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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.
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
Suggested Reading 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
Suggested Reading 33
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
Suggested Reading 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
Suggested Reading 58
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
Suggested Reading 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
Spectral Energy and Color 84
The General Strategy 86
Conclusion 88
Suggested Reading 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
Suggested Reading 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
Suggested Reading 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
Suggested Reading 130
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
Neural Function as Efficient Coding 138
Neural Function as Computation 140
Conclusion 143
Suggested Reading 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
Suggested Reading 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 cardiovascular system conveys oxygen and nutrients to body tissues, and so on. An operating 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 information 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 computer 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).
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 empirical 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, neuronal 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 physical 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 evolutionary 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.
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 perceptual 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 emergence 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 systems 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 update 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.
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 context 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 background 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, radioactive 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 education leads to. The citizens of other developed countries do a lot better.
billion years ago.
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 gradually 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 extend 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 bacteria 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
FIGURE 1.1 Sedimentary rocks in Western Australia formed by mats of bacteria that lived several
Photograph taken by Paul Harrison (Reading, UK). Stromatolithes dans la réserve naturelle marine de Hamelin.
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 various 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 concerned 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 thermodynamics 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 material 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, oxygen. 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 evolution. 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, mechanical, 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 ecological niche they occupy.
Although inanimate objects are passively influenced by energy and change accordingly (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 sometimes 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, chapter 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 somewhere 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 receipt 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 contentious issue. Adaptation refers to the general idea that evolution is based on “fitness,” 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 advantage 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 extraordinarily 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 underlying 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 structure 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 molecular genetic revolution.15 Delbruck’s establishment of a summer course on bacterial 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 reproductive 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 difficulty” for his theory. This particular enigma was resolved by the theory of kin selection 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 advantage. 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 suspect 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 passionate 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 evidence in the fossil record of invertebrates. Although basically a refinement of neoDarwinian 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 proceeds 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 informed 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 longterm 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 species 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 overwhelming 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 archaebacteria) 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 biological 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.
chemical reactions. By using the rules of plant and animal metabolism in sometimes 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 relatively little else in the way of supramolecular machinery?
Given their role in human diseases as well as those of other animals, many bacterial species have been studied intensively, a standard choice being the common gut bacterium Escherichia coli 7 (Figure 2.1). The organism’s most obvious behavior 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 metabolism and anaerobic metabolism (i.e., fermentation) depending on the local environment.
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and arouses religious feeling. Oratorio was his special gift to the world and one never hears the name of Handel without thinking of The Messiah.
Handel seemed to reunite the forms: oratorio and opera, under his massive will. At first some of his oratorios were given in costume, showing the influence of opera.
Handel had many enemies in England, but he also had friends. Although imperious, he had a sweet side, and made friends with humble folk who loved music, even though he hobnobbed with royalty. Thomas Britton, a coal heaver, his friend, is sketched by an artist of the day in a picture where Handel is playing The Harmonious Blacksmith to Alexander Pope, the Duchess of Queensbury, Colley Cibber and other famous folk. Yet he stormed at everyone and even royalty “quaked in their boots” and were forced to behave themselves at rehearsals and concerts which Handel directed.
Accused of using someone’s melody, he answered, “That pig couldn’t use such a melody as well as I could!” He helped himself to so many that he was called the “Great Plagiarist.”
His latter life was spent quietly, with a few intimate friends, drinking his beer and smoking his beloved pipe. He was always generous and as he grew older seemed to become kindlier and softer. He contributed largely to the Foundling Asylum and even played the organ there.
He wanted to die on Good Friday, “in hopes,” he said, “of meeting his good God, his sweet Lord and Saviour on the day of his resurrection,” and on Good Friday, April 6th, 1759, he died.
C W
M O 1714–1787
Now we come to the next genius, Christoph Gluck (1714–1787) born when George Frederick Handel was twenty-nine years old. He also attacked the frivolous drift of his time, but in another field from Handel and Bach, and gave the fashionable, aimless Italian opera its death blow for all time.
Gluck’s life is different from Handel, Bach, Haydn, Mozart and Beethoven as you will see later when you have read about all of these. For, until he was almost forty years old, Gluck did nothing to make him great, whereas these other men showed from their earliest years that they were unusual.
Gluck belongs to two periods for his life bridges Bach’s and Haydn’s. You will see how he first belonged to the frivolous fashionloving composers like Hasse, Jomelli and Piccinni, and how later he blossomed into the great renewer and constructor of opera and escapes into a class of his own! His is the most remarkable instance of a man who starts with an ordinary talent, and later in life grasps a vision that never came to him in his early youth and which was not caught by others in his day.
