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Talk:Club (organization)

Religious club — Preceding unsigned comment added by 2603:3023:916:4A88:111:7623:3FF3:3B9E (talk) 16:20, 19 August 2022 (UTC)

History
The reference to the Hoboken Turtle Club mentions that it existed as of 1911. In fact, the club existed much later than that -- at least until 1939. Should this be changed? My source for this is a New Yorker article that makes mention of it from that year: http://web.archive.org/web/20061211133042rn_2/www.newyorker.com/archive/content/articles/010219fr_mitchell?010219fr_mitchell —Preceding unsigned comment added by 70.112.72.88 (talk) 04:00, 15 June 2009 (UTC)

Can the paragraph beginning, "The number of regularly established clubs in London is now..." be safely omitted? It seems to be just a list of clubs current as of 1911, and thuse irrelevant to the modern day even in a historical sense. -- April

Probably. I just cut and pasted this one since I have an aesthetic aversion to empty pages, I don't know anything about the club scene myself. Bryan Derksen

And I'm grateful! The history of clubs is quite interesting, actually, and we may want to keep some sections of it. But modernizing the end of it will take work - there's everything from the club scene to chess clubs to cover. -- April

I removed three more paragraphs specific to turn-of-the-century English clubs. Historians can always go to the 11854564564897944844888845844444 Britannica if they're curious. :) -- April

The article Club rhubarbs for ages about stuffy London gentleman's clubs, but next to nothing about the sort of ordinary sport or whatever club that very many ordinary people are in. I split off the Roman clubs section and the Greek clubs section onto separate pages.Anthony Appleyard 06:29, 29 May 2005 (UTC)

In the list of famous members of the Bread Street/Friday Street Club, I'm trying to add links for them all. I can't figure out who Beaumont or Fletcher are. There aren't any famous Beaumonts of the period in WP, and there are two famous Fletchers, both literary (John and Phineas). Can anyone with a direct knowledge of the club itself help me out? --Trinite 16:43, 13 Mar 2006 (UTC)

Cleanup
I placed the tag on this page. I think it's too long and meandering--it need sections, etc. --Kewp 17:14, 21 September 2005 (UTC)

In response to the cleanup request, I split this article into sections, and rearranged some stuff to match. To balance it out, some material on ancient clubs may have to move back, and additional material on political and social clubs included. ClaudeMuncey 17:21, 12 November 2005 (UTC)

Club (disamgibuation), Club, Social Cub and their relatives have an illogical structure.
The official club disambiguation page (Club (disambiguation)) is not very good. It is incomplete and only lists one category of social club, while discussing sports clubs, country clubs, health clubs and gyms at the same level of detail. For example, social clubs and country clubs are listed as distinct categories. This page (club) does a step further and gives disambiguation at a higher level of detail, which is necessary because there are so many types of social clubs, many of which overlap. Furthermore a search for "club" takes one to club not to club (disambiguation). The whole structure of the club subject is fraught with logical errors.

The Club (disambiguation) page does not have a link to Social club! That could be corrected, but there are other problems. In reality country clubs should overlap heavily and be cited extensively in the article, social clubs, but the latter ignores many classes of social clubs and partly social clubs.

Furthermore, Club (disambiguation) is inaccurate, since Gentlemen's club is a common euphemism for a strip club.

Again, there is no need for references in this article because they are provided in the individual articles. --Zeamays (talk) 20:42, 9 December 2007 (UTC)
 * I do not agree. The fact that individual articles have references is OK for lists, but this is not a list and it should be referenced. I have not reverted your removal of the tag, but I think we should try to get consensus here. --Bduke (talk) 22:08, 9 December 2007 (UTC)

 * I agree with Zeamays that the club family is imperfect: the answer is to correct the errors and improve the overall structure. I agree with Bduke that this article, club, is not a list but a proper article, and as such should be properly referenced.  There are articles I have contributed to in which virtually every section begins with a "main" tag, and one of the chief jobs of those who watch the article is to hone the contributions of the eager many, and direct their enthusiasm to the separate articles.  One example is horse, which is long, but which nonetheless spins off as much as possible into daughter articles.  Although the club family of articles is not likely to attract the level and rate of editorial additions that horse does, I think it is a perfectly valid method of controlling and shaping a group of information.
 * Two points of fact: gentlemen's club is a legitimate term with its primary meaning the long-standing one. They are still important features of the British establishment, not merely archaisms or historical footnotes. We have encyclopedic standards to maintain; it is entirely appropriate to have an article about strip clubs, and indicate that they are also known as gentlemen's clubs, but that is a euphemistic colloquialism.
 * Secondly, social club and country club do not overlap to a great extent in Britain, where I am writing from. If the former article is missing information, please add to it.
 * I have put many hours into some of these articles, especially club and gentlemen's club. Frankly, they were pretty dire before I decided to get my hands dirty; and they are still far from adequate now, particularly with regard to references. I welcome constructive engagement with them, but I may not be able to contribute much over the next couple of weeks -- it depends on my activities outside WP, which are a bit unpredictable.  Good luck! BrainyBabe (talk) 22:14, 10 December 2007 (UTC)

 * I forgot to say, I don't know how to arrange it, but it would be great if a search for "club" would go straight to the dab page, not to this one. People might be thinking of golf clubs (the implements) or juggling clubs or any of many other meanings. BrainyBabe (talk) 22:20, 10 December 2007 (UTC)

Merge proposal (2010)
The proposal is to merge Voluntary association to here. As far as I can see, there are no significant conceptual distinctions – although there are of course differences in connotation; one wouldn't call a gentlemen's club a gentlemen's voluntary association. --Lambiam 06:25, 19 July 2010 (UTC)

 * AGAINST There is a significant legal distinction in that the word "club" is used colloquially but the term "voluntary association" is often used to distinguish a legal entity from other types such as a cooperative, domestic corporation, charity corporation or so on. Clubs may choose what type of legal entity to be, but the phrase voluntary association often has a precise and technical legal meaning. Rorybowman (talk) 19:07, 7 August 2010 (UTC)

Clubbing
It is said that clubbing is injurious to health.

— Preceding unsigned comment added by SteveM123 (talk • contribs) 01:17, 15 November 2010 (UTC)

Requested move 24 June 2015

 * The following is a closed discussion of a requested move. Please do not modify it. Subsequent comments should be made in a new section on the talk page. Editors desiring to contest the closing decision should consider a move review. No further edits should be made to this section.

The result of the move request was: Move as proposed. Cúchullain t/ c 18:28, 6 July 2015 (UTC)

– I do not believe that the organizational concept is the primary topic of "club", or that the term has any primary topic. The club as a weapon is older, and many incoming links are intended for the music genre. bd2412 T 15:42, 24 June 2015 (UTC)
 * Club → Club (organization)
 * Club (disambiguation) → Club
 * Support Club → Club (organization) per no primary topic.
 * Oppose Club (disambiguation) → Club. The title topic Club in this listing of many types of club needs itself to be disambiguated from all the many items listed within its own listing.  A redirect to Club (disambiguation) provides a suitably solution for the disambiguation of the list.  GregKaye 18:21, 24 June 2015 (UTC)

 * Support as proposed. GregKaye's proposal would result in a malplaced disambiguation page, which is something he seems to disagree with, but as yet has little support. older ≠ wiser 00:58, 25 June 2015 (UTC)
 * Bkonrad Please consider definition of malplaced which do not apply. How is a disambiguation page malplaced at a location that presents it as containing disambiguations and in a way that disambiguates the list content presented from any and all items on the list?  GregKaye 04:59, 25 June 2015 (UTC)
 * Because there is no primary topic, the disambiguation page should reside at the base name. older ≠ wiser 10:40, 25 June 2015 (UTC)
 * GregKaye: see WP:MALPLACED. --BDD (talk) 13:57, 25 June 2015 (UTC)
 * TY surely it is also a violation of WP:ASSERT to claim that an article is "malplaced" when no reasoned argument for this claim is given.  GregKaye 19:03, 25 June 2015 (UTC)
 * But there is reasoned argument for this--only you happen to disagree with the argument. That doesn't mean there is none. older ≠ wiser 19:31, 25 June 2015 (UTC)

