic
bacteria, for instance, populate deep sea trenches that admit no light and generate
extremes of heat, pressure, and terrestrial chemistry. Such organisms acquire energy
by the oxidation of inorganic minerals and other compounds that act as electron
donors.4 Other bacteria thrive many kilometers underground and derive energy
from the heat produced by radioactivity.5 Still other bacterial species are found in
niches of extreme acidity, alkalinity, and salinity, leaving hardly any locale without
some form of life.
This diversity notwithstanding, bacteria, like all other organisms, need some way
of getting information about energy sources from the environment via sensors and
some means of responding to that information by movement or other behaviors.6
How, then, do bacteria accomplish these feats without a nervous system or rela-
tively little else in the way of supramolecular machinery?
Given their role in human diseases as well as those of other animals, many bac-
terial species have been studied intensively, a standard choice being the common
gut bacterium Escherichia coli 7 (Figure 2.1). The organism’s most obvious beha-
vior is movement generated by a flagellar bundle at one end of the rod-shaped cell.
In response to local gradients of relevant chemicals (amino acids, sugars, oxygen,
carbon dioxide)8 the flagellar bundle rotates clockwise, moving the bacterium in
some direction. Periods of such swimming (called “runs” that last about a second)
are interspersed with intervals of “tumbling” when the flagellar bundle rotates
counterclockwise and becomes untangled, leaving the bacterium wandering more
or less aimlessly. Its overall progress in a given direction (up a nutrient gradient, for
example) is determined by the ratio of runs to tumbles.
To swim up rewarding gradients and down harmful ones, bacteria depend on
surface sensors that detect concentrations of nutrient molecules or potentially toxic
3
See Forterre P (2016). Microbes from Hell. (transl TL Fagan). Chicago: University of Chicago Press.
4
Some animals, even large ones such the worms and other bizarre species that live near deep sea
thermal vents, are also chemolithotrophs.
5
Li-Hung L et al. (2006). Long-term sustainability of a high-energy, low-diversity crustal biome.
Science 314:479–482. doi: 10.1126/science.1127376.
6
“Behavior” is an especially tricky word. Although often used as a synonym for motor responses,
behavior includes any response to a stimulus. The responses are diverse even in nominally simple
organisms like bacteria. In humans and many other animals, responses to stimuli include perception,
attention, emotion, memory, thinking, and homeostasis among others.
7
E. coli is a species in the genus Escherichia, in the family Enterobacteria, in the order Enterobacterias,
in the class Gammaproteobacteria, in the phylum Proteobacteria, in the kingdom Bacteria. Like it or
not, this is the way biologists describe the taxonomic status of any organism or group of organisms,
including us.
8
E. coli is a facultative anaerobe, meaning it can switch back and forth between oxidative metabo-
lism and anaerobic metabolism (i.e., fermentation) depending on the local environment.
Organisms Without Nervous Systems } 15
Cytoplasm
Pili
Flagellar
bundle
Capsule
Cell wall
Cytoplasmic
membrane
Nucleoid
Ribosomes
FIGURE 2.1 Diagram of an Escherichia coli bacterium. In addition to cytoplasmic organelles such as
ribosomes that are responsible for protein synthesis in all cells, bacteria contain a single chromosome
that forms a closed loop called the nucleoid (like all prokaryotes, bacteria have no nucleus, the
defining feature of eukaryotic organisms). A rotating flagellar bundle generates motor responses to
stimuli transduced by molecular receptors in the cytoplasmic membrane. Other extensions called pili
are used for conjugation, a process in which two bacteria join and exchange small numbers of genes
in plasmids (not shown), as opposed to the full replication of the genome that occurs during bacterial
cell division.
molecules such as antibiotics, hydrogen ions (acidity), hydroxyl ions (alkalinity),

and many other chemicals. Depending on the species, bacteria can also respond
to heat, light, mechanical forces, and attack by viruses (the bacteriophage viruses
mentioned in Chapter 1). Most is known about chemical sensors, which are proteins
that extend into the environment from the bacterial cell membrane. Determining
the links between sensory inputs and effector outputs, such as rotation of the fla-
gellar bundle, is another field of microbial study that has identified a range of intra-
cellular signaling pathways that are generally similar to (and not much less complex
than) those found in any other cell type. Some bacteria may even use changes in
membrane voltage to mediate cooperative interactions among groups of bacteria.9
If all this seems surprisingly sophisticated and not all that different from what
more complex eukaryotic cells do, right on both counts. Even at this “primitive”
level, inputs arising from the environment trigger useful outputs (behaviors) by
associations between physical and chemical causes and effects. These associations
can and do change over time by means of natural selection. Given the rate of bac-
terial replication, the times involved can be quite short: in favorable conditions, a
bacterial population can double in 30 minutes or less. The result can be rapid evo-
lution, as has become all too evident in the emergence of antibiotic-resistant strains
of pathogens that bedevil medical practice today.
Bacteria can also learn from experience and retain information (memory of a
sort), which is pretty much the stock-in-trade of animals with nervous systems (see
Chapter 3). An example is epigenetic changes in DNA arising from experience in
a particular environment that is later expressed collectively in the population.10 In
the light of such abilities, some microbiologists have pointed out that bacteria could
be considered capable of “cognition.”11 Hyperbole though this is, microbiologist
Daniel Koshland, who was responsible for many key studies of bacterial behavior,
had a point when he paraphrased Alexander Pope by saying—only partly in jest—
that “the proper study of mankind is the bacterium.”12
Whether one accepts the relevance of bacterial behavior to human cognition or
not, it is clear that bacteria are highly sophisticated organisms that obtain, store,
and use energy to associate information arising from the environment with heritable
responses that have made them highly successful for billions of years.
Protists
Protists13 form a kingdom of mostly single-celled organisms that differ from

bacteria in that they have a cell nucleus, the defining feature of eukaryotes. The
9
Prindle A, et al. (2015). Ion channels enable electrical communication in bacterial communities.
Nature 527:59–63.
10
Mathis R, Ackermann M (2016). Response of single bacterial cells to stress gives rise to complex
history dependence at the population level. PNAS 113:4224–4229.
11
Since cognitive neuroscientists have difficulty defining what human “cognition” actually means,
there is lots of leeway here. Definitions usually boil down to studying perception, memory, emotion,
attention, and decision-making under the unspoken assumption that humans enjoy these abilities to the
exclusion of most other organisms, and certainly bacteria.
12
Koshland D (1980). Bacterial Chemotaxis as a Model Behavioral System. New York: Raven Press.
Anyone who doubts that bacterial behavior and the processes underlying it are complex—and relevant
to the question of what nervous systems add to organismal talents—should also look at a recent review
of this field, such as Lyons P (2015). The cognitive cell: bacterial behavior reconsidered. Front Microbiol
6:264. doi: 10.3389/fmicb.2015.00264.
13
“Protist” is a misnomer since it means “first,” a title that clearly belongs to the bacterial kingdoms.
Macronucleus
Micronucleus
Oral groove
Food
vacuoles
Anal
pore
Contractile
Vacuole
FIGURE 2.2 Diagram of a species of the genus Paramecium. Although a single cell, these organisms
have more specializations than bacteria, including nuclei, an array of cilia, an oral groove, an anal
pore, and contractile vacuoles, all of which are absent in Escherichia coli and other bacteria.
major phyla in this kingdom are protozoa, algae, and slime molds.14 Perhaps the
most familiar example of a protist is Paramecium, a genus of ciliated organisms
many of us encountered in high school biology as one or another of these single-
cell species swimming in a drop of pond water examined under a microscope
(Figure 2.2).
Among the differences between a species of Paramecium and a bacterial species
is the presence of a cell nucleus in the Paramecium that harbors far more genes.
The single circular chromosome of a bacterium like E. coli comprises about 4,000
genes that code for proteins and somewhat more “noncoding” elements, for a total
of about 11,000 genes. Protists like Paramecia have about 40,000 coding and non-
coding genes on 50 chromosomes, far more than the single chromosome in bacteria
14
Taxonomy has always been a moving target subject to controversies whose intensity has some-
times outweighed their importance. Although modern genetics has clarified some of these arguments,
important genes tend to be conserved across taxonomic boundaries. Such debates have been ameliorated
by cladistics, a method in which organisms are categorized by characteristics that can be traced back to
a group’s most recent common ancestor but not to more ancient ones. The result is a “clade,” in which
the members are more closely related to each other than to the members of any other clade.
and about the same number that we have (humans have 46 chromosomes and about
20,000 genes).15
When a bacterial cell divides, the genome simply replicates. In contrast,
Paramecia have two nuclei, a polyploid macronucleus and a haploid micronucleus,
the latter serving as the germ line when the cell divides. The macronuclear genes
regulate the proteins underlying all the other functions of the cell, just as do
the genes in our somatic cells. These differences are more or less the forerunners
of germ line meiosis and somatic cell mitosis in us and other animals. Other
differences from bacteria include the presence of mitochondria,16 a cytoskeleton,
the absence of a cell wall, and a circumferential array of cilia that enables more
precise motility.
Although a single cell, a Paramecium is about 100–1,000 times larger than E. coli
(~100–3,000 microns in length compared ~1–3 microns) and far more specialized in
its organization, with a mouth and anus of sorts (the oral groove and the anal pore
in Figure 2.2). Like bacteria, protists have molecular sensors on their surfaces that
enable them to respond to chemical, light, and mechanical stimuli. Moreover, they
can learn and store information over time in more impressive ways than bacteria.
For example, at least one species of Paramecium can be conditioned to associate light
level with a membrane voltage.17 Finally, whereas electrical transfer of information in
bacteria is equivocal, paramecia clearly have this ability and use voltage-dependent
changes in membrane permeability to control the frequency and direction of ciliary
movement.18
Plants
A third kingdom of organisms that lack nervous systems is plants, defined by a dif-
ferent type of metabolism (photosynthesis),19 multicellularity, and (mostly) sexual
reproduction. Although we tend to think of plants as quintessentially “stupid,”
being fixed in place and lacking the sorts of sophisticated behaviors characteristic
15
These numbers in us compared to a protist underscore the point made earlier that the relation
between genotype and phenotype remains poorly understood.
16
Mitochondria are characteristic of eukaryotes and represent the wholesale incorporation of bac-
teria and the genes that those bacteria use for the production of energy, which persist in us and other
animals as a separate mitochondrial genome.
17
Armus HL, Montgomery AR, Jellison, JL (2006). Discrimination learning in Paramecia
(P. caudatum). Psychol Rec 56:489–498.
18
Eckert R (1972). Bioelectric control of ciliary activity. Science 176:473–481.
19
Photosynthesis uses light energy to drive the synthesis of carbohydrates from carbon dioxide
(CO2) and water. A “waste product” in this process is oxygen, and plants, along with some bacteria, are
the primary source of this molecule in the atmosphere. The carbohydrates created by plants are directly
or indirectly the source of nutrients for a huge range of organisms, from many species of bacteria to
humans.
FIGURE 2.3 Arabidopsis. Like all plants, wild mustard is a multicellular organism that gets its energy
from sunlight and uses it for photosynthesis. Photosynthesis is carried out by chloroplasts (analogous
to mitochondria in animal cells) that produce carbohydrates such as sugars from water and carbon
dioxide (CO2), with oxygen as the key metabolic side product (most of the oxygen we breathe
today has come from plants). In this way, solar energy is transformed and stored as starches (sugar
polymers) that are used by adenosine triphosphate, the proximate source of metabolic energy in all
cells, whether in plants, in us, and even in the chemolithotrophs mentioned earlier.
of the animal kingdom, this perspective grossly underestimates both the abilities of
plants and the cleverness of evolution.20
Take Arabidopsis, for example, a genus of wild mustard best known to many
as the central player in a grade school science project (Figure 2.3). Like nearly all
plants, species of Arabidopsis express phototropism, growing toward sources of
radiation that contain wavelengths at or beyond the short-wavelength end of the
light spectrum perceived by humans (ultraviolet radiation, typically from the Sun).
Although Darwin was the first to show by simple experiments in the nineteenth
Chamovitz D (2012). What a Plant Knows: A Field Guide to the Senses. New York: Farrar,
20
Straus, and Giroux. This delightful book provides an antidote to the conventional idea that plants are
“senseless.”
(A) (B) (C)
FIGURE 2.4 Electrically mediated motor responses in a plant (Mimosa pudica). Mechanoreceptor
cells on leaves of this species initiate electrical signals that travel along small branches (petioles) to
their intersections with the main branch (A). There, specialized cells with large water-filled vacuoles
cause the petioles to move toward the main branch when the cells lose water and shrink in response
to the electrical signal. The same process causes each leaf to collapse. The full response in (C) can be
initiated by touching a single leaf, as shown in (B).
century that the sensory receptors for phototropism are in the growing tips of
plants,21 it is now known that plants have a wide array of photoreceptor molecules,
far more than the five photopigments we humans use to sense light.22
But these are only the most obvious of the sensory abilities of plants and the
consequent plant behaviors. Plants also sense and respond to temperature and the
duration of night and day. They can detect airborne molecules (ethylene, for ex-
ample, which causes ripening and allows some parasitic plants to find their hosts in
this way). They can respond quickly to mechanical stimuli (e.g., the Venus flytrap,
native to a region of North Carolina). And some have even evolved action potentials
to respond rapidly over relatively long distances by using electrical signals, as in a
species of mimosa whose leaves collapse along an entire branch when mechani-
cally disturbed (Figure 2.4).23 Plants can also learn from experience, responding to
prior trauma by altered growth responses later, thus exhibiting a type of memory.
As plant expert Daniel Chamovitz points out, about the only thing plants don’t re-
spond to is prayer.24
Equally remarkable are interactions among the forest trees. The root tips of some
species in a stand specifically associate with other nearby members of the same
The last book Darwin wrote was on plants (The Power of Movement in Plants. London: John
21
Murray, 1880), in which he had a long-standing interest.

