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HEINZ ADOLF LOWENSTAM October 9, I 9~2-fune 7, I 993 BY JOSEPH L. KIRSCHVINK HEINZ EOWENSTAM DESCRIBED himself as a professional beach comber, but in fact he was among the twentieth century's most superb natural scientists. Builcling upon an early interest in minerals en cl fossils gainecl cluring his chilcI- hood playing on mining dumps in Germany, he was the first to blend biological and paleontological analyses to un- ravel the ecological associations of fossil communities. Af- ter being cleniec! his Ph.D. from Nazi-controllec! universi- ties for the crime of being Jewish, he flecl with his wife to the Unitecl States en cl managed to complete his degree at the University of Chicago just prior to the start of World War II. CIassifiecl as an enemy alien, he contributed to the U.S. war effort by cleveloping paleoecological techniques to locate oil-bearing coral reefs in the sub-surface of the greater Chicago area. Rather than profiting personally from his work, Heinz publishecl it freely in the open literature. In the postwar isotope frenzy at the University of Chi- cago Heinz was drafted as the "atomic paleontologist" for Harold Urey's research group. His initial role was to pro- vicle pristine fossil materials for isotopic paTeotemperature determinations, but his involvement grew rapicIly to inclucle that of identifying the most important scientific questions 95

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96 B I O G RA P H I C A L EMOIRS about Earth's past biosphere that conic! be aciciressec! for the growing fielcl of stable isotope geochemistry. In the process of fincling unalterecl fossil materials Heinz also be- gan to woncler about the process of biomineraTization it- self: How clo animals make minerals uncler biological con- trol? How clo they control the mineral composition, crystallinity, en c! particle size? Using his talents as a natural- ist en cl exploiting advances in analytical techniques, Heinz nearly cloublecl the known diversity of minerals proclucecl by organisms. One of his discoveries the biomineraTization of magnetite (Fe3O4) in the teeth of Polyplacophoran mol- lusks (the chitons) has been crucial for unclerstancling topics as diverse as the geophysics of marine sediment magnetiza- tion en cl the biophysical basis of magnetoreception in ani- mals. Heinz was born in 1912 in Upper Silesia, in what was then southeastern Germany but is now south-central Po- lancI, in the town of Siemjanowicz. This was a suburb of Laurahutte, a mining district with a steel mill. In his oral history recorclecl for the Caltech archives Heinz clescribecl his birthplace as a horrible region.... It was like Dante's Inferno. Across the whole hori- zon, you saw belching chimneys spewing out fumes from lead smelters, and steel mills. There were coal mines and iron foundries. The air was so poor that our plants in the house had to be specially tended so they didn't die from the fumes.... As a kid, I played on a mine dump you know, the stuff that goes out from a lead and zinc mine. I wasn't supposed to go there, but I went with some miners' kids, and we played. We normally picked up a rock to throw, and one day the one that I picked up was awfully heavy. I knew it couldn't be an ordinary rock, so I broke it. It looked like silver. It was galena lead ore. And that's what started me initially to collect minerals. Many of Heinz's interests in nature clearly stem from his parents (Frieda and Kurt Lowenstam), although neither

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HEINZ ADOLF LOWENSTAM 97 of them hac! a university education. Before Woric! War I his mother was the art editor of a newspaper en cl wrote poetry. She encouraged Heinz's interest in nature by taking him around and showing him things, and getting semi-popular publications on natural history. Heinz notes that she "was interested, among other things, in ancient Egypt, en cl she taught herself to react hieroglyphics. We wouic! go to muse- ums, like in Berlin, en cl she wouIcl react the inscriptions just like that en cl translate them." His father was similarly eclu- catecI, in the sense that he was a classicist. "He went to the Gymnasium the German academic high school. He always hacl pockets full of books. He was more interested in his- tory en c! literature." His grandfather wrote a six-volume his- tory of the Jewish people, en cl most members of his family stucliecl languages en cl literature. Later in life Heinz wouIcl often comment on this peculiar background! with statements like, "Ha, ha, ha, . . . you know I'm the black sheep of the family, I clon't shpeak any lankvages." Upper Silesia was also politically unstable, caught up in the ravages of the First WorIcl War. At one birthday party he remembered machine-gun fire strafing his granciparent's home while everyone hugged the floor in panic. This was the first of many "I've almost been killecI" episodes that were to punctuate Heinz's life. The Lowenstam family was hit hare! by the German economic depression in the 1920s en cl the hyperinflation that followocI. Due to his interest in the natural sciences en cl with the encouragement en cl sup- port of his maternal grandfather, Heinz enterer! what was then an experimental Hochschule focused on math, phys- ics, en cl chemistry as major subjects (in contrast to the Gym- nasium, which proviclec! a more academic education focuses! on the cIassics). Heinz founcl his new school area more conducive to collecting fossils than minerals, so he swapped collections with one of his teachers who wan tee! to become