 * Strong support a club is a blunt instrument used to bludgeon and strike things. Further, it is a suite in a deck of cards. Tertiarily, it is an organization. -- 70.51.203.69 (talk) 04:06, 25 June 2015 (UTC)
 * Exactly, a club could be many different things which is why the disambiguation page should stay at Club (disambiguation). GregKaye 05:11, 25 June 2015 (UTC)
 * Under your approach, Club would be a redirect to Club (disambiguation). That would make this title inconsistent with literally hundreds of thousands of titles that avoid such a pattern. I am not at all averse to discussing whether that is the more precise title, but we should do so in a general policy discussion, not in the context of creating a single outlier. bd2412  T 17:41, 25 June 2015 (UTC)
 * Club (disambiguation) is a perfectly sensible article title for a list of disambiguated articles that relate to the term club. No confusion is generated.  Please explain the problem.  The article is not about a club and has a different content.  There is an extremely close natural association between article titles Club (disambiguation) and Club.  How does this, I think, reasonable suggestion constitute a generation of an outlier?  Are there no other examples that follow what seems to me to be a logical and grammatically sound solution?  GregKaye 18:58, 25 June 2015 (UTC)

 * Support per nom. This is not a WP:PRIMARYTOPIC, and the disambiguation should be moved per WP:MALPLACED. kennethaw88 • talk 00:37, 26 June 2015 (UTC)
 * Reasonably strong oppose for several reasons. This seems like a pretty straightforward primary topic to me, especially since we ain't writing a dictionary. I do join the chorus of voices that Club must not redirect to a dab page. If consensus ends up deciding that there's no primary topic, the dab page must be moved here. Red Slash 05:17, 26 June 2015 (UTC)
 * The primary topic, if there is one, is the blunt object, which has historically been an element of many crimes and much warfare, and most of human pre-history and history. -- 70.51.203.69 (talk) 04:19, 27 June 2015 (UTC)
 * Support, the type of organization is not the main topic 76.120.162.73 (talk) 23:01, 30 June 2015 (UTC)
 * Support per nom. No clear primary topic. -- Necrothesp (talk) 16:23, 1 July 2015 (UTC)

 * The above discussion is preserved as an archive of a requested move. Please do not modify it. Subsequent comments should be made in a new section on this talk page or in a move review. No further edits should be made to this section.

title: Talk:Club (organization)


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Rudolf Hell

Rudolf Hell (19 December 1901 – 11 March 2002) was a German inventor and engineer.

Career
Hell was born in Eggmühl. From 1919 to 1923, he studied electrical engineering in Munich. He worked there from 1923 to 1929 as assistant of Prof. Max Dieckmann, with whom he operated a television station at the Verkehrsausstellung (lit.: "traffic exhibition") in Munich in 1925. In the same year Hell invented an apparatus called the Hellschreiber, an early forerunner to impact dot matrix printers and faxes. Hell received a patent for the Hellschreiber in 1929.

In the year 1929 he founded his own company in Babelsberg. After World War II he re-founded his company in Kiel. He kept on working as an engineer and invented machines for electronically controlled engraving of printing plates and an electronic photo typesetting system called digiset marketed in the US as VideoComp by RCA and later by III.

He has received numerous awards such as the Knight Commander's Cross of the Order of Merit of the Federal Republic of Germany, the Gutenberg Prize of the International Gutenberg Society and the City of Mainz, the Werner von Siemens Ring and the Eduard Rhein Ring of Honor from the German Eduard Rhein Foundation (1992).

His company was taken over by Siemens AG in 1981 and merged with Linotype in 1990, becoming Linotype-Hell AG.

Hellschreiber is still in use today by Amateur Radio (Ham) operators around the world. Hellverein Kiel collects Hell devices and keeps them functional. In addition, the website contains numerous documents and patents relating to the Hell devices: https://www.hell-kiel.de/en/

Death
Rudolf Hell died in Kiel on 11 March 2002.

license: Creative Commons - Attribution Share-Alike - https://creativecommons.org/licenses/by-sa/4.0/

title: Rudolf Hell


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Selfish genetic element

Selfish genetic elements (historically also referred to as selfish genes, ultra-selfish genes, selfish DNA, parasitic DNA and genomic outlaws) are genetic segments that can enhance their own transmission at the expense of other genes in the genome, even if this has no positive or a net negative effect on organismal fitness. Genomes have traditionally been viewed as cohesive units, with genes acting together to improve the fitness of the organism. However, when genes have some control over their own transmission, the rules can change, and so just like all social groups, genomes are vulnerable to selfish behaviour by their parts.

Early observations of selfish genetic elements were made almost a century ago, but the topic did not get widespread attention until several decades later. Inspired by the gene-centred views of evolution popularized by George Williams and Richard Dawkins, two papers were published back-to-back in Nature in 1980 – by Leslie Orgel and Francis Crick and by Ford Doolittle and Carmen Sapienza – introducing the concept of selfish genetic elements (at the time called "selfish DNA") to the wider scientific community. Both papers emphasized that genes can spread in a population regardless of their effect on organismal fitness as long as they have a transmission advantage.

Selfish genetic elements have now been described in most groups of organisms, and they demonstrate a remarkable diversity in the ways by which they promote their own transmission. Though long dismissed as genetic curiosities, with little relevance for evolution, they are now recognized to affect a wide swath of biological processes, ranging from genome size and architecture to speciation.

Early observations
Observations of what is now referred to as selfish genetic elements go back to the early days in the history of genetics. Already in 1928, Russian geneticist Sergey Gershenson reported the discovery of a driving X chromosome in Drosophila obscura. Crucially, he noted that the resulting female-biased sex ratio may drive a population extinct (see Species extinction). The earliest clear statement of how chromosomes may spread in a population not because of their positive fitness effects on the individual organism, but because of their own "parasitic" nature came from the Swedish botanist and cytogeneticist Gunnar Östergren in 1945. Discussing B chromosomes in plants he wrote:

"In many cases these chromosomes have no useful function at all to the species carrying them, but that they often lead an exclusively parasitic existence ... [B chromosomes] need not be useful for the plants. They need only be useful to themselves."

Around the same time, several other examples of selfish genetic elements were reported. For example, the American maize geneticist Marcus Rhoades described how chromosomal knobs led to female meiotic drive in maize. Similarly, this was also when it was first suggested that an intragenomic conflict between uniparentally inherited mitochondrial genes and biparentally inherited nuclear genes could lead to cytoplasmic male sterility in plants. Then, in the early 1950s, Barbara McClintock published a series of papers describing the existence of transposable elements, which are now recognized to be among the most successful selfish genetic elements. The discovery of transposable elements led to her being awarded the Nobel Prize in Medicine or Physiology in 1983.

Conceptual developments
The empirical study of selfish genetic elements benefited greatly from the emergence of the so-called gene-centred view of evolution in the nineteen sixties and seventies. In contrast with Darwin's original formulation of the theory of evolution by natural selection that focused on individual organisms, the gene's-eye view takes the gene to be the central unit of selection in evolution. It conceives evolution by natural selection as a process involving two separate entities: replicators (entities that produce faithful copies of themselves, usually genes) and vehicles (or interactors; entities that interact with the ecological environment, usually organisms).

Since organisms are temporary occurrences, present in one generation and gone in the next, genes (replicators) are the only entity faithfully transmitted from parent to offspring. Viewing evolution as a struggle between competing replicators made it easier to recognize that not all genes in an organism would share the same evolutionary fate.

The gene's-eye view was a synthesis of the population genetic models of the modern synthesis, in particular the work of RA Fisher, and the social evolution models of W. D. Hamilton. The view was popularized by George Williams's Adaptation and Natural Selection and Richard Dawkins's best seller The Selfish Gene. Dawkins summarized a key benefit from the gene's-eye view as follows:

"'If we allow ourselves the license of talking about genes as if they had conscious aims, always reassuring ourselves that we could translate our sloppy language back into respectable terms if we wanted to, we can ask the question, what is a single selfish gene trying to do?' — Richard Dawkins, The Selfish Gene"

In 1980, two high-profile papers published back-to-back in Nature by Leslie Orgel and Francis Crick, and by Ford Doolittle and Carmen Sapienza, brought the study of selfish genetic elements to the centre of biological debate. The papers took their starting point in the contemporary debate of the so-called C-value paradox, the lack of correlation between genome size and perceived complexity of a species. Both papers attempted to counter the prevailing view of the time that the presence of differential amounts of non-coding DNA and transposable elements is best explained from the perspective of individual fitness, described as the "phenotypic paradigm" by Doolittle and Sapienza. Instead, the authors argued that much of the genetic material in eukaryotic genomes persists, not because of its phenotypic effects, but can be understood from a gene's-eye view, without invoking individual-level explanations. The two papers led to a series of exchanges in Nature.

