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Crystal (trilobite) Eyes by Richard Fortey

Crystal (trilobite) Eyes by Richard Fortey

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06/14/2009

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Natural History: Crystal Eyeshttp://findarticles.com/p/articles/mi_m1134/is_8_109/ai_659131...1 of 38/12/2008 12:01 AM
FindArticles>Natural History >Oct, 2000> Article> Print friendly 
Crystal Eyes
Richard Fortey Five hundred million years ago, trilobites looked at the world through clear calcite glasses.One of the most difficult jobs I ever attempted was to count the number of lenses in a fossil trilobite eye. I had been fascinated by these long-extinctarthropods as a child and have since spent years studying them as a paleontologist. For the task at hand, I took several photographs from differentangles of the eye (which was about the size of a grain of rice) and then made enormous prints that allowed me to see the individual lenses (which were microscopic). Eventually I hit upon the notion of pricking each counted lens on the photograph with a pin so that it wasn't counted twice.But when I moved on to the next photo, I would be unsure of the last lens I'd counted and how the tiny hexagons linked up from one picture to theother. Was the last lens the one with the little scratch, or that one a mite larger than its neighbor? The work was undeniably suitable for anobsessive with insomnia, but it was worth it. For trilobites have the first really well-preserved visual systems in the fossil record. Furthermore,their eyes are unique in that they are made of the mineral calcite. The trilobites have many living, if remote, arthropod relatives--and trilobiteshave been extinct for 250 million years--but no other has chosen the trilobite way to see the world.Calcite is one of the most abundant minerals. The white cliffs of Dover are calcite; so are the bluffs along the Mississippi River. Surely one couldexpect no surprises from a substance so common and so familiar. But calcite has some unusual properties. Because of its natural impurities it haslong provided builders with colorful stone and decorative slabs. The purest examples of calcite, however, are transparent, with perfect crystalform and clarity. The chemical composition, Ca[CO.sub.3], is simple as minerals go. As the crystal grows, the constituent atoms stack together ina lopsided way and do not allow other, stray atoms to intrude to cloud the crystal's mineral exactitude. The clearest calcite crystal, transparentas a toddler's motives, is Iceland spar. Look into a crystal of Iceland spar and you can see the secret of the trilobite's vision. While most otherarthropods have lenses made of relatively soft, unmineralized cuticle, similar to that of the rest of their exoskeleton, trilobites used thetransparency of clear calcite as a means of transmitting light. The trilobite eye is in continuity with the rest of its shelly armor. It sits on top of the animal's cheek, an en suite eyeglass, tough as a clamshell.Clear calcite is optically complex. If you break a large piece of crystalline calcite, you are left with a regular, six-sided chunk of the mineral--arhomb--which treats light in a peculiar way. If a beam of light is shone at the sides of the rhomb, the beam splits in two, a phenomenon known asdouble refraction. The course of the two rays is determined, as is the shape of the rhomb, by the stacking of the individual atoms. There is onedirection, and one direction only, in which light does not indulge in this optical split: when a ray of light approaches along what is termed the caxis of the crystal, it is afforded free passage. Like a VIP at an international airport, this privileged ray passes straight through. If a crystal iselongated in parallel to the c axis, into the shape of a long prism, light entering from most angles will be split and the dual rays will in turn bedeflected to reach the edge of the prism and will not be "seen" by a receptor cell located at the base of the prism. But light shining along the prism'slong axis will still pass unrefracted. This is how most trilobite eyes are constructed. We know that the first trilobites already had a well-developed visual system. Indeed, the large eyes found in the genus Fallotaspis, from Morocco, prove that sophisticated vision goes back at least 540 million years to the Cambrian period.Recent laboratory work has led to the discovery of the pervasive influence of genes that control the sequence of development of the various organsas animals grow from embryo to adult. These are genes so deeply embedded in the body plans of organisms that the memory of their origin is lostfar back in Precambrian history. We can never, ever, directly sample the genetic code of the trilobite, but we can be sure that its development was under the control of the same kinds of genes we recognize in living animals. Development inexorably follows a blueprint originally drawn upin the most ancient times. It is rather wonderful to imagine this distant manifesto at work on the growing trilobite, directing the brain to beenclosed within the head and, of course, issuing instructions for the growth and development of eyes.For eyes are part of this ancient list of instructions. It seems that the making of an eye is the same impulse in fish or fly or man. Eyes are underthe control of a gene called Pax6. The end product may be very different, but the instruction "Make eyes" may be common to all animals. Thedeep language of the genes is an Esperanto of biological design that can be understood by all creatures that have light-sensitive organs. Trilobitesoffer visible evidence of the halfway point in optical history. We can feel a bond with the trilobite that would not have been apparent whennineteenth-century investigators first gazed upon the animal's stony eyes. "Look into my eyes," the trilobite now seems to say, "and you will seethe vestiges of your own history."The number of lenses in a trilobite eye varies, according to the species, from a mere one to several thousand, as in the one I attempted to count.This eye was of the compound type and, just like a fly's eye, was a honeycomb of hexagons. In most trilobite lenses, the c axis is exactly at rightangles to the surface of the lens. If you can see the whole surface of an individual lens, then the chances are that the lens can "see" you. So youcould deduce a trilobite's field of view by summing up the angles of orientation of all the individual lenses. Thirty years ago, Euan Clarkson, of theUniversity of Edinburgh, investigated the field of view of trilobite eyes for the first time. What he did was to mount several species of trilobites ina way that allowed him greater accuracy in measuring the c axis of each of hundreds of hexagonal lenses. Then he plotted the spread of directionsof these axes to see what the trilobites saw.Clarkson found that the trilobite eyes he examined looked sideways, forward, and often a little backward. Like searchlights sweeping over the
 
