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Evolution__3rd_Edition

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..<br />

Figure 18.3<br />

Around 2–3 billion years ago,<br />

prokaryotic life had evolved<br />

to live in a variety of habitats,<br />

using a variety of metabolisms.<br />

The illustration shows<br />

hydrothermal systems around<br />

structures called komatiite<br />

shields, chemotrophs in midocean<br />

ridges, and various life<br />

forms in coastal, lacustrine,<br />

and oceanic waters. Redrawn,<br />

by permission of the publisher,<br />

from Nisbet (2000).<br />

Early cellular life was prokaryotic<br />

Hydrothermal<br />

communities around<br />

andesite volcanoes<br />

Light<br />

Cu, Mo, Zn, S<br />

Stromatolites<br />

Lake<br />

communities<br />

Light<br />

year period (for instance, Knoll & Baghoorn 1977, and Schopf 1999 reviews the evidence).<br />

Cells had therefore likely evolved by 3.5 billion years ago, or soon afterwards.<br />

18.3.2 The origin of cells<br />

Coastal sediment<br />

S-microbial mats<br />

Light<br />

Organic<br />

debris<br />

S cycle and<br />

methanogens<br />

Planktonic<br />

cyanobacteria<br />

Open ocean<br />

Light Light<br />

More oxidized<br />

More reduced<br />

CHAPTER 18 / The History of Life 531<br />

Hydrothermal<br />

supply of metals<br />

and reductant<br />

in deeper water<br />

Mid-ocean ridge<br />

chemotrophic<br />

community<br />

Mg, SO 4<br />

Fe, Mn, S,<br />

CH 4, H 2<br />

Hydrothermal<br />

systems around<br />

komatiite shields<br />

Fossils of cellular prokaryotic life have been found at several sites, aged between 3.5 and<br />

2 billion years ago. They often exist in the form of stromatolites. Stromatolites are<br />

layered structures that are formed when cells grow on the sea surface, and sediments<br />

are deposited among or above the cells. The cells then grow up to the light, leaving a<br />

mineralized layer below them. As the process is repeated over time, a stromatolite<br />

builds up, consisting of many mineralized layers. Stromatolites are still formed at<br />

certain sites in the world today, but are rarer than in the past. One reason may be that<br />

grazers now consume any cell mats as they form. In the past, grazers did not exist and<br />

stromatolites could accumulate. Around 2–3 billion years ago, microbial prokaryotic<br />

life seems to have existed in several environments, and to have evolved several<br />

metabolic processes (Figure 18.3).<br />

Thus, cellular life was flourishing by 2–3 billion years ago. However, the origin of<br />

cells was probably not an evolutionarily inevitable step. Unadorned replicating molecular<br />

systems could have persisted, the molecules being replicated as their component<br />

building blocks bonded to them and formed copies, or near copies, of the whole. For<br />

the system to become more complex, it needs enzymes and metabolic systems that<br />

enable it to harvest resources more powerfully, or to exploit the resources better by<br />

converting them into the molecular units needed for replication. This step is difficult<br />

Lava<br />

Light<br />

Ni, Co, Fe,<br />

S, Mg

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