7.

Symbiotic Origin of Eukaryotic Cells 1

FIGURE 26.19
Unusual Lipids of Archaea
PHOSPHOLIPID WITH
TWO 20-CARBON ISOPRENOID CHAINS The Archaea have lipid chains
made of 5-carbon isoprenoid
OHEther units rather than 2-carbon units as
CH2 O P O CH2linkage seen in Eubacteria. The isoprenoid
chains are linked to a glycerol
HO HOCH O P O OH
CH2 CH3
HO
via an ether link rather than an
CH O (CH2 CH2 CH CH2)4 H
CH3 ester link. In some instances the
isoprenoid lipid chains may contain
Glycerol CH2 O (CH2 CH2 CH CH2)4 H 40 carbons (bacterioruberin, for
example). These longer lipids
Isoprenoid chain span the whole membrane of
the Archaea.

BACTERIORUBERIN (40-CARBON ISOPRENOID)

OH

OH
HO

OH

the Euryarchaeota. Most Crenarchaeota are thermophilic and/or possess a sulfur-

based metabolism. An example is Sulfolobus, which lives in geothermal springs, grows
best at a pH of 2–3 and a temperature of 70–80°C, and oxidizes sulfur to sulfuric acid.
The Euryarchaeota are more varied and include two distinct thermophilic lineages,
two groups of methane-generating bacteria, and the Halobacteria. The latter are
found in the super salty Dead Sea and Great Salt Lake. They are extremely salt-
tolerant and grow in up to 5 M NaCl but will not grow below 2.5 M NaCl (sea
water is only
0.6 M). They respire normally but many also trap energy from sunlight using bacterio-
rhodopsin, which is related to the rhodopsin used as a photo-detector in animal eyes.

7. Symbiotic Origin of Eukaryotic Cells

Unlike the cells of prokaryotes, eukaryotic cells are divided into compartments by
membranes (Fig. 26.20). The most important compartment is the nucleus, where the
chromosomes reside. Eukaryotes are defined by the possession of a nucleus that typi-
cally contains several linear chromosomes. Higher eukaryotes are normally diploid Eukaryotic cells are
and have pairs of homologous chromosomes, although this is not always the case divided into
for less advanced eukaryotes. compartments, including
In addition to a nucleus, almost all eukaryotic cells (animal, plant, or fungus)
contain mitochondria. Plant cells contain chloroplasts as well as mitochondria. Both
of these organelles provide the majority of energy for all cellular processes. These
organelles contain their own genomes and encode at least a few of their own
proteins. The organelle genome is prokaryotic in nature. It consists of a circular
DNA mol- ecule that is not bound by histones. Mitochondria and chloroplasts
synthesize their own ribosomes, which are more closely related to those of bacteria
than to those of the eukaryotic cytoplasm. Both mitochondria and chloroplasts are
roughly the same size and shape as bacterial cells, and the organelles grow and divide
in the same man- ner as bacteria (Fig. 26.21). When a eukaryotic cell divides, each
daughter cell inherits some of its parent’s mitochondria and chloroplasts. If either Eukaryotes are derived
organelle is lost, it can- not be reconstructed because the nucleus does not have all from the merger of two or
of the genetic information needed to assemble the entire organelle. more ancestral
2 CHAPTER TWENTY SIX • Molecular Evolution

ANIMAL CELL PLANT CELL

Endoplasmic
Ribosomes reticulum Nucleolus Cell wall

Lysosome
Vesicle
Cytoskeleton
Nucleus

Chloroplast

DNA DNA

Mitochondrion
Mitochondrion
Cytoplasmic
ribosomes

Centrioles Golgi
apparatus

Ribosomes
Endoplasmic
reticulum

FIGURE 26.20
Defining Features of Eukaryotic Cells
Eukaryotic cells have membrane bound compartments that are not found in prokaryotes. In a typical animal cell, these compartments include
the nucleus, mitochondria, endoplasmic reticulum, golgi apparatus, and lysosomes. In addition a typical plant cell also has chloroplasts, which
harvest light energy and convert it into ATP. Animal cells maintain their 3D shape with an internal cytoskeleton composed of microtubules and
microfilaments. In contrast, plant cells maintain their shape by a rigid cell wall surrounding the cytoplasm.

FIGURE 26.21
Chloroplasts arise by
Division
Transmission electron micrograph
of a dividing chloroplast in a
bean seedling. Chloroplasts and
mitochondria divide independently
of the eukaryotic cell in which
they reside. The organelles divide
by binary fission in a manner
reminiscent of prokaryotic cells. The
plastid shown here is technically an
“etioplast,” a precursor chloroplast
that has not yet developed any
green pigment. (Credit: Biochemistry
and Molecular Biology of Plants by
Buchanan, Gruissem and Jones,
The symbiotic theory proposes that the complex eukaryotic cell arose by a series
2000, American Society of Plant
Physiologists.)
of symbiotic events in which organisms of different lineages merged. The cells of
higher organisms are thus symbiotic associations. The word “symbiosis” is from the
Greek word “sym,” meaning together, and “bios,” meaning life. The nuclear genes
of eukaryotic cells are sometimes referred to as derived from the “urkaryote.”

symbiotic theory Theory that the organelles of eukaryotic cells are derived from symbiotic prokaryotes
symbiosis Association of two living organisms that interact
urkaryote Hypothetical ancestor that provided the genetic information of the eukaryotic nucleus
 7. Symbiotic Origin of Eukaryotic Cells 3