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98 B I O G RA P H I C A L EMOIRS a mineralogist, that's when he starter! his first systematic fossil collection en cl gainecl the desire to be a paleontolo- gist. A seminar in his town given by Alfrecl Wegener, who first proposer! the continental cirift theory, expanclec! his interest to include geology, not just paleontology. With continual support from his grandfather, Heinz was able to enroll in the vertebrate paleontology program at the University of Frankfurt. However, its leacling paleon- tologist cliecl suciclenly just prior to his arrival, en cl the en- tire program collapsecI. The students scatterer! to other uni- versities, en cl in the fall of 1933 Heinz chose to continue at the University of Munich, which hacl the strongest German program in paleontology with the most international out- look. Shortly thereafter Aclolf Hitler was namecl chancellor en cl the situation for German Jews became increasingly more precarious. Unfortunately, some of the professors at Munich were influenced strongly by Nazi propaganda. Others, such as Heinz's mentors Prof. Broili, Ecigar Dacque, en cl the bi- ologist Karl von Frisch, were willing en c! able to ignore the rhetoric to some extent. Von Frisch even went so far as to give Heinz a desk to work at in his laboratory suite immedi- ately after the first anti-lewish eclicts were announced. About this time Heinz met his future first wife, Elsa Weil, a stu- clent in the Luc~wig-MaximilIan University Meclical School, at a vegetarian restaurant. This Nazi influence resultecl in a most unusual twist in Heinz's choice for a Ph.D. thesis topic. During a student field! trip early in ~ 935 Heinz recallec! one of the Nazi- influenced professors (Kolbl) pounding the table and say- ing, "German things must be clone by Germans." A few minutes later he hac! the tenacity to ask Heinz what he was planning to do for his Ph.D. dissertation. In a fit of sheer impulsive rebellion Heinz announced that he was going to work on the geology of Palestine, despite the fact that he

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HEINZ ADOLF LOWENSTAM 99 hac! absolutely no personal resources to clo so. Depresses! at having shot his mouth off, he mentioned this to his friend en cl lancIlorcl later that evening, who saicI, "Don't worry. I have friencis in New York. They will take care of it." Unbe- knownst to Heinz, these friends were financial by the Iraq petroleum company, which was very interested in the geol- ogy of the MicicIle East en c! were eager to have goof! geo- logical en cl paleontological studies clone in the area. So Heinz went to Palestine for IS months. Although the area was still a British protectorate, Heinz realizer! that he wouIcl neecl to cooperate with the Bedouins to have full access to his fielcl area. With a proper introduction from the British district commissioner he was able to live with the family of the number one Sheik for several months, learning Bedouin Arabic in the process. During the intro- cluction, however, Heinz was forcer! to smoke for his first time, refusal wouIcl have been a cleacIly insult to the sheik en cl his family. (That lecl unfortunately to a 45-year tobacco acicliction en c! his ultimate clemise from lung cancer.) Dur- ing his IS months in Palestine Heinz was able to complete the first geological en cl paleontological analysis of the east- ern Nazareth Mountains, which turner! out to be one of the critical areas for unclerstancling the geology of the entire Deacl Sea rift system. During this time geologists from the Iraq petroleum company user! his geological en c! paTeonto- logical skills, as Heinz was invited repeatecIly to participate in fielcl excursions throughout the entire MicicIle East. All they asked in exchange for these trips was to have copies of his fielcl notes. At the time Heinz clicl not know that this was also the ultimate source of his fielcl support in Pales- tine. Upon completion of his dissertation research in the micicIle of 1936, Heinz returned to Munich en cl spent about a year finishing his thesis. After it was acceptec! en c! the

