Principles of Microbial Diversity. James W. Brown
2
Context and Historical Baggage
If, as described in chapter 1, evolutionary diversity underlies the various aspects of “diversity,” it is important to review what we mean by evolutionary diversity. This is especially so because even some biologists have a relatively poor understanding of modern evolutionary theory and it is important to at least dig up and expose outdated but widely held notions about how organisms are related.
The evolution of evolutionary thought
The chain of being
Well before people discovered that species changed over time, both living and nonliving things were organized (in Western culture, including Aristotle) into the “chain of being” (Fig. 2.1).
All species and substances were placed onto individual slices of a vertical scale, ascending from “inferior” to “superior.” Much thought and energy were put into deciding exactly how to order the major categories of materials, living things, species, races, social classes, and even individuals onto this linear arrangement. (It will come as no surprise to the cynical that the ethnic group placed at the top of the human region of the chain generally matched that of whoever was organizing the chain.) This was an impossible task, with lots of problematic and contradictory issues. In retrospect, it is obvious that the reason for this was that there is no linear relationship between things in nature.
As an aside, this linear arrangement of substances survives, in general form, in the periodic table, in which the elements are arranged in order of increasing
Figure 2.1 The “chain of being,” with some representative organisms shown. To the right is a closer view of one thin slice of this “chain.” doi:10.1128/9781555818517.ch2.f2.1
atomic number. The chain is broken into segments, and the segments are stacked in such a way that elements with similar properties form columns. The reason this works is because elements are structured in a linear series based on atomic number.
The evolutionary ladder
The earliest forms of evolutionary thought had each species (up to but not including humans) moving up the “evolutionary ladder,” a climbable form of the chain of being. Evolution, in this view, was linear and progressive; species evolved from one rung to the next as an effect of a progressive force (Fig. 2.2).
Although the notion of an evolutionary ladder is pre-Darwinian and hopelessly incorrect, we are discussing it here because this scheme is firmly embedded in modern biological unconscious thought. You even hear the term “evolutionary ladder” used fairly often by research scientists who actually know better (at least, one hopes they do). The terms “higher” and “lower” eukaryotes, and “missing link,” which are in common use, are relics of this view.
Early phylogenetic trees
By the time of Charles Darwin, it was clear that the evolutionary ladder was not a reasonable view of evolution. Darwin describes a much better view, which has proven to be essentially correct, in which species originate by divergence, as shown in Fig. 2.3 (this diagram is the only illustration in On the Origin of Species).
It turns out that species actually evolve by diversification, not by progression. Eukaryotes did not evolve from bacteria, animals did not evolve from ciliates, plants did not evolve from fungi, and humans did not evolve from chimpanzees. Each of these pairs of modern organisms shares a common ancestor, from which each diverged. This is a fundamental aspect of evolution that is poorly understood even by many biologists.
Figure 2.2 The “evolutionary ladder,” an evolutionary transformation of the chain of being. (Redrawn from Prothero DR, Buell CD, Evolution: What the Fossils Say and Why It Matters. Copyright 2007 Columbia University Press. Reprinted with permission of the publisher.) doi:10.1128/9781555818517.ch2.f2.2
Figure 2.3 Evolution by diversification. In this diagram, species A and I at the beginning (bottom) split many times and diverge constantly. Most of these divergences do not go anywhere (they become extinct), but some do make it, at least for a while, resulting in this case in species A splitting into three separate surviving species and species I splitting into two surviving species at time X. Species A and I no longer exist at time X. Note that most of the original species (B through H, K, and L) are in stasis, remaining unchanged through the span of time shown here (or at least the part of it during which they survive). (Reprinted from Charles Darwin, On the Origin of Species. John Murray, London, 1859.) doi:10.1128/9781555818517.ch2.f2.3
One of the best developed of the early divergent evolutionary trees was that of Ernst Haeckel. In this tree, there are three major, equivalent divisions of life—plants, animals, and protists (Fig. 2.4). This tree is a huge improvement over the evolutionary ladder. It is a tree, species are not ranked, and modern species are not considered to be the ancestors of other modern organisms. Plants and animals are not thought of as having evolved from modern prokaryotes (monerans) but are separate groups.
The Whittaker five-kingdom tree being taught in various forms in most classrooms today is a refinement of this tree (Fig. 2.5). In some ways, however, this five-kingdom tree is actually a step backward toward the evolutionary ladder. In most versions of this scheme (such as the original tree by Whittaker, above), eukaryotes are shown to be descended from within the bacteria (not true), and in many representations eukaryotic algae are shown as descendants of cyanobacteria (not true), fungi are shown as descendants of filamentous gram-positive bacteria (not true), and protists are shown as descendants of wall-less gram-positive bacteria (also not true). Also notice the implied vertical axis: either superiority (sometimes expressed as “complexity”) or time (usually labeled “time of origin”). But if this axis is complexity, what exactly is being measured? Eukaryotes are sometimes morphologically complex, but what about parasites that have simplified; why are these organisms not drawn as downward-pointing arrows? What about metabolic complexity, which would place animals close to the bottom? Is morphological complexity the only factor being considered? Why? If the vertical axis is time of origin, why are the recent emergences of bacterial families, genera, and species not considered? The genus Escherichia emerged about 100 million years ago, about the same time as the primates; why is Escherichia (along with all other bacteria), a modern organism, shown as a relic of the past?
Figure 2.4 The tree of life. (Reprinted from Ernst Haeckel, Generelle Morphologie der Organismen. G. Reimer, Berlin, Germany, 1866.) doi:10.1128/9781555818517.ch2.f2.4
Figure 2.5 The five-kingdom tree taught in most schools in the United States. (Redrawn from Whittaker RH, Science 163:150–160, 1969. Used with permission from AAAS.) doi:10.1128/9781555818517.ch2.f2.5
The reality is that the five-kingdom tree is entirely qualitative and subjective.
Molecular phylogenetic trees
If the traditional