Current views
If the selfish DNA papers marked the beginning of the serious study of selfish genetic elements, the subsequent decades have seen an explosion in theoretical advances and empirical discoveries. Leda Cosmides and John Tooby wrote a landmark review about the conflict between maternally inherited cytoplasmic genes and biparentally inherited nuclear genes. The paper also provided a comprehensive introduction to the logic of genomic conflicts, foreshadowing many themes that would later be subject of much research. Then in 1988 John H. Werren and colleagues wrote the first major empirical review of the topic. This paper achieved three things. First, it coined the term selfish genetic element, putting an end to a sometimes confusingly diverse terminology (selfish genes, ultra-selfish genes, selfish DNA, parasitic DNA, genomic outlaws). Second, it formally defined the concept of selfish genetic elements. Finally, it was the first paper to bring together all different kinds of selfish genetic elements known at the time (genomic imprinting, for example, was not covered).

In the late 1980s, most molecular biologists considered selfish genetic elements to be the exception, and that genomes were best thought of as highly integrated networks with a coherent effect on organismal fitness. In 2006, when Austin Burt and Robert Trivers published the first book-length treatment of the topic, the tide was changing. While their role in evolution long remained controversial, in a review published a century after their first discovery, William R. Rice concluded that "nothing in genetics makes sense except in the light of genomic conflicts".

Logic
Though selfish genetic elements show a remarkable diversity in the way they promote their own transmission, some generalizations about their biology can be made. In a classic 2001 review, Gregory D.D. Hurst and John H. Werren proposed two ‘rules' of selfish genetic elements.

Rule 1: Spread requires sex and outbreeding
Sexual reproduction involves the mixing of genes from two individuals. According to Mendel's Law of Segregation, alleles in a sexually reproducing organism have a 50% chance of being passed from parent to offspring. Meiosis is therefore sometimes referred to as "fair".

Highly self-fertilizing or asexual genomes are expected to experience less conflict between selfish genetic elements and the rest of the host genome than outcrossing sexual genomes. There are several reasons for this. First, sex and outcrossing put selfish genetic elements into new genetic lineages. In contrast, in a highly selfing or asexual lineage, any selfish genetic element is essentially stuck in that lineage, which should increase variation in fitness among individuals. The increased variation should result in stronger purifying selection in selfers/asexuals, as a lineage without the selfish genetic elements should out-compete a lineage with the selfish genetic element. Second, the increased homozygosity in selfers removes the opportunity for competition among homologous alleles. Third, theoretical work has shown that the greater linkage disequilibrium in selfing compared to outcrossing genomes may in some, albeit rather limited, cases cause selection for reduced transposition rates. Overall, this reasoning leads to the prediction that asexuals/selfers should experience a lower load of selfish genetic elements. One caveat to this is that the evolution of selfing is associated with a reduction in the effective population size. A reduction in the effective population size should reduce the efficacy of selection and therefore leads to the opposite prediction: higher accumulation of selfish genetic elements in selfers relative to outcrossers.

Empirical evidence for the importance of sex and outcrossing comes from a variety of selfish genetic elements, including transposable elements, self-promoting plasmids, and B chromosomes.

Rule 2: Presence is often revealed in hybrids
The presence of selfish genetic elements can be difficult to detect in natural populations. Instead, their phenotypic consequences often become apparent in hybrids. The first reason for this is that some selfish genetic elements rapidly sweep to fixation, and the phenotypic effects will therefore not be segregating in the population. Hybridization events, however, will produce offspring with and without the selfish genetic elements and so reveal their presence. The second reason is that host genomes have evolved mechanisms to suppress the activity of the selfish genetic elements, for example the small RNA administered silencing of transposable elements. The co-evolution between selfish genetic elements and their suppressors can be rapid, and follow a Red Queen dynamics, which may mask the presence of selfish genetic elements in a population. Hybrid offspring, on the other hand, may inherit a given selfish genetic element, but not the corresponding suppressor and so reveal the phenotypic effect of the selfish genetic element.

Segregation distorters
Some selfish genetic elements manipulate the genetic transmission process to their own advantage, and so end up being overrepresented in the gametes. Such distortion can occur in various ways, and the umbrella term that encompasses all of them is segregation distortion. Some elements can preferentially be transmitted in egg cells as opposed to polar bodies during meiosis, where only the former will be fertilized and transmitted to the next generation. Any gene that can manipulate the odds of ending up in the egg rather than the polar body will have a transmission advantage, and will increase in frequency in a population.

Segregation distortion can happen in several ways. When this process occurs during meiosis it is referred to as meiotic drive. Many forms of segregation distortion occur in male gamete formation, where there is differential mortality of spermatids during the process of sperm maturation or spermiogenesis. The segregation distorter (SD) in Drosophila melanogaster is the best studied example, and it involves a nuclear envelope protein Ran-GAP and the X-linked repeat array called Responder (Rsp), where the SD allele of Ran-GAP favors its own transmission only in the presence of a Rspsensitive allele on the homologous chromosome. SD acts to kill RSPsensitive sperm, in a post-meiotic process (hence it is not strictly speaking meiotic drive). Systems like this can have interesting rock-paper-scissors dynamics, oscillating between the SD-RSPinsensitive, SD+-RSPinsensitive and SD+-RSPsensitive haplotypes. The SD-RSPsensitive haplotype is not seen because it essentially commits suicide.

When segregation distortion acts on sex chromosomes, they can skew the sex ratio. The SR system in Drosophila pseudoobscura, for example, is on the X chromosome, and XSR/Y males produce only daughters, whereas females undergo normal meiosis with Mendelian proportions of gametes. Segregation distortion systems would drive the favored allele to fixation, except that most of the cases where these systems have been identified have the driven allele opposed by some other selective force. One example is the lethality of the t-haplotype in mice, another is the effect on male fertility of the Sex Ratio system in D. pseudoobscura.

Homing endonucleases
A phenomenon closely related to segregation distortion is homing endonucleases. These are enzymes that cut DNA in a sequence-specific way, and those cuts, generally double-strand breaks, are then "healed" by the regular DNA repair machinery. Homing endonucleases insert themselves into the genome at the site homologous to the first insertion site, resulting in a conversion of a heterozygote into a homozygote bearing a copy of the homing endonuclease on both homologous chromosomes. This gives homing endonucleases an allele frequency dynamics rather similar to a segregation distortion system, and generally unless opposed by strong countervailing selection, they are expected to go to fixation in a population. CRISPR-Cas9 technology allows the artificial construction of homing endonuclease systems. These so-called "gene drive" systems pose a combination of great promise for biocontrol but also potential risk.

Transposable elements
Transposable elements (TEs) include a wide variety of DNA sequences that all have the ability to move to new locations in the genome of their host. Transposons do this by a direct cut-and-paste mechanism, whereas retrotransposons need to produce an RNA intermediate to move. TEs were first discovered in maize by Barbara McClintock in the 1940s and their ability to occur in both active and quiescent states in the genome was also first elucidated by McClintock. TEs have been referred to as selfish genetic elements because they have some control over their own propagation in the genome. Most random insertions into the genome appear to be relatively innocuous, but they can disrupt critical gene functions with devastating results. For example, TEs have been linked to a variety of human diseases, ranging from cancer to haemophilia. TEs that tend to avoid disrupting vital functions in the genome tend to remain in the genome longer, and hence they are more likely to be found in innocuous locations.

Both plant and animal hosts have evolved means for reducing the fitness impact of TEs, both by directly silencing them and by reducing their ability to transpose in the genome. It would appear that hosts in general are fairly tolerant of TEs in their genomes, since a sizable portion (30-80%) of the genome of many animals and plants is TEs. When the host is able to stop their movement, TEs can simply be frozen in place, and it then can take millions of years for them to mutate away. The fitness of a TE is a combination of its ability to expand in numbers within a genome, to evade host defenses, but also to avoid eroding host fitness too drastically. The effect of TEs in the genome is not entirely selfish. Because their insertion into the genome can disrupt gene function, sometimes those disruptions can have positive fitness value for the host. Many adaptive changes in Drosophila and dogs for example, are associated with TE insertions.

B chromosomes
B chromosomes refer to chromosomes that are not required for the viability or fertility of the organism, but exist in addition to the normal (A) set. They persist in the population and accumulate because they have the ability to propagate their own transmission independently of the A chromosomes. They often vary in copy number between individuals of the same species.

