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Fidelity, Fecundity, and Longevity

By: A. Bradley Duthie

1 May 2003


This is the final essay of the three that I have tried to compile together. I recommend reading The Ecology of Memetics and The Lives of Mimics before reading this essay..
The copies building, weaving, lines
In common space. But near
We gaze at battles viewed from signs
Of micro rungs that steer
Our thoughts, our looks, our joys, and minds.
No plan, design appear,
No glowing script engraved that shines
Yet fables persevere

A single sentence on a piece of paper may hold great potential to retain the sentence's information if we apply certain properties to that sentence. If the sentence were to maximize certain characteristics, those characteristics would increase the sentence’s chances of surviving. To show  what I mean, I will provide examples on the following independent clause; I must be copied. The sentence already carries a great deal of potential in replication; the sentence itself is an instruction for the sentence’s own self-replication, but what are some ways that would maximize the time the information was retained as a whole? One very important characteristic the sentence must be found with is accurate replicating machinery. That is, the sentence must be copied well every time the sentence is copied, for an error in replication reading “I must be pie” will probably have less chance of reproduction. Occasionally a mutation may prove harmless or even beneficial, but for the most part we can safely say that a change in the instructions of the sentence would damage the overall meaning. Accurate copying machinery will boost the amount of correct copies that are manufactured. Another aspect that would ensure the survival of the sentence “I must be copied” is to increase the rate of replication. A way to do this might be to photocopy copies of the sentence continually. If the sentence can replicate quickly, the information has the benefit of being retained in many copies, as opposed to a smaller amount of copies that the sentence may produce with a slower rate of replication.
   
Yet another method of keeping the information in the sentence would be to construct the sentence so that the sentence would not easily be lost. A way to increase the amount of time a single sentence would persevere through the ages might be to carve the sentence in a strong material such as stone or steel. The information would then be stored safely for a much longer period of time than we might expect from copying the sentence down on a piece of notebook paper. If a replicator is copied accurately, we can say that the replicator has high fidelity. High fecundity is the ability to reproduce at a rapid rate, and if something has a high degree of longevity, whatever we’re dealing with will stick around for a relatively long amount of time. These three traits are very important in the world of replicators, and every replicator or set of replicators that survives well will likely have a strong balance of the characteristics that I mentioned; naturally there are some extremes. The three characteristics that determine the survival of a replicator are fidelity, fecundity, and longevity; after a clear idea of what these characteristics mean, we can apply that knowledge to develop a better understanding of the existence of replicators.

Before developing a thorough understanding about the existence and nature of replicators, we must first have a look at the properties of replicators. The three characteristics shared, to some extent at least, by all replicators are fidelity, fecundity, and longevity. The first characteristic to look at is that of fidelity. Fidelity is a replicator's "degree of accuracy in its copies" (What online). Fidelity is an important property of a replicator; if a replicator is unable to successfully replicate accurately, the copy is unlikely to last for a long period of time because the environment around the replicator with poor accuracy will soon be full of the inaccurately copied replicators. The copier with poor fidelity will not last long in the population simply because the replicator with poor fidelity will not be successful in self-propagation. Poor fidelity would dilute any benefits that a replicator has. If a replicator arose that produced an effect that gave the replicator an excellent shot of survival in the replicator's environment, the effect that benefitted the replicator might be all but lost in the next generation if the replicator could not copy itself correctly. Thus for any stability in a replicator, the replicator must be able to make accurate copies of itself. Of course, no replicator will be perfect, and errors in copying are likely to occur. Despite the importance of copying fidelity, errors in replication, or mutations, are "an important property of any copying process," (Dawkins Selfish 16). In fact, mutations are absolutely necessary in the evolutionary process. Errors in copying are essential to the evolutionary algorithm, for if all replicators were the same, the properties of the replicators could not vary in their ability to survive. Crossing over during meiosis as well as sexual reproduction have evolved to give a necessary balance between a need for copying fidelity and a need for variation in sexual reproduction. In discussing fidelity, I will use examples solely of genetic biology as the precise mechanics of memes in the brain are yet to be determined by scientists.