22
Three cone pigments, rhodopsin in rods, and melanopsin in retinal ganglion cells are the five
photopigments in the human retina.
23
Sibaoka T (1962). Excitable cells in mimosa. Science 137:226.
24
See Pollan M (2013, Dec 23). The intelligent plant. The New Yorker, for a very funny account of
ongoing debates about plant intelligence among botanists.
species for mutual support in what has been described as a “social network.”25,26
Of course, one can go overboard anthropomorphizing the abilities of plants, but,
like bacteria and protists, plants use information in their environments to generate
structural and behavioral responses that promote survival and reproduction. These
abilities seem as varied and capable as those of many animals.
The General Strategy
Thus, organisms without nervous systems—which is to say the vast majority of ex-
tant and extinct species on Earth—obviously get along quite well in their respective
niches. Although the details of the behaviors they use to do so vary enormously,
the general strategy is clear enough. All organisms, from single-cell bacteria and
protists to plants (which in many species coordinate far more cells than are pre-
sent in the human body), achieve success by associating sensory inputs with suc-
cessful responses over evolutionary and individual time. The inputs are “sensory”
responses based on the transduction of environmental energy; the outputs are the
consequences of intra-and intercellular signaling using stored energy to activate
further events in the cell or cells in question. The criteria of success are survival and
reproduction, as spelled out in Chapter 1.
Conclusion
This overview of organisms that lack nervous systems provides the background
needed to ask what nervous systems add to the biology of non-neural organisms.
In particular, do nervous systems introduce a fundamentally new strategy for sur-
vival and reproduction, or do they simply use the reflexive associations apparent in
all forms of life in the more elaborate ways made possible by neurons and synaptic
connections?
Suggested Reading
Berg HC (2004). E. coli in Motion. New York: Springer.

Chamovitz D (2012). What a Plant Knows: A Field Guide to the Senses. New York: Farrar,
Straus, and Giroux.
Darwin C (1880). The Power of Movement in Plants. London: John Murray.
25
Wohlleben P (2016). The Hidden Life of Trees. (transl. Jane Billinghurst). Vancouver: Greystone
Books.
26
Scheres B, van der Putten WH (2017). The plant perceptron connects environment to development.
Nature 543:337–345.
Eckert R (1972). Bioelectric control of ciliary activity. Science 176:473–481.

Forterre P (2016). Microbes from Hell. (transl. Fagan TL). Chicago: University of
Chicago Press.
Koshland D (1980). Bacterial Chemotaxis as a Model Behavioral System. New York:
Raven Press.
May RM (1997). The dimensions of life on Earth. In: Nature and Human Society (Raven
PH, ed), pp. 30–45. Washington, DC: National Academy Press.
Mora C, Tittensor DP, Adl S, Simpson AGB, Worm B (2011). How many species are there
on Earth and in the ocean? PLoS Biol 9 e1001127. doi:10.1371/journal.pbio.1001127.
Pollan M (2013, Dec 23). The intelligent plant. The New Yorker.
Scheres B, van der Putten WH (2017). The plant perceptron connects environment to
development. Nature 543:337–345.
Volkov AG, Foster JC, Markin VS (2010). Signal transduction in Mimosa pudica: biolog-
ically closed electrical circuits. Plant Cell Environment 33:816–827.
Whittaker RH (1969). New concepts of kingdoms or organisms. Evolutionary relations are
better represented by new classifications than by the traditional two kingdoms. Science
163:150–194.
Wohlleben P (2016). The Hidden Life of Trees (transl Jane Billinghurst). Vancouver:
Greystone Books.
3}
Organisms With Nervous Systems
Introduction
Definitions of the term “animals” (technically, members of the kingdom Animalia)

