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including Ross McIntyre, Kendall Smith, Steven Gillis, Paul Baker, Richard Robb, and Margaret Favata, all played instrumental roles in the identification, isolation, purification, and characterization of the IL-2 molecule. In 1960, Nowell made the serendipitous discovery that kidney bean extract containing a compound called phytohemagglutinin (PHA) induced the division of lymphocytes, which were previously thought to be endstage cells that lacked the ability to proliferate. Soon thereafter in 1965, two seminal papers appeared in the journal Nature describing the discovery of a blastogenic activity found in the culture media of stimulated lymphocytes that promoted their proliferation. However, the responsible agent in the lymphocyte “conditioned media” was yet to be characterized as a single molecule. In 1972, Smith, who had previously been at the National Cancer Institute (NCI), joined McIntyre’s hematology research laboratory at Dartmouth as a postdoctoral fellow. The primary research interest of the laboratory at that time had been antiviral factors called interferons. Following a loss of funding in 1973, Smith traveled to France where he continued his immunological training with Georges Mathe, who was using immunotherapy to treat leukemia. By 1974, Smith had returned to Dartmouth as an Assistant Professor of Medicine in the Division of Hematology and Oncology. As head of his own laboratory, Smith recruited Gillis to join his lab as a predoctoral fellow, and Paul Baker joined the lab in his own postdoctoral fellowship. In 1976, a report from the NCI showed that the “lymphocyte conditioned media” could support the long-term growth of T lymphocytes in culture. This activity was most likely caused by the same ingredient that had already been identified as the blastogenic factor a decade earlier in the Nature papers. With his small team at Dartmouth, Smith set out to find the active ingredient in the lymphocyte conditioned medium that was responsible for the long-term T cell growth. Several critical discoveries by Smith’s team over the next decade eventually led to the isolation and characterization of the molecule now known as IL-2. First, Steven Gillis successfully derived long-term T cell lines using mouse T cells, thereby confirming the earlier report from the NCI. Next, Paul Baker derived the first T cell clones (progeny of a single cell). These unique cellular reagents were instrumental in the construction of a new bioassay for the T cell Growth Factor (TCGF), as Smith’s team named it. By 1979, Gillis and Baker had left the Dartmouth lab, and two new people were continuing the research. Richard Robb, a postdoctoral fellow, painstakingly purified the TCGF activity from liters of lymphocyte-conditioned media, while Smith used the bioassay to quantify the purified fractions. By 1982, the recently arrived Margaret Favata had successfully 22 generated monoclonal antibodies reactive with the IL-2 molecule, enabling the purification of milligram quantities of IL-2 to be purified through a one-step affinity process. Through these steps and years, Smith and his laboratory isolated and characterized the IL-2 molecule in addition to identifying the IL-2 receptor. By 1983, they were able to publish in The Journal of Immunology on the development of monoclonal antibodies and their discovery of the IL-2 molecule. The IL-2 molecule is now known to be a 15.5 kDa globular glycoprotein of 133 amino acids. Its discovery and the characterization of its structure and function have been instrumental in the development of an entirely new class of “immunological molecules.” IL-2 serves as a prototype for the group, which currently includes 29 members. These are considered to be the “hormones of the immune system,” the immunological equivalent to those in the endocrine system. They serve to ‘communicate’ between the varying cells of the immune system and lead to their development, reproduction, and responses to invading, foreign microbes. The influential discovery of IL-2 brought research within the medical community to a new level, furthering the understanding of the complicated human immune system. The understanding of IL-2 and its functions has already aided researchers in developing new treatments for immunological diseases. Most immunosuppressive therapies used today, such as glucocorticoids and cyclosporin-A, are now known to act by blocking the production and activities of IL-2. In addition, applications of interleukins include new experimental immune-based therapies, now being pioneered by Smith and others, for the treatment of cancer and chronic viral infections, such as those due to HIV and hepatitis C virus. Researchers have also developed monoclonal antibodies reactive with interleukins and cytokines as new treatments for rheumatoid arthritis and inflammatory bowel disease. These applications provide hope for future treatments of immunological diseases, as well as for both cancer and HIV, through continuing research in the rapidly evolving field of molecular immunology. The authors would like to sincerely thank both Dr. McIntyre and Dr. Smith for their illuminating interviews as well as Dr. Smith for his kind help with