A.

The Simplest Living Things

extremely resistant to heat and further desiccation.

Under suitable conditions, the spores can germinate
and renew their vegetative growth. Spore formation is
one of several examples of the development of specialized cells or differentiation among prokaryotes.

8. Photosynthetic and Nitrogen-Fixing

Prokaryotes
It is likely that the earth was once a completely
anaerobic place containing water, ammonia, methane,
formaldehyde, and more complicated organic compounds. Perhaps the first forms of life, which may
have originated about 3.5 x 109 years ago, resembled
present-day anaerobic bacteria. The purple and green

photosynthetic bacteria may be related to organisms

that developed at a second stage of evolution: those
able to capture energy from sunlight. Most of these
gram-negative photosynthetic bacteria are strict anaerobes. None can make oxygen as do higher plants.
Rather, the hydrogen needed to carry out the reduction
of carbon dioxide in the photosynthetic process is
obtained by the splitting of inorganic compounds, such
as H2S, thiosulfate, or H2, or is taken from organic
compounds. Today, photosynthetic bacteria are found
principally in sulfur springs and in deep lakes, but at
one time they were probably far more abundant and
the only photosynthetic organisms on earth.
Before organisms could produce oxygen a second
complete photosynthetic system, which could cleave
H2O to O2, had to be developed. The simplest oxygen-

BOX 1-C IN THE BEGINNING

No one knows how life began. Theories ranging
from the biblical accounts to recent ideas about the
role of RNA are plentiful but largely unsatisfying. In
the 1800s the great physical chemist Arrhenius was
among scientists that preferred the idea held by
some scientists today that a seed came from outer
space. Until recently the only concrete data came
from fossils. Making use of a variety of isotopic
dating methods it can be concluded that cyanobacteria were present 2.2 109 years ago and eukaryotes
1.4 109 years ago. About 0.5 109 years ago the
Cambrian explosion led to the appearance of virtually all known animal phyla. Many of these then
became extinct about 0.2 109 years ago.
New insights published in 1859a were provided
by Charles Darwin. However, his ideas were only
put into a context of biochemical data after 1950 when
sequencing of proteins and later nucleic acids began.
From an astonishingly large library of sequence data
available now we can draw one firm conclusion:
Evolution can be observed;b it does involve mutation of
DNA. Comparisons of sequences among many
species allow evolutionary relationships to be
proposed.c-e In general these are very similar to
those deduced from the fossil record. They support
the idea that evolution occurs by natural selection
and that duplication of genes and movements of
large pieces of DNA within the genome have occurred often. As many as 900 ancient conserved
regions of DNA in the E. coli genome corresponding to those in human, nematode, and yeast DNA
are thought to date back perhaps 3.5 109 years.f
However, nobody has explained how life evolved
before there was DNA.
One of the first scientists to devote his career to
biochemical evolution was I. V. Oparin,g who

published a book on the origin of life in 1924.

Oparin and J. B. S. Haldane, independently, proposed
that early life was anaerobic and that energy was
provided by fermentation. In 1951 Stanley Miller
built an apparatus that circulated CH4, NH3, H2O,
and H2, compounds thought to be present in a
primitive atmosphere, past an electric discharge.
He found glycine, alanine, -alanine, and other amino
acids among the products formed.h Schrdinger
pointed out that a flux of energy through a system
will tend to organize the system. The solar energy
passing through the biosphere induces atmospheric
circulation and patterns of weather and ocean
currents.i,j Perhaps in the primordeal oceans organic
compounds arose from the action of light and lightning discharges. These compounds became catalysts
for other reactions which eventually evolved into a
rudimentary cell-less metabolism. It is a large jump
from this to a cell! Among other problems is the lack
of any explanation for the development of individual
cells or of their genomes. However, because it helps
to correlate much information we will always take an
evolutionary approach in this book and will discuss the
beginnings a little more in later chapters.
a
b
c
d
e
f
g
h
i
j

Maynard-Smith, J. (1982) Nature (London) 296, 599 601

Lenski, R. E., and Travisano, M. (1994) Proc. Natl. Acad. Sci.
U.S.A. 91, 6808 6814
Wilson, A. C. (1985) Sci. Am. 253(Oct), 164 173
Eigen, M., Gardiner, W., Schuster, P., and Winkler-Oswatitsch, R.
(1981) Sci. Am. 244(Apr), 88 118
Doolittle, R. F. (1992) Protein Sci. 1, 191 200
Green, P., Lipman, D., Hillier, L., Waterston, R., States, D., and
Claverie, J.-M. (1993) Science 259, 1711 1716
Broda, E. (1980) Trends Biochem. Sci. 5, IV V
Miller, S. L. (1953) Science 117, 528 529
Mason, S. (1993) Trends Biochem. Sci. 18, 230 231
Welch, G. R. (1996) Trends Biochem. Sci. 21, 452