Innovation (evolution)

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A phenotypic innovation is a new trait or behavior in evolutionary tribal history that cannot be explained solely as a variation of existing ancestral traits. Innovation is thus differentiated from phenotypic variation , the modulation of an already existing feature.

Differentiation between variation and innovation

Innovation in evolution is distinguished from variation, in each case related to the phenotype. Variation is explained by Charles Darwin and the synthetic theory of evolution mainly through natural selection and adaptation . The synthetic theory of evolution analyzes statistical changes in the gene frequency in populations on the basis of population genetic analyzes . Phenotypic variation is taken for granted in standard theory.

In contrast, evolutionary developmental biology ( Evo-Devo ) in particular has been investigating since the early 1990s to what extent embryonic development could explain the emergence of innovative features under ecological conditions and how their realization in the organism and their permanent anchoring (genetic / epigenetic integration) be understood.

In general, however , the term innovation is often used synonymously with variation when describing important evolutionary changes. The often blurred boundaries can lead to misunderstandings. According to Müller, innovation always means a qualitative deviation from the previous properties, and not just a quantitative one.

Examples

Fig. 1 The turtle shell is a Type II evolutionary innovation. It required complicated parallel skeletal modifications.

Real innovations occur less often in evolution than variations. However, their significance for evolution is often great, since in evolutionary history with an innovation, especially with a key innovation (see below), a long-lasting adaptive radiation at higher taxonomic levels often occurs. The following characteristics are referred to as phenotypic innovation in the literature:

Morphology:

Physiology:

  • Constant body temperature in warm-blooded animals

Behavior:

Definitions and demarcations

Ernst Mayr , one of the main proponents of synthetic theory, sees 1963 as an evolutionary innovation “every newly acquired structure or property that allows a new function to be performed, which in turn opens up a new adaptive zone”. This definition also allows a new function to come about through pure quantitative accumulation of already existing feature elements and not just through qualitative innovation. Today the latter is more likely to be equated with innovation.

Müller / Wagner 1991 stand out more on development, i.e. on the path of innovation ( novelty ). You define innovation as “a construction element in a building plan that has neither a homologous counterpart in the forerunner type nor in the same organism”. This definition excludes purely quantitative changes in existing characteristics. It allows a view of features that are either completely new or result from new combinations or subdivisions of already existing structures (e.g. the panda's thumb ). Furthermore, the Müller / Wagner definition allows it to be applied to clearly delimited, unambiguous cases not only in terms of morphology , but also in terms of physiology or behavior . Similarly, West-Eberhard speaks of “a new feature based on a qualitatively new development variant”. In order to include ecology and take into account that not only discontinuous features are dealt with, Massimo Pigliucci explains innovations in 2008 as follows: “Evolutionary innovations are new features or behaviors, or new combinations of pre-existing traits or behaviors that arise in the evolution of a lineage and that show a new function in the ecology of that lineage ”. This definition in turn makes the change of function more prominent. Furthermore, the discontinuity highlighted by Evo-Devo should not be overemphasized and rather continuous constructional changes should be included in the theory of evolution.

In the context of evolution, the term innovation is used in the following areas in addition to the fields of morphology, physiology and behavior mentioned:

Key innovation

Key innovation ( key innovation as Mayr also - -) of certain innovations in macroevolutionary process and refers to the prominent role of the adaptive radiation . Examples of key innovations are bipedality, the eye, mammary glands in mammals, and others. The concept of key innovation is always to be seen in an important reciprocal connection between its internal construction, the corresponding environmental factors and the population. In this context, they enable the utilization of new energy resources and the radiative exploration of new ecological niches. This becomes clear in the example of the birds in the wake of the innovation of the wings and the number of more than 9000 birds. If the key innovation is called the wing in a taxonomically neutral way, the meaning is even more extensive, as wings have developed independently several times ( insects , pterosaurs , birds , bats ) and have thus led to the adaptive radiation of even more niches.

Innovation therefore aims more at the constructive development, while key innovation means more the diversity as a result of the cascading effects of an innovation .

