On the Origin of Species – Chapter 3.

Chapter 3 – Struggle for Existence

This is the chapter in which Darwin really picks things up. Whereas the first two may have been more like, in Darwin’s own words, “a dry list of facts”, this one is written in a very passionate tone. Having established that individuals vary both in domesticated breeds and in nature, he now moves into the second major ingredient of natural selection: overproduction.

Some important things in this chapter:

1) Darwin strongly emphasizes adaptation. This is not very surprising, given that he considered natural selection as the main (but not only) mechanism of evolution.

“But the mere existence of individual variability and of some few well-marked varieties, though necessary as the foundation for the work, helps us but little in understanding how species arise in nature. How have all those exquisite adaptations of one part of the organisation to another part, and to the conditions of life, and of one distinct organic being to another being, been perfected? We see these beautiful co-adaptations most plainly in the woodpecker and missletoe; and only a little less plainly in the humblest parasite which clings to the hairs of a quadruped or feathers of a bird; in the structure of the beetle which dives through the water; in the plumed seed which is wafted by the gentlest breeze; in short, we see beautiful adaptations everywhere and in every part of the organic world.”

2) We may think life is well balanced and peaceful, but there is carnage going on all around, if not right this moment then at the very least during difficult seasons.

“We behold the face of nature bright with gladness, we often see superabundance of food; we do not see, or we forget, that the birds which are idly singing round us mostly live on insects or seeds, and are thus constantly destroying life; or we forget how largely these songsters, or their eggs, or their nestlings, are destroyed by birds and beasts of prey; we do not always bear in mind, that though food may be now superabundant, it is not so at all seasons of each recurring year.”

“Although some species may be now increasing, more or less rapidly, in numbers, all cannot do so, for the world would not hold them.”

3) The struggle for life can be against the environment (e.g., surviving in cold or dry habitats), but the major factor is the biotic environment. He also points out that what matters more than survival is reproduction.

“I should premise that I use the term Struggle for Existence in a large and metaphorical sense, including dependence of one being on another, and including (which is more important) not only the life of the individual, but success in leaving progeny. Two canine animals in a time of dearth, may be truly said to struggle with each other which shall get food and live. But a plant on the edge of a desert is said to struggle for life against the drought, though more properly it should be said to be dependent on the moisture. A plant which annually produces a thousand seeds, of which on an average only one comes to maturity, may be more truly said to struggle with the plants of the same and other kinds which already clothe the ground. The missletoe is dependent on the apple and a few other trees, but can only in a far-fetched sense be said to struggle with these trees, for if too many of these parasites grow on the same tree, it will languish and die. But several seedling missletoes, growing close together on the same branch, may more truly be said to struggle with each other. As the missletoe is disseminated by birds, its existence depends on birds; and it may metaphorically be said to struggle with other fruit-bearing plants, in order to tempt birds to devour and thus disseminate its seeds rather than those of other plants. In these several senses, which pass into each other, I use for convenience sake the general term of struggle for existence.”

4) Darwin considers extinction to be gradual, not catastrophic (he acknowledges what we now call background extinction but not mass extinctions).

“Battle within battle must ever be recurring with varying success; and yet in the long-run the forces are so nicely balanced, that the face of nature remains uniform for long periods of time, though assuredly the merest trifle would often give the victory to one organic being over another. Nevertheless so profound is our ignorance, and so high our presumption, that we marvel when we hear of the extinction of an organic being; and as we do not see the cause, we invoke cataclysms to desolate the world, or invent laws on the duration of the forms of life!”

5) Darwin recognizes different strategies of reproduction (what we now call r-selection and K-selection).

“A large number of eggs is of some importance to those species, which depend on a rapidly fluctuating amount of food, for it allows them rapidly to increase in number. But the real importance of a large number of eggs or seeds is to make up for much destruction at some period of life; and this period in the great majority of cases is an early one. If an animal can in any way protect its own eggs or young, a small number may be produced, and yet the average stock be fully kept up; but if many eggs or young are destroyed, many must be produced, or the species will become extinct.”

6) Competition is most intense between members of the same species because their requirements are so similar.

“But the struggle almost invariably will be most severe between the individuals of the same species, for they frequent the same districts, require the same food, and are exposed to the same dangers.”

