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Ohno presented the hypothesis in , and it is about the logic of gene duplication events in evolutionary history. When a gene gets copied or duplicated in a genome , the new gene lacks the functional constraints of the old gene. In such cases, the duplicated copy can accumulate beneficial or neutral mutations with little deleterious effect on the overall fitness of the organism. This accumulation can lead to the re-arrangement of genetic elements, so that existing structures can acquire new functions.

This hypothesis of genomic change, Jacob argues, illustrates the process of tinkering. According to Jacob, molecular biologists have shown that most morphological change in vertebrates has not resulted from new structural genes , but rather it is the consequence of a change in the regulation of genetic components, including events like heterotopy, a change in the spatial location of developmental events, and heterochrony, a change in the timing of developmental events.

Jacob argues that these events occur in embryonic development according to the precise schedule of a genetic program, suggesting that gene regulation is the key factor in the generation of animals' forms and functions. In the ninth section, Jacob outlines two consequences of the metaphor of evolution as a process of tinkering. First, if his analogy holds, then biologists should expect to find similarities in the underlying molecular elements of different species.

For example, Jacob argues, biochemists have discovered hormone peptides that trigger a variety of chemical reactions in cells from organisms in different species. Second, biologists should expect to see many imperfections or redundancies in the design of organisms.

For instance, Jacob explains, the human reproductive system illustrates a less than perfect mechanism in which almost half of the total number of conceptions result in no viable fetuses. Jacob ends his essay with a final example of tinkering, arguing that the human brain is a product of highly contingent, historical events. Jacob contends that, in humans , the addition of the neo-cortex to the rhinencephalon, a primitive part of the brain responsible for the sense of smell and theorized to control instinct, has set the conditions for the evolution of the human brain.

The human brain is thus the result of an imperfect patchwork of a structure controlling visceral or emotional drives, the rhinencephalon, and a structure controlling more sophisticated cognitive abilities, the neo-cortex. This case can be extrapolated, Jacob argues, to a general rule for evolution : evolution is the net result of a particular sequence of historical opportunities. Jacob's essay had, at first, a mixed reception. Many biologists said that the description of evolution by natural selection as a process of tinkering was blatantly obvious.

In Walter Gehring and his team at the University of Basel in Basel, Switzerland, discovered of a standard set of DNA sequences called the Homeobox in genes that controlled the embryonic development of body plans of animals. Scientists soon found the Homeobox in genes of diverse organisms from flies to humans. Given those results, scientists explicitly began referring to Jacob's essay and to his concept of tinkering.

In , Jacob published a series of lectures given at the University of Washington in Seattle, Washington, under the title The Possible and the Actual , which includes a slightly modified version of his original essay, "Evolution and Tinkering," as well as some essays expounding his philosophy of science.

In , the Novartis Foundation in London, United Kingdom, held a symposium on the concept of tinkering in evolution and development. By the second decade of the twenty-first century, scientists had cited "Evolution and Tinkering" thousands of times. Keywords: bricolage , tinkering. Sources Artmann, Stefan. Carroll, Sean B. Endless Forms Most Beautiful. The Possible and the Actual.

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