B chromosomes were first detected over a century ago. Though typically smaller than normal chromosomes, their gene poor, heterochromatin-rich structure made them visible to early cytogenetic techniques. B chromosomes have been thoroughly studied and are estimated to occur in 15% of all eukaryotic species. In general, they appear to be particularly common among eudicot plants, rare in mammals, and absent in birds. In 1945, they were the subject of Gunnar Östergren's classic paper "Parasitic nature of extra fragment chromosomes", where he argues that the variation in abundance of B chromosomes between and within species is because of the parasitic properties of the Bs. This was the first time genetic material was referred to as "parasitic" or "selfish". B chromosome number correlates positively with genome size and has also been linked to a decrease in egg production in the grasshopper Eyprepocnemis plorans.

Selfish mitochondria
Genomic conflicts often arise because not all genes are inherited in the same way. Probably the best example of this is the conflict between uniparentally (usually but not always, maternally) inherited mitochondrial and biparentally inherited nuclear genes. Indeed, one of the earliest clear statements about the possibility of genomic conflict was made by the English botanist Dan Lewis in reference to the conflict between maternally inherited mitochondrial and biparentally inherited nuclear genes over sex allocation in hermaphroditic plants.

A single cell typically contains multiple mitochondria, creating a situation for competition over transmission. Uniparental inheritance has been suggested to be a way to reduce the opportunity for selfish mitochondria to spread, as it ensures all mitochondria share the same genome, thus removing the opportunity for competition. This view remains widely held, but has been challenged. Why inheritance ended up being maternal, rather than paternal, is also much debated, but one key hypothesis is that the mutation rate is lower in female compared to male gametes.

The conflict between mitochondrial and nuclear genes is especially easy to study in flowering plants. Flowering plants are typically hermaphrodites, and the conflict thus occurs within a single individual. Mitochondrial genes are typically only transmitted through female gametes, and therefore from their point of view the production of pollen leads to an evolutionary dead end. Any mitochondrial mutation that can affect the amount of resources the plant invests in the female reproductive functions at the expense of the male reproductive functions improves its own chance of transmission. Cytoplasmic male sterility is the loss of male fertility, typically through loss of functional pollen production, resulting from a mitochondrial mutation. In many species where cytoplasmic male sterility occurs, the nuclear genome has evolved so-called restorer genes, which repress the effects of the cytoplasmic male sterility genes and restore the male function, making the plant a hermaphrodite again.

The co-evolutionary arms race between selfish mitochondrial genes and nuclear compensatory alleles can often be detected by crossing individuals from different species that have different combinations of male sterility genes and nuclear restorers, resulting in hybrids with a mismatch.

Another consequence of the maternal inheritance of the mitochondrial genome is the so-called Mother's Curse. Because genes in the mitochondrial genome are strictly maternally inherited, mutations that are beneficial in females can spread in a population even if they are deleterious in males. Explicit screens in fruit flies have successfully identified such female-neutral but male-harming mtDNA mutations. Furthermore, a 2017 paper showed how a mitochondrial mutation causing Leber's hereditary optic neuropathy, a male-biased eye disease, was brought over by one of the Filles du roi that arrived in Quebec, Canada, in the 17th century and subsequently spread among many descendants.

Genomic imprinting
Another sort of conflict that genomes face is that between the mother and father competing for control of gene expression in the offspring, including the complete silencing of one parental allele. Due to differences in methylation status of gametes, there is an inherent asymmetry to the maternal and paternal genomes that can be used to drive a differential parent-of-origin expression. This results in a violation of Mendel's rules at the level of expression, not transmission, but if the gene expression affects fitness, it can amount to a similar result.

Imprinting seems like a maladaptive phenomenon, since it essentially means giving up diploidy, and heterozygotes for one defective allele are in trouble if the active allele is the one that is silenced. Several human diseases, such as Prader-Willi and Angelman syndromes, are associated with defects in imprinted genes. The asymmetry of maternal and paternal expression suggests that some kind of conflict between these two genomes might be driving the evolution of imprinting. In particular, several genes in placental mammals display expression of paternal genes that maximize offspring growth, and maternal genes that tend to keep that growth in check. Many other conflict-based theories about the evolution of genomic imprinting have been put forward.

At the same time, genomic or sexual conflict are not the only possible mechanisms whereby imprinting can evolve. Several molecular mechanisms for genomic imprinting have been described, and all have the aspect that maternally and paternally derived alleles are made to have distinct epigenetic marks, in particular the degree of methylation of cytosines. An important point to note regarding genomic imprinting is that it is quite heterogeneous, with different mechanisms and different consequences of having single parent-of-origin expression. For example, examining the imprinting status of closely related species allows one to see that a gene that is moved by an inversion into close proximity of imprinted genes may itself acquire an imprinted status, even if there is no particular fitness consequence of the imprinting.

Greenbeards
A greenbeard gene is a gene that has the ability to recognize copies of itself in other individuals and then make its carrier act preferentially toward such individuals. The name itself comes from thought-experiment first presented by William Hamilton and then it was developed and given its current name by Richard Dawkins in The Selfish Gene. The point of the thought experiment was to highlight that from a gene's-eye view, it is not the genome-wide relatedness that matters (which is usually how kin selection operates, i.e. cooperative behavior is directed towards relatives), but the relatedness at the particular locus that underlies the social behavior.

Following Dawkins, a greenbeard is usually defined as a gene, or set of closely linked genes, that has three effects:

 * 1) It gives carriers of the gene a phenotypic label, such as a greenbeard.
 * 2) The carrier is able to recognize other individuals with the same label.
 * 3) The carrier then behaves altruistically towards individuals with the same label.

Greenbeards were long thought to be a fun theoretical idea, with limited possibility of them actually existing in nature. However, since its conception, several examples have been identified, including in yeast, slime moulds, and fire ants.

There has been some debate whether greenbeard genes should be considered selfish genetic elements. Conflict between a greenbeard locus and the rest of the genome can arise because during a given social interaction between two individuals, the relatedness at the greenbeard locus can be higher than at other loci in the genome. As a consequence, it may in the interest of the greenbeard locus to perform a costly social act, but not in the interest of the rest of the genome.

In conjunction with selfish genetic elements, greenbeard selection has also been used as a theoretical explanation for suicide.

Species extinction
Perhaps one of the clearest ways to see that the process of natural selection does not always have organismal fitness as the sole driver is when selfish genetic elements have their way without restriction. In such cases, selfish elements can, in principle, result in species extinction. This possibility was pointed out already in 1928 by Sergey Gershenson and then in 1967, Bill Hamilton developed a formal population genetic model for a case of segregation distortion of sex chromosomes driving a population to extinction. In particular, if a selfish element should be able to direct the production of sperm, such that males bearing the element on the Y chromosome would produce an excess of Y-bearing sperm, then in the absence of any countervailing force, this would ultimately result in the Y chromosome going to fixation in the population, producing an extremely male-biased sex ratio. In ecologically challenged species, such biased sex ratios imply that the conversion of resources to offspring becomes very inefficient, to the point of risking extinction.

Speciation
Selfish genetic elements have been shown to play a role in speciation. This could happen because the presence of selfish genetic elements can result in changes in morphology and/or life history, but ways by which the co-evolution between selfish genetic elements and their suppressors can cause reproductive isolation through so-called Bateson–Dobzhansky–Muller incompatibilities has received particular attention.

An early striking example of hybrid dysgenesis induced by a selfish genetic element was the P element in Drosophila. If males carrying the P element were crossed to females lacking it, the resulting offspring suffered from reduced fitness. However, offspring of the reciprocal cross were normal, as would be expected since piRNAs are maternally inherited. The P element is typically present only in wild strains, and not in lab strains of D. melanogaster, as the latter were collected before the P elements were introduced into the species, probably from a closely related Drosophila species. The P element story is also a good example of how the rapid co-evolution between selfish genetic elements and their silencers can lead to incompatibilities on short evolutionary time scales, as little as within a few decades.

Several other examples of selfish genetic elements causing reproductive isolation have since been demonstrated. Crossing different species of Arabidopsis results in both higher activity of transposable elements and disruption in imprinting, both of which have been linked to fitness reduction in the resulting hybrids. Hybrid dysgenesis has also been shown to be caused by centromeric drive in barley and in several species of angiosperms by mito-nuclear conflict.

Genome-size variation
Attempts to understand the extraordinary variation in genome size (C-value)—animals vary 7,000 fold and land plants some 2,400-fold—has a long history in biology. However, this variation is poorly correlated with gene number or any measure of organismal complexity, which led CA Thomas to coin the term C-value paradox in 1971. The discovery of non-coding DNA resolved some of the paradox, and most current researchers now use the term "C-value enigma".