7) At higher levels, closely related species experience more intense competition than distantly related species, again because their requirements are similar.

“As species of the same genus have usually, though by no means invariably, some similarity in habits and constitution, and always in structure, the struggle will generally be more severe between species of the same genus, when they come into competition with each other, than between species of distinct genera.

Some famous quotes:

“Again, it may be asked, how is it that varieties, which I have called incipient species, become ultimately converted into good and distinct species, which in most cases obviously differ from each other far more than do the varieties of the same species? How do those groups of species, which constitute what are called distinct genera, and which differ from each other more than do the species of the same genus, arise? All these results, as we shall more fully see in the next chapter, follow inevitably from the struggle for life. Owing to this struggle for life, any variation, however slight and from whatever cause proceeding, if it be in any degree profitable to an individual of any species, in its infinitely complex relations to other organic beings and to external nature, will tend to the preservation of that individual, and will generally be inherited by its offspring. The offspring, also, will thus have a better chance of surviving, for, of the many individuals of any species which are periodically born, but a small number can survive. I have called this principle, by which each slight variation, if useful, is preserved, by the term of Natural Selection, in order to mark its relation to man’s power of selection. We have seen that man by selection can certainly produce great results, and can adapt organic beings to his own uses, through the accumulation of slight but useful variations, given to him by the hand of Nature. But Natural Selection, as we shall hereafter see, is a power incessantly ready for action, and is as immeasurably superior to man’s feeble efforts, as the works of Nature are to those of Art.”

“We can dimly see why the competition should be most severe between allied forms, which fill nearly the same place in the economy of nature; but probably in no one case could we precisely say why one species has been victorious over another in the great battle of life.”

Forbes on Darwin.

Well, I certainly don’t agree with their full selection of authors (see Pharyngula and NeuroLogica), but anyway here is the set of papers in Forbes on evolution.

The Evolution Of Evolution

By Hana R. Alberts

Surveying a landscape of cognition–and controversy.


What We See When We Look In The Mirror

By Lionel Tiger

Humankind’s epic search for existential answers.


In Praise Of A Beautiful Theory

By Michael Ruse

Why evolution matters today more than ever.

The Self-Made Species

By Denis Dutton

We fail to realize just how much evolution shapes our personalities.

You Should Believe In Love At First Sight

By Helen Fisher

Charles Darwin knew about romance.


Keeping Pace With Change

By Owen D. Jones

Why it matters that behaviors evolve.

Survival Of The (Financially) Fittest

By Leo M. Tilman

Evolutionary pressures and economic fate.

We Are The Moral Animals

By Larry Arnhart

Fear of a Darwinian, animalistic hedonism is unfounded.


From Evolution Comes Literature

By Joseph Carroll

Can we prove what we read and write is rooted in biology?


Why Evolution Doesn’t Matter

By Matt Woolsey

Forget century-old debates. Biology should inspire technology for the future.

The Reticent Abolitionist

By Adrian Desmond

Slavery is hard to justify if all species–animal, black and white–share a common ancestor.


Darwin The Dog Lover

By David Allan Feller

Canine companionship inspired evolutionary observations.

A Life In 14,500 Letters

By Paul White

Glimpses of the earnest passions behind the beard.


The Debate Over Intelligent Design

By Kathryn Tabb

What would Darwin say?


Darwinian Dangers

By Ken Ham

The Creation Museum’s founder fears moral collapse and racism.


There Is No ‘Politically Correct’ Science

By John G. West

It’s impossible to isolate Darwinian theories from their societal consequences.


A Neurosurgeon, Not A Darwinist

By Michael Egnor

Why I don’t believe in atheism’s creation myth.


The Problem Of Evidence

By Jonathan Wells

If it isn’t testable, it isn’t science.

The Man In Darwin’s Shadow

By Michael Shermer

Did Alfred Russel Wallace think up evolution first?


Alfred Russel Wallace

By Michael A. Flannery

The proponent of intelligent design in Darwin’s day.

Nature discusses Canadian science woes.

Fellow Canadians,

If you’re wondering about the broader impression of the challenges facing Canadian science, you can see these stories from the prestigious UK journal Nature over the past year or so.