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00 B I O G RA P H I C A L EMOIRS ciate for his exam was schecluTec! he en c! his fiancee, Elsa, were married. One week before his thesis defense, how- ever, the Nazi government issuccl an eclict that no more Jews wouic! be allowed! to receive their doctorates at Ger- man universities. Elsa hacl aIreacly received her meclical cle- gree the previous week, but Heinz was out in the coIcI, with nothing to show for his many years of university education, not even a bachelor's degree (the German Ph.D. was an all- or-nothing affair). They hacl no options but to leave. Sev- eral of the geology faculty at Munich then clic! an extraorcli- narily risky thing, as notecl by Heinz. So Dacque wrote a letter on official university stationery, with the Nazi university seal on it, saying I had fulfilled the qualifications of the Ph.D., but due to political circumstances, they couldn't give me the diploma. He went over with me to Broili, who was the head of the paleontology depart- ment to have Broili sign where he had typed out his name. Broili sat down and signed. Within 10 minutes Kolbl [the Nazi professor] asked me to see him. I came in and [he] said, "I would like to see the letter which you just got from Broili and Dacque." I said, "What Letter?" He said, "Don't be silly." He went over to my pockets and he knew in which pocket I had it. He pulled it out, read it; his eyes popped out, he got mad. He gave it back to me and said, "Nice letter, isn't it." He knew I was going to leave within a week or two. I said, "Yes." He said, " I want to give you a letter of recom- mendation, too. But you must not show it to the Chicago Tribune, because you know what would happen to me if you did that." I said, "I don't want your recommendation." He didn't listen. He sat down and wrote the let- ter I was a good student, in general terms and gave it to me. As I went out the door he said to me, "You cannot go to America and say that we mistreated you, can you?" Fortunately Heinz and his wife had managed to get visas to immigrate to the Unitecl States, being sponsored by his wife's uncle in Chicago. The major problem was that the U.S. government consiclerec! Heinz to be a Polish citizen, given that his birthplace, Upper Silesia, was then part of Poland. (Heinz was a German citizen en cl hacl never been

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HEINZ ADOLF LOWENSTAM 101 in Poland, yet they toic! him he was Polish.) The visa queue for Polish citizens seeking to immigrate to the Unitecl States hacl been hopelessly overdrawn for 15 years. However, the U.S. consul in Stuttgart the closest one gave him one of three emergency visas. Heinz later came to suspect that this arrangement was macle possible by the silent efforts of the oil company that hac! finances! his studies in the MicicIle East. Heinz's parents en cl sister also managed to escape to Brazil, but most of his other relatives later perished in the Holocaust. His grandfather, who was not a Jew but hac! married one, chose to commit joint suicide with his wife by fire in their home rather than denounce their chilciren en cl relatives to the Nazis. Upon their penniless arrival in Chicago in June of 1937, Heinz cliscussecl his situation with several of the geology faculty at the university. At first they simply Tookoc! through his gracle sheets with little apparent interest until someone noticecl the letter from Broili en cl Dacque. As Heinz re- callec! in his oral history transcripts, I hadn't thought of it. The letter happened to be in my pile of papers. They saw that letter and said, "Could we open it?" They read it, and I'll never forget their eyes got big. "Broil), Dacque, they recommended?" Af- ter that the whole atmosphere changed dramatically. I immediately got a scholarship, in the middle of the year. I was told I would have to take a few courses, translate my thesis, and within a year or two I could get my degree. That's when I realized how important that letter was. It was a miracle that Broili and Dacque had done it, too, because without it, I don't know. After only a minor setback for failing his German lan- guage test (because he clicin't unclerstanc! the English in- structions), Heinz finally received his Ph.D. from the Uni- versity of Chicago in 1939. In one aspect Heinz was frustrates! with his experience in Chicago. Having flecl from persecution in Germany amid the destruction of his friends en cl family, he wan tell to en-

