Hopeful monster

{{Evolution}}
'''Hopeful monster''' also known as the '''hopeful monsters hypothesis''' is a [[biological]] hypothesis which suggests that major evolutionary transformations have occurred in large leaps between species due to macromutations.

==History==

===Richard Goldschmidt=== 
The German geneticist Richard Goldschmidt {{lived|1878|1958}} was the first scientist to use the term "hopeful monster". Goldschmidt thought that small gradual changes could not bridge the hypothetical divide between [[microevolution and macroevolution]]. In his book ''The Material Basis of Evolution'' (1940) he wrote "the change from species to species is not a change involving more and more additional atomistic changes, but a complete change of the primary pattern or reaction system into a new one, which afterwards may again produce intraspecific variation by micromutation." Goldschmidt believed the large changes in [[evolution]] were caused by macromutations (large mutations). His ideas about macromutations became known as the hopeful monster hypothesis which is considered a type of [[saltational evolution]].<ref>Verne Grant ''The origin of adaptations'' 1963</ref> 

According to Goldschmidt "biologists seem inclined to think that because they have not themselves seen a 'large' mutation, such a thing cannot be possible. But such a mutation need only be an event of the most extraordinary rarity to provide the world with the important material for evolution".<ref>Prog Nucleic Acid Res&Molecular Bio by By J N Davidson, Waldo E. Cohn, Serge N Timasheff, C H Hirs 1968, p. 67</ref> Goldschmidt believed that the [[neo-Darwinism|neo-Darwinian]] view of [[gradualism|gradual]] accumulation of small [[mutation]]s was important but could only account for variation within species (microevolution) and wasn't a powerful enough source for the origin of evolutionary novelty to explain new [[species]]. Instead he believed that big [[genetic]] differences between species required profound "macro-mutations" a source for large genetic changes (macroevolution) which once in a while could occur as a "hopeful monster".<ref>Nick Lane ''Power, Sex, Suicide : Mitochondria and the meaning of life'' 2005, p. 30</ref><ref>Eva Jablonka, Marion J. Lamb ''Epigenetic Inheritance and Evolution: The Lamarckian Dimension'' 1995 p. 222</ref> 

Goldschmidt is usually referred to as a [[non-Darwinian]], however he did not object to the general microevolutionary principles of the Darwinians. He only veered from the [[modern synthesis|synthetic theory]] in his belief that a new species develops suddenly through discontinuous variation, or macromutation. Goldschmidt presented his hypothesis when [[neo-Darwinism]] was becoming dominant in the 1940's and 1950's and he strongly protested against the strict [[gradualism]] of neo-Darwinian theorists. Because of this, his ideas were seen as highly unorthodox of the time by most scientists and were greatly subjected to ridicule and scorn.<ref>Donald R. Prothero ''Evolution: What the Fossils Say and Why It Matters'' 2007, p. 99</ref> However there has been a recent interest in the ideas of Goldschmidt in the field of [[evolutionary developmental biology]] as some scientists are convinced he was not entirely wrong.

===Otto Schindewolf===

Otto Schindewolf {{lived|1896|1971}} a German paleontologist also supported macromutations as part of his evolutionary theory. He was known for presenting an alternative interpretation of the [[fossil record]] based on his ideas of [[orthogenesis]], [[saltational evolution]] and extraterrestrial impacts opposed to [[gradualism]]. Schindewolf's theory was a form of orthogenesis, which stated that variation tends to move in a predetermined direction.<ref>Chunglin Kwa Styles of ''Knowing: A New History of Science from Ancient Times to the Present'' 2011, p. 237</ref> His theory became known as typostrophism and stated that evolution occurs due to a periodic cyclic model of evolutionary processes which are predestined to go through a life cycle dictated by factors internal to the organism.<ref>[https://docs.google.com/viewer?a=v&q=cache:jkGUP6Eg0lMJ:si-pddr.si.edu/jspui/bitstream/10088/7124/1/paleo_1995_DiMichele_Schindewolf_book_review_RPP.pdf+Otto+Schindewolf+orthogenesis&hl=en&gl=uk&pid=bl&srcid=ADGEESjd9GExelLt140-ZpcyC5bwymDq2iw5ZohHXJ5Eb1BJitNFXpmIiJGIuLe17hrHtQAXTX9jMzHkoJORNmumPxqVONCMZozYzLTMi6ecJQ4riGY2S5xYuvFmZKz4zXM1gtvxxkNM&sig=AHIEtbQiNcuQskMm5lRXZGVusQLcMW51Rw Book Review]</ref> 