editing. DARTMOUTH UNDERGRADUATE JOURNAL OF SCIENCE Invasion of the Red Fire Ant: A Multi-layered Metaphor for Managing Invasion through the Study of Behavior TUCKER MURPHY ʻ04 Unlimited pain. Hormiga brava (fierce ant). Formiga de fogo (fire ant). Formiga lava-pé (wash-foot ant). Solinopsis Invictus (invincible ant). These are but a few of the titles garnered by the infamous red fire ant as it cuts a swathe of invasion and decimation across the North American countryside. Beyond a show of human fear and loathing for these tiny ants (and perhaps, a fair amount of imagination), these names exhibit a true desire and passion to describe the behavior related to ant and human encounters. Admittedly, most fire ant aliases focus only on their ferocious stings and were probably coined by those who had the misfortune of coming into contact with it. But it is likely that the scientific name, at least, takes into account the “Fiery aggression, voracious feeding, and prolific nest building” characteristic of this species of ant (Taber 2000). The anthropomorphic nature of many of these names suggests that humans also tend to identify with its social ways. Because of the extreme and social character of this invasive species one cannot help but find interest in their behavior. It is no surprise, then, that fire ants have become a classic case study for behavioral ecologists as well as those biologists out to prove the severely damaging nature of invasions. The destruction and the overwhelming ecological changes wrought by red invasive fire ants and other exotic invasives with similar behavioral characteristics demonstrates to even the most skeptical of critics why a war needs to be waged against invasive species. Ironically, because they are such successful invaders and because they are so late in their stages of invasion in North America, fire ants also illustrate the near futility that is often involved in the war against invasive species. The only real weapon humans have on their side in this war lies within a thorough understanding of themselves and the enemy, which, in the case of the fire ants, includes their behavioral flexibility, their eusociality, and their human commensalism. The invasion of the red fire ant illustrates the almost inevitable consequences of global change; it is also a prime example of how human understanding of behavior can provide, at the very least, a window into the future and, possibly, a way of altering this future through changes in the behavior of both ourselves and other invasives. SPRING 2003 E.O. Wilsonʼs War Strategy: A Behavioral Approach One cannot speak of invasive red fire ants without also mentioning E.O. Wilson in the same breath. The path of the fire ant as it sweeps across Southern United States is inextricably linked with the career of this famous behavioral ecologist, evolutionary biologist and world authority on ants. A very young E.O. Wilson first discovered this invasive species--accidentally introduced in shipping from South America--in Mobile, Alabama in 1942 and was the first to publish on them in a study undertaken for the Alabama Department of Conservation. From the very beginning, Wilson placed great emphasis on examining behavior of this invasive species and noted key characteristics, such as the tendency of colonies to sometimes adopt queens from outside their own nests (Taber 2000). His original research led him to a life long passion in myrmecology and the conclusion that “Ants are premier organisms for research in behavioral ecology and sociobiology” (Holldobler and Wilson 1990). The fire ants, in the midst of a rapid spread in all directions out of Mobile at the time Wilson first encountered them, have now expanded their range to all over the southern United States and are serious pests (Taber 2000). However, Wilson’s behavioral approach has remained unchanged in its utility. The value of his original line of attack has become apparent over time, as many of his predictions regarding the invasion have come true (Taber 2000). He understood the flexible and almost invincible nature of the fire ant that would allow it to expand into many varied habitats in the western United States and Mexico despite humankind’s best efforts at control. In fact, forty years before Californians discovered a series of infestations in their own backyards Wilson had predicted their arrival (Taber 2000). Recently, weight has been added to the approach Wilson used so well; Holway and Suarez write, “This information will not only aid control efforts for particular species, but the identification of behavioral attributes common to different invasive taxa could also provide a basis for predicting which species have the potential to become problematic invaders” (1999). Essentially, through behavioral studies it is possible to gauge whether an invading species is like a live warhead 23 ready to explode upon reaching a new ecosystem or simply a blank. Perhaps E.O. Wilson’s most prophetic and telling statement was his labeling of the war against fire ants as “the Vietnam of entomology” (Taber 2000). Though those who have mistakenly found themselves in the midst of a fire ant colony and felt the full fury of their attacks might see war as an obvious