in dictionaries and textbooks are surprisingly vague. The characteristics usually
mentioned are eukaryotic, multicellular, heterotrophic (ingesting other organisms
for energy), sexually reproducing, and capable of rapid and independent movement.
But, as Chapter 2 should have made clear, some or all these properties are charac-
teristic of many organisms in the other kingdoms of life on Earth. In fact, the
major distinguishing feature of animals in most cases is the presence of a nervous
system (or a system of electrically excitable cells that plays a similar role in very
primitive animals). Most definitions don’t mention this feature, and those that do
make it seem as if nervous systems are mainly needed for rapid responsiveness.
Nonsense. Think of the responses of E. coli and paramecia to stimuli or the speed
at which a Venus flytrap can close. But if nervous systems are indeed one of the
main attributes that distinguish organisms in the animal kingdom, what exactly are
nervous systems and what advantages do they bring? Without at least some provi-
sional answers, seeking the operating principle of neural systems makes little sense.
Defining Nervous Systems
Nervous systems are made up of specialized cells called neurons and glia. Therefore,
the first order of business is to define what distinguishes these cell types from all
the many other types found in animals. This task is not entirely straightforward.
A primary characteristic of neurons is electrical excitability,1 but this feature is not
unique. Muscle cells are electrically excitable, as is the single cell of a protist like
1
The ability of the cell membrane to change the voltage across it by varying ion permeability. This
modulation is the basis of neuronal signaling. 23
Paramecium and some plant cells (see Chapter 2). Even some bacteria can com-
municate with each other by means of ion channels in their cell membranes whose
conductance can be modified.
Glial cells (the term means “glue”) are not electrically excitable in the same way
neurons are and have long been interpreted as supporting the functions of neurons.
Although interest in glia has been revived over the past decade or so by discoveries
about the many ways they interact with neurons,2 glia remain secondary players in
transmitting and storing information gleaned from the environment.
The more abstract idea that neurons are specialized for “processing informa-
tion” presents another avenue for definition, but processing information3 from the
local environment to generate biologically useful responses (ultimately successful
reproduction) is a primary function of any cell type.4 The presence of synapses—the
connections between neurons and their target cells—also contributes to the defini-
tion of neurons, but again not uniquely. Synapses are specializations that allow elec-
trochemical communication (and thus associations) between two or more neurons,
and they come in two flavors: electrical and chemical. Electrical synapses are
junctions between nerve cells that allow the direct passage of electrical current and
small molecules from one neuron to another for functional coordination. Chemical
synapses release transmitter agents in response to changes in the voltage across the
“presynaptic” cell membrane; these in turn excite or inhibit a “postsynaptic” cell
(or cells) by the release of chemical transmitter agents (see Chapter 6), amplifying
the effect on the target cell or cells in the process. (In contrast, the postsynaptic ef-
fect at an electrical synapse is always diminished.) But neither of these functions is
limited to neurons. Many cells in other organs are coupled by electrical junctions
that are anatomically and functionally the same as those in the nervous system (e.g.,
cardiac muscle cells). Moreover, other cell types can also release chemical agents
from vesicles by membrane fusion, the hallmark of a chemical synapse (e.g., pan-
creatic islet cells that release insulin, mast cells5 that release histamine). The conver-
sion of sensory or other information into all-or-none action potentials—the signals
that travel over axons to convey information from one place to another in nervous
systems—is also thought of as a property specific to neurons. But this criterion falls
short as well: skeletal muscle fibers have action potentials, whereas many neurons
do not.
2
Among these are their role in synapse formation and maintenance and some degree of mem-
brane excitability; see Káradóttir R, Hamilton NB, Bakiri Y, Attwell D (2008). Spiking and non-spiking
classes of oligodendrocyte precursor glia in CNS white matter. Nat Neurosci 11:450–456. doi: 10.1038/
nn2060.
3
Roughly defined as a physical or chemical variable that allows observers (or a cell) to extract a
signal from background noise.
4
An arguable exception is red blood cells, which have no nucleus or other features that are not
needed in a cell whose primary function is to carry oxygen (or carbon dioxide [CO2]).
5
A type of white blood cells that releases histamine in reaction to tissue injury.
Organisms With Nervous Systems } 25
The fact that neurons whose axons travel long distances need all-or-none ac-
tion potentials has also promoted the idea that the organization of nerve cell firing
over time represents a “code,” much like the strings of 0s and 1s used in computer
algorithms. Furthermore, neurons integrate the electrical effects imposed by other
nerve cells, much like the logical elements in computers. But, despite their impor-
tance in neural function, the role of action potentials as code carriers has been
exaggerated. A counterexample is the 302 neurons that constitute the nervous
system of the 1 millimeter-long roundworm Caenorhabditis elegans, the most com-
pletely documented nervous system in any animal. This set of neurons supports the
full range of the worm’s considerable behavioral repertoire without using action
potentials.6 Moreover, one of the most thoroughly studied regions of the human
brain, the retina, carries out complex neural processing without the benefit of ac-
tion potentials.7 The implication of these observations is that action potentials
are primarily an elegant solution to the problem of linking cause and effect over
distances where passive electrical conduction is inadequate (see Chapters 11 and
12 for a discussion of nervous systems as computers).8 That is not to say, however,
that the action potentials have no other functions. For example, the timing of action
potentials from the two ears impinging on neurons in the auditory brainstem is well
known to be essential for localizing the origin of sound stimuli in space.
Although the remarkable feats of human intelligence and culture might seem
impossible without some fundamental departure from the garden variety anatomy,
physiology, and cell biology evident in other human organs and in other animals,
there is not much support for this idea. The execution of elaborate behaviors in
prokaryotes and plants does not depend on nervous systems, let alone on nervous
systems as complex as ours. And since the term “intelligence” has no agreed-upon
meaning in neuroscience,9 the word can fairly be used to describe how any or-
ganism solves the problems that confront it. Even focusing on the aggregate ability
of neurons to organize and effect complex behavior fails, since the cardiovascular
6
Although neurons in C. elegans have Na+ channels in their membranes, they are not used for
the voltage-gating evident in neurons that typically support action potentials. These channels and
their effects may be an evolutionary forerunner of the full-blown action potentials in larger but oth-
erwise analogous roundworms, such as the human intestinal parasite, Ascaris, which is approximately
30 centimeters in length. This comparison implies that action potentials evolved mainly to solve the
“transmission over long distances” problem. To what extent they have been co-opted as bearers of a
code remains a largely open question.
7
Although the location of the eye suggests otherwise, the embryological origin of the retina is the
same as the rest of the brain and central nervous system (i.e., the embryonic neural tube). Other periph-
eral sensors (e.g., those in the ear and nose) derive from specializations in the outer cell layer of the early
embryo (called ectodermal placodes).
8
Passive conduction is limited to a few millimeters or less depending on the electrical characteristics
of the axon in question.
9
Intelligence is another vague descriptor that refers to problem-solving ability; the more difficult
the problems solved, the more intelligent an organism is said to be. And many of the problems that
non-neural organisms solve are clearly difficult ones.
Tentacle
Mouth
Gastrodermis Outer
muscle sheet epidermis
muscle
Stomach sheet
Theca
FIGURE 3.1 The polyp of a typical coral. The muscle cells in the animal’s epidermis generate
coordinated movements that sweep nearby microorganisms into the gut and move them along. The
evolution of such muscle sheets and the electrical conduction of pacemaker activity within them were
presumably early steps in the evolution of nervous systems proper.
system, the digestive system, the immune system, and many others operate with
little neural oversight.
In short, nervous systems employ mechanisms that are used by other cells and
cell systems for generally similar purposes. The point is simply that understanding
the operations of nervous systems may be a lot easier when they are seen as another
organ system to be explained in terms of biological functions that are commonplace.
The Emergence of Nervous Systems
If nervous systems are defined as aggregations of excitable cells that enhance the
number of behaviorally useful input–output associations an animal can make by
virtue of synaptic connections, when did they first appear, serving what sorts of
behaviors, and in what form?
No one knows when and where nervous systems appeared, but a plausible guess
is that the first excitable cells in animals were loosely organized myocytes in ani-
mals such as sponges (several classes in the phylum Porifera) that evolved sheets of
muscle cells to better coordinate movements, most likely in early Cnidaria, a phylum
of more than 10,000 species today that includes corals, sea anemones, hydra, and
jellyfish10 (Figure 3.1). The next step may have been the evolution of specialized
pacemaker regions in such sheets to further enhance their efficiency, witnessed by
10
Until recently called Coelenterates. Taxonomy is rife with changes in nomenclature as older
classifications are updated by new observations.
(A) (B) Nerve net
Radial Mouth
canal
Tentacle Nerve
Outer
net
nerve
ring
Tentacle Velum Inner nerve ring

FIGURE 3.2 The simple nerve net in a hydra (A) and the more complex network in a jellyfish (B).
The increasing complexity of such networks would have allowed increasingly complex sets of synaptic
connections (associations) between sensory inputs and behavioral outputs. The nervous systems and
behaviors even in hydra are, in fact, surprisingly complex.
many a biology student as the coordinated beating of an isolated frog or turtle

heart in a dish of saline solution. Pacemakers in the heart or gut are examples of
groups of muscle cells whose membrane potentials oscillate, spreading the effect to
adjacent muscle fibers via junctions identical to the electrical synapses mentioned
earlier. The signal continues to spread in the muscle sheet because an action poten-
tial in a given muscle cell is passively conducted to nearby cells, which are brought
to threshold and fire action potentials in turn.11
As animals diversified and were challenged to survive in more complex
environments in the sea and eventually on land and in the air, it is not much of a
stretch to imagine that they would have benefited by increasing overall control of
body parts needed to generate increasingly specialized behaviors. This progression
would eventually have given rise to the simple nerve cell networks evident in ani-
mals like hydra and jellyfish (Figure 3.2) and later to the collections of neurons in
the ganglia12 of many invertebrates that link sensory inputs with motor and other
outputs (see Figure 3.3A).13 To judge from electron microscopic evidence in such
11
Action potentials evolved in muscle fibers because efficient contraction depends on a membrane
voltage change that can spread quickly over the entire length of the fiber, which in some muscles can be
quite a distance (e.g., up to 30 centimeters in the human quadriceps muscle).
12
In both invertebrates and vertebrates, the term “ganglia” refers to collections of neurons that are
clustered together and are thus apparent anatomically. See Figure 3.3A for example.
13
“Invertebrates” is a general designation the includes all animals other than the sub-phylum of
vertebrates (i.e., animals with a backbone). More than 95% of animals are invertebrates.
species today, the first chemical synapses probably arose in animals like hydra and
jellyfish.
Considered in this general way, neurons and nervous systems are advanta-
geous in animals whose reproductive success demands ever more diverse sensory
inputs, a greater range of effectors, and a greater range of coordinated associations
among inputs and outputs as a basis for complex behaviors with increasingly
sophisticated goals.
The Emergence of Central Nervous Systems
The next step in the evolution of nervous systems is likely to have been the appear-
ance of a more centralized arrangement, an organization that is apparent today
in most invertebrates and all vertebrates (Figure 3.3).14 In many invertebrates, this
centralization is evident as conglomerations of neurons strung along the central
axis of the animal in the form of a chain of ganglia called the central nerve cord.
The largest ganglion is typically located in the equivalent of the animal’s head and
is informally referred to as the “brain.” In vertebrates, the central collection of
neurons comprises the spinal cord and a more obvious rostral enlargement—the
brain proper.15 In both instances, nerve cell axons extend from the central nervous
system via peripheral nerves to innervate muscles, glands, and other targets; the pe-
ripheral nerves also carry information to the central nervous system from sensory
receptors.
Although the differences in detail are many, the overall similarity of nervous
system organization in invertebrates and vertebrates is striking. The most obvious
difference is the relative size of the “brain” (defined in humans and other mammals
as the cerebral hemispheres, the cerebellum, and the brainstem) compared with
most invertebrates.16
What Do Brains Add?
When making comparisons among animals, one should really be talking about
nervous systems entire rather than brains as such. As shown in Figure 3.3, the brain
is not differentiated from the rest of the nervous system other than by its location
14
Vertebrates and invertebrates (animals with and without backbones) are, respectively, a sub-
phylum of the phylum of chordates (Chordata) and the phylum of invertebrates (Invertebrata).
15
The term “brain” in vertebrate animals is defined by gross anatomy. Nothing special distinguishes
the brain from the spinal cord except its greater size and anatomical complexity.
16
Given the diversity of invertebrates, there are many variations on the segmental patterns ev-
ident in worms and many insects. Variants well known to neuroscience are the nervous systems of
cephalopods such as squid and octopuses, and gastropods like the sea slug Aplysia.
(A) Leech (B) Human

