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Thinking of Biology

Asteroid impacts, microbes, and the cooling of the atmosphere


arth's surface temperature constrained microbial evolution, according to Schwartzman et al. (1993). Their hypothesis states that the maximal temperature that extant organisms of a given type tolerate is the surface temperature occurring when that type of organism arose. Schwartzman and his colleagues concluded that the temperature changed from 100C to 50C between 3.75 billion years ago (BYA) and 1 BYA. These temperatures are consistent with those derived from oxygen isotope ratios in ancient sediments (Karhu and Epstein 1986, Knauth and Lowe 1978). The 100C surface temperature they derive for 3.75 BYA is also the same as Earth's surface temperature 4.4 BYA (Kosting and Ackerman 1986). In this article, we address the cause of the delay in surface cooling until 3.75 BYA, and we explore the implications for microbial evolution of a high temperature on early Earth. We propose that three effects of the early heavy bombardment of Earth by asteroids and comets, until 3.8 BYA, could have delayed onset of surface cooling.

In the beginning . . .
Soon after Earth accreted 4.4 BYA, there was a global ocean, and Earth's atmosphere had as much as 20 atmospheres of carbon dioxide, which caused a high level of greenhouse heating. Temperatures could not have declined until the carbon dioxide was removed through weathering of continental rocks, which libcrated metal ions that combined with carbonic acid to form carbonate rocks.

by Verne R. Oberbeck and Rocco L. MancinelH


March 1994

the maximal Precambrian temperature must have been on the order of IQ^C (Kasting and Toon 1989). However, because an as-yet-unknown fraction of Earth's ancient tillites and diamictites (clastic deposits resembling glacial tills) are deposits of impact craters rather than deposits of glaciers (Marshall and Oberbeck 1992, Oberbeck and Aggarwal 1992a, Oberbeck and Marshall 1992, Oberbeck et al. 1993, Rampino 1992, Sears and Alt 1992), the existence of early glaciations and a 2O''C upper temperature limit for the Precambrian climate has been questioned (Schwartzman etal. 1993). Perry et al. (1978) proposed that the variation in oxygen isotope ratios in sediments was due to the temporal change in "*O in seawater, or alteration of sediments during burial in a closed system and not climatic variation. However, Knauth and Lowe (1978) published a study of Precambrian cherts, including those formed at the sediment/water interface, that appears to exclude the possibility of diagenesis. The temperature history of cherts between 3.5 BYA and the present is quite similar to that which has now been inferred from the evolution of the biota by Schwartzman et al. The case for a high (1993). Finally, Holmden and temperature on early Earth Muehlenbach (1993) concluded, Earth's surface temperatures, de- from the study of oxygen isotope rived from the biota (Schwartzman profiles of two-billion-year-old et al. 1993), are actually compatible ophiolite mineral deposits, that the with temperature estimates that had low oxygen isotope ratios of Protbeen derived previously from oxy- erozoic rocks and cherts do not regen isotope ratios oi sediments flect temporal depletions of '"O in (Knauth and Epstein 1976). Never- seawater. Thus, we adopt the view theless, Perry et al. (1978) rejected that oxygen isotopic data suggest a the high-temperature interpretation high-temperature Precambrian cliof isotopic data for early Earth be- mate. cause they interpreted the PrecamAdditional arguments have been brian tillites to be glacial deposits. given against early glaciation and Theoretical climatic models indi- thus for a high-temperature early cated that, if glaciation occurred, Earth. Schermerhorn (1974) chal173

