from The Textbook Letter, January-February 1999

Reviewing a science book for high-school honors courses

Fourth edition, 1996. 1206 pages + appendices. ISBN: 0-8053-1957-3.
The Benjamin/Cummings Publishing Company, 2725 Sand Hill Road,
Menlo Park, California 94025.

This Impressive Textbook
Is Strongly Recommended

William Z. Lidicker, Jr.

The fourth edition of Neil A. Campbell's Biology is a most impressive document. It is an introductory college text, but it can be recommended strongly for use in advanced-placement high-school courses and as a reference book for all high-school biology teachers. The writing is clear, scholarly and mature. Technical terms are introduced in ways that render them welcome, not abhorrent, and Campbell often helps to make new terms comprehensible by explaining their etymology and by putting them into a historical or personal context.

In various cases, Campbell anticipates misconceptions that students sometimes bring with them into the classroom, as well as questions that can arise in the students' minds as they read. For example: In his exposition of evolution, he points out that the fact of evolution is logically separable from any putative mechanism of evolution. He thus refutes the common misperception -- so often fostered by inferior middle-school and high-school books -- that evolution and natural selection are the same thing. He also calls attention to the difference between maladaptive and nonadaptive changes (page 421), and he refutes the popular misconception that evolution leads to perfection (page 433).

It may be hard to imagine that a 6.5-pound book filled with detailed material can be a joy to read, but this one is.

Chapter 1 of Biology is an introduction in which Campbell sets forth twelve themes that will recur throughout the book:

Life is organized on many structural levels

Each level of biological organization has emergent properties

Cells are an organism's basic units of structure and function

The continuity of life is based on heritable information in the form of DNA

A feeling for organisms enriches the study of life

Structure and function are correlated at all levels of biological organization

Organisms are open systems that interact continuously with their environments

Diversity and unity are the dual faces of life on Earth

Evolution is the core theme of biology

Science as a process of inquiry often involves hypothetico-deductive thinking

Science and technology are functions of society

Biology is a multidisciplinary adventure

After the introduction, Campbell provides 49 chapters that are grouped into eight units: "The Chemistry of Life," "The Cell," "The Gene," "Mechanisms of Evolution," "The Evolutionary History of Biological Diversity," "Plants: Form and Function," "Animals: Form and Function" and "Ecology."

When it isn't feasible to cover the entire book, a high-school teacher will be able to build a powerful course by using only chapter 1 plus two or three chapters from each of the eight units. This is appropriate because Campbell's recurrent themes provide coherence and connections among many discrete topics.

Having declared that "Evolution is the core theme of biology," Campbell succeeds in making it the core theme of his book. Moreover, he does this with skill. Read, for example, the opening lines of his chapter 37 ("Animal Nutrition") and see how he unobtrusively puts the entire subject of nutrition into an evolutionary context.


Another virtue of Biology is that it is surprisingly current. As examples of passages that signal the book's currency, I cite Campbell's recognition that there are several alternative concepts of what a "species" is (on pages 447-449), his discussion of endothermy in insects and in fishes (pages 901-903), his up-to-date phylogeny of bears (page 470), his reference to cloacal respiration in turtles (page 643), and his excellent treatment of contemporary investigations into the origins and diversification of major groups of organisms (see pages 494-495, page 523, and pages 541-543).

Of course, no book can treat every new bubble of intellectual activity in the biological sciences, but I am a little disappointed to find that there is no explicit description of the recent explosive developments in conservation biology and in landscape ecology. (Landscape ecology is the study of the interactions between different communities at places where they meet -- e.g., where a cornfield meets a forest, or a forest meets a marsh.) Still, Campbell's overall treatment of ecology is superior work. It is refreshing to find that he covers environmental issues at various places in the book, including all the chapters in Unit 8. He does not isolate them, as many other authors do, in an afterthought chapter about "human ecology."

Campbell is particularly successful in his efforts to portray science as a continuing human adventure. He begins each of his eight units by presenting a lengthy interview with a research biologist, and he often discusses the historical development of important concepts. (His account of Mendel's contributions, in chapter 13, is especially well done and informative.) Even more impressive are his reminders that biology is rich in unanswered questions, a point that he underscores with phrases such as "Lively debate continues" or "Currently, biologists think that . . . ."

