ASILO, Eden Beth B. DE MESA, Czarina Anne E. BS Biology III-1 Does Selection Operate Primarily On Genes?

(Genes Does Not Primarily Operate On Genes in Selection) ` Does selection act on organisms, genes, groups, colonies, demes, species, or some combination of these? Also, which of these entities might be most influenced by natural selection? Before we cite answers to the above questions we will first define selection and then gene: we define selection as repeated cycles of replication, variation, and environmental interaction so structured that environmental interaction causes replication to be differential (Glenn, Hull and Langman, p. 513). Variation is sometimes considered part of the selection process (Darden and Cain 1989), sometimes as a precondition of selection processes (Hull 1980). Either way, variation is absolutely essential for the operation of the selection processes. If there is no variation, then there are no alternative to select among. Is this variation blind, chance, random, nonprescient, nondirected, nonteleological, unforesighted, or what (Glenn, Hull and Langman, p. 513)? According to the inheritance of acquired characteristics, the environment modifies the phenotype of an organism so that it is better adapted to the environmental factors that produced this phenotypic change in the first place better adapted than those organisms that were not modified in this way. This phenotypic change is then transmitted somehow to the genetic material so that it is passed on in reproduction. Thus, according to this view, species can rapidly adapt to environmental change. In Darwinian evolution inherited variations are random with respect to (i.e., independent of) the effects that they produce, while in Lamarckian evolution they are not. Replication contains two elements; iteration (or repetition or recursion) and information. Early on Dawkins (1976) published a highly influential general account of selection that emphasized the role of replicators. They are entities whose structure contains the information that is passed on differentially in selection. it codes for the character of the individuals (or vehicles) that replicators produce. The only feature of the analysis of selection-type theories provided by Darden and Cain (1989, p. 110) with which we disagree is the demotion of iteration to an ancillary feature of selection. For them, selection is essentially a one-shot deal that can be, but need not be, repeated. They replace iteration with such evaluative notions as benefitting and suffering:

Several types of effects result from the differential interactions. In the short range, individuals benefit and suffer. 1. The transcription unit refers to the contiguous stretch of DNA that encodes the sequence in the primary transcript; this includes (a) the coding sequence of either the mature RNA or protein product, (b) the introns, and (c) the 5 leader and 3 trailer sequences that appear in mature mRNAs as well as the spacer sequences that are removed during the processing of primary transcripts of RNA coding genes. 2. The minimal sequences needed to initiate correct transcription (the promotor) and to create the proper 3' terminus of the mature RNA. 3. The sequence elements that regulate the rate of transcription initiation: this includes sequences responsible for the inducibility and repression of transcription and the cell, tissue, and temporal specificity of transcription. These regions are so varied in their structure, position, and function as to defy a simple inclusive name. Among them are enhancers and silencers, sequences that influence transcription initiation from a distance irrespective of their orientation relative to the transcription start site (pp. 461-462, see also pp. 435 ff. and 457 ff.). One response to this question, advanced by Richard Dawkins in a series of works since The Selfish Gene was published in 1976, is that there is only one level of selection, and it is the level of the gene. Appeal to any higher levels represents a misunderstanding of how natural selection works and is unnecessary, Dawkins argues. Others have argued that there may be multiple levels of selection but that they are reducible to selection at the level of the gene, or at least selection at any of these other levels can be adequately represented at the level of the gene. Still others have held the antireductionist view that there are higher levels of selection beyond the gene, and indeed beyond the individual organism. They hold that selection operates at the level of the group, population, species, and even higher levels, and it does so independently of how selection operates at some or all of the lower levels, all the way down to the gene. Dawkinss claims that all selection takes place at the level of the gene argue that his genocentric approach to evolutionary theory must be rejected because the gene has no special role in evolution at all. As Rosenberg and McShea (2008, p. 158) states that levels and units of selection are as follows: 1 reproduction, with some inheritance of traits; 2 variation arising in inherited traits; 3 differences in fitness among variants. Darwin did not know about genes in the sense we know about them today, but he suspected that natural selection had to operate above the level of the individual organism, at the level of the group. In the following quotation, B. F. Skinner (1974, pp. 4041; see also BBS special issue) says that: Darwins theory of natural selection came very late in the history of thought. Was it delayed because it opposed revealed, because it was an entirely new subject in the history of science, because it was characteristic only of living things, or because it dealt with purpose and final causes without postulating an act of creation? I think not. Darwin discovered the role of selection, a kind of causality very different from the pushpull mechanisms of science up to that time.