Natural History: Crystal Eyeshttp://findarticles.com/p/articles/mi_m1134/is_8_109/ai_659131...2 of 38/12/2008 12:01 AM
ground and low bushes but not up to the sky, the trilobites' eyes could be cast over the area surrounding the animal, but not upward or directly downward. Most trilobites lived on and around the seafloor, and this was the world they wished to appraise, a world on the sediment where, day or night, most of the events affecting their lives took place. Densely lensed eyes of trilobite type are particularly good at detecting movement. Another animal approaching across the sediment surface will trigger one lens after another as its image impinges on different parts of the field of  view. If the change is alarming, the trilobite may be stimulated to take evasive action: perhaps to roll up into a ball or to swim away as fast aspossible.The trilobite eye grew in harmony with the animal. As in all arthropods, the eye surface had to be molted along with the rest of the hardexoskeleton. In trilobites, as the new skeleton hardened after each molt, more lenses were added and new crystals generated from a zone at the topof the eye. The animals were not able to see as we see but rather appreciated the world in a thousand fragments of light, as if the brain were apointillist with a palette of prisms. The eyes may have permitted comprehension of the world in the same fashion as the similar compound eyes of living arthropods. Apposition eyes do not form complete images of their surroundings (some other arthropod eyes have lenses arranged in such a way that they are able to collaborate and produce a single, complex image).But there is another kind of trilobite eye. One of the commonest trilobites in the Devonian rocks of New York, Ohio, Ontario, Germany, andMorocco is the compact animal called Phacops. Its large, crescent-shaped eyes stand prominently atop the cheeks. Instead of lenses so minute thatthey require a microscope to be seen properly, Phacops's lenses can be recognized by our unaided eyes as tiny, perfectly formed balls, which lineup quite conspicuously in rows. These eyes seem to have been turned out by a machine, neat as billiard balls arranged in a box. When sectioned,the eyes reveal their secrets. First, the lenses are indeed nearly spherical, or perhaps slightly drop-shaped, with a disquieting resemblance toglass eyes. Second, there is usually a little "wall" between adjacent lenses, a kind of baffle that stopped light from one lens overlapping that of thenext. Often the lenses are slightly sunken, and the areas between them swollen. Clearly a very sophisticated structure (even more so than thehexagonal-lensed trilobite eye), Phacops's crystal eye is a sports coupe in the age of the boneshaker.In 1972 Kenneth M. Towe, of the Smithsonian Institution in Washington, D.C., demonstrated the efficiency of the Phacops kind of trilobiteeyes--by taking photographs through them. The fat, biconvex lenses of the phacopid eye were designed to bring bright beams to a focus. If youhold a clear glass marble up to the light and peer through it, you can get some idea of the process: you will see an upside-down world, all bent anddistorted. But in Towe's photos, the trilobite images seem to be much clearer than that. How could this be? The problem with light travelingthrough a convex lens to a focus is that different rays travel different distances through the lens according to their trajectory. This means thatthe rays are bent to different degrees. The result is a fuzzy focus. Euan Clarkson and University of Chicago physicist Riccardo Levi-Settidiscovered that something strange had happened to the calcite in the lower part of each Phacops lens: magnesium atoms were present in just theright quantity to correct the spherical aberration. For every bend to