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Eusociality: Origin and Consequences

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Eusociality: Origin and Consequences Corrections CELL BIOLOGY. For the article ‘‘The NF1 tumor suppressor criti- BIOCHEMISTRY. For the article ‘‘Engineered single-chain dimeric cally regulates TSC2 and mTOR,’’ by Cory M. Johannessen, streptavidins with an unexpected strong preference for biotin- Elizabeth E. Reczek, Marianne F. James, Hilde Brems, Eric 4-fluorescein,’’ by Filiz M. Aslan, Yong Yu, Scott C. Mohr, and Legius, and Karen Cichowski, which appeared in issue 24, June Charles R. Cantor, which appeared in issue 24, June 14, 2005, of 14, 2005, of Proc. Natl. Acad. Sci. USA (102, 8573–8578; first Proc. Natl. Acad. Sci. USA (102, 8507–8512; first published June published June 3, 2005; 10.1073͞pnas.0503224102), the authors 6, 2005; 10.1073͞pnas.0503112102), the footnotes appeared in note the following. On page 8574, the last sentence of the first the wrong order, due to a printer’s error. The ** footnote on paragraph in the left column, ‘‘Clarified lysates were normalized page 8507 should have read: ‘‘Throughout this paper, we use the for protein levels and analyzed by Western blotting with the terms monomer(ic), dimer(ic), and tetramer(ic) to refer to the following antibodies: phospho-p70S6K (T-389), phospho- number of biotin-binding domains present in a molecule, re- tuberin (S-939), phospho-tuberin (T-1462), phospho-Akt (S- gardless of the number of polypeptide chains it has.’’ In addition, 473), tuberin (C-20) from Santa Cruz Biotechnology, and the †† footnote on page 8512 should have read: ‘‘We note, protein kinase B␣͞AKT1, actin, and ␤-tubulin from Sigma- however, that extremely high-affinity mutants might not be Aldrich,’’ should read: ‘‘Clarified lysates were normalized for recovered in this procedure due to near-irreversible binding of protein levels and analyzed by Western blotting with the follow- biotin from the culture medium.’’ ing antibodies: phospho-p70S6K (T-389), phospho-tuberin (S- ͞ ͞ ͞ ͞ 939), phospho-tuberin (T-1462), phospho-Akt (S-473) from Cell www.pnas.org cgi doi 10.1073 pnas.0508348102 Signaling Technologies, tuberin (C-20) from Santa Cruz Bio- technology, and protein kinase B␣͞AKT1, actin, and ␤-tubulin from Sigma-Aldrich.’’ On page 8574, the second sentence of the PERSPECTIVE. For the article ‘‘Eusociality: Origin and conse- second paragraph in the left column, ‘‘For lentiviral production, quences,’’ by Edward O. Wilson and Bert Ho¨lldobler, which HEK293T cells were transfected with the ⌬8.2 lentiviral con- appeared in issue 38, September 20, 2005, of Proc. Natl. Acad. Sci. USA (102, 13367–13371; first published September 12, 2005; struct (encoding gag, pol, rev), VSVG, and either empty pKO ͞ vector or the pKO vector containing the following sequence 10.1073 pnas.0505858102), the authors note that Donat Agosti transcribing short-hairpin RNAs specific to NF1: 5Ј-TTATA- was omitted from the list of editors in ref. 48. The corrected reference appears below. AATAGCCTGGAAAAGG-3Ј,’’ should read: ‘‘For lentiviral CORRECTIONS production, HEK293T cells were transfected with the ⌬8.2 lentiviral construct (encoding gag, pol, rev), VSVG, and either 48. Ward, P. S. (2000) in Ants: Standard Methods for Measuring and Monitoring empty pLKO.1puro vector or the pLKO.1puro vector containing Biodiversity, eds. Agosti, D., Majer, J. D., Alonso, L. & Schultz, T. R. the following sequence transcribing short-hairpin RNAs specific (Smithsonian Institution Press, Washington, DC), pp. 99–121. Ј Ј to NF1: 5 -TTATAAATAGCCTGGAAAAGG-3 .’’ In addi- www.pnas.org͞cgi͞doi͞10.1073͞pnas.0508220102 tion, the authors would like to add the following statement to the Acknowledgments: ‘‘We would also like to thank Dr. William C. Hahn (Harvard Medical School and Dana–Farber Cancer In- stitute, Boston) and the Broad Institute (Cambridge, MA) for the pLKO.1 lentiviral constructs.’’ www.pnas.org͞cgi͞doi͞10.1073͞pnas.0508351102 www.pnas.org PNAS ͉ November 1, 2005 ͉ vol. 102 ͉ no. 44 ͉ 16119–16120 Downloaded by guest on September 27, 2021 ANTHROPOLOGY. For the article ‘‘Descent of the larynx in chim- panzee infants,’’ by Takeshi Nishimura, Akichika Mikami, Juri Suzuki, and Tetsuro Matsuzawa, which appeared in issue 12, June 10, 2003, of Proc. Natl. Acad. Sci. USA (100, 6930–6933; first published May 29, 2003; 10.1073͞pnas.1231107100), the authors note that the symbols for Cleo and Pal were switched in the legend for Fig. 2. The figure and its corrected legend appear below. These errors do not alter the conclusions of the article. www.pnas.org͞cgi͞doi͞10.1073͞pnas.0508487102 Fig. 2. SVT growth in the three chimpanzee infants: Ayumu (male, ࡗ), Cleo (female, ■), and Pal (female, Œ). (a) Growth of the oral (solid line) and oropharyngeal (dotted line) components of the SVTH.(b) Growth of the SVTH (solid line) and SVTV (dotted line). (c) Age-related changes in the ratio of SVTH to SVTV lengths. 