Survival of the fittest
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"Survival of the fittest" is a phrase that originated from
Darwinian Darwinism is a theory of biological evolution developed by the English naturalist Charles Darwin (1809–1882) and others, stating that all species of organisms arise and develop through the natural selection of small, inherited variations that ...
evolutionary theory Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
as a way of describing the mechanism of
natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the heritable traits characteristic of a population over generations. Cha ...
. The biological concept of fitness is defined as
reproductive success Reproductive success is an individual's production of offspring per breeding event or lifetime. This is not limited by the number of offspring produced by one individual, but also the reproductive success of these offspring themselves. Reproduct ...
. In Darwinian terms, the phrase is best understood as "Survival of the form that will leave the most copies of itself in successive generations."
Herbert Spencer Herbert Spencer (27 April 1820 – 8 December 1903) was an English philosopher, psychologist, biologist, anthropologist, and sociologist famous for his hypothesis of social Darwinism. Spencer originated the expression " survival of the fi ...
first used the phrase, after reading
Charles Darwin Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all species of life have descended ...
's ''
On the Origin of Species ''On the Origin of Species'' (or, more completely, ''On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life''),The book's full original title was ''On the Origin of Species by Me ...
'', in his ''Principles of Biology'' (1864), in which he drew parallels between his own economic theories and Darwin's biological ones: "This survival of the fittest, which I have here sought to express in mechanical terms, is that which Mr. Darwin has called 'natural selection', or the preservation of favoured races in the struggle for life."

^ "Herbert Spencer in his ''Principles of Biology'' of 1864, vol. 1, p. 444, wrote: 'This survival of the fittest, which I have here sought to express in mechanical terms, is that which Mr. Darwin has called "natural selection", or the preservation of favoured races in the struggle for life.'" , citing HERBERT SPENCER, THE PRINCIPLES OF BIOLOGY 444 (Univ. Press of the Pac. 2002.)
Darwin responded positively to Alfred Russel Wallace's suggestion of using Spencer's new phrase "survival of the fittest" as an alternative to "natural selection", and adopted the phrase in ''
The Variation of Animals and Plants Under Domestication ''The Variation of Animals and Plants under Domestication'' is a book by Charles Darwin that was first published in January 1868. A large proportion of the book contains detailed information on the domestication of animals and plants but it al ...
'' published in 1868."This preservation, during the battle for life, of varieties which possess any advantage in structure, constitution, or instinct, I have called Natural Selection; and Mr. Herbert Spencer has well expressed the same idea by the Survival of the Fittest. The term "natural selection" is in some respects a bad one, as it seems to imply conscious choice; but this will be disregarded after a little familiarity." In ''On the Origin of Species'', he introduced the phrase in the fifth edition published in 1869,"This preservation of favourable variations, and the destruction of injurious variations, I call Natural Selection, or the Survival of the Fittest." – intending it to mean "better designed for an immediate, local environment".