Classification of forms of innovation

Gerd B. Müller distinguishes three types of phenotypic innovation:

  • Type I Innovation : Metazoa Primary Anatomical Architecture
  • Type II Innovation : Discreet new element added to an existing blueprint
  • Type III innovation : Extreme variation of an existing building plan feature

Type I innovation is based on basic physical requirements for the different forms of aggregation of cells in multicellular organisms (adhesion, diffusion, oscillation etc.) as described by Stuart A. Newman as Dynamic Patterning Modules (DPMs). Such multicellular basic forms can be understood evolutionarily as premendelic, which emerge only after a certain scaling. Structures are not yet strictly inherited by the genotype, but their realization in each generation would be strongly determined by the prevailing environmental conditions.

Type II - From Evo-Devo's point of view, this type of innovation is particularly important because of the qualitative aspect. Examples of type II innovations are the turtle shell or the feather.

Type III innovation : Here the new trait emerges from an existing phenotypic trait, e.g. B. the narwhal tusk. The innovation can have a major new function and result in taxonomic apomorphism . If new construction elements appear in the development, standard variation in such a qualitative, phenotypic innovation cannot be seen as the cause; what is needed is a specific explanation for their origin.

Development as the main factor in phenotypic innovation

A distinction is made between three factors of evolutionary phenotypic innovation:

Initiation conditions

"The selection cannot attack features that do not yet exist and therefore do not immediately cause innovation". West-Eberhard emphasizes: In the majority of cases, it is the changed environmental conditions that give the impetus for evolutionary innovation. This applies both to the formation of the vertebrate skeleton and to the formation of the exoskeleton of marine life (sea urchins), the latter when the water is enriched with calcium. West-Eberhard sees the reason that environmental factors predominate as initiating stressors in the fact that they often last for many generations and have a broad effect, possibly long and simultaneously on the entire population. This facilitates the evolutionary development of innovation.

Possible initiating factors that can trigger phenotypic innovation:

  1. Rare genetic mutations that form the raw material for a spontaneous structure effect that emerges during development. Such mutations could possibly have been the basis for the development of the feather, new macroscopic properties based on a new material (φ- keratins ).
  2. Symbiotic union of previously separated genetic or developmental components. Endosymbiosis and horizontal gene transfer are mainly known in the evolutionary development of eukaryotes . The transfer is not limited to short gene segments, but can also include the transfer of complete genetic modules or development modules between two species.
  3. Epigenetic by-products ( side effect hypothesis ) are structures that arise when selection is based on other characteristics or levels than those that lead to innovation itself. Darwin already pointed out the side effects of selection in 1859. Selection can, for example, affect parameters such as cell division rates , cell and tissue signal exchange or the timing of different developmental processes and thereby bring the system to a threshold at which new structures automatically emerge from the reaction options of the development modules concerned. Examples of epigenetically created by-products are new skeletal elements. The origin of the bird feather is explained by Josef H. Reichholf in this way. Reichholf emphasizes z. B. the excess of food components as a possible initiator for innovation.

Realization conditions

If the initiation conditions for the innovation are unspecific and general (nutrition etc.), the implementation conditions must necessarily lie in the development. Here you can see the phenomenon of threshold effects, which means that small initial conditions can cause non-linear, phenotypic variation / innovation. The change of a single parameter leads to the response of the whole system development to the disturbing factor. This response of the system development can generate innovations. The innovation / construction process in one of the above examples will in many cases be multi-stage, i.e. H. the phases initiation - realization - integration are run through several times and function changes (see below) may occur several times.

Integration and fixation conditions

Here one asks how the new phenotypic element (additional fingers, etc.) is genetically and epigenetically fixed or assimilated so that it lasts and is completely or almost completely decoupled from the triggering (environmental) stressor. "The innovation feature must be anchored in the existing construction, development and genome system in order to ensure functionality and inheritance". "The rule seems to be that the epigenetic integration precedes the genetic integration" or as West-Eberhard puts it: "Genes are followers in evolution" "The genetic integration stabilizes and fixes newly generated characteristics and results in an ever closer mapping between Genotype and phenotype "(Müller).