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02 BIOGRAPHICAL MEMOIRS list in the U.S. Army so that he conic! go en c! fight the Nazis. When America finally joined the war, he was listecl initially as an enemy alien en cl was subjected to severe travel restrictions, even to the extent of having his camera confis- catecI. When the Arctic en cl Desert Division of the Army realizecl that they couIcl use Heinz's ability to speak Bedouin Arabic for their campaign in Egypt, they lifted his enemy status en cl rushed his citizenship papers through in record time, but before he could go, the battle in Egypt was over. At that time the German U-boats were sinking oil tankers going from Texas to the east coast, en cl the military arguccl correctly that he wouIcl be of more use to them as a civilian working on the coal en c! oil reserves for the Illinois Geo- logical Survey. Still, Heinz managed to help the war effort by interpreting aerial photographs of the Rhur district coal mines for the Army Air Corps, and recognized that modifi- cations to the coking ovens were clesignecl to extract high- octane aviation fuel. The Allies bombed the plants, putting them out of business en c! giving Heinz some measure of revenge. For a short while after graduation Heinz workocl for a small oil company en c! then mover! to the Illinois State Museum as a curator of invertebrate paleontology. Having no funds for field research, Heinz discovered that he could take the Chicago streetcar system to the end of the Stony Islancl line to reach an area rich with fossil coral reefs. His rationale for launching his studies on the paleoecology of the coral reef environments (which lee! eventually to the recovery of enormous quantities of oil) is best expressed in his own worcis from the Caltech archives. It was called Stony Island because there were fossil reefs cropping out- coral reefs. I went over there, and it was in terrible shape. Then I discov- ered next to it long dump piles that had been made when drainage canals were built to connect the Illinois River with the Great Lakes to get barges

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HEINZ ADOLF LOWENSTAM 103 through. In digging the canals the work crews had dumped all this stuff on the side. I started to walk over those old dump piles, and I found very nice fossils, all marine, and I knew they were from the Silurian period about 400 million years ago. Some people from the Field Museum had already catalogued some of these fossils the black shale type . . . but nobody had looked at the skeletal remains in dolomite. I made a big collection of the material over a period of time and then tried to identify it. I couldn't. I researched the local literature, and none of the fossil groups that had been described from the Chicago area fit what I had found at all. As Heinz expenclec! his attempts to identify the fossils he eventually cliscoverecl that they matchecl almost precisely Silurian fossils in Tennessee. Previous paleontologists who hac! stucliec! the fossils from museum drawers hac! assumer! that the two populations were from two geographically sepa- rate areas, one in the South en cl the other in the North, but Heinz hac! fount! them in the same area. The northern population was in fact composed of organisms that livecl in the reef environment, in the active wave zone. The south- ern fauna simply liver! in creeper waters en c! was composer! of smaller forms aciaptecl to a darker, less active environ- ment. Upon further stucly Heinz cliscoverecl that he couIcl identify changes in the ecological communities surrouncI- ing the reef environments that variecl systematically with distance from the reef complex, and was even able to deter- mine the direction of the prevailing wincis 400 million years ago from the horseshoe-shape atoll structures. By examin- ing subsurface cores from several localities he was able to use these distance estimates to determine the location of buriecl fossil reefs. Ultimately, Heinz cliscoverecl a massive system of Silurian reefs that stretched from the ecige of the Ozark Mountains to Greenland, it had been larger and more magnificent in Silurian time than the Great Barrier Reef of Australia is tociay. Heinz also realizer! that the porous structure of a bur- iecl reef complex was an icleal trap for oil en cl gas. Several

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04 B I O G RA P H I C A L EMOIRS major companies hac! cliscoverec! oil in the Chicago area almost by random cirilling, en cl Heinz's ability to pinpoint the locations by simply examining the cores seemed nearly miraculous. Two of the companies even went so far as to break into Heinz's office, looking for information on where to cirill, Heinz was able to identify the banclits by marking fake locations on his office map en c! watching which com- pany started cirilling there. Despite financial offers of up to ~ percent of the profits for the proprietary use of his tech- nique (which wouic! have macle him a very wealthy man) en cl to the later dismay of his chilciren, Heinz insteacl chose to publish his finclings in the open scientific literature for the benefit of all. His only compensation was the gift of a binocular dissecting microscope from one of the compa- nies. However, the title of his 1948 book on the topic (Bio- stratigraphic Studies of the Niagaran In ter-Reef Formations in Northeastern Illinois, Illinois State Museum Society) was so obtuse that it triggered a local columnist for the Chicago Tribune to complain in print about the waste of state funcis on such useless studies. This triggered a heated public re- sponse from the presidents of several major oil companies, who notes! that the work was leacling to the recovery of enormous volumes of oil. His monograph was republishecl many times by the oil industry. Immecliately after the war the University of Chicago was a hotbed of isotopic research en cl was in particular the birth- place of isotope geochemistry. HaroIcl Urey hacl recognized the importance of isotopic measurements for interpreting the past history of the Earth en cl hacl assemblecl a team focused on using deviations in stable isotope ratios to mea- sure the temperature of ancient oceans. Urey hac! obtainer! fossils of Mississippian age (about 300 million years old), extracted the calcite from the shells, and had determined a temperature of about 60C, higher than any known animal