Furthermore he was able to carry his point and not merely see the vision and let it go by. But first let us see how his life unfolded, for a man’s life helps us to understand his works.
Christoph Willibald Gluck, born July 2nd, 1714, at Weidenwang, near Nüremberg, was the son of a gamekeeper, who moved from estate to estate in the service of princes and nobles, and at the time of Christoph’s birth, was ranger to Eugene, Prince of Savoy. So, this little boy destined to become the great Chevalier von Gluck, was a child of the people even as was Haydn and others.
When he was three years old he was taken to Bohemia, (now Czecho-Slovakia), for his father entered the service of Prince Lobkowitz, a great music lover, of whom you will hear again. His parents were quite poor, yet it is remarkable that above everything else they gave Christoph a good education and at twelve he went to a Jesuit school near Eisenberg, the home of Prince Lobkowitz.
Here he learned to sing and to play the organ, the violin, the ’cello and the clavier. He was diligent and became most proficient and was loved and admired by the school fathers. But little did they dream that some day he was to write classic operas, based not on Christian stories but the pagan dramas of the Greeks!
When nearly nineteen, he left the seminary and said good-bye to the Church of St. Ignatius and went to Prague. To support himself and to carry on his scientific and musical studies he gave lessons, played for rustic festivals and earned money the best way he could, until Prince Lobkowitz became interested in him and introduced him to the musical circle at court. Here he met Count Melzi who took him to Milan, where he was taught by Giovanni Battista Sammartini, a celebrated organist and teacher of counterpoint. After four years of study he completed his musical education.
In Milan, he wrote his first opera, Artaserse which was performed in 1741. Metastasio, the popular librettist, wrote the words to Artaserse, as he did for many of Gluck’s works written in the loose style of the Italian opera. He was now twenty-eight and in the five years spent here, he composed eight operas, through which he gained great popularity. But not yet had it come to him to revolutionize opera; he simply used the old pattern which was really nothing but groups of songs, recitatives and choruses having very little connection except to give the performers the chance to do musical feats to amaze the audience with their skill. The story of these operas, meagre as it was, stopped short, for some long and elaborate cadenza, and then it went on again with no thought of the meaning of the drama but rather to tickle the taste of the audience and the performer. The orchestra, too, was a step-child, for no one
cared where it came in as long as it was politely subdued, keeping the singers on the key, and doing its best to be heard only when bidden. So, Gluck followed these ideas in the beginning and perhaps it was better that he did, otherwise he might never have realized how far opera had strayed from the ideals of Monteverde.
Having eight operas to his credit, he began to get commissions from other cities and countries, and next accepted an invitation, in 1745, to go to London as composer of opera at the Haymarket Theatre. In 1746 he wrote La Caduta de Giganti (The Fall of the Giants), with no doubt a libretto of Metastasio’s, then he gave his Artamene and was assisted in their production by Handel, who is supposed to have treated the works with contempt. He is said to have exclaimed, “Even my shoe-black can write better counterpoint than Gluck.” But we must remember that Gluck had not yet become the great Gluck. His visit to England was fruitful, for Gluck heard and digested the great oratorios of Handel, and realized that the voice and orchestra might be handled the same way in opera. No doubt his mission was beginning to dawn on him; it came, not as a great revelation, but gradually.
Another thing that gave him a push forward and shows how great people can make a success of failure: he was asked to write a pasticcio (Italian word meaning a meat-pie), or a string of melodies, very fashionable in his day. He strung together his best airs from his Italian operas, and called it Pyramus and Thisbe, but it was a dismal failure. “Ah, ha!” he must have thought, “why shouldn’t this musical drivel fail, for it is naught but trash, and with nothing that is needed to make a good literary drama.” So this was one of the experiences that led him to reform opera, making the words fit the music and not stopping a performance, so that a popular soloist could sing a meaningless trill and then start again with the other part of the word, —the way that opera was being written at that time.
After his London ups and downs he went to Paris and heard the operas of Rameau. He realized now the value of musical declamation and recitative to the meaning and action of opera if used with thought, and he was not slow in taking suggestions.