By the way, I haven’t found a colleague yet who isn’t a) glad about infrastructure funding, b) puzzled and concerned by a lack of support for people and supplies, and c) totally perplexed by the establishment of huge graduate scholarships for a small collection of individuals.

Cash concerns for Canadian scientists
Could programme cuts prompt a brain drain?

Canada’s scientists face an uncertain future
Political turmoil leaves key positions in doubt

Canada abolishes its national science adviser
After just four years, government axes post

Science in retreat
Canada has been scientifically healthy. Not so its government.

Science – speciation.


Today’s issue of Science is on speciation. The papers:

Species Uncertainties
Robert M. May and Paul H. Harvey
Science 6 February 2009: 687.

Happy Birthday, Mr. Darwin
Andrew Sugden, Caroline Ash, Brooks Hanson, and Laura Zahn
Science 6 February 2009: 727.

The Red Queen and the Court Jester: Species Diversity and the Role of Biotic and Abiotic Factors Through Time
Michael J. Benton
Science 6 February 2009: 728-732.

Adaptive Radiation: Contrasting Theory with Data
Sergey Gavrilets and Jonathan B. Losos
Science 6 February 2009: 732-737.

Evidence for Ecological Speciation and Its Alternative
Dolph Schluter
Science 6 February 2009: 737-741.

The Bacterial Species Challenge: Making Sense of Genetic and Ecological Diversity
Christophe Fraser, Eric J. Alm, Martin F. Polz, Brian G. Spratt, and William P. Hanage
Science 6 February 2009: 741-746.

Is Genetic Evolution Predictable?
David L. Stern and Virginie Orgogozo
Science 6 February 2009: 746-751.

See also:


Elizabeth Pennisi
Science 6 February 2009: 706-708.

EVOLUTION: The Overwhelming Evidence
Massimo Pigliucci
Science 6 February 2009: 716-717.

Wallace Arthur
Science 6 February 2009: 717.

Time’s Stamp on Modern Biogeography
J. Alistair Crame
Science 6 February 2009: 720-721.

Signature of the End-Cretaceous Mass Extinction in the Modern Biota
Andrew Z. Krug, David Jablonski, and James W. Valentine
Science 6 February 2009: 767-771.

Sequential Sympatric Speciation Across Trophic Levels
Andrew A. Forbes, Thomas H.Q. Powell, Lukasz L. Stelinski, James J. Smith, and Jeffrey L. Feder
Science 6 February 2009: 776-779.

Lies, Damned Lies, and Science…

I just received an email about the release of this new book

Lies, Damned Lies, and Science: How to Sort through the Noise around Global Warming, the Latest Health Claims, and Other Scientific Controversies
by Sherry Seethaler

Here is the overview:

Are antioxidants good or bad for you? Is global warming real – and if so, why it is happening? Do scientists really now think that sun exposure prevents cancer? Is organic food healthier? Do antidepressants trigger suicides? Every day, the media is packed with stories about health and science: stories that can sometimes mean the difference between life and death. Which of these stories really matter? Non-scientists desperately need tools for telling the difference between real science and misleading science.

Lies, Damned Lies, and Science: How to Sort through the Noise around Global Warming, the Latest Health Claims, and Other Scientific Controversies (FT Press Science, ISBN-13: 9780137155224, $24.99, 224 pps, hardcover, February 2009) www.ftpress.com/science helps readers sort out today’s noisiest scientific debates — and gives them common-sense techniques for evaluating any health or scientific claim. Sherry Seethaler, a science writer and educator at University of California, San Diego, works with scientists to explain their discoveries to the public. Seethaler holds an M.S. and Master of Philosophy in biology from Yale, and a Ph.D. in science and math education from UC Berkeley.

Seethaler says, “Those who promote incorrect information, either because they are trying to manipulate you, or because they themselves have been duped or are simply misinformed, rarely have more knowledge about science than you do. What they have are skills at using information to suit their purposes.”

This could be very good, or very not-good. Anyone read it?

On the Origin of Species – Chapter 2

Once again, this is just a repost of what I wrote in my book club discussion forum.

Chapter 2 – Variation Under Nature

This is a fairly short chapter, with much less information than the discussion of variation under domestication. In part, this is because a lot more was known about variation in domesticated animals and plants than in natural species. However, it wasn’t considered by everyone to be a good approach — Wallace always thought Darwin’s argument was weakened by relying so much on domestication as an analogy with natural processes.