analogy, Wilson’s comparison operates on a deeper level. The war of which he speaks is one being fought on a much broader scale across oceans and continents. It is an effort to prevent the complete deterioration of the great faunal realms (Wallace’s realms, corresponding to the main continents) and subsequent loss of biodiversity throughout the world due to introduced exotics (Elton 1958). Currently, such invaders are the second leading cause of extinction in the world. Though in the case of the red invasive fire ant most casualties are non-human native species, it is “the most comprehensive aggressive and costly conflict ever between man and bug” (Brouwer 2000). Protesters may not march in the streets over wars with insects, but there are many a doubter or critic who question whether such wars are worth the expense. There are still others who see only inevitability, since humans themselves and their disturbance and transport have facilitated the invasion. In the face of such odds, one cannot help but wonder whether this campaign will end in an outcome similar to its ill-fated namesake. Phenotypic Plasticity, High-tech Armaments, and the Super-Individual The trouble with the fire ants is that they embody almost all of the behavioral characteristics of classically good invaders with the many added benefits derived from their eusocial behavior. Ignoring for a moment the social traits particular to the fire ants, there are certain generalizations that can be made about behaviors that predispose any exotic to invasion. As outlined by Porter and Savignano S. invicta posses seven such behaviors: “(1) It prefers heavily disturbed habitats associated with human activity. (2) It tolerates a wide range of climatic conditions and (3) utilizes a wide variety of food resources. (4) It is relatively abundant in its homeland. (5) Workers are highly variable in size, a characteristic that may allow them to utilize a broader feeding niche. (6) Mated queens are capable of establishing new colonies after transport [i.e. effective dispersal]. (7) Finally, colonies of this species have a high reproductive capacity; they grow rapidly and can produce thousands of reproductives per year.” (1990) 100 90 S. invicta discovered in Puerto Rico 80 Mirex registrations withdrawn 70 HECTARES INFESTED x 10 6 EPA started public hearing on use of Mirex 60 50 Buren's Taxonomic revision Large-scale program started to control IFA with Mirex bait 40 30 20 10 Mirex belt developed IFA control with organochlorine insectcides discontinued Federal-state control program started 0 First survey of IFA distribution 1930 1940 1950 1960 YEARS 1970 1980 1990 Fig. 1. The S-shaped growth curve showing IFA infestation over time as well as some important historical events (Taber 2000). 24 DARTMOUTH UNDERGRADUATE JOURNAL OF SCIENCE Essentially, these characteristics allow the red fire ant to adapt a wide variety of habitats, making it a generalist. Of particular interest is the fourth observation: if a species is abundant in its homeland (where natural predators and competitors are present and the species is confined to a more constricted niche), then there is good indication that it would be successful if released from its native restraints. Of course, it is red fire ants’ social nature that defines their great capacity for invasion. The success of social insects across a wide variety of habitats is largely derived from the added flexibility their sociality allows them. Such flexibility is extremely beneficial to new invaders “that must quickly learn about, exploit and respond to new resources and challenges in receiving communities” (Moller 1996). Individual ant workers can survive on their own and perform most tasks as competently as individual insects of more solitary species: red invasive fire ants, each with their own powerful stings that can be used multiple times, provide a good example of this competence (Holldobler and Wilson 1990). However, sociality takes an ant’s ability to perform almost all of these tasks far beyond mere competence. It permits a fire ant to switch its behavior between two options: to act alone or to act as super-individuals within their colonies (Moller 1996). Not only is their niche broadened by their eusocialty, but ants are alone in this niche: they are the only eusocial predators occupying the soil and ground litter (Holldobler and Wilson 1990). More specifically, some of the additional benefits of a social life in red invasive fire ants include better dispersal, higher reproductive rate, greater longevity, a broader niche, more effective predator defense, enhanced competitive abilities and a lack of intraspecific competition (Moller 1996). Underlying behaviors behind all of these traits are communication, cooperation, and division of labor. A closer look into the mechanics of predator defense reveals how these behaviors result in such a wide variety of benefits and function to increase flexibility. Ants can “communicate with one another about predation risks encountered within the colony’s foraging range and then adjust their use of such areas according to feeding rewards and risks of starvation” (Moller 1996). Also, because the colony as a whole is the key reproductive unit, only