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Molecular Biology of the Cell by Bruce Alberts, Alexander Johnson, Julian Lewis, David Morgan, Martin Raff, Keith Roberts, Peter Walter by by Bruce Alberts, Alexander Johnson, Julian Lewis, David Morg

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28 Chapter 1: Cells and Genomes

early

eukaryotic cell

eukaryotic cell

capable of

photosynthesis

Figure 1–31 The origin of chloroplasts.

An early eukaryotic cell, already possessing

mitochondria, engulfed a photosynthetic

bacterium (a cyanobacterium) and retained

it in symbiosis. Present-day chloroplasts

are thought to trace their ancestry back to

a single species of cyanobacterium that

was adopted as an internal symbiont (an

endosymbiont) over a billion years ago.

photosynthetic

bacterium

chloroplasts

Many of the genes that are missing from the mitochondria and chloroplasts

have not been lost; instead, they have moved from the symbiont genome into the

DNA of the host cell nucleus. The nuclear DNA of humans contains many genes

coding for proteins that serve MBoC6 essential m1.36/1.31 functions inside the mitochondria; in

plants, the nuclear DNA also contains many genes specifying proteins required in

chloroplasts. In both cases, the DNA sequences of these nuclear genes show clear

evidence of their origin from the bacterial ancestor of the respective organelle.

Eukaryotic Genomes Are Big

Natural selection has evidently favored mitochondria with small genomes. By contrast,

the nuclear genomes of most eukaryotes seem to have been free to enlarge.

Perhaps the eukaryotic way of life has made large size an advantage: predators

typically need to be bigger than their prey, and cell size generally increases in proportion

to genome size. Whatever the reason, aided by a massive accumulation of

DNA segments derived from parasitic transposable elements (discussed in Chapter

5), the genomes of most eukaryotes have become orders of magnitude larger

than those of bacteria and archaea (Figure 1–32).

The freedom to be extravagant with DNA has had profound implications.

Eukaryotes not only have more genes than prokaryotes; they also have vastly more

DNA that does not code for protein. The human genome contains 1000 times as

many nucleotide pairs as the genome of a typical bacterium, perhaps 10 times as

MAMMALS, BIRDS, REPTILES

Fugu zebrafish

AMPHIBIANS, FISHES

Drosophila

CRUSTACEANS, INSECTS

NEMATODE WORMS

PROTOZOANS

Mycoplasma

BACTERIA

ARCHAEA

Arabidopsis

PLANTS, ALGAE

FUNGI

E. coli

yeast

malarial parasite

Caenorhabditis

human

10 5 10 6 10 7 10 8 10 9 10 10 10 11 10 12

nucleotide pairs per haploid genome

frog

shrimp

newt

wheat

lily

amoeba

Figure 1–32 Genome sizes compared.

Genome size is measured in nucleotide

pairs of DNA per haploid genome, that is,

per single copy of the genome. (The cells

of sexually reproducing organisms such as

ourselves are generally diploid: they contain

two copies of the genome, one inherited

from the mother, the other from the father.)

Closely related organisms can vary widely

in the quantity of DNA in their genomes,

even though they contain similar numbers

of functionally distinct genes. (Data from

W.H. Li, Molecular Evolution, pp. 380–383.

Sunderland, MA: Sinauer, 1997.)

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