FIGURE 3.3 The central nervous system of a typical invertebrate (A) and a far more specialized
vertebrate (B). The nervous system of the leech comprises a series of segmental ganglia (called the
central nerve cord) with a rostral enlargement (the “brain”), each with a set of peripheral nerves. The
organization of the human nervous system is, in many ways, similar. The collections of neurons that
innervate segmentally derived body units have coalesced into the spinal cord. Although the rostral
enlargement—the brain—is much greater in size, with the cerebellum as an additional component,
the overall arrangement is generally the same. The peripheral nervous system (not shown) in humans
and other mammals includes sensory ganglia as well as autonomic and enteric motor ganglia that
modulate the functions of organ systems (e.g., the cardiovascular and digestive systems).
in the head and its large number of nerve cells and their connections. Although
students spend untold hours learning the human brain anatomy, this information
is far more relevant to clinical medicine than it is to basic neurobiology and is as
likely to reflect the vagaries of evolutionary tinkering as neurological principles. In
any event, measuring the mass or volume of brains is easy, whereas measuring these
properties for entire nervous systems is not.
This limitation means that about the best one can do in comparing nervous sys-
tems across species is to assess brain size relative to body size, typically by weight.
Since nervous systems are obviously larger in larger animals, the goal is to see
whether there is a difference in the ratio of brain weight to body weight across some
group of animals and, if so, what a relatively larger brain implies. The result of such
studies is that the two metrics—brain weight and body weight—are correlated, as
one would expect. The only proviso is that, among mammals or other taxonomic
Porpoise Elephant
10 4
Human Blue
Australopithecus whale
10 3 Male gorilla
Baboon
Chimpanzee
Wolf
10 2 Opossum Lion
Brain weight (g)
10 1 Rat Alligator
Vampire
bat
Coelacanth
100 Mole
Eel
10 –1
Goldfish
10 –2
10 –3 10 –2 10 –1 100 10 1 10 2 10 3 10 4 10 5
Body weight (kg)
FIGURE 3.4 Brain-to-body weight ratios among mammals. Primates and most cetaceans—groups
thought to be more “intelligent” than other mammals—generally have greater brain-to-body weight
ratios. This observation implies that having relatively more brain mass compared to body mass
enhances the ability to solve complex problems (one definition of intelligence). Similar trends are
evident in comparisons across other animal classes as well: animals with more complex behavioral
repertoires have relatively larger nervous systems with more neurons and synapses.
After Jerison, 1973; from Purves D, Cabeza R, Huettel SA, LaBar KS, Platt ML, Woldorff M (2013). Principles of
Cognitive Neuroscience (2nd edition). Sunderland, MA: Sinauer Associates, fig 15.7A.
groups, the ratio of brain to body weight tends to be somewhat greater in species
that, by one criterion or another, seem more “intelligent” (Figure 3.4).
“Intelligence,” as already suggested, is a widely abused term applied to people
or other animals that appear to solve at least some problems better than another
individual or species.17 But that idea is clearly problematic. Einstein was certainly
a genius in physics and a talented violinist as well. But he was presumably a lousy
athlete and average (or less) in the many other ways in which human talents are
expressed. Thus, his brain—the subject of considerable intrigue and scientific sil-
liness18 —would not be expected to be larger or grossly different from anyone
17
Benson-Amram S, Dantzer B, Stricker G, Swanson EM, Holekamp KE (2016). Brain size predicts
problem-solving ability in mammalian carnivores. PNAS 112:2532–2537.
18
Paterniti M (2000). Driving Mr. Albert: A Trip Across America with Einstein’s Brain. New York:
Dial Press.
Another random document with
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nehmen würden. Somit war mit einer erheblichen feindlichen
Tätigkeit erst nach der Regenzeit zu rechnen, deren Schluß ich auf
Ende Februar annahm. Ungefähr um diese Zeit gedachte ich unsere
Streitkräfte in der Gegend von Nanungu enger zu versammeln. Bis
dahin mußten wir die Verpflegungsbestände dieses Gebietes also
schonen und so viel wie möglich von den Beständen leben, die aus
dem äußeren Umkreise unserer gesamten jetzigen Unterbringung
bezogen werden konnten. Die Jagdergebnisse waren bei Chirumba
anfangs nicht erheblich, steigerten sich aber, als auf dem östlichen
Ufer des Lujendaflusses und besonders weiter stromaufwärts
stärkere Bestände an Antilopen vorgefunden wurden. Der Verkehr
über den Fluß vollzog sich jetzt, bei dem niedrigen Wasserstande
der Trockenzeit, für die Trägerkarawanen, die ihre Lasten nach
Magazinen auf das Ostufer des Flusses schafften, durch mehrere
Furten. Außerdem waren zum Übersetzen einige große Einbäume
vorhanden. Zur weiteren Erkundung und zum Ansammeln von
Verpflegung wurden wochenlang Patrouillen entsandt. Die auf
Monate entsandte Patrouille des Leutnants von Scherbening
marschierte von Chirumba über Mtende, Mahua und schließlich
weiter nach Süden über den Luriofluß, dann den Malemafluß
aufwärts und überrumpelte die portugiesische Boma Malema. Ein
Italiener, der am Lujendaflusse Elefanten gejagt und sich gänzlich
abgerissen und verhungert bei uns eingefunden hatte, begleitete die
Patrouille des Leutnants von Scherbening. Durch verschleppte
Malaria war aber die Gesundheit des Mannes derartig untergraben
und seine Milz so riesenhaft angeschwollen, daß er aus der Gegend
von Mahua schließlich auf seine bei Malacotera gelegene Pflanzung
getragen werden mußte.
Anfang Januar 1918 begannen die Engländer, sich zu regen. Von
der Südostecke des Nyassasees her nahten sich zwei feindliche
Bataillone — das 1. und 2. der I. Kings African Rifles — der
Abteilung des Hauptmanns Goering, der herangekommen war und
dann ein festes Lager in dem spitzen Winkel zwischen Luambala-
und Lujendafluß bezogen hatte. Er sicherte die weiter oberhalb am
Lujendafluß gelegenen Verpflegungsmagazine. Am 9. Januar wurde
vormittags ein Teil des Feindes, der hier vereinzelt angriff,
geschlagen. Als der Gegner nachmittags nach Eintreffen seiner
Verstärkungen erneut vorging und zugleich feindliche Truppen auf
das Ostufer des Lujendaflusses nach Norden in Richtung auf die
Verpflegungsmagazine vordrangen, ging Hauptmann Goering mit
dem Hauptteil seiner Truppe auf das Ostufer. Im bisherigen Lager,
auf dem Westufer, blieb nur eine stärkere Patrouille, die den Feind
zurückhielt. Zu gleicher Zeit gingen feindliche Truppen — festgestellt
wurde das aus südafrikanischen Mischlingen bestehende II. Cape-
Corps — von Mtangula aus auf Mwembe vor.
Es kam nun zu einer Unzahl von kleineren Unternehmungen und
Patrouillengefechten, die uns bei der Schwierigkeit, unsere Träger,
die Verpflegung herantrugen, ausreichend zu decken, oft in eine
unangenehme Lage brachten. Die Engländer benutzten diese für
uns unangenehme Zeit geschickt zu Versuchen, die Anhänglichkeit
unserer Askari und Träger zu untergraben. Natürlich waren viele
derselben kriegsmüde. Die Strapazen waren auch wirklich recht
groß gewesen. Dazu kam bei vielen das unsichere Gefühl, wohin die
Reise nun weiter führen sollte. Die überwiegende Mehrzahl der
Schwarzen hängt an den Verwandten und an der Heimat. Sie sagten
sich: wenn wir nun weiter marschieren, dann kennen wir das Land
und die Wege nicht mehr. Von da, wo wir jetzt sind, finden wir uns
noch zurück, aber später nicht mehr. Die englischen Einflüsterungen
und Flugblätter, die Colonel Baxter in unsere Reihen tragen ließ,
fielen deshalb bei manchen auf fruchtbaren Boden, und so sind
damals eine Reihe guter Askari und selbst ältere Chargen desertiert.
Kleine Unannehmlichkeiten, wie sie immer vorkommen,
Weiberangelegenheiten und dergleichen, trugen das ihrige dazu bei,
den Leuten ihren Entschluß zum Fortlaufen zu erleichtern. Es kam
vor, daß ein alter Sol (schwarzer Feldwebel), der eine glänzende,
selbständige Patrouille geführt und eine starke Trägerabteilung mit
Lasten geschickt mitten durch die feindlichen Reihen zurückgebracht
hatte, und der wegen seiner guten Leistungen zum „Effendi“
(schwarzen Offizier) vorgeschlagen war, plötzlich verschwunden war.
Auch er war desertiert. Die Impulsivität des Schwarzen macht auch
für schlechte Einflüsterungen leicht empfänglich. Aber wenn sich der
feindliche Colonel rühmen kann, durch seine Tätigkeit in unseren
Reihen bei einigen Elementen eine moralische Krankhaftigkeit
erzeugt zu haben, so war dieser Zustand doch nur von
vorübergehender Dauer. Bald kehrten die alte Unternehmungslust
und das alte Vertrauen zurück, auch bei denen, die den Kopf hatten
hängen lassen. Das Beispiel der guten Askari, die einfach lachten
über die goldenen Berge, die ihnen der Feind versprach, wenn sie
desertierten, gewann die Oberhand. In einem so langen und
aufreibenden Kriege war eben die Stimmung gelegentlich auch
niedergedrückt. Es kam weniger darauf an, sich hierüber zu
erstaunen und zu entrüsten, als vielmehr, dem kräftig
entgegenzuwirken, und dazu waren die guten Elemente fest
entschlossen, die zahlreich unter unseren Europäern, Askari und
Trägern vorhanden waren.
Zweiter Abschnitt
Östlich des Lujendaflusses
D ie Patrouille des Hauptmanns Otto, die vom Hauptmann Tafel