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HEINZ ADOLF LOWENSTAM 109 leac! to important discoveries even tociay. The story begins in 1961 when he was sitting at low title on a wave-cut plat- form in Bermuda en cl began to woncler how the erosional processes hac! proclucec! such a level, almost bevelecI-off sub- strate. He hacl seen similar benches in Palau, where the limestone hacl eroclecl into huncirecis of nip islancis, each resembling a large mushroom with waves splashing arounc! the base en cl dense vegetation on top. At the time the dogma was that these were wave-cut benches, perhaps with some help from salt crystal formation at Tow ticle. For some rea- son this clicl not satisfy Heinz, who took out a hand lens en cl examined the limestone substrate more closely. Surprisingly the surface was coverer! with Tong strips of small chevron- shapecl groves that wanclerecl over each other en cl over- lappecl in complex patterns, something like tanglecl noocIles. While he was examining this, a chiton (a mollusk of the class Polyplacophora) wanclerecl by, leaving a fresh noocIle trail like this chiselecl into the rock surface. Heinz realizecl immecliately that the chiton was scraping off the outer (some- what greenish) layer of the rock surface, feecling on enclo- lithic algae growing in small cracks in the limestone. But for this to be the case the animaT's teeth neeclec! to be harcler than the limestone substrate it was feecling on. The biological belief at the time helcl that the teeth of mollusks were macle of a proteinaceous material like fingernails, which would not have been nearly hard enough to use as a rock chisel. A quick dissection revealecl that the teeth along the animaT's tongue plate (the raclula) were black en c! very hard, obviously mineralizecl with something but clearly not cal- cium carbonate or a calcium phosphate mineral like apa- tite (such as in human bones and teeth). The black tooth mineral was present in every incliviclual en cl every chiton species he examined. Tooth shape was even species spe-

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0 B I O G RA P H I C A L EMOIRS cific, some having several prongs en c! others curved, cup- like structures. Determining just what the hard black stuff was provecl to be more clifficult. Back in the early 1960s the best ana- lytical tool for precise mineral determination was X-ray clif- fraction, en cl the stanciarcl technique was to use a narrow beam of Cu-Koc racliation. However, when the chiton teeth were measured in this fashion the photographic emulsion came out completely fogged. The technician operating the instrument, Art Choclos, suggester! that it might be some interference or fluorescence en cl recommenclecl changing the X-ray source from Cu to a different metal like Ni or Co. That eliminates! the interference problem en c! proclucec! a nice set of diffraction lines. Unfortunately, they clicl not match any of the stanciarcl minerals that are commonly found in the reef environments. Stumped, Heinz en c! his assistant cleciclecl to search methoclically through each mineral in the stanciarcl diffraction compilation until they founcl some- thing that matched. After several days of searching, pure magnetite (Fe3O4) popped up suddenly as a perfect match. IntriguccI, Heinz then took a small hand magnet en cl clis- covered to his amazement that the entire radula stuck to it as strongly as if it were a nail: It was obviously ferromag- netic. Subsequent chemical analyses confirmed that iron was the main component in the teeth, it also explainer! the problem with the X rays, as the Cu-Koc line causes iron to fluoresce, fogging the film. It is important to put this discovery into the proper historical perspective. In 1961 magnetite was known to be a dense, inverse-spinel mineral that formecl exclusively in high- temperature, high-pressure igneous or metamorphic envi- ronments. It was thought to be terribly out ot equilibrium at room temperature en cl pressure, en cl was simply not some- thing that conic! be proclucec! in the mouth of a mollusk.