Environmental factors play an essential role in all phases of innovation. Müller therefore also speaks of epigenetic innovation . A minor disruptive factor, either an environmental stressor or a genetic mutation, can provoke a response from the entire system of development when the channeled plasticity reaches its limits. This response can be non-linear or non-gradual (discontinuous) due to the ability of the development system to organize itself. In this scenario, the specific form of expression of the morphological result is not dictated by the selection, but by the reaction norm of the development system.

The emergence of innovations is therefore not an adaptive process, but natural selection only has a secondary effect on a design result of the development. This is a core statement of the epigenetically understood Evo-Devo research and theory, as represented by Müller , Kirschner and Gerhard, West-Eberhard.

Constraints

Fig. 3 The insect wing is an evolutionary, phenotypic innovation type II. It could only serve as a wing after several changes of function (see evolution of the insect wing )

Constraints limit the phenotypic evolution and determine the direction of its course. They can be physical, morphological or phylogenetic in nature. A distinction is made between external and internal constraints. In the latter context, special mention should be made of the development constraints. Conrad Hal Waddington calls the phenomenon channeling . The type and extent of how constraints can be broken and overcome play a decisive role in how evolutionary innovation can arise in development. If developmental paths are strongly channeled, there is, in Waddington's sense, a buffering of genetic mutations that work towards maintaining the status quo in the phenotype. This can mean that development is incapable of responding with variation, even with high selection pressure, and that is precisely why it is forced to exceed threshold values, which leads to increased opportunities for innovation

Function change

Main article: Exaptation

Complex innovations often did not arise in the functions that they fulfill in recent organisms. In this way, insect wings emerged in several stages, initially presumably as rudimentary body appendages that could have grown from excess material and that in the first stage served as aquatic arthropods as lamellae. Later they could be used as sails when gliding on smooth surfaces and only building on them, after further transformations, as wings. Stephen J. Gould writes on the subject of a change in function "Every part of nature has served almost every component of every living being under slightly different conditions, probably different purposes and has been active in the living machinery of many older and very special forms."

While such examples describe long-term processes, the above-mentioned Evo-Devo concept with the phases initiation - realization - integration on the basis of threshold value effects and the non-linear responsiveness of the development systems also allows rapid morphological transitions (see Evo-Devo ).

It can be assumed that the innovation examples mentioned here, such as mammary glands or two-leggedness, represent complex, multi-stage innovation processes that each individually fulfill the spontaneous constructional characteristic of development, but only lead to what is what in cascaded stages, possibly with several function changes they can be seen in the examples. For Gerhard Vollmer , it is essential that a feature temporarily performs two or more functions at the same time. Lewontin and Gould in particular have drawn attention to how difficult it is to assign a clear evolutionary function to individual characteristics and behaviors and that there is a risk of adopting adaptionist thinking for each characteristic and its functions uncritically.

Behavioral innovation in animals

Fig. 4 In 2005 it was observed for the first time how a gorilla used a stick to feel the water depth and as a walking aid.

Animals invent innovative forms of behavior. The aim here is to understand how these can develop evolutionarily and whether there are common explanatory patterns. A distinction is made between emotional, cognitive, socially and ecologically new behavior. Raeder and Laland regard behavioral innovation as a process and a product and define innovation as "a process which leads to new or modified learned behavior and which transfers the new behavior variants into the repertoire of a population". Innovative behavior as a process includes individual and social learning. Here, social learning is seen as having a better chance of establishing itself in the population. Raeder and Laland therefore mainly report innovative behavior on primates and songbirds. The general mental and emotional makeup of an individual is relevant to whether and how often individuals appear with new forms of behavior. The Dutch behavioral scientist Carel van Schaik identifies three outstanding patterns for the occurrence of behavioral innovation in social animals:

  • the disposition of an individual
  • the social environment
  • the ecological circumstances

According to van Schaik, ecological pressure is necessary for new behaviors to emerge. It is a possible initiation condition (see above). New forms of behavior can then possibly become vital for the species. This is especially true if a species is unable to leave or enlarge its habitat. "Large ecological diffusion is likely to be positively correlated with innovation because there are new challenges and opportunities in an ecologically large area. However, innovation does not have to be correlated with extensive geographical or ecological breadth: sometimes innovation can be the migration or exploration tendency of new settlements Limiting areas because innovation expands the existing niche or opens up more efficient exploitation and thus allows individuals to remain. It is therefore assumed that there is an asymmetrical correlation between the spread of a species and innovation ".

