EFTA01043301.pdf
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Resource centre > Darwin the botanist
Darwin the botanist
Volume 5 Number 2 - October 2008
David Kohn
French title: Darwin le botaniste
Spanish title: Darwin el botanico
Summary
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In his theory of evolution, Charles Darwin proposed that living beings were "all netted together" through a
common ancestry. Yet one branch of the tree of life, the plant kingdom, was Darwin's most enduring focus. He
was a major field collector, a consummate and sustained observer of plant life, a rigorous botanical
experimentalist, and the high theorist of plant evolution.
Botany played a pivotal part in each phase of Darwin's life. As an undergraduate, he collected specimens for his
botany professor's herbarium while on a geological expedition in Wales. Voyaging for five years aboard the
HMS Beagle, he collected plants along with fossil bones and bird skins. Preparing to write On the Origin of
Species, botany became critical to the growth of his evolutionary theory. Ultimately, he turned his home and the
surrounding countryside into a botanical field station.
Darwin's six botanical books would recast large areas of plant science. His studies on the fertilization of orchids,
on insectivorous plants, and on the climbing and other plant movements were each a beautifully articulate
example of how evolution could solve the traditional mysteries of natural history. Through his work, Darwin laid
foundations for modem botany that remain firm to this day.
Darwin's botanical formation
Darwin was heir to a rich botanical tradition, one that often resonated in the specific botanical topics and
viewpoints he was to adopt. His grandfather Erasmus Darwin, for example, was an evolutionist and spread the
Linnaean doctrine that plants, just like animals, have sexes. Darwin's first botanical exposure was in his father's
ample garden at The Mount in Shrewsbury where young Charles played among apple trees bred by Thomas
Andrew Knight, President of the Royal Horticultural Society. However, the foundations of Darwin's formal
education in botany were laid at Cambridge by his dynamic professor John Stevens Henslow (1796-1861) who
later opened the way for Darwin to participate in the Beagle's scientific cruise around the world.
The Beagle voyage
The Beagle expedition concentrated on certain regions of South America: coastal Brazil, Patagonia, Tierra del
Fuego, the Andean coast as far north as Lima and the oceanic Galapagos archipelago.
Plants came before birds in the birth of Darwinian evolution. Darwin did not become an evolutionist on the
Galapagos, but the basis for a profound shift in his understanding of species was established there — and it began
with plants. When the Beagle arrived in the Galapagos in September 1835, Darwin immediately observed that
the flora appeared unique. Thereupon, he collected "all the plants in flower". This we know from a pocket field
notebook that Darwin kept when he was in the archipelago. Darwin's Galapagos plant specimens, numbering
well over 200, constitute the single most influential natural history collection of live organisms in the entire
history of science. Indeed, Darwin's plants represent the foundational collection for the entire Galapagos flora.
Settled back in London in 1837, he sketched his first evolutionary tree and began firmly applying the idea of
descent with modification to all of natural history.
In 1856, Darwin put ink to blue foolscap and began to steadily write. His seminal book, On the Origin of
Species, was finally published in November 1859.
Preserving priority
Once the Origin was published, botany became the central focus of Darwin's research for the remaining years of
his life. This botanical work was highly original, not only because of the quality of his observations, but because
it was the first attempt to apply the principles of evolution to plants.
Darwin's crucial contribution to botany was his understanding of and ability to demonstrate that the flower is a
product of evolution. By the 18th century, botanists were convinced that flowering plants have both male and
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female parts, and assumed that most plants self fertilise or inbreed. This erroneous and deeply ingrained view of
the flower continued until Darwin began publishing on the biological meaning of flowers in the 1860s. The
prevalence of cross-pollination was Darwin's single most important botanical truth.
Indeed Darwin realised there was a problem with plant sex immediately after discovering natural selection in
1838. If flowers perpetually self fertilised then both natural selection and evolution would be invalid, because
natural selection requires hereditary variation. If there is no variation, there is simply nothing to select. Evolution
stops. If flowers self fertilised for generation after generation, they would become unvarying clones.
Furthermore, without variation, where did the thousands of known plant species come from? Evolution would
not explain the diverse families of flowering plant genera and species. The world's flowers were just too big a
part of nature for Darwin to forfeit them to the creationist position that species are immutable.
In the case of flowers, he undertook decades of field and garden observations and breeding experiments, all
focused on testing and supporting one powerful hypothesis: that of natural selection.
Through his painstaking studies, such as the Primula example below, Darwin discovered that flowering plants
have evolved elaborate structures, strategies, and relations with animals and reasoned that it was all to avoid the
apparent necessity of perpetual inbreeding. Thus the Darwinian meaning of flowers became a pillar of botany
and botany became one of the strongest fields supporting evolution.
Love, Primula style
Spring 1860 was the first flowering season following Origin's publication. Darwin, bursting with experimental
energy, made several crucial botanical discoveries. In May, he observed two different kinds of flowers among
common primulas. The style, or shaft of the female part, is either tall, protruding like a pin, or it is short.
Darwin's children gathered armloads of flowers for their father, who noticed something: The two kinds of flower
occur in a 50:50 ratio. To explain these two morphs, Darwin would follow the scientific method, but after his
very own personal style. At first, the 50:50 ratio reminded him of a normal male-female sex ratio. So he believed
the two kinds of flowers were evolving into separate sexes. That is, he thought he had witnessed one step in the
evolution of separate male and female flowers. The tall-style flowers must be evolving into pure females, he
surmised. Conversely, their short stamens must be losing potency. To test this he crossed the two forms. But to
his surprise, the `males' produced abundant seeds. So the experimental method forced him to abandon his first
hypothesis. As Darwin once observed, his first explanations of things frequently proved wrong. The originality
and breakthrough would come with thinking up the next explanation.
He now realised that the maximum fertility occurs when pollen moves from one form to the other. It was always
the self fertilised flowers that had reduced fertility. Thus the two forms, each of which is both male and female,
are favoured to maintain a stable population. Darwin had in fact discovered a breeding strategy that gave a clear
advantage to cross-pollination and thus provided experimental support for his long-held interpretation of the
meaning of flowers. To think of flowers in terms of plant breeding strategies, now that is evolutionary botany.
And Darwin's primulas have been a prime example of this new way of thinking ever since.
Plant sensitivity: green adaptations
Plants don't have feet and they also don't have brains. But as far as Darwin was concerned, some plants as good
as have eyes, and they do have behaviour, often expressed in the extraordinary ways these beings can move
despite being rooted in the soil. Darwin's passion for working out the botanical adaptations that allow plants to
stay in sensitive touch with their environment occupied him for years in physiological experiments that prefigure
the biochemical and cellular studies of the early 20th century on enzymatic catalysis and plant hormones. Thus
these green adaptations in leaves and stems not only paralleled Darwin's flowers, they added a new level of
sophistication to the botanical bulwarks Darwin had built to support evolution and adaptation by natural
selection. For example, insectivorous plants that trap and digest insects with specialised leaves fascinated
Darwin. So did vines that climb up and over other plants. He saw plants as sensitive creatures whose growing
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tips or leaves or seedlings can track the movement of the sun. Indeed, he showed they could respond to the least
beam of incident light, the pull of gravity, and the slight touch of a browsing animal.
Much of this work Darwin performed in a string of hothouses that gradually sprang up along the kitchen garden
wall beginning in the late 1850s. Eventually, there were five houses heated by a boiler and offering a fair range
of conditions for the array of plants that Darwin, assisted in his last years by his son Francis, wished to study.
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