Part of his 'typostrophe' theory advocated sudden evolutionary change by macromutations but he later dropped this view (see below). His theory of [[orthogenesis]] (straight-line evolution) and eventual decay, claimed to be embedded within a cyclical view of the [[evolution]]ary process. According to (Levy, 2002) Schindewolf theory also proposed that mass [[extinction]]s, especially the ones at the end of the [[Permian]] period 225 million years ago, were the result of cosmic radiation caused by supernova explosions. Schindewolf speculated that a supernova star explosion could emit [[radiation]] which could be lethal to organisms if close enough to earth. He proposed that the radiation from a supernova could have two effects, one is extinguishing many species of life and the other that the radiation may cause macromutations which could cause new [[species]] to originate.<ref>David H. Levy ''Shoemaker by Levy: The Man Who Made an Impact'' 2002, p. 61</ref> Schindewolf was the only scientist to have speculated that the first [[bird]] may have hatched from a reptile's egg.<ref>Trevor Palmer ''Perilous Planet Earth: Catastrophes and Catastrophism through the Ages'' 2003, p.99</ref> It was only a speculation and he abandoned the view of macromutations in later publications.<ref>Otto Schindewolf ''Über den “Typus” in morphologischer und phylogenetischer Biologie''. Mainz: Akademie der Wissenschaften und der Literatur, 1969</ref> 

His book ''Basic Questions in Paleontology'' was published in German in 1950 and was translated into English in 1994 with a foreword written by Stephen Jay Gould.

===Stephen Jay Gould===

[[Stephen Jay Gould]] {{lived|1941|2002}} had attempted to update the ideas of Goldschmidt by redefining the concept of "hopeful monster" in a way that can be kept in the neo-Darwinian framework via an [[extended evolutionary synthesis|extension]].<ref>[http://books.google.co.uk/books?id=504iVZyopJ8C&pg=PA138&dq=%22hopeful+monster%22&hl=en&redir_esc=y#v=onepage&q=%22hopeful%20monster%22&f=false New Scientist 15 Apr 1982]</ref> In an article titled ''The Return of Hopeful Monsters'' (1977)<ref>[http://www.stephenjaygould.org/library/gould_hopeful-monsters.html Return of the Hopeful Monster by Stephen Jay Gould]</ref> Gould argued that the recent discovery of regulatory genes offered new evidence which supported some of Goldschmidt's postulates and that small changes in the embryological "contraint systems" can produce large morphological transformation in the adult, and possibly macro-evolutionary pathways.<ref>Stephen T. Asma ''On Monsters: An Unnatural History of Our Worst Fears'' 2011, p. 314 </ref> Gould's re-definition of the hopeful monster is different to that of Goldschmidt and they should not be confused with each other.

==Criticism==

Many [[scientist]]s rejected the hopeful monster hypothesis as [[genetic]]al research seemed to show that large [[mutation]]s would be lethal.<ref>Peter J. Bowler ''Evolution: The History of an Idea'' 1989 p. 340</ref> In response to this Goldschmidt suggested that, from time to time, big mutations can occur and although the majority of these would have been lethal, a very small number would have been compatible with survival. The most common criticism however; came from the early [[neo-Darwinism|neo-Darwinian]] theorists who asked the question "how does a hopeful monster find a mate?". These scientists were skeptical of the idea of hopeful monsters as without more than one hopeful monster there would be no possibility of breeding or establishing a new population and thus would be no chance of a new [[species]] forming.<ref>Donald R Prothero ''Evolution: What the Fossils Say and Why It Matters'' 2007, p. 99</ref> 

In response to this criticism, Patrick Bateson wrote in ''Sudden Changes in Ontogeny and Phylogeny'' (1984): 