In an animal such as a human being, there are trillions of cells. For an organism to function correctly, these cells must be working correctly together in order to maintain all the functions necessary for survival and reproduction. Mutations in any replicator are only rarely beneficial, and this statement is especially true when looking at somatic cells. A beneficial genetic mutation in a somatic cell that occurs early on in development will, by definition, add to the fitness of the organism that the mutation occurred in, but the new genes will not affect the offspring of the organism. Only the sex cells, or gametes, in an organism will pass their traits on to the next generation. Genes such as these are said to be "germ-line replicators," so a germ-line replicator can be described as "a replicator that is potentially the ancestor of an indefinitely long line of descendant replicators" (Dawkins Extended 83). For this reason, beneficial mutations in somatic cells cannot be passed, and because most mutations are not beneficial to the "germ line," any gene in the germ line that can heighten the copying fidelity in somatic cells will survive well. Of course, a gene that encourages the production of a variation in offspring will ensure that a particular species can adapt to environmental change and perhaps even take advantage of some slight variances that give the organism an edge over the organism’s competitors. A gene that encourages too much mutation will not prosper as much; as stated earlier, mutations are rarely beneficial, but a gene that encourages a healthy variation will ultimately win out over a process with near perfect fidelity. A gene encouraging variation might produce some offspring with less overall individual fitness than the offspring’s predecessor, but a gene encouraging variation should also have some slight variations that are beneficial. The process that allows a relatively small amount of variance in offspring is known as sexual reproduction. I will go more into detail regarding the dynamics of sexual reproduction, but first I would like to go more into depth about the amazing machinery that occurs in the replication of DNA.

Because somatic cells do not enter the germ line, and thus any mutations that occur are not passed, DNA replication within somatic cells must be as accurate as possible, for almost any mutation that may arise would be non-beneficial. With these conditions in effect, the selection of various cellular apparatus to increase the copying fidelity of DNA has produced an extraordinary outcome. In eukaryotes, the replication process in DNA includes a “replicative enzyme” that is called “DNA polymerase 8" (Lindahl & Wood 1898). DNA polymerase 8 has the ability to proofread during DNA replication via “exonuclease activity” (Lindahl & Wood 1898). Furthermore, after the DNA proofreading, another correction system “further minimizes replication errors by a systematic survey of newly synthesized strands” (Lindahl & Wood 1898). As a final process that has developed for error correction, “accessory factors such as the DNA helicases encoded by the genes defective in Werner syndrome and Bloom syndrome apparently serve to improve accuracy during DNA elongation” (Lindahl & Wood 1898). This improved accuracy is likely “due to resolution of stalled replication forks” (Lindahl & Wood 1898). All of the previous processes make DNA replication a process with extremely high fidelity replication. Despite this complicated and meticulous process, with trillions of somatic cells, some mutations in DNA are still inevitable.
For the germ-line, however, such an incredible degree of fidelity might not always survive as long in the gene pool. Knowing that most mutations are not beneficial, this statement might seem almost paradoxical at first, but upon further analysis we can get a better understanding of why such an assertion holds true. The environment that any species resides in is not forever a constant. Biotic and abiotic factors in an environment are always likely to be changing over time, for this reason, any gene or genes that encourage a variance in the offspring of the germ-line will give more evolutionary flexibility to the species that produces varying offspring. To add a variance in the germ-line, while still holding the necessary fidelity for genes in somatic cells, a process called crossing over, "also known as recombination," has evolved (Yount 30). Crossing over occurs in the gametes of organisms. In the process of crossing over, there is a "physical exchange of chromosome parts . . . during meiosis" (Gilman et al. 9). During the process of meiosis, chromosomes will line up inside the cell. As these chromosomes are lined up, genetic material is exchanged between different chromosomes, so a piece of nucleic acid may be swapped from one chromosome to another. This swapping of information allows for more variation to occur in the offspring of an organism. In this evolved system of crossing over, the chromosome number stays constant, and genes tend to be sort of shuffled around. Genes may be sliced in two at times, but this is more rare than one might think as the vast majority of genetic material is made up of introns, or "junk" DNA. This process allows variance in the offspring of organisms with less chance of the negative impact that would arise purely from copying errors in the genome.