According to Jablonka and Avital, difficulties can arise in the clear delimitation of new context-dependent, cognitive forms of behavior from other existing forms of behavior and their functions. The ability to learn and the willingness to innovate cannot be analyzed in isolation. Innovation results from the highly integrated interaction of complex cognitive processes and circumstances. Their underlying physiological and neurological mechanisms cannot be differentiated from other such processes, but still require the consideration of a few other processes, such as certain emotional states in specific learning situations.

The question of whether the inheritance of social learning promotes conservative or innovative behavior, or when which is not cleared today. The decisive questions seem to be about the development dimensions: At what age are innovative forms of behavior most likely? At what gender, social status or ecological stress? What requirements are there for cognitive mechanisms that not only allow the formation of new innovations that have been learned, but also lead to their functional organization? The finished arbor of the bowerbird must first be copied from older animals, then built by oneself, then used for advertising the female, in order to achieve the “goal” after several years of failures and only with increasingly improved construction technology Female is also accepted.

The use of a stick by a single gorilla (Fig. 4) is not yet an evolutionary innovation. For this, evidence must be provided that the form of behavior has been adopted in the repertoire of the population (see definition above). Jablonka and Lamb describe the inheritance (passing on) of behavior. It is necessary to recognize cognitive abilities and readiness potential, which, as mentioned above, cannot be analyzed separately. Animals do not imitate in the way and to the extent that humans imitate, as the American anthropologist and behavioral scientist Michael Tomasello was able to demonstrate in his empirical studies. The ability to imitate is necessary for the development of culture in humans (see below. Cultural evolution), so imitation can be seen as a human innovation. Animals ( e.g. bowerbird ) observe the behavioral products and less the behavioral processes of experienced animals and develop their own similar behavior from this. With the demonstration of such behavior forms and differences, it is not yet proven that or how the respective behavior works evolutionarily (adaptively) and how it contributes to biological fitness in the Darwinian sense. The evolutionary adaptive properties of innovative forms of behavior have hardly been studied in animals. The Evo-Devo view that innovation is a product of development enforces analogous to that in Chap. 6, also with innovative behaviors ( novelties ) to consider their possible non-adaptive emergence in the course of behavior development. The close connection between development and behavioral change is shown in the Evo-Devo chapter Selected empirical research results and there the section on Belyaev's Taming of Silver Foxes . Adaptive behavior, however, is the subject of Peter Richerson and Robert Boyd's theory of the evolution of human culture.

Culture as innovation

The cultural ability of humans is seen as a unique selling point of our species. Both language and culture themselves are considered evolutionary innovations. Richerson and Boyd describe how culture can evolve adaptively. They are not interested in explaining how cultural services and products have developed, such as the car, the Internet or Gothic cathedrals. That would be evolution in culture . But the authors mean the evolution of culture. Karl Eibl pointed out the difference .

Fig. 5 The interactive system of cultural evolution. Richerson and Boyd explain culture as the process of complex genome / epigenome relationships, environmental conditions, natural selection and the accumulation of inheritable knowledge.

Possible initiation conditions for the development of culture are described on a broad basis in the literature. The arguments range from the upright gait and the resulting refinement of the hands, the enlargement of the brain, the use of tools, the development of language to the invention of agriculture and sedentarism. All factors are also discussed in their interaction. For Richerson / Boyd, the ability to imitate humans and the ability to accumulate knowledge are specifically human and necessary (insufficient) prerequisites for the development of culture. Wolf Singer goes back on this question to the decisive steps of the brain evolution. For him the brain of the "pre-cultural" human being in its fundamental structure is no different from that of homo sapiens sapiens . Singer sees the differences in the "epigenetic design and fine structure" and the "cognitive abilities that are acquired and passed on through this epigenetic path" as "more as a consequence of cultural evolution and collective learning and less (as) its cause". Thus, the question of "the evolutionary development of certain cognitive functions ... which only belong to humans" applies. Singer sees this primarily in the ability to create abstract, symbolic versions. - to create so-called meta-representation of cognitive content "." We are the only species that is able to conduct dialogues in which it can say: 'I know that you know that I know' or 'I know that you know what I feel ' Singer describes the neocortical functions that make such abstract thinking and its representation in our brain possible.