vor dessen Waffenstreckung zu mir abgesandt war und die
Einzelheiten über die dortigen Vorgänge meldete, war in Chirumba
eingetroffen. Hauptmann Otto rückte jetzt mit zwei weiteren
Kompagnien nach Luambala und übernahm dort den gemeinsamen
Befehl auch über die Abteilung Goering (3 Kompagnien).
Richtigerweise erfolgte der Hauptdruck des Feindes dort bei
Luambala, und zwar auf dem östlichen Ufer des Lujenda. Es war ja
klar, daß, wenn der Feind dort flußabwärts vordrang, meine Lage bei
Chirumba, auf dem Westufer des Flusses, in einem Gebiet, dessen
Verpflegungsbestände nach und nach erschöpft wurden, und mit
dem durch die inzwischen gefallenen Regen stark angeschwollenen
Fluß in meinem Rücken äußerst ungünstig wurde.
Es war nötig, mich dieser Lage zu entziehen und meine Kräfte
rechtzeitig auf das östliche Lujendaufer zu verschieben. Leider
waren die Furten infolge Hochwassers nicht mehr passierbar, der
gesamte Uferwechsel auf die drei vorhandenen Einbäume
angewiesen.
Die Kompagnien wurden nach und nach ohne irgendwelche
Störungen auf das Ostufer hinübergesetzt. Die Ernährung fing an,
recht schwierig zu werden. Erfreulicherweise meldete Hauptmann
Koehl, der in der Gegend von M e d o und N a m u n u die sehr
verständigen Eingeborenen zum Anbau schnell reifender Feldfrüchte
angehalten hatte, daß dort schon von Mitte Februar ab auf die
Erträgnisse der neuen Ernte zu rechnen sei. Aber bis dahin war
noch ein Monat; so mußten wir mit allen Mitteln versuchen, noch
längere Zeit in der Gegend von Chirumba zu bleiben.
Erfreulicherweise halfen uns da, wie früher das Manna den Kindern
Israel, die in enormen Mengen um diese Jahreszeit
hervorschießenden Pilze aus der gröbsten Verlegenheit. Ich hatte
mich schon in Deutschland für Pilzkunde interessiert und fand bald
nahe Verwandte unserer deutschen Sorten, der Pfifferlinge,
Champignons, Steinpilze und anderer im afrikanischen Walde vor.
Ich habe sie oft in kürzester Zeit körbeweise gesammelt, und wenn
auch eine allzu einseitige Pilznahrung schwer verdaulich und nicht
allzu kräftig ist, so waren uns die Pilze doch eine wesentliche
Beihilfe.
Askarifrau
Bei strömendem Regen zogen wir dann weiter nach Osten. Die
sonst trockenen Bergschluchten waren zu reißenden Flüssen
geworden. Durch gefällte Uferbäume, die quer über den Fluß fielen,
wurden Übergänge geschaffen, ein Geländer schnell durch Stangen
oder zusammengeschlagene Baumrinde improvisiert. Mein Maultier,
das ich wegen eines Fiebers ritt, das mich befallen hatte — ich war
anscheinend besonders empfänglich für Malaria und litt häufig
darunter — sowie die wenigen anderen Reittiere, die bisher nicht in
den Kochtopf gewandert waren, schwammen hinüber. Am
Lagerplatz angekommen, bauten mir meine Leute wegen der
Feuchtigkeit des Erdbodens schnell aus Zweigen eine erhöhte
Lagerstelle, über die meine beiden Mannschaftszeltbahnen als Dach
gespannt wurden. Oberveterinär d. Res. Huber, der für das
materielle Wohl der Mitglieder des Kommandos sorgte, und unter
ihm unser tüchtiger schwarzer Koch, der bärtige alte „Baba“, waren
sogleich am Werk, und trotz regennassem Holze konnten wir uns
stets in kurzer Zeit am Feuer zu gemeinsamem Mahle einfinden. Oft
hatte es Dr. Huber fertig gebracht, hierzu in der Eile ein schützendes
Grasdach herstellen zu lassen.
An sonnigen Tagen wurde eifrig Tabak fermentiert und
geschnitten. Der tüchtige Feldintendant, Leutnant z. S. a. D. Besch,
der stets von neuem erfinderisch war, wenn es das leibliche Wohl
der Truppe galt, hatte auch hieran gedacht und den bei den
Eingeborenen vorgefundenen recht guten Tabak gesammelt. Trotz
alledem waren die Entbehrungen aber recht groß, und die
Einflüsterungen des Feindes, daß jeder Schwarze, der zu ihm
überliefe, frei in seine Heimat ziehen und dort auf eigenem Boden
behaglich leben sollte, trafen auch jetzt nicht immer auf taube Ohren.
Auch der jahrelang treu dienende Boy eines Offiziers war eines
Morgens verschwunden; wahrscheinlich hatte seine „Bibi“ (Frau) das
Kriegsleben satt bekommen.
Die Abteilung des Hauptmanns Otto rückte von Luambala aus
direkt nach Osten nach Mahua und fand dann am Luriofluß ein
reiches Gebiet mit Verpflegung vor. Abteilung Goering, die von
Luambala aus quer durch die Landschaft auf Mtende rückte, fand
unterwegs größere Verpflegungsmengen. Die Ernte war in dieser
Gegend sehr viel früher als in Deutsch-Ostafrika; der Mais fing an zu
reifen und konnte zum großen Teil schon verzehrt werden.
Das Kommando zog zunächst von Chirumba nach Mtende und
dann nach einigen Tagen weiter nach N a n u n g u ; Abteilung Wahle,
die von Chirumba aus nach Mtende gefolgt war, wurde hier von
mehreren feindlichen Kompagnien umgangen, die überraschend auf
einer Höhe im Rücken der Abteilung auftauchten und den
Botendienst und die Transporte unterbrachen. General Wahle
entzog sich durch einen Umweg dieser unbequemen Lage und
rückte in Richtung auf Nanungu näher an das Kommando heran.
Bei Nanungu fanden wir reichliche Verpflegung, und es lohnte
sich, in dem Raume von N a n u n g u , N a m u n u und weiter südlich am
Lurio wieder, wie in früherer Zeit, Aufkaufposten und Magazine
anzulegen. Die Wildbestände lieferten gute Beute, und die
Eingeborenen brachten gern Gartenfrüchte und Honig herbei, um
diese gegen Fleisch, lieber aber noch gegen Bekleidungsstücke
einzutauschen. Recht willkommen war eine wohlschmeckende,
süße, kirschartige Porifrucht, die zu Millionen in der Gegend von
Nanungu heranreifte. Ich ließ sie mit Vorliebe zu Jam verarbeiten.
Auch andere Leckereien, besonders Erdnüsse, bekamen wir
gelegentlich, und weit und breit verrieten die krähenden Hähne, daß
es in den Lagern und bei den Eingeborenen Hühner und Eier gab.
Das Einsetzen der Regenzeit stimmte nicht genau mit dem
Vorhersagen der Eingeborenen überein; es gab zwar tüchtige
Güsse, aber das Wasser lief in dem hügeligen Gelände schnell ab
und sammelte sich in der Hauptader jener Gegend, dem Msalufluß,
der zu einem starken Hindernis anschwoll. Über den Msalufluß hatte
der als Vizefeldwebel zur Truppe eingezogene Feldpostsekretär
Hartmann eine Pontonbrücke gebaut, die uns mit der Abteilung des
Generals Wahle verband. Diese war noch auf dem westlichen Ufer
des Flusses geblieben. Als schwimmende Unterstützungen der
Brücke dienten Rindenboote. Die Notwendigkeit, in dem
wasserreichen Gebiete die angeschwollenen Flüsse glatt
überwinden zu müssen, lenkte meine Aufmerksamkeit auf diese
Frage. Bisher hatten wir für alle Fälle einige Einbäume mitgetragen.
Der Transport war aber auf die Dauer zu schwierig und dieses Mittel
zu wenig leistungsfähig. Der Kriegsfreiwillige Gerth, ein Pflanzer vom
unteren Rufiji, interessierte sich besonders für diese Frage und ließ
sich von den Eingeborenen der Landschaft, die hierin besonders
sachkundig waren, im Bau von Rindenbooten unterweisen.
Nachdem die Versuche schnell zu einem Resultat geführt hatten,
wurde bei allen Kompagnien der Bau solcher Boote, für deren
Herstellung nach einiger Übung knapp zwei Stunden erforderlich
waren, mit Eifer betrieben. Diese Boote sind in größerem Maßstabe
von uns nicht benutzt worden, aber sie gaben uns das sichere
Gefühl, daß im Notfalle auch ein starkes Stromhindernis für unsere
recht unhandlichen Karawanen und Lasten nicht unüberwindlich sei.
Nach einiger Bekanntschaft mit der Gegend fanden wir im
Msaluflusse Furten, die auch bei hohem Wasserstand einen
Uferwechsel gestatteten. Unsere Kampfpatrouillen unter Sergeant
Valett und anderen gingen von unseren gesicherten Lagern bei
Nanungu aus, durchschritten den Fluß, der unser Unterkunftsgebiet
nach Westen zu als Hindernis begrenzte, und suchten den Feind in
seinen Lagern bei Mtende auf. Einer dieser Patrouillen, die
besonders stark gemacht und mit 2 Maschinengewehren
ausgerüstet war, gelang es, westlich von Mtende eine feindliche
Verpflegungskarawane zu überfallen. Die Unsrigen haben sich dann
aber nicht schnell genug den feindlichen Bedeckungstruppen
entzogen und kamen, von mehreren Seiten angegriffen, in eine
schwierige Lage. Beide Maschinengewehre gingen verloren und die
sie bedienenden Europäer fielen. Nach und nach trafen die Askari
zwar wieder vollzählig in Nanungu ein, aber der Patrouillenführer,
Feldwebel Müßlin, der sich während des Marsches allein entfernt
hatte, war in Feindeshand gefallen. Eine andere Patrouille, mit der
Hauptmann Müller nach Norden zu den Msalu überschritt, vertrieb
schnell eine englische Postierung bei Lusinje. In der Gegend von
Lusinje war auch das Lager des englischen Leutnant Wienholt
erbeutet worden, der, wie früher erwähnt, aus unserer
Gefangenschaft entlaufen und einer der besten englischen
Patrouillenführer geworden war. Die Eingeborenen wurden durch die
englischen Patrouillen gründlich bearbeitet und leisteten dem Feinde
gegen Belohnung durch Kleidungsstücke Spionagedienst. Auch der
anläßlich des Bootsbaues erwähnte Kriegsfreiwillige Gerth wurde am
Msalufluß, am Hause eines Jumben, von einer englischen Patrouille
überfallen und fand hierbei seinen Tod.
Mächtig erhoben uns damals in der zweiten Hälfte des März
1918 die Nachrichten von der gewaltigen deutschen Märzoffensive
an der Westfront, die unsere Funkenstation auffing. Ich wettete mit
dem Sanitätsoffizier beim Stabe, Stabsarzt Taute, daß Amiens bald
fallen würde. Die Zeit mehrwöchentlicher Ruhe, die jetzt beim
Stillstand unserer Operationen eintrat, benutzte ich, um meinen
rechten Fuß, der mir infolge eines Sandflohes seit einem halben
Jahr Unbequemlichkeiten machte, in Ordnung bringen zu lassen. Die
in manchen Lagern in Unmenge vorkommenden Sandflöhe bohren
sich mit Vorliebe an den Rändern der Fußnägel in das Fleisch und
verursachen dort schmerzhafte Entzündungen. Wird nicht
aufgepaßt, so greifen diese weiter um sich und nach ärztlicher
Ansicht ist die Verstümmelung vieler Füße der Eingeborenen und
der Verlust der Zehen häufig auf solche Sandflöhe zurückzuführen.
Auch ich litt unter dieser Unbequemlichkeit, und beim Gehen
bildeten sich immer wieder Entzündungen. Stabsarzt Taute konnte
mir glücklicherweise den Zeh unempfindlich machen, um dann den
Nagel herauszureißen.
Auch in anderer Weise war ich einmal etwas gehindert. Auf
einem Erkundungsgange hatte mir ein Halm des übermannshohen
Grases in mein rechtes Auge geschnitten, und bei der
nachfolgenden Behandlung war infolge von Atropin die
Anpassungsfähigkeit der Linse beeinträchtigt; ich konnte deshalb mit
dem rechten Auge nicht ordentlich sehen und keine Schrift oder
Kartenskizzen erkennen. Dieser Zustand war lästig, weil mein linkes
Auge durch eine im Jahre 1906 beim Hottentottenaufstand in
Südwestafrika erhaltene Schußverletzung so stark beschädigt
worden ist, daß ich mit diesem nur vermittelst Starbrille lesen kann.
Eine solche war aber nicht verfügbar, und so war ich gezwungen, an
verschiedene Unternehmungen heranzugehen, ohne richtig sehen
zu können.
Die Patrouillen der Abteilung Koehl waren aus der Gegend
Medo-Namunu inzwischen bis an die Küste vorgestreift, hatten am
unteren Luriofluß und weit südlich desselben portugiesische
Befestigungen erobert, einige Kanonen, vor allem aber Gewehre und