That is not to say, of course that copying errors are always detrimental to the overall fitness of an organism, but, more often than not, this is the case. The genes themselves that lie within the genome still maintain a fairly high level of fidelity in replication, for most often during the process of crossing over the genes themselves will remain accurately copied and intact. Thus we can see a high level of fidelity for the replicators themselves, but still a variance in the offspring of the organism, and this is superb example of why the replicator must be considered as the unit of selection.

The second characteristic that will be found in surviving replicators is that of fecundity. Fecundity is defined as the "speed of replication" that a replicator possesses (Dawkins Selfish 17). Replicators that can reproduce most quickly will naturally have a better chance of being found in their environment. If replicator A can reproduce itself at a rate of one copy per minute, and replicator B can reproduce itself at a rate of one copy per second, there is an obvious conclusion, other things being equal, as to which replicator will become prominent in a competitive environment that the replicators might inhabit.

The speediest replicators today belong to memetics. Memes can spread especially fast because of the incredibly efficient way in which they travel. For genes, the act of replication can only be so fast, and only so many copies can be made in a given time frame. Genes are physical bits of matter residing in cells. To replicate a gene, more material must be gathered from around the cell and the material must line up with the existing DNA. The replicator must be right next to the material to be copied. Memes are much more efficient replicators in that memes may replicate from vast distances away through different mediums and to vast amounts of material with the potential to hold copies. The inventing of the printing press allowed for a much higher degree of fecundity in replication. The printing press allowed memes to be transmitted from their source to a wealth of nearly identical artifacts. This process was many times faster, as well as more accurate, than the old method of handwriting. Today information in the mind of a particular journalist might produce the end result of a phenotype recognized as a column in a local newspaper. The phenotype of the memes that is the print on the newspaper could then be read by hundreds of people at the same time; the mental machinery that belongs to the readers of the paper would then input some amount of the information that the machinery processed. How much of the information is retained would depend on several factors, for the memes picked up from the paper would likely be competing with several other memes. The printing press allowed information to be passed much more rapidly than the old method of using handwriting for transcription, but the invention of the television allowed for greater fecundity still. With a television, memes can be broadcast almost instantaneously to millions around the world. This power that memetic replicators have achieved through the evolutionary process is taken much for granted, but to comprehend the great fecundity achieved in memetics through mediums such as televisions from a biological standpoint is amazing. As a result, any meme that is able to latch onto the airwaves of television will spread well. Notice still, that the artifact of the television is merely a vehicle for memes. Instructions for making televisions survive well largely because they aid many memes to spread.

There is a great deal of difficulty in finding a comparison to the television in the world of genetic biology. Genes replicate at a much slower pace than memes, and I believe this fact is in part due to the greater amount of horizontal transmission in memetics. In genetic biology, vertical transmission is much more commonplace. Genes are replicated at a much slower rate, although we may make some notes of vehicles with impressive fecundity. The title of highest lifetime fecundity among non-eusocial organisms has been awarded to the “Australian ghost moth . . . Tricten atripalpis,”  a moth that was found to have “laid 29,100 eggs [with] 15,000 eggs . . . found in the ovaries” (Brueland online). The amount of copies of any single gene produced by the Australian ghost moth  is extraordinary. Of course, not all of the eggs will be fortunate enough to survive and become adults; the Australian ghost moth has an “unavoidable high juvenile mortality” rate (Brueland online). Because of the environment that the Australian ghost moth in habits, Australian ghost moth genes rely heavily on maximizing the quality of fecundity. Most of the genes will not survive long, many genes will not even survive long enough to replicate themselves again down the germ-line. Thus these genes do not have a high degree of longevity.

Longevity, the third characteristic that determines the survival of a replicator, regarded as the life span of the individual replicator. Longevity is arguably the least important of the three characteristics important to the survival of any replicator. Dawkins states that the characteristic of “fecundity is much more important than longevity of particular copies” (Selfish 194). The reason that longevity is important, however, is that a replicator must survive long enough to be copied. High longevity in a replicator may allow that replicator to be copied more, but as long as the replicator is copied, the information in a replicator will be passed. In essence, replicators “need not last forever. They need only last long enough to produce additional replicators” (Dawkins, Extended 84). The idea works in practice as well, as genes themselves in any particular organism do not live long at all in comparison with the life span of the information of a particular gene in the form of copies of the gene that is, potentially, immortal.