A considerable period of time may have been required to form the first continents. Planetesimals, asteroids, and comets that hit Earth's surface during the first 700 million years of its history could have created the first land masses. However, impact basins formed during the early heavy bombardment would also have volatilized any carbonate rocks that had formed as a result of weathering of continents; this volatilization would have released carbon dioxide back to the atmosphere and would have prolonged the initial high levels of greenhouse heating even in the presence of continents. Finally, until 3.8 BYA, large impact events could have prevented the habitation of land masses by microbes that would, according to Schwartzman and Volk (1989) and Schwartzman et al. (1993), have been able to accelerate chemical weathering of continental rocks, removal of carbon dioxide from the atmosphere, and cooling. We propose that the sequence of microhial evolution consistent with the early heavy bombardment and a high temperature of early Earth is: heterotrophs, chemoautotrophs, and finally photoautotrophs.

lenged the prevailing view that Precambrian tillites atid diamictites, especially those that occur along ancient continental breakup margins, are of glacial origin. He identified the textural criteria that had, until 1974, heen used to identify tillites and diamictites as glacial deposits, and he argued that the deposits could have been produced by tectonism. Additionally, Salop (1983) pointed out that sediments bounding many ofthe ancient tillites and diamictites are those formed in warm-water environments. He postulated that the abrupt low-temperature excursions at the tilUte and diamictite horizons can only be explained by glaciations if glaciations occurred suddenly in the midst of a hot climate. Knauth and Lowe (1978) also suggested that glacial periods do not preclude hot interglacial periods. The ideas of Schermerhorn (1974), Salop (1983), and Oberbeck et al. (Oberbeck and Aggarwal 1992a,b, Oberbeck and Marshall 1992, 1993) strengthen the longstanding, and previously largely ignored, isotopic evidence for a high temperature of early Earth (Knauth and Lowe 1978). To this evidence we can now add the inferences of Schwartzmanetal. (1993). It is noteworthy that the temperature curves derived by them are quite similar to those for the same time periods derived from isotope data by Knauth and Lowe (1978). Schwa rtzman et al. (1993) pointed out that their temperature history (Figure 1; between 3.75 and 2.5 BYA) was consistent with isotopic data but higher than other estimates based on equilibrium conditions for mineral precipitation. For example. Walker (1982) cited primary evaporite gypsum precipitation in 3.5 billion-year-old sediments to determine the upper temperature limit of 58C for surface-water temperature rather than 73C (Figure 1), which would be the upper limit if the sediments were precipitated in freshwater. The limit would he even lower if gypsum was precipitated in seawater. However, Schwartzman et al. (1993) argued that metastable precipitation of gypsum is likely at temperatures far above its stability field and would
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1993) with a temperature of 100C at 4.4 BYA. The temperature history suggests that from approximately 4.4 BYA to 3.75 BYA the surface temperature was constant at approximately 100*C. This prolonged constant surface temperature has important implications for cooling of atmospheres of bombarded planets and for microbial evolution. We propose that the cause of the delay in the onset of cooling from Time (billions yrs ago) the probable 100C surface temperaFigure 1. Surface temperature history ture at 4.4 BYA until 3.75 BYA was of Earth. Temperatures after 3.75 B Y A due to three effects ofthe early heavy are from Schwartzman et al. (1993), bombardment of Earth that lasted and those hefore 3.7,S B Y A are inter- until approximately 3.8 BYA. First, preted in the text. some period of time was required to form the continents so that contibe consistent with a surface tem- nental rocks could have been available for chemical weathering and perature of 73C at 3.5 BYA. Because the isotopic data of associated cooling from removal of Knauth and Lowe (1978) give the carbon dioxide from the atmosphere. same temperature change with geo- Large-impact cratering events, oclogic history as those of Schwartz- curring during the early heavy bomman et al, (1993), because it is be- bardment, would have required some coming increasingly difficult to time to produce the first topographic explain the oxygen isotope data by dichotomy of continents and ocean temporal changes in isotopic com- basins (Frey 1980) so that weatherposition of seawater or diagenesis, ing and cooling could occur. Secand because Knauth and Lowe ond, the heavy bombardment also (1978) interpret the changes in iso- would have caused release of cartopic ratios to reflect climatic varia- bon dioxide back to the atmosphere tions, we adopt the surface tempera- by impact volatilization of carbonture change shown in Figure 1. A ate rocks and thus sustained the key feature of this curve, for this high level of initial greenhouse heatstudy, is the lOCC temperature at ing until 3.8 BYA. Finally, the heavy 3.75 BYA that declines to 73C by bombardment could also have peri3.5 BYA and defines the onset of cool- odically sterilized Earth and delayed ing of the near-surface environment. continuous habitation of microbial organisms on land masses and prevented acceleration of chemical weathering of silicate rocks and asThe onset of cooling of the sociated cooling until the end of the atmosphere bombardment. We now consider the cause of the Consider the formation of the delay in cooling of the atmosphere until 3.75 BYA and the implications oceans and the time needed to form of this delay for microbial evolu- the silicate rocks of the continents. tion. Earth's atmosphere during its Chyba (1987) calculated that, if first few hundred million years, iust comets contained 10% water, then after condensation of the oceans, during the early bombardment commay have contained up to 20 bars of ets supplied enough water to Earth carbon dioxide. Kasting and Acker- to account for the current ocean man (1986) modeled the dependence volume. Frey (1980) argued that, as of surface temperature of Earth's early as 4.4 BYA, Earth was differearly atmosphere on such an atmo- entiated into a sialic igneous outer spheric carbon dioxide partial pres- crust with a higher-density mantle sure, and they concluded that the and a glohal ocean. In his view, temperature was in the range of large asteroid impacts had, by 4.0 85-110C. !n Figure 1, we combine BYA, produced the topographic dithe inferred surface temperatures chotomy of continents and oceans. after 3.75 BYA (Schwartzman et al. Isotopic data also suggest that conBioScience Vol. 44 No. 3

