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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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18 Chapter 1: Cells and Genomes

764 genes in families with

4–19 gene members

273 genes in families

with 3 gene members

283 genes in families with

38–77 gene members

2126 genes with

no family relationship

Figure 1–20 Families of evolutionarily

related genes in the genome of Bacillus

subtilis. The largest gene family in this

bacterium consists of 77 genes coding for

varieties of ABC transporters—a class of

membrane transport proteins found in all

three domains of the living world. (Adapted

from F. Kunst et al., Nature 390:249–256,

1997. With permission from Macmillan

Publishers Ltd.)

568 genes in families

with 2 gene members

are likely to have diverged in their function—are called paralogs. Genes that are

related by descent in either way are called homologs, a general term used to cover

both types of relationship (Figure 1–21).

Genes Can Be Transferred Between Organisms, Both in the

MBoC6 m1.24/1.20

Laboratory and in Nature

Prokaryotes provide good examples of the horizontal transfer of genes from one

species of cell to another. The most obvious tell-tale signs are sequences recognizable

as being derived from viruses, those infecting bacteria being called bacteriophages

(Figure 1–22). Viruses are small packets of genetic material that have

evolved as parasites on the reproductive and biosynthetic machinery of host cells.

Although not themselves living cells, they often serve as vectors for gene transfer.

A virus will replicate in one cell, emerge from it with a protective wrapping, and

then enter and infect another cell, which may be of the same or a different species.

Often, the infected cell will be killed by the massive proliferation of virus particles

inside it; but sometimes, the viral DNA, instead of directly generating these particles,

may persist in its host for many cell generations as a relatively innocuous

passenger, either as a separate intracellular fragment of DNA, known as a plasmid,

or as a sequence inserted into the cell’s regular genome. In their travels, viruses

can accidentally pick up fragments of DNA from the genome of one host cell and

ferry them into another cell. Such transfers of genetic material are very common

in prokaryotes.

Horizontal transfers of genes between eukaryotic cells of different species

are very rare, and they do not seem to have played a significant part in eukaryote

evolution (although massive transfers from bacterial to eukaryotic genomes have

occurred in the evolution of mitochondria and chloroplasts, as we discuss below).

ancestral organism

ancestral organism

gene G

gene G

SPECIATION TO GIVE TWO

SEPARATE SPECIES

GENE DUPLICATION

AND DIVERGENCE

species A

species B

later organism

gene G 1

gene G A gene G B

gene G 2

genes G A and G B are orthologs

genes G 1 and G 2 are paralogs

(A)

(B)

Figure 1–21 Paralogous genes and

orthologous genes: two types of

gene homology based on different

evolutionary pathways. (A) Orthologs.

(B) Paralogs.

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