Two kinds of selfish genetic elements in particular have been shown to contribute to genome-size variation: B chromosomes and transposable elements. The contribution of transposable elements to the genome is especially well studied in plants. A striking example is how the genome of the model organism Arabidopsis thaliana contains the same number of genes as that of the Norwegian spruce (Picea abies), around 30,000, but accumulation of transposons means that the genome of the latter is some 100 times larger. Transposable element abundance has also been shown to cause the unusually large genomes found in salamanders.

The presence of an abundance of transposable elements in many eukaryotic genomes was a central theme of the original selfish DNA papers mentioned above (See Conceptual developments). Most people quickly accepted the central message of those papers, that the existence of transposable elements can be explained by selfish selection at the gene level and there is no need to invoke individual level selection. However, the idea that organisms keep transposable elements around as genetic reservoir to "speed up evolution" or for other regulatory functions persists in some quarters. In 2012, when the ENCODE Project published a paper claiming that 80% of the human genome can be assigned a function, a claim interpreted by many as the death of the idea of junk DNA, this debate was reignited.

Cytoplasmic male sterility in plant breeding
A common problem for plant breeders is unwanted self-fertilization. This is particularly a problem when breeders try to cross two different strains to create a new hybrid strain. One way to avoid this is manual emasculation, i.e. physically removing anthers to render the individual male sterile. Cytoplasmic male sterility offers an alternative to this laborious exercise. Breeders cross a strain that carries a cytoplasmic male sterility mutation with a strain that does not, the latter acting as the pollen donor. If the hybrid offspring are to be harvested for their seed (like maize), and therefore needs to be male fertile, the parental strains need to be homozygous for the restorer allele. In contrast, in species that harvested for their vegetable parts, like onions, this is not an issue. This technique has been used in a wide variety of crops, including rice, maize, sunflower, wheat, and cotton.

PiggyBac vectors
While many transposable elements seem to do no good for the host, some transposable elements have been "tamed" by molecular biologists so that the elements can be made to insert and excise at the will of the scientist. Such elements are especially useful for doing genetic manipulations, like inserting foreign DNA into the genomes of a variety of organisms.

One excellent example of this is PiggyBac, a transposable element that can efficiently move between cloning vectors and chromosomes using a "cut and paste" mechanism. The investigator constructs a PiggyBac element with the desired payload spliced in, and a second element (the PiggyBac transposase), located on another plasmid vector, can be co-transfected into the target cell. The PiggyBac transposase cuts at the inverted terminal repeat sequences located on both ends of the PiggyBac vector and efficiently moves the contents from the original sites and integrates them into chromosomal positions where the sequence TTAA is found. The three things that make PiggyBac so useful are the remarkably high efficiency of this cut-and-paste operation, its ability to take payloads up to 200 kb in size, and its ability to leave a perfectly seamless cut from a genomic site, leaving no sequences or mutations behind.

CRISPR gene drive and homing endonuclease systems
CRISPR allows the construction of artificial homing endonucleases, where the construct produces guide RNAs that cut the target gene, and homologous flanking sequences then allow insertion of the same construct harboring the Cas9 gene and the guide RNAs. Such gene drives ought to have the ability to rapidly spread in a population (see Gene-drive systems), and one practical application of such a system that has been proposed is to apply it to a pest population, greatly reducing its numbers or even driving it extinct. This has not yet been attempted in the field, but gene drive constructs have been tested in the lab, and the ability to insert into the wild-type homologous allele in heterozygotes for the gene drive has been demonstrated. Unfortunately, the double-strand break that is introduced by Cas9 can be corrected by homology directed repair, which would make a perfect copy of the drive, or by non-homologous end joining, which would produce "resistant" alleles unable to further propagate themselves. When Cas9 is expressed outside of meiosis, it seems like non-homologous end joining predominates, making this the biggest hurdle to practical application of gene drives.

Mathematical theory
Much of the confusion regarding ideas about selfish genetic elements center on the use of language and the way the elements and their evolutionary dynamics are described. Mathematical models allow the assumptions and the rules to be given a priori for establishing mathematical statements about the expected dynamics of the elements in populations. The consequences of having such elements in genomes can then be explored objectively. The mathematics can define very crisply the different classes of elements by their precise behavior within a population, sidestepping any distracting verbiage about the inner hopes and desires of greedy selfish genes. There are many good examples of this approach, and this article focuses on segregation distorters, gene drive systems and transposable elements.

Segregation distorters
The mouse t-allele is a classic example of a segregation distorter system that has been modeled in great detail. Heterozygotes for a t-haplotype produce >90% of their gametes bearing the t (see Segregation distorters), and homozygotes for a t-haplotype die as embryos. This can result in a stable polymorphism, with an equilibrium frequency that depends on the drive strength and direct fitness impacts of t-haplotypes. This is a common theme in the mathematics of segregation distorters:virtually every example we know entails a countervailing selective effect, without which the allele with biased transmission would go to fixation and the segregation distortion would no longer be manifested. Whenever sex chromosomes undergo segregation distortion, the population sex ratio is altered, making these systems particularly interesting. Two classic examples of segregation distortion involving sex chromosomes include the "Sex Ratio" X chromosomes of Drosophila pseudoobscura and Y chromosome drive suppressors of Drosophila mediopunctata. A crucial point about the theory of segregation distorters is that just because there are fitness effects acting against the distorter, this does not guarantee that there will be a stable polymorphism. In fact, some sex chromosome drivers can produce frequency dynamics with wild oscillations and cycles.

Gene-drive systems
The idea of spreading a gene into a population as a means of population control is actually quite old, and models for the dynamics of introduced compound chromosomes date back to the 1970s. Subsequently, the population genetics theory for homing endonucleases and CRISPR-based gene drives has become much more advanced. An important component of modeling these processes in natural populations is to consider the genetic response in the target population. For one thing, any natural population will harbor standing genetic variation, and that variation might well include polymorphism in the sequences homologous to the guide RNAs, or the homology arms that are meant to direct the repair. In addition, different hosts and different constructs may have quite different rates of non-homologous end joining, the form of repair that results in broken or resistant alleles that no longer spread. Full accommodation of the host factors presents considerable challenge for getting a gene drive construct to go to fixation, and Unckless and colleagues show that in fact the current constructs are quite far from being able to attain even moderate frequencies in natural populations. This is another excellent example showing that just because an element appears to have a strong selfish transmission advantage, whether it can successfully spread may depend on subtle configurations of other parameters in the population.

Transposable elements
To model the dynamics of transposable elements (TEs) within a genome, one has to realize that the elements behave like a population within each genome, and they can jump from one haploid genome to another by horizontal transfer. The mathematics has to describe the rates and dependencies of these transfer events. It was observed early on that the rate of jumping of many TEs varies with copy number, and so the first models simply used an empirical function for the rate of transposition. This had the advantage that it could be measured by experiments in the lab, but it left open the question of why the rate differs among elements and differs with copy number. Stan Sawyer and Daniel L. Hartl fitted models of this sort to a variety of bacterial TEs, and obtained quite good fits between copy number and transmission rate and the population-wide incidence of the TEs. TEs in higher organisms, like Drosophila, have a very different dynamics because of sex, and Brian Charlesworth, Deborah Charlesworth, Charles Langley, John Brookfield and others modeled TE copy number evolution in Drosophila and other species. What is impressive about all these modeling efforts is how well they fitted empirical data, given that this was decades before discovery of the fact that the host fly has a powerful defense mechanism in the form of piRNAs. Incorporation of host defense along with TE dynamics into evolutionary models of TE regulation is still in its infancy.

title: Selfish genetic element


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Carole Lombard

Carole Lombard (born Jane Alice Peters; October 6, 1908 – January 16, 1942) was an American actress. In 1999, the American Film Institute ranked Lombard 23rd on its list of the greatest female stars of Classic Hollywood Cinema.

Born into a wealthy family, Lombard was raised by her single mother. She made her screen debut in A Perfect Crime (1921) at the age of 12 and the experience spurred her and her mother to seek further film work, eventually resulting in a movie career.

Lombard was killed in a plane crash aboard TWA Flight 3 while returning from a war bond tour. She was 33 years old. Today, she is remembered as one of the definitive actresses of the screwball comedy genre and American comedy and as an icon of American cinema.