For, otherwise, he saw, it would be difficult to explain the persistence over evolutionary time of self-sacrificing behaviors in human beings and other species. Darwin speculated that this sort of unselfish behavior persists owing to its adaptive advantage to the groups that contain selfsacrificing individuals. He wrote: A tribe including many members who . . . were always ready to give aid to each other and sacrifice themselves for the common good, would be victorious over most other tribes; and this would be natural selection (Descent of Man: 166). So in any given environment in which groups compete, the groups with the trait of being composed of cooperating members will be fitter than those lacking the trait or having it in lesser degree. Such cooperative groups, or groups with altruistic members, will persist longer and presumably give rise to more new groups (by splitting or spawning new colonies) than those characterized by less cooperation. Indeed groups of selfish individuals should be destined for extinction. This line of thought has been described as showing that there is a level of selection above the individual organism on which selection operates. Finally, it suggests that if we cannot reductively explain the groups traits in terms of the traits of the individuals that compose it, then we have to accept groups as distinct and irreducible entities, undergoing selection at higher levels (Rosenberg and McShea, p. 159). Numerous biologists and philosophers of biology have presented analyses of gene-based selection in biological evolution (e.g., Dawkins 1976; Hull 1980; Lewontin 1970; Lloyd 1988; Sober 1984; Sober & Wilson 1998; Vrba & Gould 1986), but relatively few have tried to present a general account of selection to see which processes in addition to gene-based biological evolution are genuine selection processes and which are not. (The chief exception is Darden & Cain 1989.) Are selection processes sufficiently different from other sorts of causal processes to warrant a separate analysis? The sort of selection that goes on in biological evolution is surely an instance of selection, but how about other putative examples of selection, for example, the reaction of the immune system to antigens, operant learning, the development of the central nervous system, and even conceptual change itself (Cziko 1995)? Broadly speaking, there are two kinds of gene concepts. Here is a schematic formulation of a referentially indefinite functional gene concept: A gene for trait x is any stably inherited factor that causes an organism [or certain cells of the organism], given the rest of what it has in common with conspecifics, to have the potential for manifesting x, where x will (or can be made to) appear under the appropriate developmental plus environmental circumstances. While discontinuous gene concepts is more specific concepts of the gene, though they may still allow further specification, are committal, at least to some degree, about the structure or location of genes. What is typically required is a mixed mode of identification in terms of both structure and function (Burian, 1995).

The conclusion of the previous question is that biologists cannot explain the persistence of individual traits of altruism, cooperation, or other fitness-reducing dispositions by appeal to their aggregated effects on the fitness of the groups such individuals compose. In other words, natural selection cannot operate on units larger than reproducing individual organisms. If a group has a trait that no individual member of it can have, such as the trait of being composed of mostly altruists, or mostly less-virulent virions, then it is a trait the group will not have for long, a trait without evolutionary significance, and a trait Darwinism need not take seriously. In the levels of selection debate, this argument against higher levels of selection actually went beyond the denial that natural selection requires us to postulate groups with traits irreducible to those of their individual members. The argument concluded that, from the Darwinian point of view, we really do not even need to take individual organisms seriously. For the only real level at which selection operates is the level of the gene. The gene is the only real subject of selection. The argument for this view can be conveniently formulated using the distinction due to Dawkins (1982), and independently to the philosopher David Hull (1988), between replicators and interactors, introduced in Chapter 2. Recall that a replicator is a thing whose structure is copied in the next generation. Thus DNA sequences are paradigmatic replicators. An interactor, or in Dawkinss term, a vehicle, is a thing that interacts with the environment, well or poorly, for better or worse. A replicator may well be its own interactor, or the interactor may be the vehicle that carries around the replicator (hence Dawkinss term, vehicle). Evolution by natural selection can be economically expressed as the differential perpetuation of replicators owing to fitness differences among interactors. The claim Dawkins made in The Selfish Gene is in effect that so far as Darwinian evolution is considered, the only real replicators and interactors are the genes. A great deal of the attractiveness of Dawkinss view is due to the fact that genes are far more faithfully copied across generations than phenotypes, or the traits of groups and individuals for which genes presumably code. Further, the genes seem to many to be the ultimate determinants of organismal form and function, and of group capacities. The genes are the cause of development, for both individuals and groups. And together these seem to some like a solid basis for holding that they are the ultimate beneficiaries and victims of natural selection, and therefore that they are the entities whose traits are ultimately selected for and against. In other words, the genes seem like real targets of selection, real interactors or vehicles. For these reasons, Dawkins argues that evolutionary theory should not take organismal interactors seriously. Compared with genes, which are passed more or less intact from generation to generation, organisms are ephemeral. They are born and they die. They come and go. But genes are forever, or at least their DNA sequences are almost perfectly copied over and over again, and they persist for very long periods as Dawkins referred it on his The Selfish Gene: But genes are denizens of geological time: genes are forever. Genes, like diamonds, are forever, but not quite in the same way as diamonds. It is an individual diamond crystal that lasts, as an unaltered pattern of atoms. DNA molecules don't have that kind of permanence. The life of any one physical DNA molecule is quite shortperhaps a matter of months, certainly not more than one lifetime. But a DNA molecule could theoretically live on in the form of copies of itself

for a hundred million years. Moreover, just like the ancient replicators in the primeval soup, copies of a particular gene may be distributed all over the world. The difference is that the modern versions are all neatly packaged inside the bodies of survival machines.

So we can think of organisms as mere extensions of the genes, as what he called extended phenotypes. Dawkinss thesis is a species of qualified eliminativism. It is not that organelles, cells, tissues, organs, organisms, and groups do not exist. Rather, it is that they have no ultimate explanatory role in evolutionary biology. At most referring to them enables us to abbreviate our descriptions of evolutionary processes, which in fact all transpire only on the level of genes. All of the higher level entities, from cell to organism to groups, along with their productionsbee hives, beaver dams, and spider websare built by the genes to ensure their own survival. When it looks like selection is operating at a higher level, selecting individual giraffes for neck length, it is really selecting genes the protein products of which contribute to neck length. In Dawkinss view, so far as evolution is concerned, there really are no interactors above and beyond the genetic replicators.