Gluck was probably the most all round man of his day, for he knew literature and science as did few musicians. He knew all the influential people in the arts, sciences, and music in London, Hamburg, Dresden and Vienna, and his home was a center of learned and delightful people. When in Vienna but a short time, he was commissioned to write an opera and he produced, with success, La Semiramide, after which he went to Copenhagen. His next opera Telemacco in which he began to work out his new ideas was well received, in Rome and Naples.
In 1750 after many disappointments, he was married to a lady he had long adored. They lived happily together, for Marriane Pergin not only brought him money which was a great joy, but was always his devoted and understanding help-mate. She was an accomplished woman, and a companion that many might envy. But, sad to say, they had no children, so they adopted a niece of Christoph’s, a lovely little girl with great musical talent. The three lived lovingly together until the poor little child sickened and died, making the Glucks most
unhappy, for they adored her, as is often the case, even more than if she had been their own child.
In 1751 Gluck journeyed to Naples. Didn’t he travel a lot in the days of the stage coach and brigands! In the same year he became conductor to Prince Frederick at Vienna and in 1754 was officially attached to the opera, and Maria Theresa made him court chapel master.
Soon after, the Pope, pleased with what he had done in Rome, made him Chevalier of the Golden Spur and from that time he always styled himself Ritter (Chevalier) von Gluck.
In Il re pastore (The Shepherd King), we see the dawning of Gluck’s best period of writing (1756). The overture is better music than he had written before, and from this time on, Gluck became the genius in the opera world for which he is known. From 1756 to 1760 he lived apart from the world studying and after this he began to broadcast his ideas in writing and composing.
When the Archduke Joseph of Austria, afterwards the Emperor, married Isabella of Bourbon, Gluck wrote Tetede which was performed with great pomp. After this he wrote the ballet Don Giovanni, or The Libertine, particularly interesting, for it certainly gave Mozart an idea for his own great work Don Giovanni.
Again our “wandering minstrel” moved, this time to Bologna where he conducted a new opera which, strange to say, showed not a sign of his new ideas!
“O E ” B
Soon he met Calzabigi, another librettist, with whom he wrote his first epoch-making opera Orpheus and Euridice. Although in some parts it is written like the older operas, he used many of his new ideas. The public at first were bewildered but they liked it. The next opera written with his new librettist was Alceste, so different was it, and so full of his best thought that the public did not like it. The pleasure-loving people went to be amused and heard music almost as serious as oratorio. It was austere, and its climax was not satisfactory. Yet it and Orpheus and Euridice mark the birth of music drama which Mozart and Wagner developed further.
In Orpheus and Euridice the chorus was an important part of the drama as it had been in the old Greek drama from which Gluck took many of his stories; and was not something dragged in to fill up space. Instead, too, of the over-embroidered arias they were simple and expressive, and the characters were real living beings, instead of figures on which to drape showy melodies. Naturally, the composers were jealous of him and went so far as to say that the principal singer had written Orpheus and Euridice.
Gluck said of his Alceste: “I seek to put music to its true purpose; that is, to support the poem, and thus to strengthen the expression of the feelings and the interest of the situation without interrupting the action.... In short, I have striven to abolish all those bad habits which sound reasoning and true taste have been struggling against now for so long in vain.” He abolished the unnecessary cadenza, a fancy, trilly part composed by the soloist himself and used just before the close of a piece. You will see in a later chapter how Beethoven dealt with it.
Happily Gluck and Calzabigi still continued working together and in 1770 he wrote Paride and Elena (Paris and Helen) which proved Gluck to be a writer of beautiful romantic song.
By now Vienna and Paris were enthusiastic about him, yet he was severely criticized because he dared to write and compose differently from everyone else. The adventurer into new paths must always expect trouble from those who have not caught up with him.
Now our traveler goes to Paris where he presents Iphigenia in Aulis. The story was taken from a play of the French dramatist Racine. Although this was the fourth work in Gluck’s new style it was not as good as the others. His enemies did their utmost to hurt him as they resented his coming into Paris to reform French opera. And as the musicians and singers were not good artists, it was almost impossible to give it well, and probably it would never have reached the stage had it not been for Marie Antoinette the French Queen who was later guillotined. She had been a real friend and pupil of Gluck, when a young princess in Vienna. Nevertheless the opera pleased its audiences, and it paid well, and Gluck was given a new court office in Vienna.