To his credit, Campbell doesn't attempt to avoid the philosophically difficult issue of reductionism versus holism. Instead he addresses it with understanding, and links it to the theme of emergent properties, in his opening chapter:

Because the properties of life emerge from complex organization, scientists seeking to understand biological processes confront a dilemma. One horn of the dilemma is that we cannot fully explain a higher level of order by breaking it down into its parts. A dissected animal no longer functions; a cell dismantled to its chemical ingredients is no longer a cell. According to a principle known as holism, disrupting a living system interferes with the meaningful explanation of its processes. The other horn of the dilemma is the futility of trying to analyze something as complex as an organism or a cell without taking it apart. Reductionism -- reducing complex systems to simpler components that are more manageable to study -- is a powerful strategy in biology. For example, by studying the molecular structure of a substance called DNA that had been extracted from cells, James Watson and Francis Crick deduced, in 1953, how this molecule could serve as the chemical basis of inheritance. The central role of DNA was better understood, however, when it was possible to study its interactions with other substances in the cell. Biology balances the pragmatic reductionist strategy with the longer-range objective of understanding how the parts of cells and organisms are functionally integrated. [page 4]

The concept of emergent properties in the chemistry of life makes an impressive reappearance in chapter 6, "An Introduction to Metabolism." This chapter also has an excellent treatment of thermodynamics -- a topic which has fundamental importance in biology but which is badly muddled (or ignored altogether) in most introductory textbooks.

Like all the biology books that we see these days, this one is elaborately illustrated, but the illustrations in Biology are not mere adornments. They have been designed to be informative and explanatory. Figure 12.2, for example, is clever, absorbing and educational at multiple levels. Four small photographs show two sets of human parents, while four more photographs show sons and daughters that those parents have produced -- and now the student is asked to match the parents to their respective offspring. In trying to do so, the student confronts several aspects of heredity and sexual reproduction.

Another notable illustration is figure 4.8. Here, structural diagrams of the sex hormones estradiol and testosterone are flanked by photos in which male and female wood ducks display their striking sexual dimorphism. Here again there are multiple lessons, the most obvious of which is: Subtle differences in molecular architecture can produce profound differences in development, physiology and behavior.


In this excellent book, outright errors are so infrequent that it seems disrespectful to point them out. I must mention some of them, however, in keeping with my duties as a reviewer:

Bigger Issues

Looking beyond omissions and factual errors, I find that I have a few disagreements with Campbell over difficult ecological issues. As examples:

While focusing on ecological matters, I want to add that Campbell's chapter on population biology (chapter 47) is, overall, remarkably good. He includes immigration and emigration in the population-growth equation, he makes a clear and correct distinction between r (a population's actual per-capita rate of growth) and rmax (the population's potential rate of growth under ideal conditions), he recognizes the irony in applying the name "life table" to what is really a tally of deaths, he offers an excellent treatment of human population growth, and he recognizes that density regulation, in most situations, depends on multiple factors.


Some other introductory biology texts are good, but this one is exceptional. It combines comprehensiveness with currency, obvious scholarship, and literary achievement. Did I mention the short but wonderful chapter about water, the solvent of life? Or the five-page mini-course on climatology (in chapter 46)? Or the succinct, intelligent essay about the substrates of selection (in chapter 21)? Whether as a classroom text or as a teacher's reference book, Campbell's Biology is adaptable to all levels of instruction. You will be glad that you chose it.

In This Worthy Textbook,
Nothing Is Dumbed Down

Michael T. Ghiselin

A striking feature of this fourth edition of Neil A. Campbell's Biology is the series of "Interview" articles in which Campbell talks with outstanding biologists. There are eight such articles, occupying two to four pages each, and Campbell has done a good job of selecting his interviewees: Eloy Rodriguez (a plant chemist who specializes in studies of secondary metabolites), Shinya Inoué (a cell biologist who has conceived major innovations in microscopy), David Satcher (a medical geneticist who was the director of the federal Centers for Disease Control and Prevention when Campbell interviewed him, and who is now the Surgeon General of the United States), John Maynard Smith (an evolutionary biologist who is particularly interested in the evolution of sex), Edward O. Wilson (who studies social organization in animals and is a prominent advocate for the preservation of biodiversity), Adrienne Clark (the botanist who directs Australia's Commonwealth Scientific and Industrial Research Organization), Patricia Churchland (a philosopher and neuroscientist who tries to interpret the human mind in terms of scientific findings about the human brain), and Margaret Davis (a forest ecologist who uses fossil pollen to learn about the biogeography of trees in prehistoric times).