We must not, of course, confuse the longevity of a replicator with the longevity of a vehicle.  In discussing the longevity of a replicator, we are discussing how long an individual replicator will be available for replication. The life span of a strand of DNA in the nucleus of a cell is an example of the property of longevity in a replicator; this is not likely to have anything to do with the life span of the vehicle that the replicator helps produce. Although most genes in an organism are “well protected. . . [and] can sometimes survive as long as the lifetime of the organism” (Blackmore 101). The information encoded in the replicator, along with obvious environmental factors, determine the longevity of any particular vehicle. Of course, vehicle longevity is important to replicator longevity for similar reasons, and there is good reason to expect replicators to produce vehicles with long life spans. Vehicles with a high degree of longevity have a longer lifetime to reproduce and pass their genes on to another generation. While humans are used to somewhere between 40 years to a century of life, the rest of the genetic world of biology varies greatly. The mayflies are renowned in the world of entomology for "their one-day adult life span" (Mountains 6). Mayflies make up for their lack of life span with a high degree of fecundity. In a female mayfly's short lifetime, the mayfly "can lay up to 8,000 eggs" (Seewer A7). All of these eggs are laid in their short life span. In essence, the genes that build a mayfly use the vast majority of the resources that can be acquired to reproduce quickly and efficiently. Any resources that would have otherwise gone to extending the life span of the mayfly are, for whatever reason, trumped by high reproductive fecundity. In other words, in the ecological niche of the mayfly, there is a great benefit to reproducing quickly and in great quantities. Of course, there are other ecological niches, some which are filled by vehicles with a much longer life span than that of the mayfly. The bristlecone pine is accepted by many biologists to be the oldest living organism on earth. Bristlecone pine trees have "been known to live at least 4,900 years" of age (Babowice 1). The oldest of these trees, which grew in Nevada's Great Basin National Park and was granted the name "Prometheus," unfortunately was "cut down in 1964" (Babowice 1).

Prometheus represents the extremity of longevity in the genetic world, but what artifacts in the world have stood the test of time? The example that I will use is perhaps a rather obvious example. The pyramids of ancient Egypt have stood the test of time for over four thousand ears. The Egyptian ruler Khufu ordered the building of “his 480-foot-high pyramid. [The pyramid's] purpose was to preserve his body for eternity” (Art-i-facts 3). Although the body of Khufu was not preserved for eternity because his tomb and its chambers “were plundered in ancient times,” the pyramid of Khufu stands as a monument to longevity in artifacts; the pyramid is the only one of the “seven ancient wonders that has survived” to this day (Art-i-facts 3). Thus we can see that both memes and genes may construct vehicles with high degrees of longevity. Prometheus as an organism survived for thousands of years, and the great pyramids may still have some more years left in them.

With the knowledge about what enables a replicator to survive, we press on to get a picture of how life might have originated. For life to originate there must be enough energy and resources in the area, and, most importantly, a replicator of some kind. Life on the planet earth almost certainly at least 3.5 billion years old. Scientists know this "because they have found fossils of microbes in rock of about that age" (Conner). Current scientists are now giving "greatest weight to the family of theories based on an organic 'primeval soup'" (Dawkins Watchmaker 148). This model of life's origin comes from the work of Stanley Miller. Miller conducted an experiment in 1952 where he "attempted to reproduce the conditions that first produced life on earth" (Reville 2). After his simulation had finished, Miller noted that several organic compounds as well as "several amino acids" could be found in the chemical analysis" (Reville 2). Because amino acids make up proteins, the "primeval soup" model holds a firm ground among biologists.