History of the phrase

By his own account,
Herbert Spencer Herbert Spencer (27 April 1820 – 8 December 1903) was an English philosopher, psychologist, biologist, anthropologist, and sociologist famous for his hypothesis of social Darwinism. Spencer originated the expression " survival of the fi ...
described a concept similar to "survival of the fittest" in his 1852 "A Theory of Population". He first used the phrase – after reading
Charles Darwin Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all species of life have descended ...
's ''
On the Origin of Species ''On the Origin of Species'' (or, more completely, ''On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life''),The book's full original title was ''On the Origin of Species by Me ...
'' – in his ''Principles of Biology'' of 1864 in which he drew parallels between his economic theories and Darwin's biological, evolutionary ones, writing, "This survival of the fittest, which I have here sought to express in mechanical terms, is that which Mr. Darwin has called 'natural selection', or the preservation of favored races in the struggle for life." In July 1866 Alfred Russel Wallace wrote to Darwin about readers thinking that the phrase "
natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the heritable traits characteristic of a population over generations. Cha ...
" personified nature as "selecting", and said this misconception could be avoided "by adopting Spencer's term" ''Survival of the fittest''. Darwin promptly replied that Wallace's letter was "as clear as daylight. I fully agree with all that you say on the advantages of H. Spencer's excellent expression of 'the survival of the fittest'. This however had not occurred to me till reading your letter. It is, however, a great objection to this term that it cannot be used as a substantive governing a verb". Had he received the letter two months earlier, he would have worked the phrase into the fourth edition of the ''Origin'' which was then being printed, and he would use it in his next book on "Domestic Animals etc.". Darwin wrote on page 6 of ''
The Variation of Animals and Plants Under Domestication ''The Variation of Animals and Plants under Domestication'' is a book by Charles Darwin that was first published in January 1868. A large proportion of the book contains detailed information on the domestication of animals and plants but it al ...
'' published in 1868, "This preservation, during the battle for life, of varieties which possess any advantage in structure, constitution, or instinct, I have called Natural Selection; and Mr. Herbert Spencer has well expressed the same idea by the Survival of the Fittest. The term 'natural selection' is in some respects a bad one, as it seems to imply conscious choice; but this will be disregarded after a little familiarity". He defended his analogy as similar to language used in chemistry, and to astronomers depicting the "attraction of gravity as ruling the movements of the planets", or the way in which "agriculturists speak of man making domestic races by his power of selection". He had "often personified the word Nature; for I have found it difficult to avoid this ambiguity; but I mean by nature only the aggregate action and product of many natural laws,—and by laws only the ascertained sequence of events." In the first four editions of ''
On the Origin of Species ''On the Origin of Species'' (or, more completely, ''On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life''),The book's full original title was ''On the Origin of Species by Me ...
'', Darwin had used the phrase "natural selection". In Chapter 4 of the 5th edition of ''The Origin'' published in 1869, Darwin implies again the synonym: "Natural Selection, or the Survival of the Fittest". By "fittest" Darwin meant "better adapted for the immediate, local environment", not the common modern meaning of "in the best physical shape" (think of a puzzle piece, not an athlete). In the introduction he gave full credit to Spencer, writing "I have called this principle, by which each slight variation, if useful, is preserved, by the term Natural Selection, in order to mark its relation to man's power of selection. But the expression often used by Mr. Herbert Spencer of the Survival of the Fittest is more accurate, and is sometimes equally convenient." In ''The Man Versus The State'', Spencer used the phrase in a postscript to justify a plausible explanation of how his theories would not be adopted by "societies of militant type". He uses the term in the context of societies at war, and the form of his reference suggests that he is applying a general principle.
"Thus by survival of the fittest, the militant type of society becomes characterized by profound confidence in the governing power, joined with a loyalty causing submission to it in all matters whatever".
Though Spencer's conception of organic evolution is commonly interpreted as a form of Lamarckism, Herbert Spencer is sometimes credited with inaugurating
Social Darwinism Social Darwinism refers to various theories and societal practices that purport to apply biological concepts of natural selection and survival of the fittest to sociology, economics and politics, and which were largely defined by scholars in We ...
. The phrase "survival of the fittest" has become widely used in popular literature as a catchphrase for any topic related or analogous to evolution and natural selection. It has thus been applied to principles of unrestrained
competition Competition is a rivalry where two or more parties strive for a common goal which cannot be shared: where one's gain is the other's loss (an example of which is a zero-sum game). Competition can arise between entities such as organisms, indiv ...
, and it has been used extensively by both proponents and opponents of Social Darwinism. Evolutionary biologists criticise the manner in which the term is used by non-scientists and the connotations that have grown around the term in
popular culture Popular culture (also called mass culture or pop culture) is generally recognized by members of a society as a set of practices, beliefs, artistic output (also known as, popular art or mass art) and objects that are dominant or prevalent in a ...
. The phrase also does not help in conveying the complex nature of natural selection, so modern biologists prefer and almost exclusively use the term
natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the heritable traits characteristic of a population over generations. Cha ...
. The biological concept of fitness refers to both
reproductive success Reproductive success is an individual's production of offspring per breeding event or lifetime. This is not limited by the number of offspring produced by one individual, but also the reproductive success of these offspring themselves. Reproduct ...
( fecundity selection), as well as survival ( viability selection), and is not prescriptive in the specific ways in which organisms can be more "fit" by having phenotypic characteristics that enhance survival and reproduction (which was the meaning that Spencer had in mind).


Critiquing the phrase

While the phrase "survival of the fittest" is often used to mean "
natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the heritable traits characteristic of a population over generations. Cha ...
", it is avoided by modern biologists, because the phrase can be misleading. For example, survival is only one aspect of selection, and not always the most important. Another problem is that the word "fit" is frequently confused with a state of physical fitness. In the evolutionary meaning " fitness" is the rate of reproductive output among a class of genetic variants.