In 1776 the trouble that had been brewing with Gluck’s opponents came to a climax. Piccinni was his great Italian rival and the city of Paris was torn as to who was the better composer. All the literary men and the court were divided into factions, one for and one against Gluck. Some great men, including Jean Jacques Rousseau were Gluckists, while others of importance were Piccinnists. Never had there been so great a contention for musical glory or struggle against new ideas. It was a most extraordinary thing, but it does show that there was great musical interest or people would never have wasted so much time in argument and in writing for or against these men. Finally it came to a head, and it was decided to give them both the same libretto of Iphigenia in Tauris to see who could write the better opera. Gluck completed his within the year and after nearly three years, Piccinni finished his. They were both performed and needless to say Gluck won the award and even Piccinni said himself that Gluck’s was the better. It is nice to know that after Gluck’s death, Piccinni tried to collect funds to raise a memorial as a tribute to him!
So artistic rivalry need not dim admiration.
In Iphigenia in Tauris again the master rises to great heights. His overture was splendid, his orchestral color was superb. He pictured the different characteristics of the various groups of people and of
the individuals themselves in word and music as it never had been done before.
He wrote Armide in 1777. It did not succeed although it was very lovely and dreamy and in it, he suggested the sounds of babbling brooks and the song of the nightingales.
Gluck wrote thirty operas, seven of which are in his new style: Don Giovanni, Orpheus and Euridice, Paris and Helen, Alceste, Iphigenia in Tauris, Iphigenia in Aulis and Armide.
P
And thus this great path-breaker advanced opera seria (grand opera).
The old sinfonia in three movements which opened the opera, disappeared, and instead came the introduction or overture, suggesting the opera itself. He taught and wrote that composers could do anything to assist the action of the opera; he elevated the story to an important place; the characters in the plot were thought of as people and not as puppets, and they were studied individually and not as machinery only. The situations in the story governed the kind of music he used and he tried hard to make the orchestra a main part of the opera. It seems odd that nobody had thought of this before. Yet you have seen how much time had been given to the voice throughout the ages, and how long it had taken instruments to arrive at their full importance. So we see Gluck improving as he worked with a better librettist. From now the opera writer had to use thought in composition, as he would in writing a play.
But Gluck had trouble with the singers on account of his innovations. He was the crossest conductor of his time, would allow no one to dictate to him, and scolded the singers as they had never been scolded before.
He must have looked droll conducting, for he used to take off his wig during rehearsals, and wrap a cloth about his head to keep the draughts from fanning him! He would rage if the singers tried to do what they had been permitted to do in other operas! Some singers demanded extra pay when Gluck conducted. Sometimes he would repeat a passage twenty or thirty times and no pianissimo was soft enough and no fortissimo loud enough! Someone said of him while he was conducting, “He lives and dies with his heroines, he rages with Achilles, weeps with Iphigenia and in the dying scene of Alceste throws himself back in his chair and becomes as a corpse.”
Otherwise he was always the kind soul who attracted everybody from Marie Antoinette down. She used to receive him in her boudoir so that they could enjoy conversation without court formalities.
One day two prima donnas refused to obey him when rehearsing Iphigenia, and he said: “Mesdemoiselles, I have been summoned here to Paris especially to produce Iphigenia. If you sing, well and good, but if not, that is your business; only I shall then seek an audience with the Queen, and inform her that the opera cannot be performed, and I shall put myself into my carriage and straightway leave for Vienna.” You may know that the ladies did their best!
In closing let us tell you what Berlioz, a master of orchestration, said of Gluck’s orchestration in Alceste: “Of its kind I know nothing more dramatic, nothing more terror-inspiring.” And this was said of a man who had only the simplest orchestra with which to work. After much fighting, he was the first to introduce into the orchestra the kettle-drums and cymbals, which moderns have used with grandeur.
Gluck lived to see his own success, but the Piccinni strife and the jealousies may have weakened his constitution, for he died rather suddenly in 1787, a few weeks after the first performance of Mozart’s Don Giovanni.
There are many memorials in Europe to Gluck, not the least being his bust which stands beside Lully and Rameau in the Grand Opera of Paris.
P C
It is very hard to realize that time was when there were no public concerts. Music was confined for so many centuries to the churches, to the public squares, to the King’s Chamber, or to the ball rooms of wealthy nobles, that it had not become the democratic art that it is now. Of course the first opera houses in Italy had been steps in the direction of bringing music to the people. The concerts begun by the Danish organist, Buxtehude, in Lübeck about 1673, and the Tonkünstler-societät in Vienna of the same period were the first public concerts. In England, John Banister started concerts at about the same time, which were the first to admit an audience by payment of a fee. Handel’s friend, Thomas Britton, the coal-heaver, gave concerts at his home for 10 shillings the series!