While Miller's model is quite plausible, there is another rival theory that is also gaining ground. This rival theory is that "of the Glasgow chemist Graham Cairns-Smith" (Dawkins Watchmaker 148). Glasgow claims that "the random nature [of the primeval soup] would make it highly probable that some of the molecules produced would poison the mixture" (Connor). Instead, Cairns-Smith proposes that the original process of replication may have been much different from the process we view today in studying the cells of different organisms. Cairns-Smith even suggests that the process "may have been inorganic, [and] he uses the analogy of crystals growing in a watery solution" (Connor). Cairns-Smith suggests that crystals in a watery solution might replicate themselves in different formations, and the crystals might have used clay to form "the basis of life's origin" (Connor). What is important, however, is the inorganic molecules ability to retain information and make copies, or have copies made, of themselves. There is another idea of the origin of life. This idea is called the "panspermia hypothesis," the idea that life began "elsewhere in the universe and was subsequently seeded on earth" (Reville 3). This idea is not irrational; indeed evidence shows that the "Murchison meteorite which landed in Australia in 1969 contains about 80 amino acids, including at least eight of the 20 amino acids that are present in proteins in earth organisms" (Reville 3). I will not go into detail about this idea though, for although the hypothesis is a possibility that should not be overlooked, a replicator still must have originated somewhere, and that means somewhere along the line there must be a self replicating entity that formed either organically, or inorganically. The replicator must have been able hold together long enough to be copied, copy itself accurately, and copy itself as often as possible.

I have explained the popular theories of how a replicator might have originated, but now I would like to focus more specifically on memes. Perhaps the best speculation on the origin of memes comes from Susan Blackmore in her book The Meme Machine. In Dr. Blackmore’s book, Blackmore places heavy emphasis on imitation as the origin of the meme as a replicator. Imitation might have originally come about as a method of learning, for imitation “is certainly a ‘good trick’ if you can acquire [the trait of imitation]. If [a human‘s]  neighbor has learned something really useful. . .[their may be a benefit] (in biological terms) to copy [the neighbor]” (Blackmore 75). Imitation could get the replication process started for the meme. When the “new replicator was born,” the memes that would be selected were “those with high fidelity, high fecundity, and longevity” (Blackmore 102). This process would have little effect on genes at first. The effects would likely be confined mainly to early humans building small artifacts, but eventually the meme would gain more ground based on the memes higher degrees of fidelity, fecundity, and longevity over the gene.

This brings up the topic of scaffolding. The removal of scaffolding occurs in biology when some characteristic of life is no longer needed, so evolution removes the characteristic. An example of scaffolding removal that is easy to visualize is the construction of a Roman Arch. A Roman arch is built in such a way that, if one were to remove any single piece of the arch, the entire arch would fall to the ground. A Roman arch, however, can be constructed quite easily with the use of a scaffold. When the arch is fully complete, and the scaffold is no longer of use, the scaffold is removed to reveal a full and free standing Roman arch. The example of a Roman arch is much like what happens in evolutionary biology. Often pieces of biological machinery evolve to serve one purpose, but when parts of the organism are no longer needed they are removed because components are a waste of energy for an organism to have. This removal might leave other bits of biological machinery that had evolved off of the previous component, but show little or no sign of the previous component that the new biological machinery was built off. The idea of scaffolding removal does not just work for the biological machinery of an organism, however, but replicators themselves as well. If Cairns-Smith's idea of inorganic replicators is correct, inorganic replicators are a type of scaffold for organic replicators. Once inorganic replicators started to employ the use of organic molecules such as RNA for replication and survival, the organic molecules would soon be able to replicate even faster than the crystals until the organic material surpassed their predecessors altogether. Thus even if life could not arise organically, we can see how a type of scaffolding could be employed to give us the end result of something very different. I do feel obliged at this time to remind the reader that evolution has no foresight or plan, and if there is to be some sort of scaffolding through the process of evolution, every intermediate of the scaffolding must have been selected as profitable to posses at one time in evolutionary history.

This scaffolding leads me to what may seem to be a rather silly question, but in taking into account my own ideas of memetic ecology and the idea's implications I have yet to see any faults in my line of reason. If there is indeed the possibility that our current genetic material emerged from the scaffolds of a much slower inorganic replicator, is there a possibility that the new, speedy, memes overtake their genetic predecessors? What is stopping the meme from overtaking the gene as a replicator? With, on average, a much higher degree of fidelity, fecundity, and longevity, the meme leaves any other replicating entity in the dust. Genes just can not keep up. Just think about how fast the ecological counterparts of genetics have set themselves up, and in only thousands of years, a blink of the geological time scale. Memes are surpassing genes by building cities, highways, and farmlands. This is, of course, not because of any conscious foresight of the replicator; the result is not even the conscious foresight of humans, but a mere algorithm of evolutionary biology. I would like to leave the reader with this idea in mind, that memes are rapidly overtaking their predecessors because of their superior fidelity, fecundity, and longevity. My prediction as of now, is quite radical, so I present the position as humbly as possible as only a student of biology. I predict that memes will one day completely overtake their predecessors. I would, to be honest, enjoy being wrong on this prediction. Some replicators are spreading that result in the saving of genetic ecosystems, but I fear they may be too little too late. Whatever the case may be, I hope that the reader can now approach the idea of replicators, taking into account the principles of fidelity, fecundity, and longevity.