Interpreted as expressing a biological theory

The phrase can also be interpreted to express a theory or hypothesis: that "fit" as opposed to "unfit" individuals or species, in some sense of "fit", will survive some test. Nevertheless, when extended to individuals it is a conceptual mistake, the phrase is a reference to the ''transgenerational'' survival of the heritable attributes; particular individuals are quite irrelevant. This becomes more clear when referring to
Viral quasispecies A viral quasispecies is a population structure of viruses with a large number of variant genomes (related by mutations). Quasispecies result from high mutation rates as mutants arise continually and change in relative frequency as viral replicatio ...
, in ''survival of the flattest'', which makes it clear ''to survive'' makes no reference to the question of even being alive itself; rather the functional capacity of proteins to carry out work. Interpretations of the phrase as expressing a theory are in danger of being tautological, meaning roughly "those with a propensity to survive have a propensity to survive"; to have content the theory must use a concept of fitness that is independent of that of survival."Stephen Jay Gould
Darwin's Untimely Burial
, 1976; from Philosophy of Biology:An Anthology, Alex Rosenberg, Robert Arp ed., John Wiley & Sons, May 2009, pp. 99–102.
von Sydow, M. (2014)
‘Survival of the Fittest’ in Darwinian Metaphysics – Tautology or Testable Theory?
(pp. 199–222) In E. Voigts, B. Schaff & M. Pietrzak-Franger (Eds.). ''Reflecting on Darwin''. Farnham, London: Ashgate.
Interpreted as a theory of species survival, the theory that the fittest species survive is undermined by evidence that while direct competition is observed between individuals, populations and species, there is little evidence that competition has been the driving force in the evolution of large groups such as, for example, amphibians, reptiles, and mammals. Instead, these groups have evolved by expanding into empty ecological niches. In the punctuated equilibrium model of environmental and biological change, the factor determining survival is often not superiority over another in competition but ability to survive dramatic changes in environmental conditions, such as after a meteor impact energetic enough to greatly change the environment globally. The main land dwelling animals to survive the K-Pg impact 66 million years ago had the ability to live in tunnels, for example. In 2010 Sahney et al. argued that there is little evidence that intrinsic, biological factors such as competition have been the driving force in the evolution of large groups. Instead, they cited extrinsic, abiotic factors such as expansion as the driving factor on a large evolutionary scale. The rise of dominant groups such as amphibians, reptiles, mammals and birds occurred by opportunistic expansion into empty ecological niches and the extinction of groups happened due to large shifts in the abiotic environment.


Interpreted as expressing a moral theory


Social Darwinists

It has been claimed that "the survival of the fittest" theory in biology was interpreted by late 19th century capitalists as "an ethical precept that sanctioned cut-throat economic competition" and led to the advent of the theory of "
social Darwinism Social Darwinism refers to various theories and societal practices that purport to apply biological concepts of natural selection and survival of the fittest to sociology, economics and politics, and which were largely defined by scholars in We ...
" which was used to justify
laissez-faire ''Laissez-faire'' ( ; from french: laissez faire , ) is an economic system in which transactions between private groups of people are free from any form of economic interventionism (such as subsidies) deriving from special interest groups ...
economics, war and racism. However, these ideas pre-date and commonly contradict Darwin's ideas, and indeed their proponents rarely invoked Darwin in support. The use of the term "social Darwinism" as a critique of capitalist ideologies was introduced in Richard Hofstadter's ''Social Darwinism in American Thought'' published in 1944.


Anarchists

Russian zoologist and anarchist Peter Kropotkin viewed the concept of "survival of the fittest" as supporting co-operation rather than competition. In his book '' Mutual Aid: A Factor of Evolution'' he set out his analysis leading to the conclusion that the fittest was not necessarily the best at competing individually, but often the community made up of those best at working together. He concluded that
In the animal world we have seen that the vast majority of species live in societies, and that they find in association the best arms for the struggle for life: understood in its wide Darwinian sense – not as a struggle for the sheer means of existence, but as a struggle against all natural conditions unfavourable to the species. The animal species, in which individual struggle has been reduced to its narrowest limits, and the practice of mutual aid has attained the greatest development, are invariably the most numerous, the most prosperous, and the most open to further progress.
Applying this concept to human society, Kropotkin presented mutual aid as one of the dominant factors of evolution, the other being self-assertion, and concluded that
In the practice of mutual aid, which we can retrace to the earliest beginnings of evolution, we thus find the positive and undoubted origin of our ethical conceptions; and we can affirm that in the ethical progress of man, mutual support not mutual struggle – has had the leading part. In its wide extension, even at the present time, we also see the best guarantee of a still loftier evolution of our race.