tinents first existed at some time between 4.3 and 4.0 BYA (Bowring etal. 1989). The initial global ocean, a product of comet impacts, would have prohibited removal of atmospheric carbon dioxide because rocks were not exposed to weathering. However, if land masses were in place after 4.0 BYA, cooling of the atmosphere by removal of carbon dioxide through weathering of surface silicate rocks could, in the absence of other adverse factors, have been started at this time. One such adverse factor is impact volitalization of carbonate rocks, which would have recycled carbon dioxide to the atmosphere and helped to sustain the high initial surface temperatures on Earth through greenhouse heating until the end of the heavy bombardment. This mechanism might have kept enough carbon dioxide in the atmosphere of Mars until the end of the heavy bombardment of the inner planets at 3.8 BYA, so that surface temperatures remained above freezing, liquid water existed, and surface runoff channels formed (Carr 1989). Still another impact mechanism could have retarded cooling until 3.8 BYA. Microbial enhancement of continental rock weathering would have facilitated surface cooling (Schwartzman and Volk 1989} if microbes were present on continents at 4.0 BYA under the suggested habitable temperature conditions (Figure 1). However, another adverse effect of impacts may have delayed the weathering of continental rocks in place by 4.0 BYA. Recent developments in the field of impact catastrophism indicates that microbial life may not have existed continuously or for significant periods of time on continental land masses until 3.8 BYA. Maher and Stevenson (1988) advanced the notion that impacts of large planetesimals, asteroids, and comets during the heavy period of bombardment of Earth, until 3.8 BYA, could have periodically sterilized Harth, frustrated the origin of life, or killed any existing organisms. Sleep etal. (1989) calculated that the last impacts that completely vaporized the ocean could have occurred as early as 4.4 BYA and as late as 3.8 BYA. Oberbeck and March 1994