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HEINZ ADOLF LOWENSTAM 111 Mineralogists and petrologists assured Heinz that the chi- tons had to be picking up grains of magnetite from the sancl the same way that sharks en cl rays were known to accu- mulate heavy minerals in their inner ear for their balance organs. But, by simply dissecting out the raclula en cl look- ing at it carefully, Heinz was able to show that the iron was of biological origin. It accumulates! first as the iron pro- tein, ferritin, in epithelial cells that were tightly attached to a proteinaceous but unmineralizecl embryonic tooth. The iron was then transporter! rapidity into the young teeth in the form of the mineral ferrThycirite Hydrous Fe2O3), forming a few rows of bright reel teeth. At a very sharp, suciclen transition most of the tooth volume was converter! into black magnetite, with graclual aciclition of more ferrThycirite (con- verting to magnetite) as the teeth matured. This simple series of observations was able to shut up the most severe critics instantly. Magnetite was being formecl at low tem- peratures en cl pressures, in an animal, no less. Although it is now well known that magnetite can be precipitates! from aqueous solution under strongly reducing conditions, it was not appreciated in 1961. Of aciclitional importance was the fact that the raclular teeth stuck strongly to a magnet. That was the first clear, macroscopic, en cl easily reproclucible effect of a magnetic field on a biological structure, and in one sense earns Heinz the title of father of biomagnetism. (This was actually a much simpler biomagnetic effect than Linus Pauling's 1933 discovery that cleoxyhemagiobin is paramagnetic.) In his seminal 1962 paper reporting this discovery Heinz notecl that chitons were known to have a local homing instinct, with inclivicluals returning to their own preferrer! clepres- sions in the rock cluring low ticle. Interestingly enough he clicl not suggest explicitly in that paper that they might be using a magnetite compass as a navigational aicI, but it is

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112 B I O G RA P H I C A L EMOIRS clearly implied from the context. It is a pity that the paper was publishecl in the Bulletin of the Geological Society of America, because not many biologists react it. Numerous claims of apparent magnetic field! sensitivity in animals hacl been macle prior to 1960. Biophysicists, how- ever, were vociferous in denouncing those studies for the simple reason that they knew of no plausible mechanism through which the weak magnetic fielcl of the Earth couIcl influence the diamagnetic en cl paramagnetic materials present in living organisms, en c! magnetic induction was too weak to be of use with an electrical detection system. Prominent neurobiologists hacl even statecl flatly in print that there were no physiological ferromagnetic materials and hence, magnetoreception was impossible. Heinz's discovery of mag- netite in the chiton teeth obviously unclerminecl the basis of this biophysical argument (anc! paver! the roar! for much of my research). Subsequent discoveries have confirmed the central role of magnetite as the biophysical transducer of the magnetic field! in living organisms spanning the evolu- tionary spectrum from the magnetotactic bacteria to mam- mals, with a fossil record extending back at least 2 billion years on Earth en c! perhaps 4 billion years on Mars. (As of this writing the best evidence for ancient life on Mars is the presence of probable biogenic magnetite in the ALHS4001 meteorite. ~ In the vertebrates, chains of uniform-size mag- netite crystals, optimized for their magnetic properties, have been founcl recently in specializecl cells connected to the ophthalmic branch of the trigeminal nerve, this nerve is now known as the main conduit of magnetic fielcl informa- tion to the brain. This magnetite system is one of the few truly novel sensory mociaTities cliscoverec! in the past 50 years, en cl Lowenstam's discovery in the chiton teeth was the first hint that anything like this might be possible. Rather than pursue the neurophysical aspects of the

