Evolutionary Biology

Charles Darwin III: Descent with modification

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Did you know that Charles Darwin preferred the phrase descent with modification over the simpler term evolution? In his groundbreaking book On the Origin of Species, Darwin chose his words very carefully. "Evolution" was used in different ways at the time, and Darwin wanted to convey the important concept that life forms descended from a common ancestor.

The first edition of Charles Darwin's groundbreaking book, On the Origin of Species by Means of Natural Selection, had only one illustration in it – a picture of a family tree, or descent, also called phylogeny. For the book publisher, this must have been an expensive investment and a somewhat worrying choice. The diagram was printed on oversized paper that had to be unfolded out of the volume to be seen, an expensive printing task. In addition, to have a single drawing in a book was unusual in the middle 1800s because realistic illustrations of plants and animals were considered to be highly artistic. Illustrations were an important selling point of popular books about natural history such as Origin, a non-technical work written for the general public to read. Yet, all 1,250 copies of Origin sold in a day. For Darwin to have placed only a single picture in the book, he must have considered it crucial to his discussion.

Figure 1: This was the only illustration that appeared in Origin of Species.

Darwin's single picture was a chart, not a portrait of exotic species or even a map. With it, Darwin sought to explain a new concept in science: how various pieces of biology fit together to explain the origins and evolution of species. This involved many details and a lot of ideas. His one drawing was meant to illustrate all of the following:

  • How natural selection works over generations to promote structural variation in the physical form or behavior of an organism.
  • How these variations accumulate to change a species over time.
  • How populations within a species tend to become different from one another.
  • How structural change eventually produces new species.
  • How several species can arise over time from a single ancestral species.
  • How a new genus can evolve from a line of new species.
  • How extinction is a natural part of the evolutionary process.
  • How all species are actually related to one another.
  • How clusters of similar species can form because they have a common origin.

Evolution is a complex, multi-faceted process, and this list is a complex set of ideas to relate. So it is no wonder Darwin focused on producing a graphic to help explain them to the world. He also wrote about five pages explaining how the diagram was to be read.

Darwin chose his words carefully. Here and elsewhere in Origin he used a certain phrase – "descent with modification" – over and over again as an expression for "evolution". Why this instead of the simple one-word term "evolution"?

Part of the answer must be that evolution was a still a fuzzy concept, and it was Darwin's job to make it clear.

"Evolution" vs. "Descent with modification"

At that time, scientists were commonly using the term evolution in discussing physical growth, the changes an individual goes through as one matures. The other meaning of evolution referred to structural change in a species that took place over time, which some, including Darwin, also called transmutation. So, there was a reason for his preference for "descent with modification" over "evolution". First, he wanted to make clear that his discussion of evolution dealt with transmutation (modification), not growth and development. Second, Darwin meant to emphasize that the big picture of biological evolution was far more complicated than the image of the fur of a fox transmuting from reddish brown to white as an adaptation to life in the Arctic. It also involved the production of a pedigree that linked species because they are genetically related, through the process of descent.

Figure 2: The beaks of four species of Galapagos finches, from Darwin's Journal of Researches, 1839. Darwin found that the beaks of finches on islands throughout the Galapagos were specialized to optimize the diet available to them. Thus, finches on islands where large, hard-shelled nuts were prevalent developed robust beaks (far left), and finches on islands where insects or flowers were available developed delicate, pointy beaks (far right).

The concept of phylogenetic descent was a new idea that made Darwin's theories of evolution more sensible than previous proposals that tried to explain certain observations. For many decades before Origin appeared, natural scientists had wrestled with a puzzling problem of biodiversity. While taxonomists who classified organisms never intended to find patterns, it was clear to all those who studied taxonomy that there was a "natural order" that grew out of the process of classifying animals and plants (see our Taxonomy I: What's in a name? module).

Scientists wondered why, when classified, groups of species seemed to form clusters, as if some sort of biodiversity magnet pulled them together and put them in one place. Within clusters, species tended to be similar to one another by different degrees. Surely it was no coincidence that all species of cats are alike, from the alley cat to the lion to the prehistoric saber tooth that roamed the western United States. Chance could not be the reason why dogs, wolves, and coyotes are all variations on the theme of "Dog." Similarly, chance could not explain the similarities and differences of the Galapagos finches that Darwin collected while he was with the Beagle expedition. What was behind the repeated pattern of species clusters that was so common in nature?

Comprehension Checkpoint
Darwin preferred the term "descent with modification" over "evolution" because

Biodiversity as a staircase: A "macro" view of life

Before Darwin, there was only one available model that naturalists used to explain species' similarities and differences, and that would not work to solve this problem. Scientists had thought that the most important pattern of biodiversity was what they called the Scale of Nature. This was the notion that the vast range of living organisms – say, from snail to ant to fish to mouse to monkey to man – was a feature of divine creation meant to highlight our own superiority. Life, they thought, was arranged like a set of stairs, with "lower" forms situated on the bottom and "higher" forms, humans, appearing at the top. This idea can be traced as far back as Aristotle, more than 2,000 years ago, and it was popularized in the 1800s by writers like Robert Chambers. Of course, it was all based on assumption. No evidence was provided to support the model, but it was generally accepted by tradition.

