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Lesson3 BW Letter - en

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Lesson3 BW Letter - en

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Shadrack Rabaloi
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© © All Rights Reserved
We take content rights seriously. If you suspect this is your content, claim it here.
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3.

Lamarck and the Inheritance of Acquired Characters


If we look at change in the natural world, it is hard not to be surprised by some of the
peculiar adaptations that we see. To take what might be the most common example,
think of the neck of a giraffe. Obviously it’s a great advantage for the giraffe to be able
to eat leaves from high in the trees. And obviously, too, the giraffe would like to be able
to get more food. So a perfectly natural way to describe what might have happened
to giraffes would be to say that they kept stretching their necks, over and over again,
desperately trying to eat the highest leaves, and as a response, their necks grew. We
even see cases like this in our own lives: if you lift lots of weights, your muscles grow
in response. Why wouldn’t evolution be like this?

THINK[1]: For many scientists over hundreds of years, this conclusion seemed
obvious; yet for us now it might seem strange. What do you think about it?
More broadly, when is our everyday experience useful as a source of scientific
knowledge, and when might it mislead us? How can we tell the difference?

As it turns out, evolution isn’t like this (or, at least, it’s al-
most always not like this, though more about that later),
but this is one of the oldest and most common misun-
derstandings about the evolutionary process. Many of
the most prominent scientists for more than a century
were convinced that it must be the case that, to describe
the problem as they would have, acquired characters
are inherited – that is, that characters that organisms
A giraffe helps itself to some very
develop during their lives (like stretched necks or big high food, Kruger National Park,
biceps) will be passed on to their offspring. Darwin South Africa (CC-BY-SA; by Escu-
himself even argued that what he called “the effects of lapio at Wikimedia Commons)
use and disuse,” or the ways in which using a part or disusing another part could
affect how characters were passed from parents to offspring, was an important factor
in cases like the disappearance of eyes from species that live in caves.
One of the first authors to really develop a view of the organic world like this – and
whose name has been attached to it ever since – was the French zoologist Jean-Baptiste
de Lamarck (1744–1829). His view of the development of life is sometimes called
“transformism,” to indicate that while he did believe that species changed, he didn’t
have a theory of “evolution” in the modern sense. On the contrary, he had a picture
on which life was constantly being created all around us, starting with very simple
microorganisms, and steadily evolving upwards in progress and complexity (recall our
first reading from Darwin about concepts of “higher and lower”). So microorganisms,
in part because they are striving for better lives, will become jellyfish, which will
become worms, and onward through insects, molluscs, and finally vertebrates like us.

THINK[2]: Lamarck is also interesting because of how he relates scientific pride


and national pride. For many years in France, it was thought that Lamarck had
largely invented the theory of evolution, and Darwin had in some sense stolen
the credit. What’s different between Darwin’s view and the picture of Lamarck’s

1
theory illustrated here? How might the question of inventing scientific theories
become a matter of national pride or a dispute between countries?

In his central work on biology, the Zoological Philosophy published in 1809, Lamarck
himself described the inheritance of acquired characters in terms of what he called
two laws of biology:

Lamarck, Zoological Philosophy (1809)

First Law
In every animal which has not passed the limit of its development, a more frequent
and continuous use of any organ gradually strengthens, develops, and enlarges that
organ, and gives it a power proportional to the length of time it has been so used;
while the permanent disuse of any organ imperceptibly weakens and deteriorates it,
and progressively diminishes its functional capacity, until it finally disappears.
Second Law
All the acquisitions wrought by nature on individuals, through the influence of the en-
vironment in which their race has long been placed, and hence through the influence
of the predominant use or permanent disuse of any organ; all these are preserved by
reproduction to the new individuals which arise, provided that the acquired modifi-
cations are common to both sexes, or at least to the individuals which produce the
young.
Here we have two permanent truths, which can only be doubted by those who have
never observed or followed the operations of nature. . .1

The first law, then, argues that whatever parts an organism uses frequently, as a result
of doing the same kinds of actions over and over again, will cause those parts to
become larger and stronger. Disuse will cause other parts to shrink or even disappear.
The second law says that these changes in parts will be inherited, such that after long
enough they would become a permanent part of the species. So the stretched neck of
the giraffe would, with enough time, become a permanent part of all giraffes.
While the inheritance of acquired characters wasn’t the only mechanism of species
change for Lamarck (like Darwin, he also believed that changes in the environment
of an organism would be extremely important), it was identified closely enough with
Lamarck’s theory that the inheritance of acquired characters itself would come to be
called Lamarckism, even today.
Even though, as already mentioned, Darwin himself did think that acquired characters
were inherited, in the years after the Origin of Species was published, this idea came
under increasing attack. The German biologist August Weismann (1834–1914) was the
1Lamarck, Jean-Baptiste. 1914 [1809]. Zoological Philosophy. Translated by Hugh Elliot. New York:
Hafner Publishing Co., p. 113.