{{cquote|Hopeful monsters were disparaged on the grounds that even if a big change in the phenotype could occur as a result of a mutation, the hopeful monster would be a novelty on its own with no possibility of finding a mate. Without a mate there would no new species. However, if we suppose that, somehow or other, there were enough hopeful monsters to breed successfully with each other, the possibility exists of competition between the hopeful monsters and the stock from they sprang. It is not at all difficult to suppose that, by the process of natural selection, hopeful monsters could quickly replace their competitors if they were better adapted to the environment. No new fancy principles of evolution are involved here.<ref>Patrick Baeson ''Sudden Changes in Ontogeny and Phylogeny'' in Gary Greenberg, Ethel Tobach ''Behavioral Evolution and Integrative Levels: The T.C. Schneirla Series'' 1984, p. 162</ref>}}

==Modern science==

Some modern scientists have written that hopeful monsters are neither impossible nor should be seen as anti-Darwinian because even if proven to exist would not replace the evidence for gradual evolution by [[mutation]] but supplement it. The early neo-Darwinian synthesis theorists had rejected hopeful monsters due to lack of evidence; however there is now evidence that Goldschmidt was not entirely wrong.

[[Richard Dawkins]] wrote in his book ''Climbing Mount Improbable'' (1996):

{{cquote|My suggestion is that Scyllarus may actually present an example in the wild of a homeotic mutation, analogous to antennapedia in Drosophila in the laboratory. Unlike antennapedia, this mutation has been incorporated into an actual evolutionary change in nature. My tentative conjecture is that an ancestral Scyllarid mutated homeotically, slipping the developmental subroutine appropriate to a uropod into a segment where an antenna ought to be, and that the change conferred some benefit. If I am right, it would constitute a rare example of a macro-mutation’s being favoured by natural selection: a rare vindication of the so-called ‘hopeful monster’ theory that we met in Chapter 3.<ref>Richard Dawkins ''Climbing Mount Improbable'' 1996, pp. 232-233</ref>}}

Donald R. Prothero in his book ''Evolution: What the Fossils Say and Why It Matters'' (2007) has written:

{{cquote|The past twenty years have vindicated Goldschmidt to some degree. With the discovery of the importance of regulatory genes, we realize that he was ahead of his time in focusing on the importance of a few genes controlling big changes in the organisms, not small-scales changes in the entire genome as neo-Darwinians thought. In addition, the hopeful monster problem is not so insurmountable after all. Embryology has shown that if you affect an entire population of developing embryos with a stress (such as a heat shock) it can cause many
embryos to go through the same new pathway of embryonic development, and then they all become hopeful monsters when they reach reproductive age.<ref>Donald R Prothero ''Evolution: What the Fossils Say and Why It Matters'' 2007, p. 100</ref>}}

In 2008 [[evolutionary biologist]] Olivia Judson in her article ''The Monster Is Back, and It’s Hopeful'' listed some examples which may support the hopeful monster hypothesis<ref>[http://opinionator.blogs.nytimes.com/2008/01/22/the-monster-is-back-and-its-hopeful/ The Monster Is Back, and It’s Hopeful by Olivia Judson]</ref> and an article published in the nature journal in 2010 titled ''Evolution: Revenge of the Hopeful Monster'' reported that studies in stickleback populations in a British Columbia lake and [[bacteria]] populations in a Michigan lab have shown that large individual genetic changes can have vast effects on organisms "without dooming it to the evolutionary rubbish heap". According to the article "Single-gene changes that confer a large adaptive value do happen: they are not rare, they are not doomed and, when competing with small-effect mutations, they tend to win. But small-effect mutations still matter — a lot. They provide essential fine-tuning and sometimes pave the way for explosive evolution to follow."<ref>[http://www.nature.com/news/2010/100217/full/463864a.html Evolution: Revenge of the hopeful monster by Tanguy Chouard]</ref>

A paper by (Page ''et al.'' 2010) have written that the Mexican axolotl (''Ambystoma mexicanum'') could be classified as a hopeful monster as it exhibits an adaptive and derived mode of development that has evolved rapidly and independently among tiger salamanders. According to the paper there has been a recent interest in aspects of the hopeful monster hypothesis in recent years:

{{cquote|Goldschmidt proposed that [[mutation]]s occasionally yield individuals within populations that deviate radically from the norm and referred to such individuals as "hopeful monsters". If the novel phenotypes of hopeful monsters arise under the right environmental circumstances, they may become fixed, and the population will found a new species. While this idea was discounted during the [[Modern synthesis]], aspects of the hopeful monster hypothesis have been substantiated in recent years. For example, it is clear that dramatic changes in phenotype can occur from few mutations of key developmental genes and phenotypic differences among species often map to relatively few [[genetic]] factors. These findings are motivating renewed interest in the study of hopeful monsters and the perspectives they can provide about the evolution of development. In contrast to mutants that are created in the lab, hopeful monsters have been shaped by [[natural selection]] and are therefore more likely to reveal mechanisms of adaptive evolution.<ref>[http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2900274/ Microarray analysis of a salamander hopeful monster reveals transcriptional signatures of paedomorphic brain development]</ref>}}

===Evolutionary developmental biology===

Goldschmidt presented two mechanisms for how hopeful monsters might work. One mechanism, involved “systemic mutations”, rejected the classical [[gene]] concept and is no longer considered by most modern scientists however, his second mechanism involved “developmental macromutations” in “rate genes” or “controlling genes” that change early development and thus cause large effects in the adult phenotype. These kind of mutations are similar to the ones considered in contemporary [[evolutionary developmental biology]].<ref>[http://www.palaeodiversity.org/pdf/03Suppl/Supplement_Theissen.pdf Homeosis of the angiosperm flower: Studies on three candidate cases of saltational evolution]</ref>

===Guenter Theissen===

Guenter Theissen a professor of [[genetics]] has classified homeotic mutants as hopeful monsters and documented many examples for animal and plant lineages that may have originated as hopeful monsters in his scientific publications (Theissen, 2005 and Theissen ''et al.'' 2006).<ref>[http://evolocus.com/Publications/Theissen2006.pdf The proper place of hopeful monsters in evolutionary biology]</ref><ref>[http://jxb.oxfordjournals.org/content/57/13/3531.full.pdf Catching a ‘hopeful monster’: shepherd’s purse (Capsellabursa-pastoris) as a model system to study the evolution of flower development]</ref>

==Creationist quote mining==

[[Creationist]]s are known for notoriously [[quote mining]] Goldschmidt, misrepresenting his views on purpose and for setting up a [[straw man]] definition of the hopeful monster. Thousands of creationist books since the 1960's have described the hopeful monster hypothesis as a bird hatching from a dinosaur egg or a reptile laying an egg with a bird popping out. The creationist [[Duane Gish]] was known for promoting this misrepresentation thoughout his books which he described as believing in a miracle.

In ''Abusing Science: The Case Against Creationism'' (1983) Philip Kitcher has discussed this issue, and wrote:

{{cquote|So far as I know, nobody is currently defending the idea that the birds evolved through the emergence of a single individual bird from a reptilian egg. The principal suggestion has been that some mutations — perhaps mutations in regulatory genes — might produce large effects by altering the timing of developmental events. By modifying the pattern of development, they could produce organisms with a different form from that of the parents.<ref>Philip Kitcher ''Abusing Science: The Case Against Creationism'' 1983, p. 149</ref>}}

Unfortunately creationists out of dishonesty never choose to really study what Goldschmidt actually said and continue to peddle the lie that he promoted the view that birds popped out of reptile eggs. According to [[Stephen Jay Gould]] the scientist who speculated that the first bird may have hatched from a reptile's egg was Otto Schindewolf who linked cosmic radiation to mutational rates from supernova explosions.<ref>[http://books.google.co.uk/books?id=WzDfOuHgBzMC&pg=PR11&dq=Schindewolf+bird+egg&hl=en#v=onepage&q=Schindewolf%20bird%20egg&f=false Basic Questions in Paleontology]</ref> Creationists have obviously confused the ideas of Goldschmidt with Schindewolf.

==See also==
*[[Directed mutation]]

==External links==
*[http://ag-evolutionsbiologie.de/app/download/5785604171/kutschera+niklas5_08.pdf Macroevolution via secondary endosymbiosis: a Neo-Goldschmidtian view of unicellular hopeful monsters and Darwin’s primordial intermediate form]
*[http://www.evolocus.com/publications/theissen2009.pdf Saltational evolution: hopeful monsters are here to stay]

==References==
{{reflist|2|80%}}

[[Category:Evolution]]
[[Category:Biology]]

Categories: Evolution, Biology