nestcentered predation is a substantive threat to established colonies (Moller 1996). Death or mutilation of single ant means little to the functioning and propagation the super-organism of the colony. This truth is even more apparent in fire ant colonies where masses of ants resting or patrolling function as reserves, an activity which permits “fine-tuning to the inevitable wide shifts that occur in the requirements of the brood” (Holldobler and Wilson 1990). SPRING 2003 Genetic Plasticity, More High-tech Armaments and the Super-Colony As if the red invasive fire ants did not already possess enough firepower and behavioral flexibility, they also have an history of genetic plasticity. They have undergone two metamorphoses of particular concern since their arrival in the United States. The first such genetic alteration was for greater cold tolerance (James et al. 2002). This new and improved invader arose from the hybridization of the black (S. richteri) and red imported species (S. invicta) of fire ants. S. richteri arrived in the United States before S. invicta, but has since been largely overwhelmed by its more vigorous cousins (Taber 2000). The additional ability of fire ants to withstand temperatures below freezing would prove catastrophic, since as E.O. Wilson originally predicted, one of the few (and therefore important) factors limiting fire ants’ distribution to the southern States is their inability to tolerate severe cold (Taber 2000). The second, perhaps even more serious, genetic change undergone by red imported fire ants harks back to one of E.O. Wilson’s original observations that “IFA [invasive red fire ant] colonies sometimes adopt queens from outside the nest” (Taber 2000). With this apparently minor insight into behavior Wilson hit on one of the most intensely studied topics today regarding fire ants: the steadily increasing abundance of polygynous and unicolonial colonies among S. invicta in the USA since the 1970s (Bourke 2002). Though multiple-queen colonies do exist in the red fire ants native habitat, they differ consistently and dramatically “in their social organization and contain relatively few queens that are close relatives, whereas colonies in the United States contain high numbers of unrelated queens” (Ross et al. 1996). The extreme social nature of the polygynous form makes it much hardier and more hazardous than the original monogynous invader. Polygyny takes the concept of super-individual one step further with the creation of a “super-colony,” in which the flexibility is amplified. Since there is no intraspecific territoriality between colonies this new form of ant is able to exist at densities often ten times as great as the monogyne form (Porter and Savigno 1990). Such high numbers lead to a much improved ability to compete against and repel individuals of other species. Porter and Savigno report that polygynous ants are more capable of wreaking havoc on an ecosystem and decimating native ants than are their monogyne predecessors (1990). Perhaps, most alarmingly, polygyny increases the ants invincibility to human attack since “the presence of two or more queens seems to help colonies recover from an attack by pesticides” (Unlimited Pain 2000). In polygynous colonies, if one queen is killed there is still another to keep producing more fire ants. 25 MONOGYNOUS COLONY POLYGYNOUS COLONY Queen number One Many Genotype of queen(s) BB Bb Queen phenotype High fat reserves, rapid oogenesis Low fat reserves, slow oogenesis (advantageous under monogyny) Worker phenotype Invasion of the Red Fire Ant: Bb workers tolerate multiple queens, but A Multi-layered Metaphor for only if they bear the b allele; Bb workers Managing Invasion through the detect and kill BB queens; all bb quenns Study of Behavior BB workers in (and workders) die early because b is a absence of recessive lethal. Bb workers are intolerant of multiple queens Table 1. Genetic and phenotypic traits of monogynous and polygynous colonies in introduced Solenopsis invicta fire ants (Bourke 2002). If the consequences of the spread of the polygyne form are as disagreeable as they appear, the question for invasion biologists attempting to look into the future becomes: why is it increasing in frequency and can the tide of increase be stemmed? The answer can only be discovered through an interdisciplinary approach combining genetics, behavior, and the ecology surrounding S. invicta. In a noteworthy paper, Krieger and Ross found a rather simple feature underlying a fairly complex behavior (1998). They established the genetic basis of gyny variation in fire ants and pinpointed the shifting of two alleles (B and b) at a particular locus (see table 1). According to Krieger and Ross, all fire ants known to be polygynous contain b-like alleles, while those that are monogynous only have the B allele (1998). In discovering the often searched-for bridge between genetics and social behavior, molecular biologists have demonstrated the true complexity of this invasion and developed predictive models. As to whether they are any closer to stopping the spread of polygynous ants is debatable. Herein lies the problem with a behavioral approach: each time the picture gets more complete it also