2
first to propose that sex cells, like eggs and sperm – the germ line – are separate from
the rest of the cells of the organism – the somatic cells – and that no changes present
in the somatic cells could be inherited, as only the germ cells were responsible for
producing the next generation of organisms. While this was hotly debated at the time
(until future advances in cellular biology made it possible to understand the way in
which embryos develop from gametes), it seriously weakened the credibility of the
theory of the inheritance of acquired characters among at least some biologists.
But not all biologists were dissuaded (we’ll look at some of those in this reading, and
some in the next reading). One group, largely American, defended Lamarckism as
a way to address what they thought were two crucial flaws in Darwin’s system: the
randomness of variation and the long amount of time that it took for those variations
to accumulate.
Let’s consider the two problems they are responding to. First, they argue that there is
evidence that variation is in fact not random – that organisms are, as Lamarck claimed,
responding to their environments, and hence varying in precisely the direction that
was needed to deal with the environmental challenges which they were facing. This
kind of variation, Darwin argued, was impossible; natural selection, he claimed, is the
only feature in evolution that is directed toward increased fitness. The second worry,
about the speed of evolution, was a common one in the years just after the Origin. It
was thought that the earth was much younger than we now believe it to be – because
radioactivity was not yet understood, it was thought that the earth would have cooled
off much more quickly from its initial molten state. That made biologists worry that
there wouldn’t have been enough time for natural selection to act in order to produce
the kinds of organisms that we see today.

THINK[3]: We saw that an important part of evaluating evolutionary theory was


the calculation of the age of the earth that had been derived in physics (and, later,
the discovery of radioactivity). How do you think the knowledge generated in
physics might be related to biological claims? When would these be useful? Can
you think of circumstances where appeals to other sciences might be unhelpful
or problematic?
THINK[4]: The idea of “random variation” has always been difficult to interpret
in evolutionary theory. By it, Darwin means that variations are not biased in
the direction of what the organism “wants,” but rather occur without regard to
whether they will be helpful or harmful. Natural selection, on the contrary, is not
at all random, in its ability to drive populations toward increased fitness.
What kinds of misunderstandings do you think could arise from the description
of variation as “random?” What difference is made by the presence of natural
selection within the evolutionary process? In general, what might be some differ-
ences between scientific theories that describe their results as probabilities versus
those, like Newtonian physics, that describe precisely what will happen?

These scientists – who would soon be known as neo-Lamarckians – argued that


Lamarck’s two laws, or something like them, could solve both of these problems. One,
Alpheus Hyatt (1838–1902), drew much of his experience from marine invertebrates.

3
He argued that the evolution of cephalopods (like today’s squid, octopuses, and
cuttlefishes) was the best evidence of the truth of Lamarck’s principles. Let’s look at
his text first, as he wrote in a journal article from 1884:

Hyatt, from the journal Science (1884)

The efforts of the Orthoceratite to adapt itself fully to the requirements of a mixed habi-
tat gave the world the Nautiloidea: the efforts of the same type to become completely a
littoral crawler developed the Ammonoidea. The successive forms of the Belemnoidea
arose in the same way; but here the ground-swimming habitat and complete fitness
for that was the object, whereas the Sepioidea represent the highest aims as well as the
highest attainments of the Orthoceratites, in their surface-swimming and rapacious
forms.2

An Orthoceratite (top left), an ammonite (Asteroceras obtusum, bottom left), and a nautilus (Nautilus
pompilius, right; left two images CC-BY; by Nobu Tamura; right image public domain; all Wikimedia
Commons)

The first example that he offers us here, and the one that we will unpack, concerns the
following evolutionary story. Some of the most commonly found fossils in the fossil
record are a sort of spiral-shaped cephalopod called the ammonites, which lived for
hundreds of millions of years before going extinct at the same time as the dinosaurs.
They are such commonly found fossils that they were described by the ancient Roman
author Pliny, and finding an ammonite can be a way for paleontologists to tell just
how old a layer of rock actually is. With their closely related group the nautiluses (a
few of which are still around today), they would have been a common sight in the
ocean for a vast amount of the earth’s history.
2Hyatt, Alpheus. 1884. “The Evolution of the Cephalopoda.—I.” Science 3 (52): 122–7, p. 125. https:
//doi.org/10.1126/science.ns-3.52.122.