16120 ͉ www.pnas.org Downloaded by guest on September 27, 2021 PERSPECTIVE Eusociality: Origin and consequences Edward O. Wilson*† and Bert Ho¨ lldobler‡§ *Museum of Comparative Zoology, Harvard University, 26 Oxford Street, Cambridge, MA 02138-2902; ‡School of Life Sciences-LSC 274, Arizona State University, Tempe, AZ 85287-4501; and §Theodor-Boveri-Institut fu¨ r Biowissenschaften (Biozentrum), Universita¨t Wu¨ rzburg, Lehrstuhl fu¨ r Zoologie II, Am Hubland, D97074 Wu¨ rzburg, Germany Contributed by Edward O. Wilson, July 12, 2005 In this new assessment of the empirical evidence, an alternative to the standard model is proposed: group selection is the strong binding force in eusocial evolution; individual selection, the strong dissolutive force; and kin selection (narrowly defined), either a weak binding or weak dissolutive force, according to circumstance. Close kinship may be more a consequence of eusociality than a factor promoting its origin. A point of no return to the solitary state exists, as a rule when workers become anatomically differenti- ated. Eusociality has been rare in evolution, evidently due to the scarcity of environmental pressures adequate to tip the balance among countervailing forces in favor of group selection. Eusociality in ants and termites in the irreversible stage is the key to their ecological dominance and has (at least in ants) shaped some features of internal phylogeny. Their colonies are consistently superior to solitary and preeusocial competitors, due to the altruistic behavior among nestmates and their ability to organize coordinated ac- tion by pheromonal communication. n eusociality, an evolutionarily ad- of groups will spread if the positive in- theory are difficult to relate to the com- vanced level of colonial existence, tergroup component of the altruists’ fit- plexities of tangible social phenomena. adult colonial members belong to nesses exceeds the negative within-group We suggest that each force can be two or more overlapping genera- component of the altruists’ fitnesses heuristically classified as binding or dis- I solutive in its effect on colony cohesion tions, care cooperatively for the young, (4–6). Altruism is defined as behavior and are divided into reproductive and that benefits others at the cost of the and either strong or weak in its relative nonreproductive (or at least less-repro- lifetime production of offspring by the power. The degree of relatedness, the ductive) castes. The phenomenon is well altruist. The forces that determine this similarity across the whole genome of marked and nearly confined to insects, outcome are group selection, the differ- individuals as a result of recent common especially ants, bees, wasps, and ter- ential survival and reproduction of en- ancestry, is a factor that biases the di- mites, where it has been subject to a tire cooperative groups as a result of the rection and strength of the forces. When large body of mostly specialized research frequency of the eusociality alleles in elevated, say by lower individual dis- scattered across disciplines from genet- each; individual direct selection, accru- persal rates, relatedness can bring alleles ics to paleontology. It has long been the ing from the differential personal sur- for presociality and eusociality together conviction of researchers on social in- vival and reproduction of each of the more quickly. If it brings such genes to- sects that common principles exist at the colony members; and kin (indirect) se- gether more frequently due to shared organismic and superorganismic levels, lection, which we define here in the microhabitat preference, mate choice, or thus between individual insects and the original narrow sense as the favoring or other biological traits, it can have the tightly integrated colonies they compose disfavoring by individuals of collateral same effect. Relatedness can also in- (1, 2). Parallels have been persuasively relatives, i.e., relatives other than per- crease variance in presocial and eusocial drawn between the self-construction of sonal offspring. The inclusive fitness of alleles among groups, thus quickening organisms from molecules and tissues the prescribing genotype, of individual the pace of colony selection. But relat- and that of superorganisms from inter- colony members and hence statistically edness is relevant only insofar as it acting entire organisms (3). The princi- the colonies they compose, is the nonad- affects the frequency of alleles that pre- ples can be further parsed into two ditive product of the three forces. The scribe social behavior. Eusociality arises segments of the time scale: the develop- definition of kin selection in the com- by the superiority of organized groups mental decision rules that assemble or- monly used broad sense, which folds in over solitaires and cooperative preeuso- ganisms and colonies in each generation individual direct selection as well, pro- cial groups. It can, in theory at least, be and the origin of these rules through duces results that are consistent with initiated by group selection in either evolutionary time. those from the narrow sense but intu- the presence or absence of close related- Focusing here on the second princi- itively less clear and less practicable in ness and, when close relatedness exists, ple, evolutionary process, we suggest application.
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