Tautology

"Survival of the fittest" is sometimes claimed to be a tautology. The reasoning is that if one takes the term "fit" to mean "endowed with phenotypic characteristics which improve chances of survival and reproduction" (which is roughly how Spencer understood it), then "survival of the fittest" can simply be rewritten as "survival of those who are better equipped for surviving". Furthermore, the expression ''does'' become a tautology if one uses the most widely accepted definition of "fitness" in modern biology, namely reproductive success itself (rather than any set of characters conducive to this reproductive success). This reasoning is sometimes used to claim that Darwin's entire theory of evolution by natural selection is fundamentally tautological, and therefore devoid of any explanatory power. However, the expression "survival of the fittest" (taken on its own and out of context) gives a very incomplete account of the mechanism of natural selection. The reason is that it does not mention a key requirement for natural selection, namely the requirement of ''heritability''. It is true that the phrase "survival of the fittest", in and by itself, is a tautology if fitness is defined by survival and reproduction. Natural selection is the portion of variation in reproductive success that is caused by ''
heritable Heredity, also called inheritance or biological inheritance, is the passing on of traits from parents to their offspring; either through asexual reproduction or sexual reproduction, the offspring cells or organisms acquire the genetic informa ...
'' characters (see the article on
natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the heritable traits characteristic of a population over generations. Cha ...
). If certain heritable characters increase or decrease the chances of survival and reproduction of their bearers, then it follows mechanically (by definition of "heritable") that those characters that improve survival and reproduction will increase in frequency over generations. This is precisely what is called "
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
by
natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the heritable traits characteristic of a population over generations. Cha ...
". On the other hand, if the characters which lead to differential reproductive success are not heritable, then no meaningful evolution will occur, "survival of the fittest" or not: if improvement in reproductive success is caused by traits that are not heritable, then there is no reason why these traits should increase in frequency over generations. In other words, natural selection does not simply state that "survivors survive" or "reproducers reproduce"; rather, it states that "survivors survive, reproduce and ''therefore'' propagate any ''heritable'' characters which have affected their survival and reproductive success". This statement is not tautological: it hinges on the
testable Testability is a primary aspect of Science and the Scientific Method and is a property applying to an empirical hypothesis, involves two components: #Falsifiability or defeasibility, which means that counterexamples to the hypothesis are logicall ...
hypothesis that such fitness-impacting heritable variations actually exist (a hypothesis that has been amply confirmed.) Momme von Sydow suggested further definitions of 'survival of the fittest' that may yield a testable meaning in biology and also in other areas where Darwinian processes have been influential. However, much care would be needed to disentangle tautological from testable aspects. Moreover, an "implicit shifting between a testable and an untestable interpretation can be an illicit tactic to immunize natural selection ... while conveying the impression that one is concerned with testable hypotheses". Skeptic Society founder and ''
Skeptic Skepticism, also spelled scepticism, is a questioning attitude or doubt toward knowledge claims that are seen as mere belief or dogma. For example, if a person is skeptical about claims made by their government about an ongoing war then the ...
'' magazine publisher
Michael Shermer Michael Brant Shermer (born September 8, 1954) is an American science writer, historian of science, executive director of The Skeptics Society, and founding publisher of ''Skeptic'' magazine, a publication focused on investigating pseudoscientifi ...
addresses the tautology problem in his 1997 book, ''
Why People Believe Weird Things ''Why People Believe Weird Things: Pseudoscience, Superstition, and Other Confusions of Our Time'' is a 1997 book by science writer Michael Shermer. The foreword was written by Stephen Jay Gould. Summary In the first section, Shermer discusses t ...
'', in which he points out that although tautologies are sometimes the beginning of science, they are never the end, and that scientific principles like natural selection are
testable Testability is a primary aspect of Science and the Scientific Method and is a property applying to an empirical hypothesis, involves two components: #Falsifiability or defeasibility, which means that counterexamples to the hypothesis are logicall ...
and falsifiable by virtue of their predictive power. Shermer points out, as an example, that population genetics accurately demonstrate when natural selection will and will not effect change on a population. Shermer hypothesizes that if hominid fossils were found in the same geological strata as
trilobite Trilobites (; meaning "three lobes") are extinct marine arthropods that form the class Trilobita. Trilobites form one of the earliest-known groups of arthropods. The first appearance of trilobites in the fossil record defines the base of the ...
s, it would be evidence against natural selection. Shermer, Michael; ''
Why People Believe Weird Things ''Why People Believe Weird Things: Pseudoscience, Superstition, and Other Confusions of Our Time'' is a 1997 book by science writer Michael Shermer. The foreword was written by Stephen Jay Gould. Summary In the first section, Shermer discusses t ...
''; 1997; Pages 143–144


See also

* * * * * * * * * * * * * * * * * * * *
Red Queen hypothesis The Red Queen hypothesis is a hypothesis in evolutionary biology proposed in 1973, that species must constantly adapt, evolve, and proliferate in order to survive while pitted against ever-evolving opposing species. The hypothesis was intended ...
* *


Notes


References


External links


Origins of the phrase


AboutDarwin.com — Darwin's Timeline





Tautology links



by Stephen Jay Gould
Evolution and Philosophy: A Good Tautology is Hard to Find
by John Wilkins, part of the talk.origins archive.
CA500: "Survival of the fittest is a tautology"
from the talk.origins index to creationist claims by Mark Ridley.

by Don Lindsay.

by the Doubting Thomas


Morality link



* ttps://web.archive.org/web/20051122134556/http://www.anselm.edu/homepage/dbanach/dnr.htm David Hume — Dialogues Concerning Natural Religion
Evolution and philosophy — Does evolution make might right?
by John S. Wilkins.


Kropotkin: Mutual Aid

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