4
THINK[5]: The fossil record is an important source of data for our understanding
of evolution, but it is also quite strange. What do you think might be some
advantages and disadvantages of using fossil data to support a conjecture in
evolution? How might we correct some of those disadvantages with other kinds
of experiments to produce a more robust hypothesis?

How did they evolve? The ancestor to all of the am-


monites and nautiluses was a group of straight-shelled
organisms, called the Orthoceratites. It is this transition,
from the straight-shelled Orthoceratites which lived in
open water, to the much more diverse ammonites and
nautiluses, that interests Hyatt. He argues that the evo-
lution of these two groups could not have been caused
by natural selection. A better explanation, he argues, is
the direct interaction between organisms and their en-
vironment. As the environment changed around them,
or the ancestor organisms had reasons to move to dif-
ferent environments, they began to struggle to survive
in their new homes. The fact that the former, straight-
Alpheus Hyatt, in an engraving
from around 1885 in Popular
shelled form, which had been very efficient in open
Science Monthly (public domain; water, needed (in one case) to be able to live in a more di-
Wikimedia Commons) verse environment (sometimes swimming, sometimes
at the bottom of the ocean, sometimes in shallower water) led it to evolve into the
nautilus, and the fact that it needed (in another case) to live in shallow, coastal areas
led it to evolve into the ammonites.
Of course, Hyatt doesn’t think that a marine invertebrate is actually thinking about
what it wants to do – it’s not sitting in the deep water thinking “I really wish I could
live near the shore.” As he puts it:

Hyatt, from the journal Science (1884)

We cannot seriously imagine these changes to have resulted from intelligent effort;
but we can fully join Lamarck, Cope, and Ryder, in imagining them as due to efforts
induced by the physical requirements of the habitat, and think this position to be
better supported by facts than any other hypothesis. (Footnote: A noted French writer
well known to embryologists, Lacaze-Duthier, has lately asked, “Who, at the present
time, supports Lamarck?” The author can answer, that some of our leading scientific
men consider Lamarck’s hypothesis to contain more fundamental truths than Darwin’s
or any other.)3

3Hyatt, p. 125.

5
THINK[6]: Hyatt is openly making an appeal to persuasion here. In placing
himself in opposition to Darwin, he explicitly lists the names of authorities –
Lamarck himself, as well as Cope and Ryder – who agree with him, calling them
“some of our leading scientific men.”
What do you think this says about Hyatt’s theory and his position within the
scientific establishment? How should we understand these arguments that appeal
directly to the authority of other scientists? When do you think that they might
be valid?
What the Orthoceratites are doing, though, is struggling against their habitat. Some
of them found themselves in shallow water, and thus began causing changes in them-
selves that, when passed on to offspring for many generations, created the ammonites.
Hyatt is very clear, however, about what this means: Lamarck, he argues, has described
“more fundamental truths” than Darwin has. The force of the organism’s struggle
against its environment is much more powerful, Hyatt thinks, than natural selection.
In addition to Lamarck, Hyatt mentions two other fig-
ures as proponents of neo-Lamarckism, one of whom
was Edward Drinker Cope (1840–1897). Cope is an
amazing figure, worth studying for lots of reasons.
Among other things, he was one of the first major Amer-
ican paleontologists, and a dispute between him and
fellow fossil-hunter Othniel Charles Marsh (1831–1899)
kicked off a competition to find dinosaur fossils in the
American West that became known as the Bone Wars.
This radically changed our understanding of the nature
and evolution of dinosaurs, and helped to create the
contemporary discipline of paleontology. Edward Drinker Cope, in an en-
graving from 1881 in Popular
But in addition to that, Cope was another promi- Science Monthly (public domain;
nent supporter of neo-Lamarckism. For him, neo- Wikimedia Commons)
Lamarckism was a way to understand a different feature
of the fossil record than the one that was important for Hyatt. In his words:

Cope, Origin of the Fittest (1887)

It is sufficiently well known that the essential features of a majority of genera are not
adaptive in their natures, and that those of many others are so slightly so, as to offer
little ground for the supposition that this necessity has preserved them.
Both laws [that is, natural selection, and another evolutionary force proposed by
Cope] must be subordinate to that unknown force which determines the direction
of the great series. If a series of suppressions of the nervous and circulatory systems
of beings of common birth produced the “synthetic” predecessors of the classes of

6
Vertebrata, the direction toward which the highest advanced, or its ultimate type, can
be only ascribed as yet to the divine fiat. So far as we can see, there is no reason or
law to produce a preference for this direction above any other direction.4

For Cope, then, the important thing that neo-Lamarckism lets us understand is the
fact that many of the “essential features” of various groups of organisms seem not to
be adaptive, and therefore not to have been created by natural selection. Despite this,
however, they have been preserved, sometimes for a very long time, in the history
of life. (We will return to other explanations for non-adaptive features in the next
reading, on the related concept of orthogenesis.) For Cope, then, the best way to
explain this, in the same manner that Lamarck had done, is to interpret the series of
organisms across evolutionary history as having a sort of inherent “direction.” Unlike
Lamarck, Cope does not want to say that this direction points toward “progress” or
“complexity” or “higher” organisms. It is often, as he says here, hard for us to even
clearly understand how that direction should be explained, unless by “divine fiat.”
What we need to do, he argues, is attempt to understand the “unknown force” that
drives the Lamarckian, non-adaptive change in different groups of organisms. That,
rather than natural selection, is the most important process in evolutionary theory.

THINK[7]: This leaves us three alternatives in play for the explanation of these
non-adaptive characters. One is a classic, Lamarckian force of upward progress
caused by struggle with the environment (as supported also by Hyatt). One is
an unknown force driving organisms in particular directions, but without any
invocation of adaptation (as supported by Cope). Lastly we have Darwin, who
would presumably have appealed to natural selection to explain these changes.
How should we compare these possibilities? What kinds of data does each ex-
planation have to support it? What features of the natural world does each leave
unexplained or unconsidered? Which do you think would have made the most
sense had you been asked to choose in the late-19th century?

Let’s think first a bit about the positive view of the neo-Lamarckians, before we turn
to what their perspective might have to tell us about our view of evolution today. It is
no accident that many of the most prominent neo-Lamarckians were paleontologists,
looking at long-term changes in groups in the fossil record. When we look at series of
fossils as their characteristics change over time, we often are struck that there seems
to be linear progression in certain features, in a way that doesn’t seem necessarily to
be connected with any selective pressure. Whatever the force would be that controlled
the appearance of progression like this – Hyatt, with Lamarck, though that it was the
active efforts of organisms to respond to the needs they had within their environments,
while Cope didn’t know what it was, but hoped we would one day find out – it is the
main driver of change in species, especially over geologic time.
4Cope, Edward Drinker. 1887. The Origin of the Fittest. New York: D. Appleton and Company,
pp. 106–7.

7
Darwin, for his part, knew
about cases like this. He
had argued that they
should be understood
in a very different way.
Certain parts, he wrote in
the Origin, are correlated
with one another in
the way that they grow
and develop. So natural
selection, working on one
part that was important
for an organism’s survival,
Cope’s sketch of four dinosaurs. Problems abound, including, for
the dinosaur on the front in the water, the head’s being placed on might “accidentally” (so
the end of its tail! From an article in the journal American Naturalist to speak) cause changes in
(public domain; Wikimedia Commons) a different part, if the two
parts’ growth were linked. That could mean that natural selection might appear to
be working on characters that weren’t actually relevant for an organism’s success
in its current environment, which might produce the appearance of non-adaptive
evolution. (Much more on this in the next reading.)
But if the neo-Lamarckian explanation was valid, why are we reading about their
theories as a misunderstanding of evolution? What happened to neo-Lamarckism?
In fact, the real reason that it declined in importance was a persistent failure to
replicate its results experimentally. If organisms are really responding to the needs
of their environments in a way that is inherited by their offspring, then placing a
population of organisms in a new environment, over a long period of time, should
cause the descendants to eventually be born with modified features, more suitable to
the new environment in which they were placed, variations that weren’t present in
the population before, and only exist now because of the organisms’ having struggled
in their new environment. This kind of experimental evidence, however, simply never
materialized, though the experiments were tried across a wide variety of different
species.