Such role models are appropriate here, for Biology should be regarded as a book for students who are interested in biology as a career and who need a solid knowledge of the fundamentals before they undertake specialized courses in college. This is a worthy text in which nothing is dumbed down and nothing is glossed over as too difficult.

Is the book suitable for high-school students in an honors course or an advanced-placement course? The answer depends in part upon the students. Campbell has put a great deal of biology into Biology, and some students -- even though they are bright, diligent and qualified to take an honors course -- may feel overwhelmed by it.

Wherever I have checked Campbell's material in detail, I have found it to be unusually accurate. There are few errors, and I have not encountered any egregious blunders of the sort that so often occur in inferior books. (Just to prove that I have looked carefully, let me describe one of the few mistakes that caught my eye: In figure 30.2, on page 629, the artist has added an imaginary piece to the gut of a larval tunicate, making the gut extend through the entire length of the animal's tail.)

In general, the mistakes and missteps in this book arise not because Campbell has got things wrong but because he hasn't got things quite right. His historical material, for example, includes a common anachronism: He makes Gregor Mendel's work look more like modern genetics than it really was.

One of the great merits of Biology is Campbell's effort to tie things together by using a few major themes. This helps the reader to strive for something more than the brute memorization that can make biology courses deadly. Evolution is definitely the central theme of the whole book, and Campbell repeatedly draws attention to the utility of evolutionary thinking in making sense of otherwise unintelligible phenomena.

Teaching About Classification

One sector of biology that has become more militantly evolutionary in recent years is systematics, the science of classification. Our classifications are becoming increasingly genealogical, as Darwin predicted they would, and they are increasingly based on inferences about common ancestry. Campbell endorses this trend, but he occasionally runs into trouble when he tries to explain the technicalities. I suppose that we can forgive him for treating homology and analogy as two kinds of "similarity" (page 472), since nearly everyone else makes the same mistake -- but it is indeed a mistake. Homology and analogy are two kinds of correspondence, and it is important to recognize that similarity and correspondence are different phenomena. Let me illustrate this difference by considering the malleus and the incus, two tiny bones in the mammalian inner ear. The malleus and the incus have arisen from, and hence are homologous to, the articular bone and the quadrate bone in the jaws of the mammal-like reptiles: The malleus is a modified articular, and the incus is a modified quadrate. The modifications have been so extreme, however, that the original bones have been transformed beyond recognition. Even though the malleus and the articular display the kind of correspondence that we call homology, they don't display any discernible similarity -- and the same is true of the incus and the quadrate. As Campbell himself points out, the relationship between the malleus and the articular, or between the incus and the quadrate, becomes obvious only when one examines fossils that show the intermediate stages.

Similarity can be a rather tricky concept, and judgments about similarity are often highly subjective. This leads to the question of how and whether similarity is useful in tracing genealogies and in recognizing common ancestries. Systematists are not of one mind about how we should go about the task of classifying organisms, and the systematists' controversies can be confusing to outsiders. Campbell provides an adequate treatment of these controversies and tries to explain three major schools of systematic thought. Unfortunately, however, he calls the three schools "phenetics," "cladistics" and "classical evolutionary systematics." This nomenclature isn't quite right.

Phenetics isn't a kind of systematics. Phenetics is the study of similarity. It is a field of scientific inquiry, like genetics or embryology. When Campbell says "phenetics," he actually is referring to pheneticism, a school of systematics which had its heyday during the 1960s and which revolved around the doctrine that classifications should be based strictly upon observed similarities.