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HEINZ ADOLF LOWENSTAM 113 magnetite discovery, Heinz wonclerec! what other weirs! min- erals living organisms might form. Within a few months he cliscoverecl goethite (~-FeOOH) capping the teeth in an- other primitive group of mollusks, the Archaeogastropocis. During the 1960s en cl 1970s the mineral list grew steaclily beyond apatite, carbonates, en cl opal to inclucle lepiclocrocite, vaterite, ferrihycirite, wociclellite, ciahIlite, en c! a variety of amorphous iron en cl phosphate minerals, to name a few. In aciclition Heinz began a systematic compilation of the phyI- etic distribution of these materials, as well as efforts to track the time of their evolutionary origin. In this process he macle another funciamental observation concerning the bio- Togical processes that different organisms user! to form biominerals there was a clear spectrum of biological con- trol. Some organisms actively direct every aspect of the min- eral formation process, inclucling chemical purity, crystal- linity, crystal orientation, en cl crystal shape en cl size. By precipitating the minerals inside the cell they produce min- eral products that are unlike anything proclucec! inorgani- cally. Because of the complex assemblage of biomolecules involvecl in this type of mineralization, Heinz termed this process "matrix mecliatecI," or "biologically controllecI," biomineralization. On the other hand some minerals sim- ply form as an indirect result of biological activity, associ- atec! with metabolic by-proclucts, these he termec! "biologi- cally induced." By standing back and looking both at the temporal distribution of fossil forms en cl their phyletic clis- tribution, Heinz was able to observe new patterns in the data relating to the underlying biochemistry. Of particular importance was his observation that virtually all the min- eral products that appeared! nearly simultaneously in the Early Cambrian (the Precambrian-Cambrian boundary in- terval) in approximately 40 phyletic-level groups involved the use of calcium minerals (phosphates en c! carbonates).

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4 B I O G RA P H I C A L EMOIRS In a seminal paper coauthored! with Lynn MarguTis in 1980 he notecl that all of the requisite biochemical transport systems for this process hacl to have been present in the last common ancestor of all animals, as all eukaryotic cells rely on the precise control of calcium ion concentrations to regulate the mitotic processes (through microtubule poly- merization) en c! for seconcI-messenger systems. Hence, most of the clifficult evolutionary prerequisites neeclecl for the wiclespreacl biomineralization of evolving animal groups were aireacly present Tong before something associates! with the Cambrian Explosion (like a runaway preciator/prey interac- tion) triggered the biomineralization cascade. This concept certainly is the foundation of a "grant! unifies! theory" of biomineralization, which may help to unravel the complex genetics en cl biochemistry of biomeclically important pro- cesses like tooth en c! bone formation. Despite the pain en cl the suffering that Heinz experi- encecl as a youth in Germany, California life en cl profes- sional beachcombing calmed him. For many years after World War II he hacl severe aversions to all things German, in- clucling a sincere inability to speak the language en cl a strict injunction against traveling there. How conic! he? German citizens in his age group bore responsibility for the Holo- caust that destroyed his family, even though his ancestors hac! liver! in Germany for at least the previous 400 years. Those of us who knew Heinz well were therefore stunned when the Faculty in Munich presented Heinz with an hon- orary Ph.D. in 1980, en cl he accepted it. This apparently took several years of careful advance preparation by Lynn Margulis, Dolf Seilacher, en cl Wolfgang Krombine, who graclu- ally managed to persuade Heinz that it would be a good signal to the younger German scientists who bore no re- sponsibility for the errors of their parents. Even so, Heinz rememberer! the experience as troubling, particularly when _~ . .. . .

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HEINZ ADOLF LOWENSTAM 115 he saw elclerly Germans catching the bus in Munich en c! wondering, "What were they cloing cluring the war? Were they responsible?" To Heinz's academic chiTciren he was a quiet intellec- tual giant who spoke with a soft Milwaukee-German accent, which for many years was muffled severely by the use of cigarettes. During class lectures we hac! to sit quietly near the front simply to hear him, but no one ever complainecI, as he was a stimulating en cl fascinating lecturer. In one episode in the early 1970s we counter! no less than five cigarettes lit at the same time scattered along the chalk tray, as Heinz wouIcl become so excited en cl immersed in his subject that he wouic! forget that he aireacly hac! some lit. Even on his fielcl trips particularly those memorable excursions to Baja California shared with Leon T. Silver- Heinz wouic! grab our attention for hours on ens! en c! amaze us with his ability to see subtle relationships between form, function, chemistry, en cl biology of natural en cl ancient eco- systems. In the evenings arounc! the campfire uncler the protection of beautiful groves of California oak trees, he wouIcl tell us encIless stories of the South Pacific, Palau, Japan, South America, en c! his chiTc~hooc! in a fractures! en cl war-torn Europe. We at first thought most of these were fairy tales, until friends en cl family confirmecl them later. Heinz inspirer! all of us to pursue our own intellec- tual interests wherever they wouIcl leacI, with total clisregarcl for personal fame, fortune, or personal safety. We miss him dearly. Heinz is survivecl by three talentecl en cl caring women who shared his life, three chilciren, many talentecl grancI- chiTciren, en c! many more academic offspring, inclucling the present author.