Fogleman (1989b, 1990) found that the last ocean-vaporizing impact could have occurred at 3.8 BYA. Therefore, life may not have been continuously present, even in the deep oceans, until after 3.8 BYA. If so, the lack of continuous life on Earth's land surfaces until after 3.8 BYA, due to the heavy bombardment, may be another cause for the delay in onset of cooling of the nearsurface environment. Heterotrophic and chemoautotrophic organisms could have existed continuously in the deep oceans after 3.8 BYA, because there would then have been no ocean-vaporizing impacts that killed deep sea life but only ones that sterilized the surface. Therefore, microbes could have continuously evolved and colonized the land surfaces after 3.8 BYA from these continuously available stocks. The sustained decline in Earth's surface temperature may have been delayed until 3.75 BYA (Figure 1) if ocean-vaporizing impacts occurred until the end of the bombardment, if intermittent but extensive land ecosystems were created only after 3.8 BYA, and if these microbial ecosystems were effective in accelerating removal of carbon dioxide from the atmosphere. All of the impact mechanisms for delay in cooling seem consistent with the onset of cooling at 3.75 BYA as inferred here from the isotopic data (Knauth and Lowe 1978), inf^erences from biology (Schwartzman et al. 1993), and the probable temperature 4.4 BYA (Kasting and Ackerman 1986). Impacts initially produced a global ocean that prohibited rock weathering and cooling; they could also have been responsible for a variety of other processes that kept the 10-20 bars of carbon dioxide present in the atmosphere until 3.8 BYA. Then, 50 million years later, cooling began when continents existed, no ocean-vaporizing impacts occurred, continental microbial ecosystems were possible, and there was minimal impact recycling of carbon dioxide.

then cooled rapidly to 73''C by 3.5 BYA, and if continental microbes accelerated this cooling of the surface (Schwartzman et al. 1993), organisms must have occupied the surface of continental land mass soon after the impact bombardment ended 3.8 BYA. The implication is that the surface organisms evolved rapidly from deep-sea organisms that would only have been present continuously after the last ocean-vaporizing impact no later than 3.8 BYA. No direct evidence exists for the evolution of life before 3.8 BYA, leading to controversy over the sequence of microbial evolution. We now link the impact history of early Earth (Chyha et al. 1990, Maher and Stevenson 1989, Oberbeck and Fogleman 1989a,b, 1990, Sleep et al. 1989), 16S rRNA phylogenetic trees (Woese and Pace 1993), climatology (Kasting and Ackerman 1986), and the hypothesis that the upper temperature limit for growth of microbial groups corresponds to the actual surface temperature of Earth at the time of the groups' first appearance (Schwartzman et al. 1993) to show that each of these methodologies independently suggests the same sequence of microbial evolution on Earth from 4.4 to 3.5 BYA. We propose that this sequence of evolution is: anaerobic heterotrophs, followed by chemoautotrophs, and finally photoautotrophs. The early bombardment of Earth before 3.8 BYA prevented life from gaining a foothold at Earth's surface and constrained it to the dark, deep ocean (Maher and Stevenson 1988, Oberbeck and Fogleman 1989a,b, 1990, Sleep et al. 1989). On the other hand, the supply of organic mass from impacting comets (Clark 1988, Oro 1961) was also greatest during this period (Chyba etal. 1990, Oberbeck and Aggarwal 1992b, Oberbeck et al. 1989). We propose that comets were the source of organic nutrients that sustained a small biomass of organisms, limited by the distribution of organics, in the hot, deep ocean just after the heavy bombardment. Thus, we suggest that the earliest microbial forms were thermophilic anaerobic heterotrophs. After 3.8 BYA, the rate of impacts 175