Early life and education (1908–1920)
Lombard was born in Fort Wayne, Indiana, on October 6, 1908, at 704 Rockhill Street. Christened Jane Alice Peters, she was the third child and only daughter of Frederic Christian Peters (1875–1935) and Elizabeth Jayne "Bessie" (Knight) Peters (1876–1942). Her two older brothers, with whom she was close all her life, were Frederic Charles (1902–1979) and John Stuart (1906–1956). Lombard's parents both came from wealthy families, and biographer Robert Matzen called her early years her "silver spoon period". Her parents' marriage was strained, and in October 1914, her mother took the children and moved to Los Angeles. Although the couple did not divorce, the separation was permanent. Her father's continued financial support allowed the family to live comfortably, and they settled into an apartment near Venice Boulevard.

At Virgil Junior High School, Lombard played tennis, volleyball, and swimming, and won trophies in athletics. At the age of 12, her passion for sports landed Lombard her first screen role. While playing baseball, she caught the attention of film director Allan Dwan, who later recalled seeing "a cute-looking little tomboy... out there knocking the hell out of the other kids, playing better baseball than they were. And I needed someone of her type for this picture." With the encouragement of her mother, Lombard took a small role in the melodrama A Perfect Crime (1921). She was on set for two days, playing the sister of Monte Blue. Dwan later said "She ate it up".

Career beginnings and Fox contract (1921–26)
Though A Perfect Crime was not widely distributed, the experience spurred Lombard and her mother to audition for more film work, but she was unsuccessful. While appearing as the queen of Fairfax High School's May Day Carnival at the age of 15, Lombard was scouted by an employee of Charlie Chaplin and offered a screen test to appear in The Gold Rush (1925). Lombard did not win the role, but her test was seen by the Vitagraph Film Company, which expressed interest in signing her. Although this did not materialize, their condition that she adopt a new first name led to her selecting the name "Carole" after a girl with whom she played tennis at Virgil Jr. High School.

In October 1924, 16-year-old Lombard signed a contract with the Fox Film Corporation. Lombard's mother contacted gossip columnist Louella Parsons, who arranged a screen test. According to biographer Larry Swindell, Lombard's beauty convinced studio head Winfield Sheehan to sign her to a $75-per-week contract, and she abandoned her schooling to pursue the new career. Fox disliked her surname and she was renamed Carole Lombard, the surname of a family friend.

Most of Lombard's appearances with Fox were bit parts in low-budget Westerns and adventure films. She later said, "All I had to do was simper prettily at the hero and scream with terror when he battled with the villain." However, she enjoyed other aspects of film work such as photo shoots, costume fittings, and socializing with actors on the studio set. Lombard embraced the flapper lifestyle and became a regular at the Cocoanut Grove nightclub, where she won several Charleston dance competitions.

In March 1925, Lombard landed a leading role in the drama Marriage in Transit with Edmund Lowe. A reviewer for Motion Picture News wrote that Lombard displayed "good poise and considerable charm". However, the studio heads were unconvinced that Lombard was leading-lady material, and her contract was not renewed. Gehring has suggested a facial scar resulting from a car crash was a factor in this decision, but that incident occurred nearly two years later on September 9, 1927. According to historian Olympia Kiriakou, on the night of the crash, Lombard was on a date with a man named Harry Cooper. On Santa Monica Boulevard, Cooper hit another car; the windshield shattered and shards of glass cut "Lombard's face from her nose and across her left cheek to her eye." Lombard underwent reconstructive surgery and faced a long recovery period. For the remainder of her career, Lombard learned to hide the mark with makeup and careful lighting. At the time of the crash, Lombard was already under contract with Mack Sennett. In October 1927, Lombard and her mother Bess sued Cooper for $35,000 in damages, citing in the lawsuit that "where she formerly was able to earn a salary of $300 monthly as a Sennett girl, she is now unable to obtain employment of any kind." The lawsuit was settled out of court, and Lombard received $3,000. Although Lombard feared that the incident would end her career, Sennett pledged to help her recover. He afforded her "lucrative film roles and ample publicity", including the nickname "Carole of the Curves". Kiriakou explains, "the nickname simultaneously drew audiences' focus away from her facial scars and worked harmoniously with the physicality and female sensuality that were emblematic of Lombard's performances" in Sennett's films.

Breakthrough and early success (1927–29)
Although Lombard initially had reservations about slapstick comedies, she became one of Sennett Bathing Beauties and appeared in 18 short films (all as Lillian Smith in the Smith Family series) from September 1927 to March 1929, Lombard's first experiences in comedy provided valuable training for her future comedic work. In 1940, she called her Sennett years "the turning point of [my] acting career".

Sennett's productions were distributed by Pathé Exchange, and the company began casting Lombard in feature films. She had prominent roles in Show Folks and Ned McCobb's Daughter (both 1928), and reviewers observed that she made a "good impression" and was "worth watching". The following year, Pathé elevated Lombard to a leading lady. Her success in Raoul Walsh's picture Me, Gangster (also 1928), with June Collyer and Don Terry in his film debut, finally eased the pressure that her family had been exerting for her to succeed. In Howard Higgin's High Voltage (1929), Lombard's first sound film, she played a criminal in the custody of a deputy sheriff, both of whom are among bus passengers stranded in deep snow. Her next film, the comedy Big News (1929), cast her with Robert Armstrong and was a critical and commercial success. Lombard was reunited with Armstrong for the crime drama The Racketeer, released in late 1929. The review in Film Daily wrote: "Carol Lombard proves a real surprise, and does her best work to date. In fact, this is the first opportunity she has had to prove that she has the stuff to go over."

Paramount contract and first marriage (1930–33)
Lombard returned to Fox for a one-off role in the Western The Arizona Kid (1930). It was a big release for the studio, starring the popular actor Warner Baxter, in which Lombard received third billing. Following the success of the film, Paramount Pictures recruited Lombard and signed her to a $350-per-week contract, gradually increasing to $3,500 per week by 1936. They cast her in the Buddy Rogers comedy Safety in Numbers (also 1930), and one critic observed of her work, "Lombard proves [to be] an ace comedienne." For her second assignment, Fast and Loose (also 1930) with Miriam Hopkins, Paramount mistakenly credited the actress as "Carole Lombard". She decided she liked this spelling and it became her permanent screen name.

Lombard appeared in five films released during 1931, beginning with the Frank Tuttle comedy It Pays to Advertise. Her next two films, Man of the World and Ladies Man, both featured William Powell, Paramount's top male star. Lombard had been a fan of the actor before they met and they were soon in a relationship. The differences between the pair have been noted by biographers: She was 22, carefree, and famously foul-mouthed, and he was 38, intellectual, and sophisticated. Despite this, Lombard married Powell on June 26, 1931, at her Beverly Hills home. Talking to the media, she argued for the benefits of "love between two people who are diametrically different", claiming that their relationship allowed for a "perfect see-saw love". The marriage to Powell increased Lombard's fame, while she continued to please critics with her work in Up Pops the Devil and I Take this Woman (both 1931). In reviews for the latter film, which co-starred Gary Cooper, several critics predicted that Lombard was set to become a major star. She went on to appear in five films throughout 1932. No One Man and Sinners in the Sun were not successful, but Edward Buzzell's romantic picture Virtue was well received. After featuring in the drama No More Orchids, Lombard was cast as the wife of a con artist in No Man of Her Own with Clark Gable The film was a critical and commercial success, and Wes Gehring writes that it was "arguably Lombard's finest film appearance" to that point. It was the only picture that Gable and Lombard made together. There was no romantic interest at this time, however, as she recounted to Garson Kanin: "[we] did all kinds of hot love scenes ... and I never got any kind of tremble out of him at all".

In August 1933, Lombard and Powell divorced after 26 months of marriage, but they remained friends until the end of Lombard's life. At the time, she blamed it on their careers, but in a 1936 interview, she admitted that this "had little to do with the divorce. We were just two completely incompatible people".

She appeared in five films that year, beginning with the drama From Hell to Heaven and continuing with Supernatural, her only horror vehicle. After a small role in The Eagle and the Hawk, a war film starring Fredric March and Cary Grant, she starred in two melodramas: Brief Moment, which critics enjoyed, and White Woman, where she was paired with Charles Laughton.

Lombard was involved romantically with Russ Columbo, the famous crooner killed in a tragic accident in 1934. Lombard had been guiding Columbo's movie and radio career and told Sonia Lee of Mirror magazine in 1934 that they had been engaged. Other press outlets had reported on their relationship earlier that year; Screenland Magazine declared, "the Russ Columbo and Carole Lombard romance is one of Hollywood's most charming."