Richerson / Boyd describe how culture can be realized as follows: Humans are capable of cumulative cultural evolution. "People can add one innovation to another until the results resemble organs with extreme perfection, such as an eye." Michael Tomasello referred to this as the jack effect . Second, humans combine individual learning and social learning (imitation). Knowledge can be accumulated through either of the two forms of learning. However, it is only through the possibility of selective learning , that is the combination of the two forms of learning in a decision-making situation that is advantageous from a cost-benefit perspective, but thirdly, there is accumulated knowledge that can be an object of natural selection. Learning per se cannot be interpreted as adaptive. In other words, learning alone does not improve fitness. Even if knowledge is constantly increased and passed on through individual or school learning, according to Richerson and Boyd this is not demonstrably adaptive. At the level of the human species, this can only be achieved by selection, which selects the individuals with the most advantageous combinations of both forms of learning, whereby the population adapts.

Richerson / Boyd define evolution through culture as follows: "By cultural evolution we understand behavior or artifacts that are transmitted and modified over many generations and that lead to complex artifacts or behavior". Through culture, people change their own living conditions (environment). The changed environment also affects the genome through epigenetic processes. Thus, through their own actions, humans also change and shape their genome and thus in turn their morphological and behavioral evolution (see niche construction ). Both mechanisms are closely interwoven and cannot be separated from one another. Thus the authors come to the conclusion: "Adaptation through cumulative cultural evolution is not a by-product of intelligence and social life".

Culture itself is an evolutionary human characteristic. In essence, this is the result of Richerson and Boyd's view of the interactive processes involved (see Fig. 5). Culture is an evolutionary process in the sense that people decide through selective learning whether they learn more individually or more socially through school, training, etc. and thus imitate others, if that is more cost-effective and efficient. The result of the combination of the two forms of learning in the interaction of human - environment - genome is that, in the Darwinian sense according to Richerson and Boyd, is subject to selection and can adaptively improve the biological fitness of humans at the population level. Temporary mismatches ( maladaptations ) as falling birth rate, destruction of resources, etc. can not be excluded in and contradict not the understanding of Darwinian evolution principle.

Fig. 6 Javelin of the
Anazasi culture. Even the production of a spear like this one, as well as its practiced handling, require to a high degree the inheritance of accumulated knowledge from individual and social learning over many generations. The shaft must be drawn exactly straight, a hewn, slender stone tip must be permanently mounted, and the length, alignment, balance and throwing technique of the weapon must be patiently tested. The evolutionary prerequisites for such innovative cultural, human achievements are described in Fig. 5 and in the text.