Some things I found interesting in this chapter:

1) Species are hard to define. We talked in class about how there is no clear definition of species and how this causes problems in biology. Well, Darwin recognized the difficulty very early (p.67).

“Nor shall I here discuss the various definitions which have been given of the term species. No one definition has as yet satisfied all naturalists; yet every naturalist knows vaguely what he means when he speaks of a species.”

2) Darwin begins to suggest that the variation within species (e.g., varieties) is the same stuff that turns into differences among species. He introduces the term “incipient species” to indicate this. However, not all incipient species will become species — some may go exinct and some may not change further (p.76).

“Hence I believe a well-marked variety may be justly called an incipient species; but whether this belief be justifiable must be judged of by the general weight of the several facts and views given throughout this work.

It need not be supposed that all varieties or incipient species necessarily attain the rank of species. They may whilst in this incipient state become extinct, or they may endure as varieties for very long periods.”

3) Sometimes daughter species can co-exist with parental species — it is not always a gradual change of one species into another. In other words, Darwin recognizes cladogenesis (indeed, something compatible with punctuated equilibria) and not only anagenesis (p.76) .

“If a variety were to flourish so as to exceed in numbers the parent species, it would then rank as the species, and the species as the variety; or it might come to supplant and exterminate the parent species; or both might co-exist, and both rank as independent species.”

4) Nevertheless, Darwin does not think that species are real. They are just convenient constructs (p.76).

“From these remarks it will be seen that I look at the term species, as one arbitrarily given for the sake of convenience to a set of individuals closely resembling each other, and that it does not essentially differ from the term variety, which is given to less distinct and more fluctuating forms. The term variety, again, in comparison with mere individual differences, is also applied arbitrarily, and for mere convenience sake.”

UPDATE: John Wilkins does not think that this and similar passages indicate that Darwin considered species as mere conveniences, only that he considered the distinction between varieties (which he called “incipient species”) and species to be mainly arbitrary. He may have a point.

5) Darwin suggests that widespread, numerous species are likely to produce more daughter species (p.77). This would seem to contradict later models of speciation involving geographic isolation, especially ones based on drift in small isolates.

“Hence it is the most flourishing, or, as they may be called, the dominant species,— those which range widely over the world, are the most diffused in their own country, and are the most numerous in individuals,—which oftenest produce well-marked varieties, or, as I consider them, incipient species.”

6) Darwin has compiled a series of data (he doesn’t show them here) comparing genera that are diverse (lots of species) and those that are not, and argues that larger genera include species that themselves include more varieties. He argues on this basis that dominant lineages will become more dominant, since these varieties are incipient species. However, he also notes that this does not continue indefinitely because some previously dominant lineages disappear and some small genera can expand.

An open letter to American universities and agencies.

Subject: An open letter to American universities and agencies
Jan. 31, 2009

Dear Sir or Madam,

You are probably aware that Canadian researchers have long made do with comparatively small amounts of funding, particularly relative to colleagues in the United States, but that they have nevertheless been very productive (Nature 430: 311-316). To be sure, you are familiar with many of the current international projects of which Canada is a leader. You may also have noticed that many Canadians have returned after working in the United States and elsewhere, and that prominent researchers from the United States and elsewhere have also opted to move to Canada to pursue their research interests.

You also are no doubt aware that the recent federal budget released by the minority government of Canada contains no new funding for Genome Canada and requires the three major funding councils (CIHR, NSERC, and SSHRC) to find savings of $87 million. You may also be aware that this government has begun imposing “priority areas” for research support, the majority of which are short-term and applied in nature, and that they have decided to launch scholarships for a small number of students that are, in effect, significantly larger than the average starting grants for primary investigators.

In contrast, your new President has chosen to make a major investment in scientific research as part of an economic stimulus package. He clearly recognizes that scientific expertise and knowledge will be crucial for the long-term health of your citizens and your economic future. In particular, it is likely that you will be expanding your efforts in important areas of cutting edge science, such as genomics.