the left, there was a compensating bend to the right. This corrective layermade a bowl within the lens; the trilobite had thus manufactured what modern opticians term a doublet. The animals with these eyes may haveseen more complete images of an object than their hexagonal-lensed fellows. All this 400 million years ago.The trilobite whose lenses I started counting had the hexagonal lens design and was a particularly goggle-eyed species, with peepers puffed up likelittle bladders. The eyes bulged out on either side of the head in the manner of those slightly grotesque ornamental goldfish that have such athyroidal look. I named this shrimp-sized animal Opipeuter, having recruited the help of a classicist friend to find out the Greek for "one whogazes." The lenses of Opipeuter's eyes were tiny, but unlike those in the crescent-shaped eyes of most trilobites that lived on the seafloor,Opipeuter's lenses faced in all directions, even downward. This trilobite must have been a free swimmer rather than a bottom dweller. Ancientoceans could have swarmed with trilobites, just as krill throng in modern-day seas. These elongated trilobites were the remote ecologicalequivalents of shrimp, moving through the water column or even on the surface. A number of different trilobites proved to have this free-swimming design. Fossils of one, the Cyclops-eyed Pricyclopyge, are common in dark Ordovician mudstones 400 million years old. These sediments were originally deposited in relatively deep water, to which these swimmers wereapparently confined. Could one somehow test the difference in life habits between shallow- and deep-dwelling trilobites by examining their eyes?Tim McCormick, now at the University of Glasgow, and I, following techniques used to determine how light intensity influences the eyes of livingarthropods, were able to insinuate ourselves into the daily lives of our fossil trilobites by making a series of careful measurements on eyeconstruction. We were able to show that Pricyclopyge had eyes constructed in the same fashion as crustaceans that still live in the deeper part of the water column in oceans today. So it seems that trilobites were indeed able to swim at various depths.Blind, totally eyeless trilobites have given us another indication of the range of trilobite habits and habitats. My colleague Bob Owens and Icollected trilobite fossils in some localities in Wales where ten or more blind or nearly blind species lived together. They must have crawled aboutthe seafloor in a dark world. Bottom dwellers lost their eyes not because they were degenerate but because, like living crustaceans that inhabitcaves, these species simply did not need eyes in their specialized environment. Yet other trilobites in the same Welsh rocks were huge-eyedswimmers. It did not take us long to deduce that the swimmers had swum well above the lightless seafloor on which the blind animals dwelled,and had joined their sightless fellows only when they drifted to the seafloor in death and, fortunately for us, were fossilized.Richard Fortey ("Crystal Eyes") brings us his great enthusiasm for ancient fossil trilobites in a chapter he has adapted from his new book Trilobite!Eyewitness to Evolution (Knopf, November 2000). The volume is something of a bridge between his other writings--learned and popular. On theone hand, he is at work as "trilobite coordinator" for the upcoming tome Treatise of Invertebrate Paleontology, and on the other hand, heconfesses, he "must be one of the few paleontologists to have written two humorous books pseudonymously." A Merit Researcher at the NaturalHistory Museum in London and visiting professor of paleobiology at Oxford University, Fortey, like many other paleontologists, collected fossils as

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