Implications for microbial evolution


If the earth's temperature was constant at 100C until 3.75 BYA and

decreased and the surface could be inhabited. This development had two effects: it decreased influx of organic matter, and it allowed the surface of Earth to become continuously inhabited. The limited supply of organic matter in the presence of abundant carbon dioxide at the surface then gave a selective advantage to chemoautotrophs over heterotrophs. According to the temperature at which these types of organisms now live (up to 110C; Kristjansson and Stetter 1992), this change in advantage may have occurred at approximately 3.75 BYA (Figure 1). Removal of carbon dioxide from the atmosphere through chemoautotrophy then allowed Farth to cool more rapidly than by abiotic processes alone. We suggest that chemoautotrophy evolved earlier than photoautotrophy because chemoautotrophs can live in deeper, dark regions of the ocean, which would have been habitable before surface environments, and chemoautotrophs can tolerate higher temperatures than photoautotrophs. Thus, impacts together with chemoautotrophy may have determined the time of onset of cooling of Earth's atmosphere at 3.75 BYA. Then photoautotrophy evolved. Surface-sterilizing impacts capable of vaporizing the photic zone of the oceans occurred after the last ocean-vaporizing impact, perhaps as late as 3.5 BYA (Oberbeck and Fogleman 1989a,b, 1990, Sleep et al. 1989). Thus, photoautotrophs could have been intermittently present after 3.8 BYA and continuously present at Earth's surface after 5.5 BYA. This presence was also implied by the microbial evolution scheme put forward by Schwartzman et al. (1993), in which the temperature tolerance of existing organisms reflects Earth's surface temperature when that type of organism first evolved. The 73''C maximal tolerance observed in existing photoautotrophs (Meeks and Castenholz 1971) is thus the temperature of Earth's surface at 3.5 BYA, just after the last surface-sterilizing impact. It would not be surprising if modern photoautotrophs would have the same upper limit of tolerance to temperature as thecyanobacterialike organisms of 3.5 BYA represented

by the fossil record (Schwartzman et al. 1993). We propose that the sequence of microbial evolution, which is consistent with impact constraints and the hypothesis that temperature constrained microbial evolution (Schwartzman et al. 1993), is as follows: anaerobic heterotrophs followed by chemoautotrophs (extant examples include Pyrococcus: maximal temperature, 1O5''C; and Pyrodictium: maximal temperature, 11OC; Kristjansson and Stetter 1992), with the photoautotrophs arising later (e.g., Synechococcus lividus: maximal temperature, 73C; Meeks and Castenholz 1971). The branching order depicted by 16s rRNA phylogenetic trees also suggests that photoautotrophy arose after heterotrophy and chemoautotrophy (Woese and Pace 1993). We suggest that heterotrophy arose no later than 3.8 BYA, chemoautotrophy arose approximately 3.75 BYA, and photoautotrophy arose approximately 3.5BYA. The climate history and path of evolution of the biosphere of any habitable planet may be fixed, in part, by its unique early-impact history. The decline in surface temperature may have been determined by the impact history of Earth and by microbial evolution. Consider the possibilities if the impact history had been different. If the heavy bombardment of Earth had been more prolonged, the plateau of constant temperature (Figure 1) may have extended in time. In that event, the surface temperature would not have reached habitable levels for many types of organisms until much later. If so, the evolution of chemoautotrophy and photoautotrophy from the heterotrophs may have proceeded later, which could have delayed the evolution of more complex life forms. The implied synergism between the astrophysical environment and the biosphere suggests that global change in climate and the biosphere has its roots in astrophysics.

release of COj. Icarus 79: 311. Chyba, C. F. 1987. The cometary contribution to the oceans of primitive Earth. Nature 330: 632-635. Chyba, C. F., P. J. Thomas, I.. Brookshaw, and C. Sagan. 1990. Cometary delivery of organics lo the early Earth. Science 249; 366-373. Clark, B.C. 1988. Primeval procreative comet
pond. Origins Life Evot. Biosphere IS:

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Bowring, S. A. 1989. 3.966a gneisses from the Stave province. Northwest Territories, Canada. Geology 17; 971-975. Carr, M. H. 1989. Recharge of the early atmosphere of Mars by impact-induced