Cladistics, too, is a field of study -- the study of biological lineages or genealogies. When Campbell says "cladistics," he really is referring to cladism, a style of systematics in which classification is strictly genealogical.

As for "classical evolutionary systematics": This is a school of systematics that represents a sort of middle position between pheneticism and cladism. Its practitioners incorporate both phenetic and cladistic criteria into decisions about classification.

The classification system that usually is taught to beginning students presents a real problem because it is based on a mixture of cladistic criteria, phenetic criteria, and old traditions. One of these traditions is the notion that we should divide the animals into two equivalent groups called the vertebrates and the invertebrates. This fancy, which goes back to Lamarck, is rather like dividing the residents of the United States into two groups called "New Yorkers" and "others." The invertebrates are the "others" that remain after we subtract the vertebrates -- the group to which we ourselves happen to belong -- from the rest of the animals. They don't have any common feature that is scientifically interesting or meaningful. Although Campbell emphasizes that more than 95% of all the animal species are invertebrates, he adheres to the equal-space tradition that textbook-writers have always followed: one chapter for the invertebrates, then one chapter for the vertebrates.

In his chapter about the invertebrates, Campbell makes a heroic effort to get away from the usual, dull listing of taxonomic groups, and he tries to give a précis of the comparative anatomy and comparative embryology that underlie the classification of animals. This leaves him little room for telling about the animals' other properties or their ecological roles, and his attempt to deal with embryology is oversimplified to the point of outright error. In particular, he confuses the distinction between the schizocoela and the enterocoela with the distinction between protostomes and deuterostomes. Let me explain this:

In some animals, the body cavity is a true coelom, i.e., a cavity lined with an epithelial layer made of mesoderm. These animals are called the coelomates, and they are divided into two groups -- the schizocoela and the enterocoela -- according to how the coelom arises during embryogeny. In the schizocoela the coelom forms as a split in the mesoderm, but in the enterocoela the coelom forms as an evagination of the gut. Campbell recognizes that the distinction between the schizocoela and the entercoela is fundamental in classification, but he muddles it with another distinction that is equally important, i.e., the distinction between protostomes and deuterostomes. (This latter distinction, based on how the mouth originates during embryogeny, is so powerful that the deuterostomes -- which comprise the phyla Chordata, Echinodermata and Hemichordata -- are viewed as a separate branch of the animal kingdom.) In trying to make things simple, Campbell treats the protostomes as if they all were schizocoela, but this is just wrong. Many protostome groups do not even have a true coelom.

When he focuses on our own phylum, the Chordata, Campbell rightly cites three diagnostic characters -- the notochord, the dorsal, hollow nerve cord, and the postanal tail. But then he adds a fourth character: the presence of pharyngeal slits. This is a mistake, for pharyngeal slits aren't unique to the Chordata. Pharyngeal slits are also present in the Hemichordata, a phylum that isn't even mentioned in Campbell's book.

All in all, the chapter on vertebrates is more satisfactory than the one about invertebrates: It does not concentrate so heavily on classification, and it gives more information about form and function. For example, Campbell explains how vertebrate jaws have arisen through modification of earlier structures. But even so, this book -- like so many others -- perpetuates the impression that the essential reason for studying biology is the need to understand humans.

In his efforts to describe biodiversity, Campbell is more effective when he deals with the plants and the fungi than when he deals with animals, because he conveys a better appreciation of the various roles that the plants and fungi play in the economy of nature. If he wanted to present the animals in a comparable way, he would have to add another chapter or two about the invertebrates, emphasizing how these immensely diverse animals live their lives and how they fit into the natural world.

William Z. Lidicker, Jr., is a professor emeritus in the Department of Integrative Biology at the University of California at Berkeley, a curator emeritus of mammals in that institution's Museum of Vertebrate Zoology, and a fellow of the California Academy of Sciences. His research deals with the evolution, ecology and conservation biology of vertebrates.

Michael T. Ghiselin is a biologist, a senior research fellow at the California Academy of Sciences, and chairman of the Academy's Center for the History and Philosophy of Science. His research has emphasized comparative anatomy and the evolution of modes of reproduction. His books include The Triumph of the Darwinian Method and The Economy of Nature and the Evolution of Sex.


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