Munition, meist von altem Modell, und erhebliche Mengen an
Verpflegung erbeutet. Die Eingeborenen erwiesen sich hierbei
freundlich gegen unsere Leute und sahen in ihnen auch hier die
Befreier von der portugiesischen Bedrückung. Auch Patrouillen der
Abteilung Otto waren von Mahua her in das Gebiet südlich des
Lurioflusses gestreift, und der in Eingeborenenangelegenheiten so
erfahrene Oberleutnant d. Ldw. Methner, erster Referent unseres
Gouvernements, rühmte die Tüchtigkeit und Klugheit der
portugiesischen Eingeborenen und den verständigen und weiten
Blick ihrer Ortsoberhäupter.
Leutnant von Scherbening, der mit seiner Patrouille die Boma
Malema genommen hatte, berichtete von dem großen Reichtum
dieser Gegend. Um auch uns hiervon etwas zukommen zu lassen,
schickte er ein erbeutetes Schwein nach Nanungu. Da es nicht
laufen wollte, wurde es 500 km weit getragen. Bei seiner Ankunft
stellte sich leider heraus, daß es gar kein europäisches Schwein
war, sondern ein Porischwein, wie wir es selbst im Walde häufig
erlegten.
Wieder war eine Zeit gekommen, wo es schwer war, Nachrichten
über den Feind zu erhalten. Aber aus den unvollkommenen Karten,
die uns zur Verfügung standen, war doch manches herauszulesen.
Ich konnte nicht zweifeln, daß die sicher bevorstehenden feindlichen
Operationen mit ihren Hauptkräften von der Küste, aus der Gegend
von P o r t o A m e l i a , ansetzen würden. Das Auftreten stärkerer
feindlicher Kräfte bei M t e n d e sowie die allerdings sehr unsichere
Nachricht, daß auch feindliche Truppen von Südwesten her auf
M a h u a im Anmarsch waren, zeigten mir, daß zugleich mit dem
bevorstehenden Vorrücken der feindlichen Hauptkräfte andere
Truppen von Westen her operieren wollten. Es schien eine Lage
heranzureifen, in der es möglich war, die innere Linie, auf der ich
stand, auszunutzen und den einen oder den anderen Teil des
Feindes vereinzelt zu fassen. Die Nachschubverhältnisse des
Gegners bedingten es, daß diejenigen seiner Kolonnen, die von
Westen kamen, nicht allzu stark sein konnten. Hier also bot sich
auch voraussichtlich die von mir gesuchte günstige Chance. Mit dem
Hauptteil meiner Truppen blieb ich deshalb in der Gegend von
Nanungu und zog nach hier auch die Abteilung des Hauptmanns
Otto vom Lurio heran. Mit diesen Kräften wollte ich angriffsweise
verfahren und zwar nach Westen zu. Dem Hauptmann Koehl, der
seine Abteilung bei M e d o sammelte, fiel die Aufgabe zu, die von
Porto Amelia heranrückenden Hauptstreitkräfte des Feindes
hinzuhalten und allmählich schrittweise auf mich zurückzugehen.
Hauptmann Müller, der nach jahrelanger Tätigkeit beim
Kommando eine selbständige Abteilung von 2 Kompagnien
übernommen hatte, wurde aus der Gegend von Nanungu auf Mahua
vorgesandt, um dort dem Feinde nach Möglichkeit Abbruch zu tun.
Er umging Mahua und überraschte südwestlich dieses Ortes die
befestigte Verpflegungsstation Kanene. Die verteidigenden
englischen Europäer sahen ein, daß die angesammelten Bestände
verloren waren. Um dies wenigstens teilweise zu verhindern,
machten sie sich über die Alkoholbestände des Lagers her und
fielen recht angeheitert in unsere Hände.
Ich selbst rückte Mitte April gleichfalls in Richtung auf Mahua vor
und hörte beim Anmarsch schon aus weiter Entfernung von dort her
heftige Detonationen. Hauptmann Müller war nordöstlich Mahua bei
Koriwa auf ein feindliches Bataillon unter Colonel Barton gestoßen,
das einen Streifzug unternommen hatte und nun von den Unsrigen
während des Marsches sogleich angegriffen wurde. Trotzdem auf
unserer Seite kaum 70 Gewehre im Gefecht waren, gelang es doch,
den Feind auf seinem rechten Flügel zu umfassen und ihn von dort
aus von einem Termitenhügel (großen Ameisenhügel) so energisch
unter wirksames Maschinengewehrfeuer zu nehmen, daß er in
wilder Flucht davonlief. Er verlor dabei über 40 Mann. Oberleutnant
z. S. Wunderlich, der einen schweren Schuß durch den Unterleib
erhalten hatte, mußte zu dem 2 Tagemärsche entfernten Lazarett
von Nanungu geschafft werden und starb nach kurzer Zeit.
Der Schlag, zu dem ich das Gros angesetzt hatte, war durch die
schwache Abteilung Müller bereits erfolgreich ausgeführt worden. Ich
wandte mich deshalb mit meinen Hauptkräften wieder in die Gegend
dicht westlich Nanungu. Dort war inzwischen ein stärkerer Feind am
Msaluflusse eingetroffen und hatte diesen mit stärkeren Patrouillen
überschritten. Meine Berechnung, einen stärkeren feindlichen Körper
unmittelbar nach Überschreiten des Flusses überraschend fassen zu
können, traf nicht zu; die erhaltenen Meldungen waren irrtümlich
gewesen. Aber in einer ganzen Reihe kleinerer Zusammenstöße am
Msalufluß und westlich desselben wurden dem Gegner durch unsere
Kampfpatrouillen doch nach und nach erhebliche Verluste zugefügt,
und seine Streifabteilungen räumten bald das östliche Msalu-Ufer.
Unsere Verpflegungspatrouillen, deren Aufgabe es war, in der
Richtung auf Mahua weiter Verpflegung zu beschaffen, stießen am
3. Mai überraschend auf stärkere feindliche Abteilungen in der
Gegend von S a i d i , die unser Feldlazarett und unsere
Verpflegungsmagazine bei M a k o t i stark gefährdeten.
Nach Makoti war zur Vorbereitung der zukünftigen, mehr nach
westlicher Richtung geplanten Operationen ein Teil unserer
Bestände gebracht worden. Unsere sofort entsandten
Kampfpatrouillen hatten am Kirekaberge bei Makoti mehrere
Zusammenstöße mit dem Feinde. Ich glaubte zunächst nur an
Patrouillen des Gegners, entsandte deshalb den Hauptmann Schulz
mit einer starken Patrouille dorthin zur Verstärkung und rückte selbst
am 4. Mai mit dem Gros an die Straße Nanungu-Mahua. Von hier
aus glaubte ich, schnell gegen irgendwo überraschend auftretende
feindliche Kräfte eingreifen zu können. Die allgemeine Lage klärte
sich dadurch, daß unsere Patrouillen im Laufe des Tages am
Kirekaberge auf einen neuen Gegner gestoßen waren. Eine
feindliche Abteilung war zurückgeworfen worden, und es war
wahrscheinlich, daß stärkere Kräfte in verschanzten Lagern dahinter
standen. Am 5. Mai morgens marschierte ich aus meinem Lager ab,
auf Makoti zu. Während des Anmarsches wünschte ich sehnlichst,
daß der Feind uns den Angriff auf seine befestigte Stellung ersparen
möchte, und hoffte, was ja auch nach der ganzen Lage gar nicht
unwahrscheinlich war, daß er aus seinen Schanzen herauskommen
und sich so ein Kampf im freien Felde entwickeln würde. Gelang es
uns bei dieser Gelegenheit, mit unseren Hauptkräften einzugreifen,
ohne daß der Feind von unserem Anmarsch eine Ahnung hatte, so
war ein erheblicher Erfolg nicht unwahrscheinlich.
Um 11 Uhr vormittags langte ich am Kirekaberge an und begab
mich nach vorn zum Hauptmann Schulz, der mit seinen Patrouillen
einige in dem Stangenholz befindliche Felsengrotten besetzt hatte.
Als ich gerade angekommen war, erhielt ich von einem Sol
(schwarzen Feldwebel), der von einem Patrouillengang
zurückkehrte, die Meldung, daß der Feind in großer Stärke
ausgeschwärmt vorrücke und sofort auf nahe Entfernung auftauchen
müsse. Ich benachrichtigte hiervon den Oberleutnant Boell, der mit
seiner Kompagnie soeben hinter der Abteilung Schulz eingetroffen
war, und beauftragte ihn, im Falle eines feindlichen Angriffes
sogleich einzugreifen. Dann ging ich zurück und ordnete den
Aufmarsch unserer weiteren, nach und nach eintreffenden
Kompagnien. Inzwischen ging vorn das Gefecht los. Der Feind hatte
in dichten Schützenlinien vorgehend unsere Patrouillen schnell aus
ihren Steingrotten zurückgeworfen, war dann aber zu seiner größten
Überraschung in das wirksame Maschinengewehrfeuer der
Kompagnie Boell geraten und teilweise zurückgegangen. Die
demnächst eintreffende Abteilung Goering wurde sogleich zum
rechts umfassenden Angriff angesetzt, der Feind dadurch völlig
überrascht und mit recht schweren Verlusten in großer Eile
zurückgeworfen.
So langten wir nach mehreren Kilometern heftigen Nachdrängens
vor den feindlichen Verschanzungen an. Auf unserem linken Flügel,
wo zwei weitere Kompagnien eingesetzt wurden, ging das Gefecht
mehrfach hin und her, und es war mir im Busch schwer, Freund und
Feind zu unterscheiden. Es dauerte dadurch einige Zeit, bis ich mir
von den Verhältnissen auf dem linken Flügel ein klares Bild machen
konnte, und erst die Meldung des Majors Kraut, der von mir zur
Erkundung dorthin gesandt war, ließ mich erkennen, daß das
Vorgehen unseres linken Flügels auf einer Waldlichtung in recht
wirksames feindliches Feuer geraten und dadurch zum Stocken
gekommen war. Ein Gegenangriff, den der Feind unternahm und der
schon ziemlich dicht an den Standplatz des Kommandos gekommen
war, hätte uns recht unangenehm werden können. Zu unserem
großen Glück traf aber gerade in diesem Augenblick Oberleutnant
Buechsel, der mit seiner Kompagnie detachiert gewesen war und
deshalb verspätet ankam, auf dem Gefechtsfelde ein und konnte der
Gefahr begegnen.
Auf dem rechten Flügel hatte inzwischen Hauptmann Goering
erkannt, daß ein frontales Anstürmen auf die feindlichen
Verschanzungen keine Aussicht auf Erfolg bot. Er hatte deshalb
Oberleutnant Meier mit einer starken Patrouille die Stellung des
Gegners umgehen lassen, um von rückwärts her einen dort
befindlichen feindlichen Minenwerfer zu beschießen und womöglich
fortzunehmen. Diese Wegnahme gelang nicht, da der Feind
unerwarteterweise noch über Reserven verfügte, die die Patrouille
Meier fernzuhalten imstande waren. So kam das Gefecht zum
Stillstand. Bei Einbruch völliger Dunkelheit lagen wir dem Feinde
dicht gegenüber. Beiderseits fielen nur noch ab und zu Schüsse.
Während des Gefechts wurden die büromäßigen Arbeiten — es
wurde auch in Afrika geschrieben, wenn auch nicht so viel als sonst
üblich — erledigt. Eine Anzahl Eingänge, wie Beschwerden und
sonstige Unannehmlichkeiten lagen vor. Mit den Kompagnieführern
konnte ich mich von Zeit zu Zeit mündlich besprechen und ließ sie
zu diesem Zweck zu mir kommen. Ich selbst wechselte meinen
Standplatz so wenig wie möglich, um bei der Übermittlung der
eingehenden Meldungen Schwierigkeiten und unliebsame
Verzögerungen zu vermeiden. Abgekocht wurde weiter rückwärts,
wo auch der Verbandplatz eingerichtet worden war. Auch wir, die
Angehörigen des Kommandos, erhielten durch unsere schwarzen
Diener wie gewöhnlich die zubereitete Verpflegung in die
Gefechtslinie vorgebracht.
Um die Truppe zu weiterer Verwendung wieder in die Hand zu
bekommen, wurden Teile derselben aus der vorderen Gefechtslinie
zurückgezogen und gesammelt. Ich überlegte, daß es zweckmäßig
sei, die Nacht über so liegen zu bleiben, um am nächsten Tage das
Gefecht wieder aufnehmen zu können und vor allem, um zu
versuchen, den Feind von seinem Wasser abzuschneiden, das
irgendwo außerhalb seines Lagers liegen mußte.
Da traf gegen Mitternacht die Meldung ein, daß eine unserer
Patrouillen an der Straße N a n u n g u - M a h u a auf einen stärkeren
Feind gestoßen sei. Ich mußte befürchten, daß dieser Gegner, den
ich der Selbständigkeit seines Auftretens wegen für stark hielt, weiter
auf Nanungu vordringen und sich so in den Besitz unserer an dieser
Straße gelagerten Kompagnielasten (Munition, Verbandzeug,
Verpflegung, Kranke usw.) sowie der Magazine von Nanungu setzen