See also

Evolutionary developmental biology

Evolution of the bird feather

Evolution of the insect wing

Hominization

Individual evidence

  1. a b c d e f Gerd B. Müller: Novelty and Key Innovations. In: Mark Pagel (Ed.): Encyclopedia of Evolution. Oxford University Press, 2002, pp. 827-830.
  2. Gerd B. Müller: Epigenetic Innovation. In: Massimo Pigliucci, Gerd B. Müller (Ed.): Evolution - The Extended Synthesis. MIT Press, 2010, p. 311.
  3. ^ Massimo Pigliucci: What, if anything, Is an Evolutionary Novelty? In: Philosophy of Science. Volume 75, December 2008, p. 896.
  4. a b Carel van Schaik: The evolution of innovation. In: Technology Review Archives. 2/2006, p. 88.
  5. ^ Carel van Schaik: Groups make you smart. In: The time. November 6, 2008, No. 46.
  6. a b c d e J. R. Richerson, R. Boyd: Not by Genes Alone. How Culture Transformed Human Evolution. University of Chicago Press, 2005.
  7. ^ Massimo Pigliucci: What, if anything, Is an Evolutionary Novelty? In: Philosophy of Science. Volume 75, December 2008, p. 898.
  8. ^ Gerd B. Müller, Günter P. Wagner: Novelty in Evolution: Restructuring the Concept. 1991, p. 243.
  9. Gerd B. Müller: Epigenetic Innovation. In: Massimo Pigliucci, Gerd B. Müller (Ed.): Evolution - The Extended Synthesis. MIT Press, 2010, p. 312.
  10. ^ Mary Jane West-Eberhard: Development Plasticity and Evolution. Oxford University Press, 2003, p. 198.
  11. Massimo Pigliucci: What, if Anything, Is an Evolutionary Novelty? In: Philosophy of Science. Volume 75, 2008, p. 890.
  12. See also: Nick Lane: Amazing Inventions of Evolution . 2nd edition Darmstadt: Konrad Theiss Verlag 2017.
  13. Karel F. Liem: Evolutionary strategies and morphological innovations: Cichlid pharyngeal jaws. In: Systematic Zoology. Volume 22, No. 4, December 1973, pp. 425-441.
  14. ^ F. Galis, EG Drucker: Pharyngeal Biting Mechanics in Centrarchid and Cichlid Fishes: Insights into a Key Evolutionary innovation. In: Journal of Evolutionary Biology. Volume 9, 1996, pp. 641-670. (A detailed study of a key innovation in fish evolution).
  15. Gerd B. Müller: Epigenetic Innovation. In: Massimo Pigliucci, Gerd B. Müller (Ed.): Evolution - The Extended Synthesis. MIT Press 2010, p. 314ff.
  16. ^ Stuart A. Newman: Dynamic Patterning Modules. In: Massimo Pigliucci, Gerd B. Müller (Ed.): Evolution - The Extended Synthesis. MIT Press, 2010, pp. 281ff.
  17. ^ G. Müller, S. Newman: The innovation triad. An Evo-Devo agenda. In: Journal of Experimental Zoology. 304B, 2005, pp. 387-503.
  18. Gerd B. Müller: Epigenetic Innovation. In: M. Pigliucci, Gerd B. Müller (Ed.): Evolution The Extended Synthesis. MIT Press, 2010, p. 313.
  19. Gerd B. Müller: Epigenetic Innovation. In: M. Pigliucci, Gerd B. Müller (Ed.): Evolution - The Extended Synthesis. MIT Press, 2010, p. 314ff.
  20. ^ Gerd B. Müller, Stuart A. Newman: The innovation triad. On EvoDevo agenda. In: Journal of Experimental Zoology. Volume 304B, 2005, p. 491.
  21. ^ MJ West-Eberhard: Development Plasticity and Evolution. Oxford University Press, 2003, pp. 500ff.
  22. ^ MJ West-Eberhard: Development Plasticity and Evolution. Oxford University Press, 2003, p. 501.
  23. Gerd B. Müller: Epiegentic Innovation. In: Massimo Pigliucci, Gerd B. Müller (Ed.): Ecolution - The Extended Synthesis. MIT Press, 2010, p. 307.
  24. Josef H. Reichholf: The invention of the pen. In: Andreas Sentker, Frank Wigger (Hrsg.): The driving force of evolution - diversity, change and becoming human. 2008, p. 99ff.
  25. ^ Gerd B. Müller, Stuart A. Newman: The innovation triad. On EvoDevo agenda. 2005, p. 492.
  26. ^ Gerd B. Müller, Stuart A. Newman: The innovation triad. On EvoDevo agenda. 2005, p. 493.
  27. ^ A b c Gerd B. Müller, Stuart A. Newman: The innovation triad. On EvoDevo agenda. 2005, p. 494.