THINK[8]: Imagine that you were a neo-Lamarckian, and you were presented
with the following experimental result. Rats have been trained to solve a maze for
multiple generations in the lab, and in spite of this, their descendants don’t wind
up solving the maze any faster than their ancestors. What kinds of responses
might you have? How could you still defend your theory? More broadly, when
do you think a scientist would have to admit defeat, to concede that their theory
had failed? How much evidence, and evidence of what kinds, would be required?

What can we learn from the episode of neo-Lamarckism that could be helpful for
understanding evolutionary theory today? First, it cautions us that it can be extremely
difficult to interpret the patterns that we see in the fossil record. There are many
different ways in which such a pattern might have been generated, different targets of
natural selection, and different modes of interaction between organisms and their

8
environments. Sometimes, something that looks like a pattern to us might not even
have been evolutionarily generated at all: it might, for example, just be the result of
the forces of physics to which an organism is subject while it is growing.
One way of understanding the critiques of the neo-Lamarckians demonstrates a point
about evolution that has become more and more important in recent years. Something
that neo-Lamarckian evolution was supposed to give us was the ability for organisms to
respond quickly when changes in their environments made their previous form of life
difficult or impossible. The Orthoceratites, that is, saved themselves from extinction
by successfully struggling with their environment to change their features, turning
into ammonites or nautiluses. Some recent work in evolution has returned to this
insight, focusing on the ways in which a concept now known as phenotypic plasticity
might play a similar role. The idea here is that an organism might not be selected for a
particular, permanent solution to a problem – rather, in some circumstances, it might
be better to simply remain flexible, particularly if the environment is highly variable.
Only later, when the changing environment “settles down,” would natural selection
arrive to make some of these flexible outcomes more permanent. By taking this
worry more seriously, we were attuned to a variety of really interesting phenomena
connecting evolution, molecular biology, and organismic development. In that sense,
the neo-Lamarckians were right – environmental change is a real and important
problem that organisms have to be able to solve, and there are interesting theoretical
questions at work about how natural selection has managed to solve it.

THINK[9]: We not infrequently see examples like this, where theories that were
thought to be long since disproven gain a new life as they are reinterpreted in light
of fresh data. What do you think this might tell us about the relationship between
the history of science and the practice of science today? Should we encourage
scientists to learn more about history, or would that be a waste of time?

Finally, it’s worth noting that contemporary scientific research has offered us some
insights into very particular ways that, in fact, acquired characters might actually
be passed on to offspring. To take just one example, some behaviors in life might
cause certain kinds of changes in DNA, other than mutations in the actual genetic
sequence – such as changes in methylation patterns – that could still be passed on and
govern certain future outcomes in the lives of offspring. While nothing like Lamarck’s
complete theory can be vindicated, perhaps the examples here will need to be updated
soon!
THINK[10]: Do you think it is helpful to refer to contemporary results in molec-
ular biology in terms of old theoretical names like “neo-Lamarckism?” Give some
reasons that this identification might be both confusing and helpful.

THINK: NOS Reflection Questions:


What does the story of neo-Lamarckism tell us about the following features of the
nature of science?
• evidential relevance (empiricism)
• role of cultural beliefs (national identity)

9
• interdisciplinary thinking
• role of probability in inference
• role of imagination and creative synthesis
• resolving disagreement

Further Reading
• Bowler, Peter J. 1992. The Eclipse of Darwinism: Anti-Darwinian Evolution
Theories in the Decades around 1900. Baltimore, MD: Johns Hopkins University
Press.
• Bowler, Peter J. 2017. “American Palaeontology and the Reception of Darwinism.”
Studies in History and Philosophy of Biological and Biomedical Sciences 66: 3–7.
https://doi.org/10.1016/j.shpsc.2017.06.002.
• Jaffe, Mark. 2000. The Gilded Dinosaur: The Fossil War between E. D. Cope and
O. C. Marsh and the Rise of American Science. New York: Crown.
• Haig, David. 2007. “Weismann Rules! OK? Epigenetics and the Lamarckian
Temptation.” Biology & Philosophy 22 (3): 415–28. https://doi.org/10.1007/s10539-
006-9033-y.

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