würde. Ich rückte deshalb noch in der Nacht mit dem Hauptteil
meiner Kräfte über Makoti wieder an die Straße Nanungu-Mahua ab.
Dicht am Feinde blieben nur starke Patrouillen, die aber nicht
bemerkten, daß der Gegner ebenfalls noch während der Nacht seine
Stellung räumte und in Richtung auf Mahua abzog. Am 6. Mai stellte
sich heraus, daß die Meldung von den starken feindlichen Kräften an
der Straße Nanungu-Mahua, die meinen Abmarsch veranlaßt hatte,
verkehrt gewesen war; es befand sich dort überhaupt kein Feind.
Hauptmann Müller, der das Schießen der englischen Minenwerfer
gehört hatte, war in vortrefflicher Initiative aus seinem nordöstlich
Mahua gelegenen Lager im Eilmarsch sofort auf den Gefechtslärm
losmarschiert und anscheinend für Feind gehalten worden.
Als er auf dem Gefechtsfeld ankam, stellte er fest, daß der
Gegner abgerückt war. Der Feind, der aus 4 Kompagnien und einer
Maschinengewehrkompagnie bestand, und, nach seinen
Befestigungsanlagen zu urteilen, 1000 Mann stark war, war durch
unsere wenig mehr als 300 Gewehre — wir waren 62 Europäer und
342 Askari gewesen — vollständig geschlagen worden. Auf seiner
Seite waren 14 Europäer und 91 Askari gefallen, 3 Europäer und 3
Askari hatte er an Gefangenen verloren. Außerdem war sein
Hospital mit etwa 100 Verwundeten in unsere Hand gefallen; andere
Verwundete hat er nach Aussage von Eingeborenen noch
mitgenommen. Unsere Verluste betrugen: 6 Europäer, 24 Askari, 5
andere Farbige gefallen, 10 Europäer, 67 Askari und 28 andere
Farbige verwundet.
Während diese für uns so erfreulichen Erfolge gegen die
westlichen feindlichen Kolonnen erzielt wurden, hatte Abteilung
Koehl gegen die feindliche Division, die von Porto Amelia auf
Nanungu vordrang, andauernd Gefechte, manchmal von
erheblichem Umfange, zu bestehen. Bei Medo hatte der Feind nach
seiner eigenen Angabe recht schwere Verluste; in einem Gefecht,
das westlich von Medo stattgefunden hatte, war es dem Hauptmann
Spangenberg mit seinen 2 Kompagnien gelungen, den Feind sehr
gewandt zu umgehen, von rückwärts her an seine leichte
Feldhaubitzbatterie heranzukommen und diese im Sturm zu
nehmen. Fast die gesamte Bedienung und Bespannung fiel. Leider
war es nicht möglich, die Geschütze und die Munition mitzunehmen.
Sie wurden unbrauchbar gemacht. Aber trotz solcher Einzelerfolge
mußte Abteilung Koehl weiter zurückweichen. Es nahte der
Augenblick, wo sich vielleicht durch rechtzeitiges Eingreifen meiner
Hauptkräfte bei der Abteilung Koehl ein durchschlagender Erfolg
gegen General Edwards erzielen lassen würde.
Wieder einmal war aber die Verpflegungsfrage ein Bleigewicht für
die Bewegungen. Die Feldfrüchte der Landschaft waren im
wesentlichen aufgezehrt bis auf den Mtama, der in dieser Gegend
früher heranreift als in Deutsch-Ostafrika. Aber er war noch nicht reif.
Um nicht rein aus Verpflegungsgründen abrücken zu müssen, halfen
wir uns dadurch, daß wir den Mtama durch Trocknen notreif
machten. Die Frucht war auch auf diese Weise gut verwertbar, und
da in der Gegend sehr viel wuchs, so konnte im allgemeinen jeder
so viel bekommen, wie er haben wollte, und keiner litt Not.
Der Bestand der Felder veranlaßte mich, mit den Hauptkräften
der Truppe mehr nach Südwesten, in der Richtung auf Mahua, zu
marschieren und in der Gegend des Timbaniberges, am
Koromaberg, Lager zu beziehen. Von hier aus wollte ich im Notfall
nach Süden weiterziehen, um in den fruchtbaren Gegenden der
Vereinigung des Malema- und Lurioflusses die dortigen reichen
Verpflegungsgebiete auszunutzen. Westlich des Timbaniberges war
das Gelände günstig, um ein entscheidendes Gefecht gegen
General Edwards aufzunehmen, der der Abteilung des Hauptmann
Koehl von Nanungu weiter in südwestlicher Richtung folgte. Das
außerordentlich felsige und zerrissene Gelände am Timbaniberg und
6 km nordöstlich davon bis zu der Stelle, auf die die Abteilung Koehl
zurückgewichen war, war nicht günstig für ein von mir beabsichtigtes
entscheidendes Gefecht. Am 21. Mai verrieten sich neue feindliche
Lager westlich der Stellung der Abteilung Koehl durch ihren Rauch.
Ich vermutete, daß dieser neue Gegner am 22. Mai der Abteilung
Koehl von Westen her in den Rücken marschieren würde. Da habe
ich leider versäumt, der Abteilung Koehl den ganz bestimmten
Befehl zu geben, sogleich mit ihren Hauptkräften aus dem
ungünstigen Gelände heraus bis südwestlich des Timbaniberges zu
rücken. Statt des unzweideutigen Befehls gab ich eine Anweisung,
die zu viel Freiheit des Handelns ließ.
So kam es, daß die Abteilung Koehl ihre Träger mit den
Munitions- und Bagagelasten erst am 22. Mai vormittags in Marsch
setzte. Auch das wäre noch gut gegangen, wenn nicht
unglücklicherweise an ihrem Anfange der Gouverneur marschiert
wäre, der sich bis dahin bei der Abteilung Koehl aufgehalten hatte. In
Verkennung des Ernstes der Lage machte der Gouverneur mitten in
dem ungünstigen Gelände, wo er jeden Augenblick der
Überraschung durch den Feind ausgesetzt war, ohne sich
wirkungsvoll verteidigen zu können, einen längeren Halt. Hierdurch
ließen sich die Bagagen der Abteilung Koehl trotz dem ihnen vom
Hauptmann Koehl erteilten ausdrücklichen Befehle verleiten,
ebenfalls zu halten. Ich selbst erkundete an diesem Tage vormittags
noch einmal das südwestlich des Timbaniberges gelegene, recht
günstige Gelände und traf hierbei unter anderem den Leutnant
Kempner, der tags vorher bei Abteilung Koehl verwundet worden war
und zurückgetragen wurde. Bei der Abteilung Koehl selbst, wo seit
dem Morgen mehrere Angriffe des Feindes abgeschlagen waren,
war Gefechtslärm in weiter Ferne zu hören. Mit Hauptmann Koehl
bestand telephonische Verbindung, und ich kehrte, ohne eine
Ahnung von den Verhältnissen seiner Bagage zu haben, gegen 11
Uhr vormittags in das Koromalager zurück.
Um 12 Uhr mittags war ich gerade im Lager eingetroffen, als
plötzlich Minenwerferfeuer in großer Nähe erscholl, zweifellos
zwischen uns und der Abteilung Koehl. Unmittelbar darauf war die
Fernsprechverbindung dorthin unterbrochen. Jetzt blieb keine Wahl,
als sofort vom Koromalager her mit allen Kräften auf diesen neuen
Gegner vorzumarschieren, wobei ich die stille Hoffnung hatte, daß
es trotz der Ungunst des Geländes vielleicht gelingen würde, ihn zu
überraschen und entscheidend zu schlagen. Nach einer knappen
Stunde trafen wir am Timbaniberge ein und warfen vorgeschobene
feindliche Abteilungen schnell zurück. Einige versprengte Leute von
uns meldeten, daß der Gouverneur und die Bagage des Hauptmann
Koehl vom Feinde überraschend angegriffen und alle Lasten
verloren seien. Der Gouverneur selbst sei mit genauer Not
entkommen, andere sagten, er sei gefangen genommen worden.
Der Feind schoß ziemlich lebhaft mit mehreren Minenwerfern und
wurde durch unsere Kompagnien von mehreren Seiten angegriffen.
Er hatte aber eine gute Stellung eingenommen, in der er sich
verschanzt und einen Teil der erbeuteten Lasten geborgen hatte.
Leider nahmen wir ihm nur wenige wieder ab. Aber die feindliche
Stellung wurde doch umstellt und unter konzentrisches, für den
Feind recht verlustreiches Feuer genommen. Nach einer später
erbeuteten Nachricht haben die I. King’s African Rifles allein hierbei
etwa 200 Mann verloren.
Bei dieser Einschließung des Feindes unterstützten uns mehrere
Kompagnien und Patrouillen des Hauptmanns Koehl. Auch dieser
hatte sich mit seinen Hauptkräften gegen den neuen in seinem
Rücken auftretenden Feind gewandt und hoffte, denselben schlagen
zu können, während eine starke Patrouille, mit der Front nach
Nordosten, seinen bisherigen Gegner hinhielt. Diese Patrouille war
aber viel zu schwach. Sie wurde zurückgedrängt und mußte von
neuem durch Truppen der Abteilung Koehl verstärkt werden. Wenn
der Feind auch zweifellos im ganzen erhebliche Verluste erlitten
hatte, so war ein durchschlagender Erfolg für uns doch nicht
erreichbar. Das Gefecht wurde bei Eintritt der Dunkelheit
abgebrochen, und wir rückten in das von mir erkundete günstige
Gelände zwischen Timbani- und Koromaberg ab.
Im Lager am Koromaberg hatte sich inzwischen der Gouverneur
eingefunden. Er hatte bei dem Abenteuer sämtliche Lasten verloren
und wurde durch Unteroffizier Reder, dem bewährten und
umsichtigen Führer einer Kolonne, verpflegt. Auch ich steuerte dazu
bei, dem Gouverneur aus seiner Verlegenheit zu helfen und verehrte
ihm ein paar blaue Strümpfe, die seine Gattin mir im Anfang des
Krieges angefertigt hatte, die aber leider abfärbten.
Außer dem sehr fühlbaren Verlust von etwa 70000 Patronen
hatten wir auch den Verlust eines größeren Bestandes von
Papiernoten — ich glaube, es waren 30000 Rupien — zu beklagen.
Meinem Wunsche, statt mit Papiernoten zu bezahlen, lieber
Requisitionsscheine auszugeben und dadurch eine Menge
Sicherheitstransporte zu ersparen und unnötige Verluste zu
vermeiden, war früher nicht stattgegeben worden. Es waren
Millionen Rupien Papiernoten gedruckt worden. Das Mitschleppen
derselben war in dem jetzigen Stadium des Krieges eine besondere
Last. Um in der Zukunft wenigstens weitere Verluste zu vermeiden,
hat auf meine Anregung dann der Intendant einen großen Teil der
früher mühsam hergestellten Noten wieder vernichtet.
Dritter Abschnitt
Im Gebiet des Lurio- und Likungoflusses
A m 23. Mai wurden vom Koromalager auf einem quer durch den
Busch nach Koriwa abgesteckten Wege der Rest unserer Lasten
und der Hauptteil der Truppe in Bewegung gesetzt. Der Hauptteil
unserer Trägerkolonnen und die Kranken waren vorangegangen. Die
Nachhut unter Hauptmann Otto blieb noch einige Tage am
Koromaberge und wies dort mehrere Angriffe des Feindes
erfolgreich ab. Es schien, als ob der Gegner wieder einmal nach der
Beendigung einer konzentrischen Operation dort bei Timbani den
Hauptteil seiner Truppen vereinigt hätte und vor Antritt des
Weitermarsches einiger Zeit zur Regelung seines Nachschubs
bedurfte. Zurückkehrende Patrouillen meldeten starken Autoverkehr
auf der Straße Nanungu-Timbaniberg. Andere Patrouillen
berichteten von dem Vormarsch feindlicher Kräfte von Osten her auf
dem nördlichen Lurioufer.
Vom Feinde unbelästigt marschierte ich zunächst in die reiche
Gegend von Kwiri, südlich Mahua, und dann von dort aus weiter zum
Luriofluß. Dabei stellte es sich aber heraus, daß ein Teil unserer
Schwerverwundeten und Kranken diese mehrtägigen Märsche in
ihren „Maschillen“ (Tragbahren) nicht würde durchhalten können. Da
war es nicht leicht, für ärztliche Pflege zu sorgen. Es waren zu wenig
Pfleger da, um die Kranken einzeln von Fall zu Fall zurücklassen zu
können. So blieb nichts anderes übrig, als die Kranken von Zeit zu
Zeit zu sammeln und dann gemeinsam unter einem Arzt als
vollständiges Lazarett zu etablieren und sich endgültig von ihnen zu
trennen. Auch der Chefarzt der Schutztruppe, Generaloberarzt Dr.
Meixner war bei Kwiri mit einem solchen Lazarett liegen geblieben.
Von Leutnant d. Res. Schaefer, der uns bei den Vorbereitungen zum
Gefecht von Jassini so ausgezeichnete Dienste geleistet hatte, und
der jetzt an Schwarzwasserfieber schwer erkrankt war, nahm ich bei
dieser Gelegenheit Abschied. Der erfahrene Afrikaner war sich über

