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Library of human understanding

Biology: Evolution and Life

One family tree, four billion years deep.

Every living thing on Earth reads the same genetic code and descends from common ancestors. Biology is the study of how that one inheritance became bacteria, oak trees, and the people reading this page. But genes alone do not make life: they work within cells, tissues, organisms, and ecosystems, each level shaping what comes next.

Life's Structure: DNA Helix

Adenine (A)
Thymine (T)
Guanine (G)
Cytosine (C)

DNA is a double helix of nucleotides bonded by complementary base pairing: adenine pairs with thymine, and guanine pairs with cytosine. This structure allows cells to replicate their genetic information accurately.

Genes are sections of DNA that code for proteins. During replication, the two strands separate and each serves as a template for a new strand, creating two identical DNA molecules.

Evolution occurs when mutations introduce variation in DNA sequences, and natural selection acts on this variation over generations, shaping populations and species.

The gene-centric view of evolution

In The Selfish Gene (1976) and The Extended Phenotype (1982), Richard Dawkins reframed evolution around genes as the units of selection and replication. In this view, organisms are vehicles for genes, and natural selection operates at the genetic level. Genes replicate with occasional errors, and those errors that improve a gene's chances of copying itself become more common.

This perspective helped explain many evolutionary puzzles: why genes would build large bodies and complex brains, why organisms often act against the interests of their species, and how traits that seem to damage an organism can persist if they benefit the genes that code for them. The gene-centric view unified Darwinian evolution with Mendelian genetics and became central to the modern evolutionary synthesis.

Critics argue that this view, though powerful, can overlook the hierarchical nature of biological organization: genes do not act in isolation, but within cells, tissues, organs and organisms. The gene-centric model explains inheritance but struggles with how complex systems arise from genetic instructions, and with the role of regulatory mechanisms, epigenetics, and the organism's environment in shaping what genes do.

Systems biology and biological relativity

Denis Noble and other systems biologists propose that genes are necessary but not sufficient to explain life. In this view, causation flows in multiple directions: genes influence cells, but cells also regulate which genes are active; tissues shape cell behaviour; organisms interact with their environments. Life is a multi-level system in which no single level holds all the answers.

Noble's principle of biological relativity states that the level at which causation appears to operate depends on the question asked and the scale at which we look. A mutation in a gene affects the protein it codes for, the cell containing it, the organ it helps build, the organism's behaviour, and the ecosystem it inhabits. Each level has real causal power.

This view explains epigenetics, in which the same DNA sequence is read differently depending on chemical modifications that are reversible and can be inherited; developmental plasticity, in which the same genes build different forms in different environments; and the canalization of development, in which multiple genetic and environmental perturbations can be buffered by the system's robustness.

Genes and environment: Two streams of inheritance

Genes carry information copied from parent to offspring, but so does the environment. An organism inherits DNA from its parents, but also the cell's epigenetic state, maternal effects such as hormones and antibodies, its ecological niche, and the behaviour of other organisms around it.

Recent work in developmental biology, epigenetics and environmental science shows that the boundary between 'nature' and 'nurture' is porous. Gene expression is controlled by regulatory sequences that respond to the environment. Histone modifications and DNA methylation, which affect which genes are read, can be inherited for generations without changing the DNA sequence. Microbes in and around an organism influence its development and health.

Both perspectives agree that genes are crucial to life. The disagreement is about what 'genes' do and at what level we need to look to understand inheritance, development and evolution. Genes interact with regulatory networks, other genes, the cell, the organism, the ecosystem, and history. Understanding life means understanding all these interactions.

Pioneers of Biology and Evolution

From Darwin's voyage to modern systems biology, these thinkers have reshaped how we understand life. Two recent figures—Richard Dawkins and Denis Noble—represent opposing perspectives on a fundamental question: At what level of biological organization does life's complexity arise?

Charles Darwin

1809 – 1882 · Biology, Geology

  • Spent five years on the voyage of HMS Beagle collecting the evidence that shaped his thinking.
  • Explained the diversity of life by natural selection in On the Origin of Species (1859); Alfred Russel Wallace reached the same idea independently.
“There is grandeur in this view of life, with its several powers, having been originally breathed into a few forms or into one; and that, whilst this planet has gone cycling on according to the fixed law of gravity, from so simple a beginning endless forms most beautiful and most wonderful have been, and are being, evolved.”
On the Origin of Species, first edition (1859), closing paragraphLater editions add 'by the Creator' after 'breathed'.

Gregor Mendel

1822 – 1884 · Biology

  • Bred about ten thousand pea plants in his monastery garden in Brno and found that traits pass on in discrete units in predictable ratios.
  • Published in 1866; the work was ignored until it was rediscovered in 1900, and it became the basis of genetics.

Barbara McClintock

1902 – 1992 · Biology

  • Discovered transposons, 'jumping genes', in maize in the 1940s, showing that genomes are not fixed.
  • Received the Nobel Prize in Physiology or Medicine alone in 1983.

Theodosius Dobzhansky

1900 – 1975 · Biology

  • Joined Mendel's genetics to Darwin's selection in Genetics and the Origin of Species (1937), a founding book of the modern synthesis.
“Nothing in biology makes sense except in the light of evolution.”
Title of an essay in The American Biology Teacher, March 1973

Rosalind Franklin

1920 – 1958 · Biology, Chemistry

  • Made the X-ray diffraction images of DNA, including Photograph 51, that showed its helical form.
  • Her data were shown to Watson and Crick without her knowledge and were decisive for their model.
  • Went on to work out the structure of viruses, including tobacco mosaic virus.

Francis Crick

1916 – 2004 · Biology, Physics

  • Proposed the double-helix structure of DNA with James Watson in 1953.
  • Stated the central dogma of molecular biology and helped crack the genetic code.
“It has not escaped our notice that the specific pairing we have postulated immediately suggests a possible copying mechanism for the genetic material.”
Watson and Crick, Nature 171, 737–738, 25 April 1953Written jointly with James Watson.

Richard Dawkins

born 1941 · Biology, Philosophy

  • Introduced the concept of the gene as the unit of natural selection in The Selfish Gene (1976), framing evolution from the perspective of genes rather than organisms.
  • Extended the gene-centred view in The Extended Phenotype (1982), proposing that genes influence the world beyond the body, including the behaviour of other organisms.
  • Advocated for the modern synthesis and gene-centric view of evolution in numerous works and public debates.
“The gene is the unit of selection. Genes are the things that replicate, with occasional errors, in the process of life. They are the basis of inheritance.”
The Selfish Gene, first edition (1976), Ch. 1
“A gene's-eye view of evolution sees life as genes controlling robots whose job is to preserve and propagate the genes.”
The Selfish Gene, first edition (1976), Ch. 1

Denis Noble

born 1936 · Biology, Physiology

  • Pioneered systems biology and cardiac electrophysiology, building the first mathematical model of the heart's electrical activity in 1962.
  • Developed the biological relativity principle and 'Third Way' of evolution (alongside others), emphasizing multi-level causation and the role of complex biological systems beyond genes.
  • Authored The Music of Life (2006) and Dance to the Tune of Life (2016), arguing that genes require biological context and organizational hierarchies to function.
“Life is the result of interactions between genes and the rest of the organism and the environment. The genes are not the 'drivers' of the organism. They are part of a network that is regulated from the top down, and from the bottom up, and sideways.”
The Music of Life: Biology Beyond the Genome (2006), Ch. 2
“The notion that genes themselves drive evolution is not supported by what we know about the working of biological systems.”
Dance to the Tune of Life: Biological Relativity (2016), Ch. 5