Success in screwball comedies (1934–35)
The year 1934 marked a high point in Lombard's career, beginning with Wesley Ruggles's musical drama Bolero, where George Raft and she showcased their dancing skills in an extravagantly staged performance to Maurice Ravel's Boléro. Before filming began, she was offered the lead female role in It Happened One Night, but turned it down because of scheduling conflicts. Bolero was favorably received, while her next film, the musical comedy We're Not Dressing with Bing Crosby, was a box-office hit.

Lombard was then recruited by the director Howard Hawks, a second cousin, to star in his screwball Twentieth Century which proved a watershed in her career and made her a major star. Hawks had seen the actress inebriated at a party, where he found her to be "hilarious and uninhibited and just what the part needed", and she was cast with John Barrymore. In Twentieth Century, Lombard plays an actress who is pursued by her former mentor, a flamboyant Broadway impresario. Hawks and Barrymore were unimpressed with her work in rehearsals, finding that she was "acting" too hard and giving a stiff performance. The director encouraged Lombard to relax, be herself, and act on her instincts. She responded well to this tutoring, and reviews for the film commented on her unexpectedly "fiery talent"—"a Lombard like no Lombard you've ever seen". The Los Angeles Times' critic felt that she was "entirely different" from her formerly cool, "calculated" persona, adding, "she vibrates with life and passion, abandon and diablerie".

The next films in which Lombard appeared were Henry Hathaway's Now and Forever (1934), featuring Gary Cooper and the new child star Shirley Temple, and Lady by Choice (1934), which was a critical and commercial success. The Gay Bride (1934) placed her with Chester Morris in a gangster comedy, but this outing was panned by critics. After reuniting with George Raft for another dance picture, Rumba (1935), Lombard was given the opportunity to repeat the screwball success of Twentieth Century. In Mitchell Leisen's Hands Across the Table (1935), she portrays a manicurist in search of a rich husband, played by Fred MacMurray. Critics praised the film, and Photoplay's reviewer stated that Lombard had reaffirmed her talent for the genre. It is remembered as one of her best films, and the pairing of Lombard and MacMurray proved so successful that they made three more pictures together.

Critical recognition (1936–37)
Lombard's first film of 1936 was Love Before Breakfast, described by Gehring as "The Taming of the Shrew, screwball style". In William K. Howard's The Princess Comes Across, her second comedy with MacMurray, she played a budding actress who wins a film contract by masquerading as a Swedish princess. The performance was considered a satire of Greta Garbo and was widely praised by critics. Lombard's success continued as she was recruited by Universal Studios to star in the screwball comedy My Man Godfrey (1936). William Powell, who was playing the eponymous Godfrey, insisted on her being cast as the female lead; despite their divorce, the pair remained friendly and Powell felt she would be perfect in the role of Irene, a zany heiress who employs a "forgotten man" as the family butler. The film was directed by Gregory LaCava, who knew Lombard personally and advised that she draw on her "eccentric nature" for the role. She worked hard on the performance, particularly with finding the appropriate facial expressions for Irene. My Man Godfrey was released to great acclaim and was a box-office hit. It received six nominations at the 9th Academy Awards, including Lombard for Best Actress. Biographers cite it as her finest performance, and Frederick Ott says it "clearly established [her] as a comedienne of the first rank."

By 1937, Lombard was one of Hollywood's most popular actresses, and also the highest-paid star in Hollywood following the deal which Myron Selznick negotiated with Paramount that brought her $450,000, more than five times the salary of the U.S. president. As her salary was widely reported in the press, Lombard stated that 80% of her earnings went in taxes, but that she was happy to help improve her country. The comments earned her much positive publicity, and President Franklin D. Roosevelt sent her a personal letter of thanks.

Her first release of the year was Leisen's Swing High, Swing Low, a third pairing with MacMurray. The film focused on a romance between two cabaret performers, and was a critical and commercial success. It had been primarily a drama, with occasional moments of comedy, but for her next project, Nothing Sacred, Lombard returned to the screwball genre. Producer David O. Selznick, impressed by her work in My Man Godfrey, was eager to make a comedy with the actress and hired Ben Hecht to write an original screenplay for her. Nothing Sacred, directed by William Wellman and co-starring Fredric March, satirized the journalism industry and "the gullible urban masses". Lombard portrayed a small-town girl who pretends to be dying and finds her story exploited by a New York reporter. The film was Lombard's only Technicolor feature-length production, and she later praised it highly as one of her personal favorites.

Lombard continued with screwball comedies, next starring in True Confession (1937), what Swindell calls one of her "wackiest" films,. She played a compulsive liar who wrongly confesses to murder. Lombard loved the script and was excited about the project, which reunited her with John Barrymore and was her final appearance with MacMurray. Her prediction that it "smacked of a surefire success" proved accurate as critics responded positively, and it was popular at the box office.

Dramatic efforts and second marriage (1938–40)

True Confession was the last film Lombard made on her Paramount contract, and she remained an independent performer for the rest of her career. Her next film was made at Warner Bros., where she played a famous actress in Mervyn LeRoy's Fools for Scandal (1938). The comedy met with scathing reviews and was a commercial failure, with Swindell calling it "one of the most horrendous flops of the thirties".

Fools for Scandal was the only film Lombard made in 1938. By this time, she was devoted to her relationship with Clark Gable. The pair had reunited at a Hollywood party and began a romance early in 1936. The media took great interest in their partnership and frequently questioned if they would wed. Gable was separated from his wife, Maria, but she did not want to grant him a divorce. As his relationship with Lombard became serious, Maria eventually agreed to a settlement. The divorce was finalized in March 1939, and Gable and Lombard eloped in Kingman, Arizona on March 29. The couple bought a 20 acre ranch in Encino, California, where they kept barnyard animals and enjoyed hunting trips. Almost immediately, Lombard wanted to start a family, but her attempts failed; after two miscarriages and numerous trips to fertility specialists, she was unable to have children. In early 1938, Lombard officially joined the Baháʼí Faith, which her mother had been a member of since 1922.

While continuing with a slower work-rate, Lombard decided to move away from comedies and return to dramatic roles. She appeared in Made for Each Other (1939) with James Stewart playing a couple facing domestic difficulties. Reviews for the film were highly positive, and praised Lombard's dramatic effort; financially, it was a disappointment. Lombard's next appearance came with Cary Grant in the John Cromwell romance In Name Only (1939), a credit she personally negotiated with RKO Radio Pictures upon hearing of the script and Grant's involvement. The role reflected her recent experiences, as she played a woman in love with a married man whose wife refuses to divorce. She was paid $150,000 for the film, continuing her status as one of Hollywood's highest-paid actresses, and it was a moderate success. At the 12th Academy Awards ceremony in February 1940, Lombard was quoted as comforting Gable after his loss as Rhett Butler from Gone with the Wind, with the comment "Don't worry, Pappy. We'll bring one home next year". Gable replied that he felt this had been his last chance to which Lombard was said to have replied, "Not you, you self-centered bastard. I meant me."

Lombard was eager to win an Academy Award, and selected her next project with the expectation that it would bring her the trophy. Vigil in the Night (1940), directed by George Stevens, featured Lombard as a nurse who faces a series of personal difficulties. Although the performance was praised, she did not get her nomination, as the sombre mood of the picture turned audiences away and box-office returns were poor. Despite the realization that she was best suited to comedies, Lombard completed the drama They Knew What They Wanted (1940), co-starring Charles Laughton, which was mildly successful.

Final roles (1941–42)

Accepting that "my name doesn't sell tickets to serious pictures", Lombard returned to comedy in Mr. & Mrs. Smith (1941), about a couple who learns that their marriage is invalid, with Robert Montgomery. Lombard was influential in bringing Alfred Hitchcock, whom she knew through David O. Selznick, to direct one of his most atypical films. It was a commercial success, and audiences were happy with what Swindell calls "the belated happy news ... that Carole Lombard was a screwball once more."

It was nearly a year before Lombard committed to another film, as she focused instead on her home and marriage. Determined that her next film be "an unqualified smash hit", she was also careful in selecting a new project. Through her agent, Lombard heard of Ernst Lubitsch's upcoming film: To Be or Not to Be (1942), a dark comedy that satirized the Nazi takeover of Poland. The actress had long wanted to work with Lubitsch, and felt that the material—although controversial—was a worthy subject. Lombard accepted the role of actress Maria Tura, despite it being a smaller part than she was used to, and was given top billing over the film's male lead Jack Benny. Filming took place in the fall of 1941, and it was reportedly one of the happier experiences of Lombard's career.