  28. ^ MJ West-Eberhard: Development Plasticity and Evolution. Oxford University Press, 2003, p. 157.
  29. Gerd B. Müller: Epigenetic Innovation. In: Massimo Pigliucci, Gerd B. Müller (Ed.): Evolution - The Extended Synthesis. MIT Press, 2010, pp. 307-332.
  30. Gerd B. Müller: Epigenetic Innovation. In: Massimo Pigliucci, Gerd B. Müller (Ed.): Evolution - The Extended Synthesis. MIT Press, 2010, p. 323.
  31. C. H. Waddington: Canalisation of development and the inheritance of acquired characters. In: Nature. Volume 150, 1942, p. 563 ff.
  32. ^ Gerd B. Müller, Stuart A. Newman: The innovation triad. On EvoDevo agenda. 2005, p. 493.
  33. ^ Gerd B. Müller, Stuart A. Newman: The innovation triad. On EvoDevo agenda. 2005, p. 494. (with reference to Averof & Cohen 1997 and Marden & Kramer 1994)
  34. Stephen J. Gould: The Panda's Thumb - Reflections on Natural History. Suhrkamp TB Science, 1987 (original 1980).
  35. ^ Gerd B. Müller, Stuart A. Newman: The innovation triad. On EvoDevo agenda. 2005, p. 497.
  36. Gerhard Vollmer: How can we see the world? In: Peter Fischer, Klaus Wiegandt (Ed.): Evolution. History and future of life. Fischer Taschenbuch, 2003, p. 286.
  37. ^ Richard Lewontin, Steven Jay Gould: The spandrels of San Marco and the Panglossian paradigm: a critique of the adaptionist program. 1979.
  38. PJ Richerson, R. Boyd: Not by genes alone. How culture transformed human evolution. University of Chicago Press, 2005, p. 137.
  39. ^ Eva Jablonka, Eytan Avital: Animal Innovation: The Origins and Effects of New Learned Behaviors. In: Biology and Philosophy. 21, 2006, p. 135; book review of: SM Reader, KN Laland (Ed.): Animal Innovation. Oxford University Press, 2003.
  40. SM Reader, KN Laland (Ed.): Animal Innovation. Oxford University Press, 2003, cit. based on: Eva Jablonka, Eytan Avital: Animal Innovation: The Origins and Effects of New Learned Behaviors. 2006, p. 136.
  41. ^ Eva Jablonka, Eytan Avital: Animal Innovation: The Origins and Effects of New Learned Behaviors. 2006, p. 135.
  42. ^ Eva Jablonka, Eytan Avital: Animal Innovation: The Origins and Effects of New Learned Behaviors. 2006, p. 138 with reference to Raeder / Laland chap. 2-4
  43. ^ Eva Jablonka, Eytan Avital: Animal Innovation: The Origins and Effects of New Learned Behaviors. 2006, p. 137.
  44. ^ Eva Jablonka, Eytan Avital: Animal Innovation: The Origins and Effects of New Learned Behaviors. 2006, p. 139.
  45. ^ A b Eva Jablonka, Marion Lamb: Evolution in Four Dimensions. Genetic, Epigenetic, Behavioral, and Symbolic Variation in the History of Life. MIT-Press, 2006, p. 169.
  46. ^ Eva Jablonka, Marion Lamb: Evolution in Four Dimensions. Genetic, Epigenetic, Behavioral, and Symbolic Variation in the History of Life. MIT-Press, 2006, chap. 5: The Behavioral Inheritance System, pp. 155-191.
  47. ^ Michael Tomasello: The Chimpanzee Culture. 2007.
  48. to this chap. see also: Michael J. O'Brien , Stephen J. Shennan (Eds.): Innovation in Cultural Systems: Contributions from Evolutionary Anthropology (Vienna Series in Theoretical Biology). MIT Press, 2009.
  49. ^ Karl Eibl: Culture as an intermediate world. An evolutionary perspective. Suhrkamp, ​​2009, p. 99.
  50. Wolf Singer: The evolution of culture. In: Ernst Peter Fischer, Klaus Wiegandt: Evolution. History and future of life. Fischer TB, 2003, p. 301.
  51. Wolf Singer: The evolution of culture. 2003, p. 303.
  52. a b Wolf Singer: The evolution of culture. 2003, p. 304.
  53. a b Wolf Singer: The evolution of culture. 2003, p. 305.
  54. ^ Karl Eibl: Culture as an intermediate world. An evolutionary perspective. Suhrkamp, ​​2009, p. 38.
  55. ^ JR Richerson, R. Boyd: Not by Genes Alone. How Culture Transformed Human Evolution. University of Chicago Press, 2005, p. 109.