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https://en.wikipedia.org/wiki/Spandrel_(biology)

The term spandrel was an architecture term originally thought up during the Roman period to explain the triangle area between two arches that come together. Stephan Jay Gould, a paleontologist at Harvard, and Richard Lewontin, a population geneticist, originally coined this term for the use of explaining secondary byproducts of adaptions and were not necessarily adaptations themselves. The spandrel was an architectural space created as part of the gap between the area where two arches came together and the ceiling of the building. These spandrels as they were named did not actually come into use into later on when artists realized they could make designs and paint in these small areas adding to the overall design of the building.

It is thought in the scientific community today that everything an animal has developed that has a positive effect on that animal’s fitness was due to natural selection or some adaptation. Gould and Lewontin propose an alternative hypothesis to this saying that due to adaptation and natural selection, byproducts are formed in addition. These byproducts of adaptations that had no real relative advantage to survival were termed as spandrels.

Gould and Lewontin rebuted certain counter arguments that stated spandrels were just small unimportant byproducts but stated that, “we must not recognize that small means unimportant. Spandrels can be as prominent as primary adaptions” (Gould and Lewontin).

A main example used by Gould and Lewontin is the example of the human brain. It is explained that the human brain is the area in humans that is thought to have the most spandrels. So many secondary processes and actions come in addition to the human brain and its main functions. These secondary processes and thoughts are the spandrels of the human body, which eventually through thought and action can turn into an adaption or fitness advantage to humans. Just because something is a secondary trait or byproduct of an adaption does not mean it has no use because it has the possibility to eventually be used as something beneficial towards the animal.

Final Draft
Spandrel Biology The term spandrel was an architecture term originally thought up during the Roman period to explain the triangle area between two arches that come together. Stephan Jay Gould, a paleontologist at Harvard, and Richard Lewontin, a population geneticist, originally coined this term for the use of explaining secondary byproducts of adaptions and were not necessarily adaptations themselves. Gould and Lewontin explained and introduced their newly formed idea in their controversial paper, “The Spandrels of San Marco and the Panglossian Paradigms: A Critique of the Adaptionist Programme.” The spandrel was an architectural space created as part of the gap between the area where two arches came together and the ceiling of the building. These spandrels as they were named did not actually come into use into later on when artists realized they could make designs and paint in these small areas adding to the overall design of the building. The example mentioned above is the main example Gould and Lewontin used to argue their hypothesis. It is thought in the scientific community today that everything an animal has developed that has a positive effect on that animal’s fitness was due to natural selection or some adaptation. Gould and Lewontin propose an alternative hypothesis to this saying that due to adaptation and natural selection, byproducts are formed in addition. These byproducts of adaptations that had no real relative advantage to survival were termed as spandrels. Gould and Lewontin later expanded their hypothesis to state that eventually these spandrels, or secondary byproducts, could be of use to the organism thus improving its fitness and becoming an adaptation of itself. This is important in providing another explanation for traits and phenotypes to arise in animals, even those that may eventually become beneficial to the animal, through other means than just adaption. This theory proposed by both Gould and Lewontin has been a topic of high controversy and continues to be today forty years after its original proposal. One major argument against the use of spandrels in the evolutionary biology standpoint is the argument of “Nooks and Crannies.” This argument goes into discussion about that fact that yes, spandrels are bound to arise as unintended consequences of adaptions. What if these byproducts are just tiny bits and pieces that lay in the nooks and crannies of an animal’s primary structure? These spandrels do exist but the argument is that these do not matter in the overall structure of the animal. Gould and Lewontin rebut this claim stating that, “we must not recognize that small means unimportant. Spandrels can be as prominent as primary adaptions” (Gould and Lewontin). The area covered by any of the spandrels under any dome in San Macro does not differ from the dome itself, both are substantial to the overall design. Gould and Lewontin compare the byproducts of animals to the spandrels. These byproducts are small and originally thought to be of no use just as the spandrels were. Then artists realized they could paint and design these spaces causing them to add to the overall design of the building just as the byproducts may be small things but can contribute to the overall design of the animal. Another more popular argument addressed by Gould and Lewontin is the argument that “Secondary Means Unimportant” (Gould and Lewontin). In this argument, critics argue that spandrels can never be important to the overall structure and fitness of animals because they arise secondly as consequences to more prominent adaptations. Gould and Lewontin counter argue this point by example of the human brain. It is explained that the human brain is the area in humans that is thought to have the most spandrels. So many secondary processes and actions come in addition to the human brain and its main functions. These secondary processes and thoughts are the spandrels of the human body, which eventually through thought and action can turn into an adaption or fitness advantage to humans. Just because something is a secondary trait or byproduct of an adaption does not mean it has no use because it has the possibility to eventually be used as something beneficial towards the animal. Additional criticism claimed that Gould and Lewontin’s spandrels were nothing more than pendentives, which is another architectural term that refers to a space leftover. Gould and Lewontin responded by “outlining the consideration for grounds of assigning or denying a structure of a spandrel” (Hagan). A spandrel could have originated as a spandrel through a secondary trait of adaption, which was the most common case. On rare occasion though some spandrels primary function were not seen or adapted for its current use for multiple generations. Gould and Lewontin came to the conclusion that how well a structure is crafted for a specific function cannot be used for telling whether a spandrel is really a spandrel because their true use could not have been seen yet. The failure to recognize the importance and central role of spandrels in animals and the importance of other methods of gaining fitness advantages by ways other than adaptation in relation to evolutionary biology is the principal point of spandrel biology. This has been the basis of a new evolutionary theory of thought in the hopes of explaining secondary traits and the true importance of them in animal adaption. Spandrels are important promoters in addition to that of adaption and natural selection. Gould and Lewontin state, “ We are not trying to derail the effort to establish a true “evolutionary psychology” on genuine Darwinian principles or even to overthrow the centrality of adaptation in evolutionary theory. We wish, rather, to enrich evolutionary theory by a proper appreciation of the interaction between structural channeling and functional adaptation” (Gould and Lewontin). The theory of spandrel biology itself is like that of an actual spandrel it is a secondary thinking to that of adaption and evolution trying to expand on the evolutionary thought process. Stephan Jay Gould and Richard Lewontin say something is missing when it comes to the process of natural selection and adaption. They say secondary characteristics and byproducts that most just threw aside calling them worthless of unimportant because they had no direct fitness advantage towards the animal. What happens when these secondary traits or byproducts became productive towards the overall survival of the animal? Gould and Lewontin focused the majority of their research on these traits coining the term spandrel. The term spandrel described the area between where two arches came together that was essentially thought to be wasted space until artist decided to decorate those areas. This is the thought of the secondary characteristics and byproducts of animals. Even though these traits might not be important now they could possibly be of advantage to the animal in the future. Gould and Lewontin argued that these spandrels were therefore not all new adaptions themselves but the byproducts of past adaptions that were just becoming of use to the animal. Ever since Gould and Lewontin have released their controversial theory of spandrel biology they and their theory have been the subject of a lot of criticism. Spandrel Biology is a very confusing subject when first learning about it. It is very hard to find articles on the web about Spandrel Biology as most have just tossed it aside. Most works on or about the subject of spandrel biology are papers or rebuttals to criticism by Gould and Lewontin themselves. Not many evolutionary biologists have elaborated or dug deeper into this proposed theory and those who have wrote on this topic have been the criticizer. Overall Gould and Lewontin make a good argument in their presenting of their theory the spandrel that still today is a fairly new and unknown topic in the field of evolutionary biology.