In light of this, we ask that you kindly refrain from recruiting our best scientists, who may soon become open to moving their programs to your fine nation. As you can understand, we require these individuals to maintain Canada’s scientific productivity and credibility, as well as to teach in our universities and to train our future generations of investigators. We also ask, if you would be so kind, that you not take over major international initiatives in which Canada has heretofore assumed the leading role. Finally, we ask for your patience as we struggle to live up to our commitments in joint research efforts with our colleagues in the USA.

We are deeply hopeful that this represents only a temporary setback, but we thank you in advance for your understanding during this difficult time.

Yours sincerely,

Canadian science

More on the disaster in Canadian science.

Thanks are due to the Globe and Mail (one of Canada’s national newspapers) for continuing to report on the lack of new support for Canadian science in the new budget by the Conservative government. In a time when the US is injecting major new funds into science and technology, when Canada is now in a leadership position in many major initiatives following a reversal of the “brain drain”, and when the future of the economy after the current challenges will clearly be based in large part on biotechnology, the Conservatives have decided to cut off the development of new research and to exert increased control on the science that does receive funding. This administration is a disaster for Canadian science, and for all elements of society affected by science, from the economy to citizens’ health.

Here is what’s going on, folks. The Conservative government is investing in infrastructure but is cutting off support for new initiatives by not supporting the equally important investment in people and operating costs. They are requiring the three federal granting councils to cut $87 million from their already disproportionately small budgets (hint: I received considerably more from NSERC as a postdoc than as a primary investigator). They are focusing support for students into a smaller number of insanely large scholarships ($50K/year, twice the average starting grant!). They are imposing their own priority areas on funding programs, including mostly applied issues (e.g., forestry, automotive, fisheries) and destroying support for basic research. And they have dissolved the national science advisor’s office.

If you’re not upset about this, fellow Canadians, you will be when we lose our best and brightest and rapidly fall behind other countries in the new biotech economy.

Genome Canada again.

Here is the official word from Genome Canada:

Federal Budget 2009

Key Messages from the Board of Directors of Genome Canada

  • Genome Canada is pleased with the federal government’s 2009 budget in which millions will be invested in research infrastructure over the next two years. This is good news for the scientific community across the country that needs to be at the cutting-edge of research infrastructure and new technologies in order to maintain Canada’s competitiveness at the national and international level.
  • Although Genome Canada did not receive funding in the 2009 federal budget to fund new genomics research projects, this will not impact Genome Canada’s current projects that received a full commitment of funding from previous federal government investments in 2007 and 2008.
  • Genome Canada has in place two five-year funding agreements with the Government of Canada for a total of $240M:
    $100 M (2008-2012)
    $140 M (2009- 2013)
  • These investments flow to Genome Canada on a cash requirement basis. Thus, a total of $107M has been invested in 2008-09; and a total of $106.5M will be invested in 2009-10, creating and maintaining over 2,350 HQP positions per year.
  • Over the same period of time, Genome Canada has raised over $225M from other strategic partners in the private, public and philanthropic sectors to support genomics research in Canada.
  • Since its inception in 2000, Genome Canada has provided operating funds to Canadian genomics researchers, while complementing other sources of funds for infrastructure coming from such agencies as Canada Foundation for Innovation, to allow them to be among world leaders in their respective fields such as human health, agriculture, environment, forestry, fisheries, new technology, and GE3LS (ethical, environmental, economic, legal and social issues).
  • Genome Canada is confident that the Government of Canada and its other financial strategic partners will do everything possible over the coming years to secure additional funding to support new initiatives in genomics research in Canada while increasing Canada’s productivity, wealth and well-being of all Canadians.

In a recent story in GenomeWeb Daily News, President of Genome Canada Martin Godbout said,

“Genome Canada is not in the federal budget this year, and there’s good reason for it,” Godbout explained.

“Genome Canada is not founding any infrastructure or equipment,” explained Godbout, who described this year’s federal budget as one focused on projects that will feed those objectives. “It is a decision that I respect,” he told GWDN. “It’s hard everywhere.”

This conflicts with his earlier reaction (here and here), but it does bear noting that no current projects will be cut and that there is significant funding for infrastructure through other agencies.

The problem is that this does not allow new initiatives to be developed or productive ones to be expanded. It’s still very bad news in that regard. More importantly, it is essential that the ministry realize that infrastructure alone can’t do anything. You also have to support people and research expenses if you desire results.