becomes more complex. Even so, it is the best weapon humans currently have in their conflict with fire ants. Ravages of Invasion Long before the fire ant invasion, the often overwhelming effects of ants on an ecosystem were known. It does not necessarily take a lifetime devoted to the study of myrmecology to learn that ants “alter their physical environment profoundly” (Holldobler and Wilson 1990). The fire ant has more than lived up to this billing. In an aptly titled article “When Fire Ants Move In, Others Leave” looking at the impact of an invasive species on a broad ecological scale, Elizabeth Pennisi reports that S. invicta severely disrupts the gradient of ant species across the East Coast of the United States (2000). Biologist expected to find an increase in the number of ant species 26 with decreasing latitude (diversity tends to increase closer to the tropics) and they did in all places except those where the red invasive fire ants were present. Here, native ants were either locally extinct or very rare. In a much smaller scale study, limited to a single field, Porter and Savigno determined that competitive replacement is the primary mechanism behind the decimation of native ants (1996). The fire ant also directly (through predation) and indirectly (through induced behavioral changes) disrupts arthropod, invertebrate and vertebrate communities; of particular concern are the many endangered species negatively affected by S. invicta (Forys et al. 2001). Such an overpowering invasion can only end in the simplified environment in which S. invicta becomes the dominant population regulator. This domination and simplification, in turn, might lead to complete ecosystem collapse or greater susceptibility to invasion. Human Fortifications The only hope for humans in their battle against red invasive fire ants is management and control. Recently, several good candidates for this purpose have been brought forward, including a microsporidian disease (Thelohania solenopsae), a parasitic ant (Solenopsis dagerri), and probably the most promising candidate to date, a parasitic phorid fly (Pseudacten curvatus) (Lockley 1996). Essential to the implementation of any of these agents of biocontrol is a thorough behavioral analysis of the interactions between them, their hosts, and the rest of the ecological community. For example, Oi and Williams show that the introduction of the disease T. solenopsae chronically debilitates S. invicta queens (2002). Studies on S. dagerri have found that this workerless, social parasite redirects fire ant workers to its own brood to the detriment of the colony’s larvae (Lockley 1996). Porter’s analysis of behavioral risk is so exhaustive as to address P. curvatus’s possible negative effects on agriculture and on native ants (2000). Based on observations of fly behavior, DARTMOUTH UNDERGRADUATE JOURNAL OF SCIENCE he concludes that it poses only a small risk. However, even this limited risk needs to be balanced against “potential benefits to numerous other native organisms and a high probability that release of this fly will actually benefit native fire ants because impacts on imported fire ants will almost certainly be much greater than those on native fire ants” (Porter 2000). In spite of the optimism generated by Porter’s study and the subsequent release of phorid flies, humans should not begin their victory dances across abandoned fire ant mounds too soon because there is always the chance that fire ants will return. This possibility was the main concern of Oi and Williams , who suggested that “long-term suppression of S. invicta populations under field conditions may be dependent on increasing the ratio of infected to uninfected colonies and/or limiting the rate of reinfestation by uninfected colonies” (2002). Because of their invincible social nature and many modes of rapid dispersal, once fire ants gain a foothold they are almost impossible to extinguish or expel. Even Porter qualifies his results with the statement, “A concern is that no one has ever successfully used classical biocontrol agents against any social insect” (2000). Such reservations bring into question what management and control really entail. In the end, behavioral studies of biocontrol agents in the laboratory can provide invasion biologists with predictions only up to a certain point. After that, scientists must take the leap of faith necessary to introduce the control agent and continue their experiments on a much larger scale in which post-control field observations become of paramount importance. Combating the Enemy Within Matters of control are further complicated by fire ants’ close relationship with humans. In fact, many biologists have observed that fire ants thrive in exactly the areas from which we are trying to eradicate them (Forys et al. 2001). Walter Tschinkel, a biologist with a good grasp of insect behavior, provides a definitive opinion on the issue: “The fire ant is clearly and dramatically associated with ecologically disturbed habitats, created mostly by man. . .It is abundant in old fields, pastures, lawns, roadsides, and any other open, sunny habitats. It shares these habitats with many other ‘weedy’ plant and animal species . . . Man is the fire