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THE ORIGIN OF LIFE  195















                             (a)


                                                 (b)









                                                                               (c)

             Figure 8.8  Prokaryote fossils from the Gunflint Chert of Ontario, Canada (c. 1.9 Ga): (a) Eosphaera,
             (b) Kakabekia, and (c) Gunfl intia. Specimens are 0.5–10 μm in diameter. (Redrawn from photographs
             in Barghoorn & Taylor 1965.)


             least 2.7 Ga, but it is also the oldest hint of   transfer,  fl agella for movement and  chloro-
             the occurrence of eukaryotes, long before any   plasts in plants for photosynthesis. There are
             fossils of that major life domain.              also many major biochemical differences
                                                             between prokaryotes and eukaryotes.
                                                               The origin of eukaryotes is mysterious
             LIFE DIVERSIFIES: EUKARYOTES                    because they are in many ways so different
                                                             from prokaryotes. The most attractive idea
             Eukaryote characters
                                                             for their origin is the endosymbiotic theory,
             Evidence about the earliest evolution of the    proposed by Lynn Margulis in the 1970s.
             three domains is scant. It has long been        According to this hypothesis (Fig. 8.9c), a
             assumed that prokaryotes (i.e. Archaea and      prokaryote consumed, or was invaded by,
             Bacteria) were the sole life forms for a billion   some smaller energy-producing prokaryotes,
             years or more, and that eukaryotes came         and the two species evolved to live together
             much later. This evidence is much more          in a mutually benefi cial way. The small invader
             blurred now (Embley & Martin 2006), and         was protected by its large host, and the larger
             the fossils, biomarkers and molecular evi-      organism received supplies of sugars. These
             dence suggest that eukaryotes might be as old   invaders became the mitochondria of modern
             as one or other of the prokaryote domains.      eukaryote cells. Other invaders may have
             The appearance of eukaryotes was important,     included worm-like swimming prokaryotes
             whenever it happened, because they are          (spirochaetes) that became motile fl agella,
             complex and include truly multicellular and     and photosynthesizing prokaryotes that
             large organisms.                                became the chloroplasts of plants.
               Eukaryotes are distinguished from pro-          The endosymbiotic model is immensely
             karyotes (Fig. 8.9a, b) by having a nucleus     attractive, and some aspects have been con-

             containing their DNA in chromosomes (pro-       firmed spectacularly. Most notable is that the
             karyotes have no nucleus, and they have only    mitochondria and chloroplasts in modern
             a circular strand of DNA) and cell organelles,   eukaryotes are confi rmed  as  prokaryotes,
             that is, specialized structures that perform key   the mitochondria being closely related to
             functions, such as  mitochondria for energy     α-proteobacteria and the chloroplasts to
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