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116 B I O G RA P H I C A L S E L E C T E D EMOIRS B I B L I O G RAP H Y 1942 Geology of the eastern Nazareth Mountains. 7. Geol. 50~7~:813-45. 1946 With E. P. DuBois. Marine pool, Madison County, a new type of oil reservoir in Illinois. Report of Investigations Illinois State Geo- logical Survey 45 (36) :30-55. 1950 Niagaran reefs of the Great Lakes area. 7. Geol. 58~4~:430-87. 1951 With S. Epstein, R. Buchsbaum, and H. C. Urey. Carbonate-water isotopic temperature scale. Geol. Soc. Am. Bull. 62~4~:417-25. With H. C. Urey, S. Epstein, and C. R. McKinney. Measurement of paleotemperatures and temperatures of the upper Cretaceous of England, Denmark, and the southeastern United States. Geol. Soc. Am. Bull. 62 (4) :399-416. 1953 With S. Epstein. Temperature-shell-growth relations of Recent and interglacial Pleistocene shoal-water biota from Bermuda. 7. Geol. 61 (5) :424-38. 1954 Environmental relations of modification compositions of certain carbonate secreting marine invertebrates. Proc. Natl. A cad. Sci. U. S. A. 40~1~:39-48. With S. Epstein. Paleotemperatures of the post-Aptian Cretaceous as determined by the oxygen isotope method. 7. Geol. 62~3~:207- 48. 1957 With S. Epstein. On the origin of sedimentary aragonite needles of the Great Bahama Bank. 7. Geol. 65~4~:364-75.

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HEINZ ADOLF LOWENSTAM 1958 117 With R. N. Ginsburg. The influence of marine bottom communities on the depositional environment of sediments. 7. Geol. 66 ~ 3) :310- 18 it. 1961 Mineralogy, O 18/O 16 ratios, and strontium and magnesium con- tents of recent and fossil brachiopods and their bearing on the history of the oceans. 7. Geol. 69~3~:241-60. 1962 Magnetite in denticle capping in recent chitons (Polyplacophora). Geol. Soc. Am. Bull. 73 (4) :435-38. Goethite in radular teeth of recent marine gastropods. Science 137~3526) :279-80. 1964 Sr/Ca ratio of skeletal aragonites from the recent marine biota at Palau and from fossil gastropods. In Isotopic and Cosmic Chem- istry, eds. H. Craig, S. L. Miller, and J. G. Wasserberg, pp. 114-32. Amsterdam: North-Holland. 1971 Opal precipitation by marine gastropods (Mollusca). Science 171 (3970) :487-90. 1974 Impact ot lile on chemical and physical processes. In The Sea: Ideas and Observations on Progress in the Study of the Seas, ed. E. D. Goldberg, pp. 715-96. New York: J ohn Wiley. 1975 With D. P. Abbott. Vaterite: A mineralization product of the hard tissues of a marine organism (Ascidiacea) . Science 188 (4186) :363- 65. With G. R. Rossman. Amorphous, hydrous, ferric phosphatic der- mal granules in Molpadia (Holothuroidea): Physical and chemi- cal characterization and ecologic implications of the bioinorganic fraction. Chem. Geol. 1 5 ~ 1 ): 1 5-5 1.

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118 B I O G RA P H I C A L 1978 EMOIRS Recovery, behaviour and evolutionary implications of live Monoplacophora. Nature (London) 273~5659~:231-32. 1979 With J. L. Kirschvink. Mineralization and magnetization of chiton teeth. Paleomagnetic, sedimentologic, and biologic implications of organic magnetite. Earth Planet. Sci. Lett. 44~2~:193-204. With S. Weiner, B. Taborek, and L. Hood. Fossil mollusk shell or- ganic matrix components preserved for 80 million years. Paleobiology 5 (2) :144-50. 1980 What, if anything, happened at the transition from the Precam- brian to the Phanerozoic? Precambrian Res. 11~2~:89-91. With L. Margulis. Evolutionary prerequisites for early Phanerozoic calcareous skeletons. BioSystems 12:27-41. 1981 Minerals formed by organisms. Science 211~4487~:1126-31. 1989 With S. Weiner. On Biomineralization. Oxford: Oxford University Press.

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