Death
When the U.S. entered World War II, Lombard traveled to her home state of Indiana for a war bond rally with her mother and Clark Gable's press agent, Otto Winkler. Lombard raised more than $2 million in defense bonds in a single evening. Her party had been scheduled to return to Los Angeles by train, but Lombard was eager to reach home more quickly and wanted to travel by air. Her mother and Winkler were afraid of flying and insisted that the group follow their original travel plans.

In the early morning hours of January 16, 1942, Lombard, her mother and Winkler boarded a Transcontinental and Western Air Douglas DST (Douglas Sleeper Transport) aircraft to return to California. After refueling in Las Vegas, TWA Flight 3 took off at 7:07 p.m. and crashed into Double Up Peak near the 8300 ft level of Potosi Mountain, 32 smi southwest of the Las Vegas airport. All 22 aboard, including Lombard, her mother, Winkler and 15 U.S. Army soldiers, were killed. Lombard was 33 years old. The crash's cause was attributed to the flight crew's inability to properly navigate over the mountains surrounding Las Vegas. As a precaution against the possibility of enemy Japanese bomber aircraft coming into American airspace from the Pacific, safety beacons normally used to direct night flights had been turned off, leaving the pilot and crew of the TWA flight without visual warnings of the mountains in their flight path.

Aftermath
When The Jack Benny Program aired on January 18, Jack Benny did not attend the live radio broadcast. At its opening, announcer Don Wilson stated Benny would not appear that night, but did not explain why. The show that night did not feature any comedy, just musical numbers. Lombard had been scheduled to appear on the following Sunday's broadcast.

Lombard's funeral was January 21 at Forest Lawn Memorial Park Cemetery in Glendale, California. She was interred beside her mother under the name of Carole Lombard Gable. Despite remarrying twice following her death, Gable was interred beside Lombard when he died in 1960.

To Be or Not to Be, Lombard's final film, was in post-production at the time of her death. Allegedly, the film's producers decided to cut a line in which Lombard's character asks "What can happen on a plane?" out of respect for the circumstances surrounding her death. Although, there is no indication that this line existed and was removed posthumously, the film's script as filed with the Production Code Administration included the addendum:. "This certificate is issued with the understanding that Anna's speech: 'No, not at all...' down to and including: 'She might hit an air pocket.' has been replaced; also that Sigorsky's speech '-- maybe you'll want to take care of her after my departure.' has been omitted."

At the time of her death, Lombard had been scheduled to star in the film They All Kissed the Bride; when production started, she was replaced by Joan Crawford. Crawford donated all of her salary for the film to the Red Cross, which had helped extensively in the recovery of bodies from the air crash.

Shortly after Lombard's death, Gable, who was inconsolable and devastated by his loss, joined the United States Army Air Forces. Lombard had asked him to do that numerous times after the United States had entered World War II. After officer training, Gable headed a six-man motion picture unit attached to a B-17 bomb group in England to film aerial gunners in combat, flying five missions himself. In December 1943, the United States Maritime Commission announced that a Liberty ship named in her honor would be launched. Gable attended the launch of the SS Carole Lombard on January 15, 1944, the two-year anniversary of Lombard's war bond drive. The ship was involved in rescuing hundreds of survivors from sunken ships in the Pacific and returning them to safety.

In 1962, Jill Winkler Rath, widow of publicist Otto Winkler, filed a $100,000 lawsuit against the $2 million estate of Clark Gable in connection with Winkler's death. The suit was dismissed in Los Angeles Superior Court. Rath, in her action, claimed Gable promised to provide financial aid for her if she would not bring suit against the airline involved. Rath stated she later learned that Gable settled his claim against the airline for $10. He did so because he did not want to repeat his grief in court, and subsequently he provided her no financial aid in his will.

Legacy

Author Robert D. Matzen has cited Lombard as "among the most commercially successful and admired film personalities in Hollywood in the 1930s", and feminist writer June Sochen believes that Lombard "demonstrated great knowledge of the mechanics of film making". George Raft, her co-star in Bolero, was extremely fond of the actress, remarking "I truly loved Carole Lombard. She was the greatest girl that ever lived and we were the best of pals. Completely honest and outspoken, she was liked by everyone".

Historian Olympia Kiriakou identifies Lombard as a progressive, feminist studio-era star. She describes Lombard's politics as "proto-feminist", explaining that "many of her political and social statements pre-date the second-wave feminist movement, yet were very much in line with the second wave's focus," particularly her views about women's roles in the home and workplace. Lombard's independent star persona balanced her femininity and screen glamour with "male business sense". She was described by Photoplay columnist Hart Seymore as the "perfect example of a modern Career Girl", which was based on Lombard's capability to "live by the logical premise that women have equal rights with men." In 1937, Photoplay published an article about Lombard's business acumen entitled "Carole Lombard tells: 'How I Live by a Man's Code'," in which she offers readers rules for how to be successful in business and at home such as "play fair [with men]...don't burn over criticism—stand up to it like a man." Notably, in the article Lombard tells readers that she "doesn't believe in a man's world," and encourages women to "work—and like it," adding: "All women should have something worthwhile to do, and cultivate efficiency at it, whether it be housekeeping or raising chickens. Working women are interesting women." But as Kiriakou explains, such an article was published in order "to elicit a specific response from the fan magazine readers—namely, to view Lombard's independent star as indistinguishable from the Lombard heroines they saw on screen."

Moreover, according to scholar Emily Carman, Lombard's independent female star persona was able to emerge only when she "attained greater professional autonomy in the mid-1930s," ultimately leading her to become one of the first stars of the studio-era to go freelance. Freelancing gave Lombard more autonomy over her career decisions, and the types of roles she was able to play. Additionally, Lombard was the first Hollywood star to propose profit participation: in 1938, she negotiated with Selznick International Pictures to take a reduced salary of $100,000 in exchange for a 20 percent cut of the distributor's gross of $1.6 to $1.7 million, and subsequent smaller percentages as the gross increased. Carman explains that this contract also included a "no-loan out" clause, the right to employ Travis Banton as her costume designer of choice, as well as all legal rights to her image. Carman concludes that Lombard's strategic business sense and easy-going nature were central to her independent star persona, and the control she maintained over her career was a challenge to the "paternalistic structure" of the studio system.

Lombard was particularly noted for the zaniness of her performances, described as a "natural prankster, a salty tongued straight-shooter, a feminist precursor and one of the few stars who was beloved by the technicians and studio functionaries who worked with her". Life magazine noted that her film personality transcended to real life, "her conversation, often brilliant, is punctuated by screeches, laughs, growls, gesticulations and the expletives of a sailor's parrot". Graham Greene praised the "heartbreaking and nostalgic melodies" of her faster-than-thought delivery, whereas The Independent wrote "Platinum blonde, with a heart-shaped face, delicate, impish features and a figure made to be swathed in silver lamé, Lombard wriggled expressively through such classics of hysteria as Twentieth Century and My Man Godfrey."

In 1999, the American Film Institute ranked Lombard 23rd on its list of the 25 greatest American female screen legends of classic Hollywood cinema, and she has a star on the Hollywood Walk of Fame at 6930 Hollywood Blvd. Lombard received one Academy Award for Best Actress nomination for My Man Godfrey. Actresses who have portrayed her in films include Jill Clayburgh in Gable and Lombard (1976), Sharon Gless in Moviola: The Scarlett O'Hara War (1980), Denise Crosby in Malice in Wonderland (1985), Anastasia Hille in RKO 281 (1999) and Vanessa Gray in Lucy (2003). Lombard's Fort Wayne childhood home has been designated a historic landmark. The city named the nearby bridge over the St. Mary's River the Carole Lombard Memorial Bridge.

Lombard's star at the Hollywood Walk of Fame is shown in the movie Pretty Woman.

title: Carole Lombard


text:
Selfish gene

Selfish gene may refer to:
 * Selfish gene theory, the gene-centered view of evolution
 * The Selfish Gene, a book by Richard Dawkins discussing the selfish gene theory
 * Selfish genetic elements, which are genetic segments that can enhance their own transmission at the expense of other genes in their host genome, most commonly by creating new copies of themselves within that genome.
 * "Selfish Gene", a song by Panda Bear from the album Panda Bear Meets the Grim Reaper

title: Selfish gene