ant’s best friend’” (Brouwer 2000). This “friendship” is dependent on two main aspects of human interaction with the environment. First, transport by humans, including the movement of colonies with mated queens in plant nursery stock or construction fill and transportation of newly mated queens in vehicles and even ballast tanks, creates a dispersal corridor for fire ants (Forys et. al 2001, Taber 2000). Second, the result of human disturbance is a habitat with reduced native predators and competitors that can be easily colonized by SPRING 2003 human commensals or early succession species such as the fire ant (Forys et al. 2001). Solenopsis Invicta’s affinity for human habitats adds to both the element of futility and of hope in the fight against fire ants, for the success or failure of a management program is largely dependent on how we behave. Not only do fire ants seek human company, it is no secret that these ants are mirrors of ourselves in many aspects of their social behavior and in their effects on an ecosystem. E.O Wilson recognized this truth and took much of what he learned in his years of studying ants and applied it to humans in books such as, On Human Nature (1978). Consider the homogenization of the environment by fire ants greatly amplified; that is an idea of what the future could hold in store should humans continue their current destruction of habitat and not attempt to control and prevent invasions. The picture of the earth converted into a giant fire ant mound should be reason enough for why “refuge, beauty and interest, and security” (Elton 1958) are worth fighting for. Luckily, unlike fire ants, we are not so genetically programmed in our actions and tasks that we do not have behavioral flexibility to alter our damaging ways and provide the possibility for a better future, less full of fiery stings. REFERENCES Bourke, A.F.G. (2002). Genetics of social behaviour in fire ants. Trends in Genetics, 18, 221-223. Brittingham, M.C. and Temple, S.A. (1983). Have cowbirds caused forest songbirds to decline? Bioscience, 33, 31-35. Brouwer, G. (2000, August). Them: California takes on the Fire Ant. LA Weekly, available at http://www.laweekly.com/ink/00/37/featuresbrouwer.php. Carlton, J.T. and Geller, J.B. (1993). Ecological roulette: the global transport of nonindigenous marine organisms. Science, 261, 78-82. Forys, E.A., Allen, C.R., and Wojcik, D.A. (2001). Influence of the proximity and amount of human development and roads on the occurrence of the red imported fire ant in the lower Florida Keys. Biological Conservation, 108, 27-33. Gilbert, L. (2002). Fire Ants, Armadillos and Phorid flies--Answers to Frequently Asked Questions. National Teachers Association, available at http://uts.cc.utexas.edu/~gilbert/research/fireants/ faquans.html#import. Holway, D.A., and Suarez, A.V. (1999). Animal behavior: an essential component of invasion biology. Trends in Ecology and Evolution, 14, 328-330. Holldobler, B. and Wilson, E.O. (1990). The Ants. Cambridge: Harvard UP. James, S.S., Pereira, R.M., Vail, K.M. and Ownley, B.H. (2002). Survival of imported fire ant (Hymenoptera: Formicade) species 27 subjected to freezing and near-freezing temperatures. Environmental Entomology, 31, 127-133. of Competitive Replacement. Annals of the Entomological Society of America, 81, 913-918. Lockley, T.C. (2002). Imported Fire Ants. University of Minnesota, available at http://ipmworld.umn.edu/chapters/lockley.htm. Perlman, D.L., and Wilson, E O. (2002). Introduction to Exotic Species, available at http://www.earthscape.org/t1/wie01/exotic_species_ intro.html. Mack et al. (2000). Biotic Invasions: Causes, Epidemiology, Global Consequences, and Control. Ecological Applications, 10, 689-710. Moller, H. (1996). Lessons for Invasion Theory from Social Insects. Biological Conservation, 78, 125-142. Oi, D.H. and Williams, D.F. (2002). Impact of Thelohania solenopsae (Microsporidia:Thelohaniidae) on plygyne colonies of red imported fire ants (Hymenoptera: Formicade). Journal of Economic Entomolgy, 95, 558-562. Penninsi, E. (2000). When fire ants move in, others leave. Science, 289, 231. Porter, S.D. (2000). Host Specificity and Risk Asssesment of Releasing the Decapitating Fly Pseudacteon curvatus as a Classical Biocontrol Agent for Imported Fire Ants. Biological Control, 19, 35-47. Porter, S.D. and Savigno, D.A. (1990). Invasion of polygyne fire ants decimates native ants and disrupts arthropod community. Ecology, 71, 2095-2106. Porter, S.D., Vaneimeren, B., and Gilbert, L.E. (1988). Invasion of Red Imported Fire Ants (Hymenoptera: Formicade) - Microgeography Rodda, G.H., Fritts, T.H., and Chiszar, D. (1997). The disappearance of Guam’s wildlife. Bioscience, 47, 565-574. Ross, K.G. and Keller, L. (1998). Genetic control of social organization in an ant. PNAS USA, 95, 14232-14237. Ross, K.G., Vargo, E.L., and Keller, L. (1996). Social evolution in a new environment: The case of introduced fire ants. PNAS USA, 93, 3021-3025. Taber, S.W. (2000). Fire Ants. College Station: Texas A&M UP. Unlimited Pain: The fire ants are coming. (2000, April 15). The Economist, p. 28-29. Vinson, B. (1994). Impact of Invasion of Solenopsis Invicta (Buren) on Native Food Webs. Exotic Ants: Biology, Impact, and Control of Introduced Species. ( D.F. Williams, Ed.). San Francisco: Westview P. Wilson, E.O. (2002). Only Humans Can Halt the Worst Wave of Extinction Since the Dinosaurs Died, available at http://raysweb.net/ specialplaces/pages/wilson.html. FACULTY: Have an undergraduate working in your lab? Has a student of yours produced an especially well-written class paper? Encourage him or her to submit to the DUJS. 28 DARTMOUTH UNDERGRADUATE JOURNAL OF SCIENCE The Self: The Cruise Control of Social Interaction CAROLINE TARNOK ʻ02 Introduction During the last hundred years, the self has become an increasingly important topic of research in psychology. Although it has been studied extensively and should be a subject with which we are personally familiar, the actual nature of the self is still widely debated. Given the number of subtopics that have arisen and the paths that this research has taken, Higgins asserts the value of adopting an explicit perspective on the self and exploring the implications of that perspective (1996). In this paper, I develop an evolutionary framework of the self based on the idea that we rely on others in our social groups for protection and reproduction. I propose that the self is a false exterior created by the brain. It is a system that exists to facilitate presentation strategies, which in turn secure an individual’s place in society. Higgins presents the idea of a self-digest that summarizes a body of information about the individual for the purpose of serving selfregulatory functions (1996). My theory embraces the idea of the self as a survival tool, but I hypothesize that the self functions not to regulate an individual’s role with respect to the general environment, as Higgins states, but with respect to other individuals. Evolutionary Background Dawkins puts forth several ideas that might help explain an evolutionary view of the self. The first idea is that of the selfish gene. He maintains that “. . . we, and all other animals, are machines created by our genes. . . . Our genes have survived, in some cases for millions of years, in a highly competitive world. This entitles us to expect . . . that a predominant quality to be expected in a successful gene is ruthless selfishness” (1989). Animals, as the carriers of these genes and having evolved by natural selection, also exhibit selfish behavior. A selfish act is defined as an act that increases the chance of survival of the agent, no matter how slim, and regardless of whether or not it is to the advantage of others (Dawkins 1989). This idea says nothing about the motives of the individual; it simply implies that animals act so as to increase the survival prospects of their genes. In order to propagate our genes, however, we must not only protect ourselves, but also succeed in reproduction. This struggle for existence includes being dependent on the environment and on other beings (Darwin 1859). SPRING 2003 Human ancestors were social beings, and membership within the complex group structures probably aided survival. Improved predation, defense against predators, and defense against rival groups were all beneficial for these individuals, who may have been “underequipped to deal with their environment” (Sedikides & Skowronski 2000). As in other animal groups, members differ in social status, where those of higher status have access to more resources and higher-quality mates. To propagate their genes, then, humans and other animals rationally (although not necessarily consciously) strive to maintain or advance their status within the group (de Waal 1998). In humans, I propose that the Self developed to facilitate the self-presentation that this position-maintenance requires. The Evolution of the Self Other animals are also able to self-regulate in response to the environment, albeit using simpler and more automatic mechanisms. Still, they influence others’ opinions of themselves and engage in social protection and reciprocation. Yet we do not say that they have a self in the same way that humans do. The key difference may be that the social structures of lower animals are less complex, with status often being more obvious within a comparatively stable hierarchy. Sedikides and Skowronski argue that only humans developed a symbolic self, which includes the mental representation of one’s traits, executive functions, and reflexive potential (2000). This self was the outgrowth of (1) a general evolution in cognitive capacity due to environmental changes and (2) the evolution of a complex, flexible social structure. The second development was arguably the crucial antecedent for the self humans possess today. Other animals can determine their current social standing with relative ease. With a more complex and flexible social organization, however, it is much more difficult for an individual to determine his status at any one time. In addition, it is even more difficult to determine—and remember—the status of others with whom he will need to interact (Sedikides & Skowronski 2000). Facing numerous and complex costbenefit analyses that would require more brain power and time than is typically available, an organism would benefit from developing automated agendas and scripts as soon 29