Figure 3a: Scale of Nature Model

Figure 3b: How the Scale of Nature and Phylogenetic Models interpret the anatomical similarities and differences between apes and humans. The Scale of Nature model assumes a hierarchy of lower and higher organisms, while the Phylogenetic model does not. The current phylogenetic relationship among chimps, gorillas, and humans is different than that believed to be true in Darwin's day and is shown in the green inset.

The Scale of Nature, which was actually more of a "macro" view of life, would not work for Darwin because it did not relate to the clusters of similar species that he had observed. Why would so many types of cats or finches exist? The Scale of Nature suggested an unchanging, linear quality to evolution, but that surely could not explain the explosive variety of adaptations that Darwin saw among the finches he found on the Galapagos Islands during the Beagle voyage. Darwin observed finches that were adapted to feeding on different things: birds with beaks that were specialized to eat seeds, leaves, insects, or nectar. The list of items could not be interpreted as a scale-like linear climb involving a worse-adapted food source to better-adapted food source, or from poor-food to rich-food.

Comprehension Checkpoint
The __________ model assumes a hierarchy of higher and lower life forms.

Biodiversity as a tree: A "micro" view of life

Instead of the macro view offered by the Scale, Darwin was focused on a "micro" view of biodiversity: What could explain the small variations distinguishing species that actually resembled one another? He came to see that evolutionary changes on the micro level would add up to the differences that were obvious at the macro level. So, instead of a stairway or ladder as a metaphor for understanding the cluster pattern of biodiversity, Darwin pictured a tree.

This was a brilliant insight. Rather than being arrow-like and linear, a tree has many elements that spread out in different directions. Rather than being static, it is dynamic. It grows over time, just as evolution is embedded in time. It sprouts branches, as if it were generating new varieties and new species. Or, it may have branches that do not subdivide. Some branches grow straight up, parallel to the trunk, while most head off in different directions as they develop, resembling alternative adaptations. Some branches grow into stumps and die out, becoming extinct. Others may grow long and last for generations, thousands of years, tens of thousands of years, and even longer. None of the branches of a tree is judged to be any better than others; none is superior and none is inferior. They are all simply different.

Crucially important is the fact that all the branches of a tree are interconnected. You can trace their origins from their endpoints to the parent shoots from which they grew, just as you might trace the roots of dogs, or cats, or Galapagos finches to their original ancestral species.

The Origin tree diagram illustrates how a branching pattern of evolution can produce a greater number of species over time than what was there to begin with. It shows how some lines of species, or lineages, split more frequently than others. It shows that some lineages do not split at all but evolve almost like a column. It shows that extinction is a basic property of descent: Many populations are left behind and do not reach the top because they have died out.

Comprehension Checkpoint
Darwin saw evolution as a

Darwin's early model

Figure 4: Darwin's earliest depiction of the tree of life, showing how many species, closely or distantly related, might evolve from a single ancestor.

Coming up with this tree-model of evolutionary process and pattern was not easy. In fact, Darwin's personal notebooks reveal how his own understanding grew over the years. In one notebook, which he began writing soon after returning from the Beagle expedition, he drew a crude stick-like diagram to show that many species could evolve from a single ancestral species by somehow splitting apart.

This early graphic, shown in Figure 4, is shaped like a cross between a tree and a starburst. It seems as if Darwin was trying to form an idea of how the great diversity of species could come about naturally from a single origin, rather than having each species being specially created. It is a flat image, as if he were drawing the diversity of Galapagos finches on a map of the islands. That, in fact, was key to his figuring out that evolution had occurred on the Galapagos, that is, how the birds had evolved across space. But his 1859 drawing, clearly for the first time, provided the blueprint of evolution though time. It illustrated his notion of descent with modification, how natural selection produces change and also a pedigree of connections between species that shows where they came from historically, meaning phylogenetically.

Darwin's finches

To better understand the one illustration that Darwin included in Origin, it is helpful to picture the finches that he studied on the Galapagos. What would this diagram look like if he had illustrated it with species of finch? At the bottom of the figure we would see the ancestral finches. As the lines diverge and branch out higher in the diagram, new species of finch would appear, leading to the array of modern birds at the top of the picture. To better illustrate this idea, work your way through "Darwin's Finches," the interactive animation linked below.

Darwin's Finches interactive animation

Interactive Animation:
Darwin's Finches interactive animation

With the tree of life as a metaphor for evolution, Darwin changed the way both scientists and the public view the origin of species. There would no longer be a need to interpret the biodiversity of nature as a ladder or scale, with some species better or worse than others due to the details of their size, fur, and teeth or as measured by their intelligence or ancestry. All are adapted to their specific environments, even though some might not survive. And all, at some level, share a common source of origin.

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