The 18th century saw a great development in giving public concerts. In France, the Concerts Spirituels were begun in 1725. The object of these were to give music to the people on the days of religious festivals when the opera house was closed. There were about 24 concerts a year; the political events of 1791 put an end to the society but it had already given the people a taste for concerts, and many new societies grew out of it. The festivals of Three Choirs in West England (see page 190) were founded in 1724, and the Academy of Ancient Music in 1710. The Musikverein in Leipsic was founded in 1743 and was later turned into the famous Gewandhaus concerts in 1781.
This movement for public concerts went hand in hand with the development of instruments and the perfecting of performers. In fact the word concert came from “consort—the union or symphony of various instruments playing in concert to one tune.”
T M S
The symphony came to life in Germany. Paul Landormy in his History of Music tells us that it was the time of the “poor scholars” who were educated free from expense in the schools with the understanding that they were to learn the “musician’s trade” and take part in the concerts organized by the cities and the courts. Thus symphony orchestras grew up all over Germany,—Munich, Stuttgart, Dresden, Darmstadt, Hamburg where Telemann conducted, in Leipsic, Berlin and Mannheim.
In Mannheim appeared the most important group of composers, known as the Mannheim School, and many wrote the early symphonies which led from the works of Bach to those of Haydn and Mozart. The best known of these composers are: Johann Stamitz (1717–1757), Franz Xavier Richter (1709–1789), Anton Filtz, Christian Cannabich, Ignaz Holzbauer, Ernst Eichner and Giovanni Battista Toeschi. Under the direct influence of the Mannheim School were: François Joseph Gossec (1734–1829), a Belgian living in Paris who wrote many symphonies; Luigi Boccherini (1743–1805) known as one of the first writers of chamber music in the form used by the classic writers; Giovanni Battista Sammartini (1701–1775) of Milan; the sons of Bach, Karl Ditters von Dittersdorf, and Joseph and Michael Haydn.
From the painting by J. B. Greuze, in the Louvre, Paris.
Chevalier Christoph Willibald von Gluck. Father of Modern Opera.
From a statue by Barrias, in the Luxembourg Gallery, Paris.
The Boy Mozart.
CHAPTER XX
“Papa” Haydn and Mozart—the Genius
F J H 1732–1809
About the time in history when Franz Joseph Haydn was born, the world was very much upset. No one knew what to think or how. It was a time of battle and struggle as he was born in the midst of the Seven Years’ War and lived during the French Revolution. Everyone except for a few great persons felt bitter and discontented and doubt was everywhere. This seems to be the way wars and conflicts affect all peoples and it is why wars are so damaging.
Yet out of this mixture of feeling and thinking, the great classic period of music was created by such men as Bach and Haydn and Mozart and the finishing touches were put on it by Beethoven, the colossus.
Franz Joseph Haydn was born in Rohrau (1732), a little town in Austria near Vienna. His father was a wheelwright and his mother was a very good cook. Beethoven’s mother, too, was a professional cook.
These simple parents, his brothers and sisters, measuring not a baker’s, but a wheelwright’s dozen, had an hour or two of music every evening after the hard day’s work, and Mathias, the father, played the harp and sang. It was during these evenings that little Joseph’s father noticed that at the age of six he was passionately fond of music.
One time at a festival the drummer failed to appear and there was no one who could play for the choristers who were to march through the town. His teacher, Frankh, called Joseph and showed him how to make the drum stroke and told him to practice it. When he was left by himself he found a meal tub, over which he stretched a cloth, put it on a stool and drummed with such vigor that the whole thing toppled over and he and his drum were covered with meal! But he learned to drum! And the people laughed when in this solemn church festival, the little six year old Joseph was seen drumming the big
drum carried by a hunchback in front of him. The drums on which he played are still at Hainburg. But, we forget, we have not brought him from Rohrau!