Works Cited

"Art-i-facts: Khufu's Grand Gallery." Dig Jan. 2002: 3.

Babowice, J. H. "Tough Bristlecone Pine Takes Title of Oldest Tree." Daily Herald 19 Mar. 2003, Lake ed.: 1.

Blackmore, Susan. The Meme Machine. Oxford: Oxford UP, 1999.

Brueland, Henning. University of Florida Book of Insects Records Chapter 18 Highest Lifetime Fecundity. 15 Dec. 1995. Zoological Institute, Dept. of Animal Ecology, University of Bergen. 27 Feb. 2003 <http://ufbir.ifas.ufl.edu/Chap18.htm>.

Connor, Steve. "Science." The Independent 21 Nov. 1993, 3rd ed.: NOPGCIT.

Dawkins, Richard. The Blind Watchmaker. 2nd ed. New York: W.W. Norton & Company Inc., 1996.

Dawkins, Richard. The Extended Phenotype. 2nd ed. Oxford: Oxford UP, 1999.

Dawkins, Richard. The Selfish Gene. 2nd ed. Oxford: Oxford UP, 1989.

Gilman, Michael, et al. Recombinant DNA. 2nd ed. New York: W.H. Freeman and Company, 1992. 1-10.

Lindahl, Tomas, and Richard D. Wood. "Quality Control by DNA Repair." Science 3 Dec. 1999: 1897-1905

"Mountains of Mayflies Mean it's Time to Compost." BioCycle July 1997: 6.

Seewer, John. "Mayflies Signify a Healthy Lake Erie, But They're Yucky!" Seattle Times 5 July 2000: : A7.

Reville, William. "How it all Started, Just After the Soup." Irish Times 5 Oct. 1997, City ed.: 2.

What is a Replicator? 2000. Thinkquest.org. 26 Feb. 2003 <http://library.thinkquest.org/C004367/ef1.shtml>.

Yount, Lisa. Genetics and Genetic Engineering. New York: Facts on File Inc., 1997. 29-31.

Works Consulted

Nicole, Dyer. "Older than Aerosmith." Science World 10 Dec. 2001: 5.

Hull, David, "Replication", The Stanford Encyclopedia of Philosophy (Winter 2001 Edition), Edward N. Zalta (ed.), URL = <http://plato.stanford.edu/archives/win2001/entries/replication/>.

Ince, Martin. "Life on Earth - and beyond: Never Mind Monkeys, How About Martians as Ancestors, Writes Martin Ince:." Financial Times 29 Aug. 1998, London ed.: 6.

In Search ofthe Secrets of Aging: The Genetic Connection. 1996. National Institute on Aging. 25 Mar. 2003 <http://nia.nih.gov/health/pubs/secrets-of-aging/p2.htm>.

McMullin, Barry. Replicating Confusion. 4 Mar. 1996. 21 Feb. 2003 <http://www.eeng.dcu.ie/~alife/ecal95/rpl-l2h/node2.html>.

Romney, Lee. "Group Hopes to Clone Oldest Living Tree." Milwaukee Journal Sentinel 13 Oct. 2002, Early ed.: 3A.

Romney, Lee. "The State; Effort to Clone Bristlecone Pine Fails; Hopes of Replicating the 4,768-year-old Methuselah Live On, However, in the Seedings Growing from its Cones." The Los Angeles Times 6 Feb. 2003, Home ed.: B6.

What is a Meme? Comp. Christy Buzzard, et al. Beloit College. 21 Feb. 2003 <http://www.beloit.edu/~fyi/meme%20site/What.html>.
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