Reference Section 1.	Gould, Stephan J., and Richard Lewontin. "The Exaptive Excellence of Spandrels as a Term and  prototype." The Exaptive Excellence of Spandrels as a Term And prototype. The National Academy of Science, 30 Sept. 1997. Web. 28 Oct. 2014. 2.	Buss, David, Martie Haselton, Todd Shackelford, April Bleske, and Jerome Wakefield. "Adaptations, Exaptations, and Spandrels." Adaptations, Exaptations, and Spandrels. American Psychologist. Web. 11 Sept. 2014. 3.	Hagan, Edward. "What about Spandrels?" What about Spandrels? Institute For Theoretical Biology. Web. 13 Sept. 2014. 4.	Gould, Stephan, and Richard Lewontin. "The Spandrels of San Marco and the Panglossian Paradigm." The Spandrels of San Marco and the Panglossian Paradigm. The Royal Publishing Society, 21 Sept. 1979. Web. 13 Sept. 2014. 5.	Taylor, Christopher. "Catalogue of Organisms." Inevitable Spandrels. Catalogue of Organisms, 28 Aug. 2008. Web. 14 Sept. 2014. 6.	"Thoughts Explained: The Biological Spandrel." The Biological Spandrel. BlogSpot, 9 Feb. 2012. Web. 28 Oct. 2014. 7.	"It's a Spandrel Sort of . . .!" Why Evolution Is True, 29 Nov. 2009. Web. 28 Oct. 2014. 8.	Fox, Jeremy. "Why "The Spandrels of San Marco" Isn't a Good Paper." Oikos, 26 Aug. 2011. Web. 28 Oct. 2014.

October 1st Project
Article: https://en.wikipedia.org/wiki/Spandrel_(biology)

Three Suggestions: This article goes into detail about the naming of Spandrel, who named it, and the criticism of the thinking. This article must go into more detail on what Spandrel Biology actually is rather than going into its criticisms. This article is more about the criticisms rather than the actual topic. I would go into more detail on the Spandrel and instead of what it is being compared to what actually Spandrel Biology is. Also the language used in this article makes it very difficult to read and comprehend even as a college student. The language should be more general and use terms everyone can understand. This article could also use some examples of Spandrel Biology in the real world making it easier to comprehend and understand.

One Sentence: Evolutionary biology uses the term Spandrel for features arising as byproducts, rather than adaptations, that have no clear advantage towards and organisms fitness or survival.

Citation: Stephen Jay Gould (1997). "The Exaptive Excellence of Spandrels as a Term and Prototype" (http://www.pnas.org/content/94/20/10750.full)