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Brains as Engines of Association: An

Operating Principle for Nervous

Understanding Operating Systems 8th Edition

Survey of Operating Systems 6th Edition Jane Holcombe

Survey of Operating Systems, 7th Edition Jane Holcombe

Survey of Operating Systems 5th Edition Jane Holcombe

Modern Operating Systems 5th Edition Andrew S.

Survey of Operating Systems, 5e 5th Edition u2013 Ebook

Understanding Operating Systems, 8th ed. 8th Edition

Exergy as a useful tool for the performance assessment

Published in the United States of America by Oxford University Press

© Oxford University Press 2019

All rights reserved. No part of this publication may be reproduced, stored in

You must not circulate this work in any other form

Library of Congress Cataloging-​in-​Publication Data

Printed by Sheridan Books, Inc., United States of America

PART I } What Nervous Systems Do for Animals

2. Organisms Without Nervous Systems 13

3. Organisms With Nervous Systems 23

PART II } Nervous Systems as Engines of Association

PART III } Evidence that Neural Systems Operate Empirically

Spectral Energy and Color 84

8. Evidence from Geometry 89

9. Evidence from Motion 104

10. Evidence from Audition 117

PART IV } Alternative Concepts of Neural Function

Neural Function as Efficient Coding 138

12. Summing Up 145

I am deeply grateful to a host of excellent collaborators over the past 20 years,

What Nervous Systems

Putting the Question in Perspective

Energy is needed to do any kind of work, whether in terms of ordinary conversation

In addition to the rapid growth of “classical genetics” in the early twentieth

prevalence of particular genes (thus Dawkins’s phrase “the selfish gene”).18,19 In

Organisms Without Nervous Systems

According to current estimates, the kingdoms of life on Earth comprise at least

As mentioned in Chapter 1, the two bacterial kingdoms (eubacteria and archae-

molecules such as antibiotics, hydrogen ions (acidity), hydroxyl ions (alkalinity),

Protists13 form a kingdom of mostly single-​celled organisms that differ from

(A) (B) (C)

Murray, 1880), in which he had a long-​standing interest.

The General Strategy

Berg HC (2004). E. coli in Motion. New York: Springer.

Eckert R (1972). Bioelectric control of ciliary activity. Science 176:473–​481.

Organisms With Nervous Systems

Definitions of the term “animals” (technically, members of the kingdom Animalia)

Defining Nervous Systems

The Emergence of Nervous Systems

(A) (B) Nerve net

Tentacle Velum Inner nerve ring

many a biology student as the coordinated beating of an isolated frog or turtle

The Emergence of Central Nervous Systems

What Do Brains Add?

(A) Leech (B) Human

D ie Patrouille des Hauptmanns Otto, die vom Hauptmann Tafel

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Library of Congress Cataloging-in-Publication Data

Protists13 form a kingdom of mostly single-celled organisms that differ from

Murray, 1880), in which he had a long-standing interest.

Eckert R (1972). Bioelectric control of ciliary activity. Science 176:473–481.