209-238. Frey, H. 1980, Crustal evolution of the early earth: the role of maior impacts.Precaw/j. Res. 10: 195-216, Holmden, C , and K. Muehtenbach. 1993. The "*O/"'O ratio of 2-billion-year seawater inferred from ancient oceanic crust. Science 259: 1733-1736. Karhu, J., and S. Epstein. 1986. The implication of the oxygen isotope records in coexisting cherts and phosphates. Geochim. Cosmochim. Acta 50: 1745-1756. Kasting, j . F., and T. P. Ackerman. 1986. Climatic consequences of very high carbon dioxide levels in the Earth's early atmosphere. Science 234: 1383-1385. Kasting, J. F., and O. B. Toon. 1989. Climare evolution on the terrestrial planets. Pages 423-449 in S. K. Atreya, J. B. Pollack, and M. S, Matthews, eds. Origin and Evolution of Planet and Satellite Atmospheres. University of Arizona Press, Tucson. Knauth, L. P. and S. Epstein. 1976. Hydrogen and oxygen isotope ratios ui nodular and bedded cherts, Ceoffcim. Cosmochim. Acta 40: 1095-1108. Knauth, P. L. and D. R. Lowe, 1978. Oxygen isotope geochemistry of cherts from the onverwitcht group (3.4 billion ye.irs), Transvaal, South Africa, with implications for secular variations in the isoropit composition of cherts. Earth and Planetary Science Letters 41: 209-222. Kristjansson, J. K., and K. O. Stetter. 1992. Thermophilic Bacteria. CRC Press, Boca Raton, FL. .Maher, K. A., and D. J. Stevenson. 1988. Impact frustration of the origin of life. Nature 331: 612-614. .Marshall, J. R., and V. R. Oberbeck. 1992. Textures of impact deposits and the origin of rillites. American Geophysical Union EOS Suppi. 73(43): 324 (abstract P31B-4). Meeks, J. C . and R. W. Castenholz. 1971. Growth and photosynthesis in an extreme thermopbile, Synechococcus lividus (Cyanophyta). Arch. Mikrobiol. 78: 25-41. Oberbeck, V. R., and H. Aggarwal. 1992a. Impact crater deposit production on Earth. Pages 1011-1012 in Lunar and Planetary Science Conference Abstracts. Lunar and Planetary Science Institute, Houston, TX. . 1992b. Comet impacts and chemical evolution on the bombarded eartb. Origins Life Evol. Biosphere 21: 317-338. Oberbeck, V. R., and C. Fogleman. 1989a. Estimates of the maximum time required to originate life. Origins Life Evol. Biosphere 19: 549-560. . 1989b. Impacts and tbe origin of life. Nature 339: 434. . 1990. Impact constraints on the en-

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continuity of life. Origins Life Evol. Biosphere 20: 181-195.

Oberbeck, V. R., and J. R. Marshall. 1992. Impacts, flood basalts, and continental breakup. Pages 10K?-t0l4 in Lunar and Planetary Science Conference Abstracts. Lunar and Planetary Science Institute, Houston, TX. Oberbeck, V. R., J. R. Marshall, and H. R. Aggarwal. 1993. Impacts, tillites, and the breakup of Gondwanaland. Journal of Geology 101: 1-19. Oberbeck, V. R., C. P. Mckay, T. W. Scattergood, G. C. Carle, and J. R. Valentin. 1989. The role of cometary particle coalescence in chemical evolution. Origins Life Evol. Biosphere 19: S49-560. Oro, J. 1961. Comets and the formation of biochemical compounds on the primitive earth. Nature 190: 389-390. Perry, E. C. Jr., S. N. Ahmad, and T. M. Swulius. 1978. The oxygen isotope composition of 3,800 m.y. old metamorphosed chert and iron formation from Isukasia West Greenland, journal of Geology 86: 223-239. Rampino, M. 1992. Ancient "glacial" deposits are ejects of latge impacts: the ice age paradox explained. American Geophysical Union EOS Suppl. 73(43): 99 (abstract A32C-1). Salop, I. J. 1983. Geologic Evolution of the Earth During the Precambrian. SpringerVerlag, New York. Schermerhorn, L. J. G. 1974. Later Precambrian mixtites: glacial or nonglacial?/\"J. ;. Sci. 274: 673-824. Schwartzman. D., M. McMenamin, and T. Volk. 1993. Did surface temperatures constrain microbiai evolution?6/<)Sc(V(:r 43: 390-393. Schwartzman D. W., and D. Volk. 1989. Riotic enhancement of weathering and tbe babitability of Earth. Nature 340; 457-460. Sears, J. W., and D. Alt. 1992. Impact origin of large intercratonic basins in the stationary Proterozoic crust and the transition to modern plate tectonics. Pages 385-392 in M. J. Barthlomew, D. W. I lyndman. D. W. Moqk. and R. Masson, eds. Basement Tectonics: Characterization and Comparison of Ancient and Mesozoic Continental Margins. Kluwer Academic, Dordrecht, The Netherlands. Sleep, N. H.. K. J. Zahnle, J. F. Kasting, and H. J. Morowitz. 1989. Annihilation of ecosystems by large asteroid impacts in the early earth. Nature 342: 139-142. Walker, J. C. G. 1982. Climatic factors on Archean Earth. Pataeogeogr. Palaeoclimatol. Palaeoecoi 4: 1-1 I. Woese, C. R., and N. R. Pace. 1993. The RNA World. Cold Spring Harbor Laboratory Press, Cold Spring Harbor, NY.