Not long before J. M. Frankh, a relative, came to visit the Haydns, and it was decided that he should take Joseph to Hainburg to teach him. The excitement, of course, was great and little Joseph felt very important with all the hustle and bustle preparing for his departure. Little did Saperle (his nickname) realize what a hard master he was getting in Frankh, who only cared for the pay he received from Joseph’s father. Nevertheless he learned much and showed great talent while at Hainburg and one day a great thing happened. Reutter, the organist of St. Stephen’s in Vienna, visited Frankh and as they talked of music the conversation turned to the choir school which Reutter directed. Frankh sent for Joseph, a slight, dark haired, dark eyed little boy, and Reutter asked him to read a piece of music at sight. Joseph looked at it and said: “How can I, when my teacher couldn’t?” Yet, Joseph did sing it sweetly and he entered the choir school. Here his life was a misery, for Reutter was harsh and unsympathetic, but soon Joseph’s hard life in the choir school was over, for one very cold winter night, he felt a little frisky, as many a healthy lad does, and pulled off the wig of a man in the choir. Reutter, who had wanted an excuse to rid himself of Joseph, because his voice had begun to break, threw him out into the cold. Poor Saperle had no other place to go and wandered about all night, until he met his acquaintance Spangler, a tenor who was very poor and so had sympathy with Haydn. He took him home to live with him and his wife and child in his attic,—one small room with no comfort and no privacy. All this time young Haydn was forced to earn his daily bread by teaching as much as he could, playing for weddings, baptisms, funerals, festivals, dances and street serenadings. This street serenading was a sweet and pretty custom of the time.
One night Haydn and some other youths serenaded Kurz, a prominent comedian. Kurz, pleased by the music below his window, called to the lads: “Whose music is that?” “Joseph Haydn’s,” called back Haydn. “Who is he and where?” asked Kurz. “Down here, I am Haydn,” said Joseph. Kurz invited him upstairs and Haydn, at the age of seventeen, received a commission for a comic opera, which had two special performances.
All this time he mixed with the poor and laboring people, and their songs became his songs, and his heart was full of their frolics and their pains. He was of the people and was so filled with their humor that later he was called the father of humor in music.
Soon, in order to be alone, and to work in peace, he took a room in another attic, and bade good-bye to his very good friends. His room was cold in winter and let in the rains and snows, but it did have a spinet on which Haydn was allowed to play, and fortunately Metastasio the librettist lived in this house. Here Haydn studied the works of Karl Philip Emanuel Bach, Fuchs’ Gradus ad Parnassum (Steps to Parnassus, Parnassus meaning the mountain upon which the Greek Muses lived and so comes to mean the home of learning). He practised too, during this time, on any instrument he could find and learned so much that he became the founder of the modern orchestra.
When Metastasio discovered that there was a hard working musician in his house he met him and then introduced him to Porpora the greatest Italian singing teacher in Vienna. Not long after meeting him, Porpora entrusted to his care Marianne Van Martines, his ten year old pupil, the future musical celebrity. At seventeen Marianne wrote a mass which was used at St. Michael’s Church and she became the favorite singer and player of Empress Maria Theresa. You see women even in those days composed and performed!
So began Haydn’s successes. Porpora engaged him as accompanist, and treated him half way between a valet and a musician, but Haydn’s sweet nature carried him through all unpleasantnesses and he was so anxious to learn and to earn his six ducats that he did not care if he did have to eat with the servants.
In 1751–2, he wrote his first mass, his first string quartet, and his first comic opera for Kurz, The Crooked Devil, the music of which has been lost. Soon after he met Gluck at the concerts of the Prince of Hildburghausen, where Haydn acted as accompanist; at the prince’s house too, he met Ditter von Dittersdorf, the violinist. The princes and nobles of these days did much for music for it was usually at their homes and under their guidance that the composers received opportunities to work.
Nevertheless, we see Haydn during these days slaving to make his daily bread, but with the money he made he bought books on music
theory and held himself sternly down to hard work, morning, noon, and night.
In 1755 Baron von Fürnburg, a music amateur, who gave concerts at his home, asked him to compose for him, and he wrote eighteen quartets, six scherzandi for wind instruments (the ancestors of his own symphonies), four string quartets, to be played by the village priest, himself, the steward, and the ’cellist Albrechtsberger.
All these pieces show how much happier he was since becoming part of the Baron’s staff, for they are merry and jolly, and filled with that humor which Haydn was the first to put into music.
Here, too, he met the cultivated Countess Thun, who was so interested in his struggle for success, and in the youth himself that she became his pupil. From this time on he began to earn more and to live more comfortably.
Everything seemed to be clearing up for him now. The Countess introduced him to Count Morzin, a Bohemian nobleman of great wealth, and in 1759 he became his musical director. His orchestra had eighteen members and here he wrote his first Symphony (the first of one hundred and twenty-five!)
All this time he kept up his teaching and very soon married the daughter of a wig-maker, who did not understand him and with whom he was very unhappy, but he lived with her like the good man he was until within a few years of his death.