Call for Nominees for the 1995 AIBS Distinguished Service Award
Since 1972, the AIBS Distinguished Service Award has been presented to individuals who have contributed significantly in the service of hiology. The principal criteria for this award are that the recipients shall have made an outstanding contribution toward: advancing and integrating the biological disciplines, applying biological knowledge to the solution of world problems, introducing pertinent biological considerations that improve public policy and planning. Emphasis is placed on distinguished service. Scientific discovery per se is not included as a criterion for this award., although some nominees carry this distinction as well. Previous recipients of the award have been:
1972- -Harve Carlson, George Miller, Detlcv Bronk 1973- -Theodosius Dobzhansky, Rene Dubos 1974- -James G. Horsfall 1975- -W. Frank Blair, Theodtjre Cooper 1976- -Paul B. Sears, Edward O. Wilson 1977- -Paul J. Kramer, Elvin C. Stakman, William C. Steere 1978- -Eugene P. Odum, Howard T. Odum, George Gaylord

Simpson
1979- -Theodore C. Byerly, H. C. Chiang, Lee M. Talhot 1980- -Arthur D. Hasler, A. Starker Leopold, Ruth Patrick 1981- -Peter H. Raven 1988Donald E. Stone 1982- -George M. Woodwel! 1989Alfred E. Harper 1983- -Karl Maramorosch 1990Gene E. Likens 1984- -Arnold B. Grobman 1991John S. Niedcrhauser 1985- -Sayed Z. El-Sayed 1992Ruth Huhhard 1986- -Garrett Hardin 1993John A. Moore 1987- -Perry L. Adkisson

Verne R. Oberbeck is a research scientist at the NASA Ames Research Center, Moffett Field, CA 94035. Rocco L. Mancinelli is a research scientist at the SETl Institute, Mountain View, CA 94043. /994 American Institute of Biological Sciences.

AIBS members are invited to submit nominations for this award, which will he presented at the 1995 AIBS Annual Meeting. Each nomination must be accompanied hy a complete curriculum vitac and a statement of the individual's service to the hiology profession. In particular, the supporting statement should highlight the nominee's accomplishments in each of the three award criteria given ahove. Nominators should note that traditional academic vitae often omit contributions to puhlic affairs. Because this area is considered equally important in the overall consideration, care should he taken to bring out the nominee's relevant accomplishments. Since 1981, recipents have been limited to single individuals, but nominations remain active for three consecutive years (e. g., for the 1995, 1996, and 1997 awards). Send nominations (with biographies) to the AIBS Executive Director, 730 11th Street, NW, Washington, DC 20001-4521, by 1 October 1994.

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