The urkaryote is the hypothetical ancestor that provided the genetic information
found in the present-day eukaryotic nucleus. Mitochondria are
According to the symbiotic theory, mitochondria are descended from bacte- derived from ancestral
ria that colonized the ancestors of modern eukaryotic cells. These bacteria received bacteria that specialized
in respiration, whereas
shelter and nutrients, and in return, devoted themselves to generating energy by res-
chloroplasts are
piration. During the following eons, these bacteria became narrowly specialized for descended from
energy production, lost the ability to survive on their own, and evolved into mito-
chondria. The term endosymbiosis is sometimes used for symbiotic associations where
one partner is physically inside the other (“endo” is from the Greek for inside), as in
the present case. Similarly, chloroplasts are descended from photosynthetic bacteria
that took up residence in the ancestors of modern-day plants. The term plastid refers
to all organelles that are genetically equivalent to chloroplasts, whether functional
or not. Since fungi do not contain chlorophyll, the green light-absorbing pigment of
plants, fungi were once thought to be degenerate plants that had lost their chloro-
phyll during evolution. However, fungi contain no trace of a plastid genome, mean-
ing they were never photosynthetic, confirming the evolutionary tree based on
rRNA sequences (see Fig. 26.17).

7.1. The Genomes of Mitochondria and Chloroplasts

Both mitochondria and chloroplasts contain a genome consisting of a circular DNA
molecule, presumably derived from the ancestral bacterial chromosome. Over
evolu- tionary time, these organelle genomes lost many genes that were unnecessary
for life as an organelle inside a host cell. In addition, many genes that are still necessary
have been transferred to the chromosomes in the nucleus. Consequently, the
mitochondria of animals have very little DNA left. For example, human
mitochondrial DNA has only 13 protein-encoding genes, together with genes for
several rRNA and tRNA molecules (Fig. 26.22). However, about 400 different
proteins are present in mito- chondria. The genes for most of these reside in the
nucleus and these polypeptides are imported into the organelle after synthesis on Chloroplasts and
the ribosomes of the eukaryotic mitochondria possess small
cytoplas
The chloroplasts of higher plants retain rather more DNA than mitochondria—
approximately enough for a hundred genes—but this is still much less than their Defects due to
bacterial ancestors. It is estimated that 1000 or more genes from the ancestral photo- mutations in
synthetic bacterium have been transferred to the plant cell nucleus. mitochondrial genes are
During sexual reproduction, mitochondria and chloroplasts are inherited mater-
nally. When a sperm fertilizes an egg cell to create a zygote, the organelles of the
sperm are lost. The new individual retains the organelles from the egg cell (i.e., those
from the female parent only). Certain inherited defects of humans are due to muta-
tions in the mitochondrial DNA. These affect the generation of energy by respi-
ration and affect the function of muscle and nerve cells in particular. These defects
are passed on through the maternal line because all children with the same mother
inherit the same mitochondria.
Partial exceptions to the rule of maternal inheritance for organelles occur in a
few single-celled eukaryotes. Chlamydomonas is a single-celled green alga whose
cells contain a single chloroplast. During mating, about 5% of the zygotes receive two
chloroplasts (one from each parent) rather than one. In these cells recombination
can occur between the two different chloroplast genomes. Division of the zygote
gives cells with only a single chloroplast each. These may be examined to
determine the outcome of the genetic crosses.

endosymbiosis Form of symbiosis where one organism lives inside the other
plastid Any organelle that is genetically equivalent to a chloroplast, whether functional in photosynthesis or not
4 CHAPTER TWENTY SIX • Molecular Evolution

FIGURE 26.22 EVOLUTION OF MITOCHONDRIAL GENOME

Genetic Map of Human
Mitochondrial DNA Estimated Ancestral Protozoa and Fungi Animals
3,000,000 bp 50,000 bp 15,000 bp
A) During evolution, the
mitochondrial genome has been
streamlined. Many of the genes
necessary for mitochondrial function
have moved to the nucleus, causing
the mitochondrial genome to shrink
in size. B) The mitochondrial DNA
of humans contains the genes for
ribosomal RNA, transfer RNA, Extra genes lost or To nucleus
and some proteins of the
A captured by nucleus
electron transport chain.
HUMAN MITOCHONDRIAL DNA

F PT

V E

0/16569
14747

L
3229

HUMAN mt 12336 L
I DNA (16,569
Q bp) HS
M

5512
10766
W ND4L
A
N ND3
C R
Y
7445 G
ATP8

SD
B ATP6
K

7.2. Primary and Secondary Endosymbiosis

A symbiotic relationship where one organism lives inside the other is known as
endo- symbiosis. Primary endosymbiosis refers to the original internalization of
prokaryo- tes by an ancestral eukaryotic cell, resulting in the formation of the
mitochondria and chloroplasts. Two membranes surround mitochondria and
chloroplasts. The inner one is derived from the bacterial ancestor and the outer
“mitochondrial” or “chloroplast” membrane is actually derived from the host-cell
membrane. However, several line- ages of protozoans appear to have engulfed other
single-celled eukaryotes, in particu- lar algae. Several groups of algae therefore have
chloroplasts acquired at second-hand by what is termed secondary endosymbiosis.
In contrast to the typical two membranes of primary organelles, four membranes
surround chloroplasts obtained by secondary endosymbiosis. In most cases, the
nucleus of the engulfed eukaryotic alga has disappeared without trace. Occasionally,

primary endosymbiosis Original uptake of prokaryotes by the ancestral eukaryotic cell, giving rise to mitochondria and chloroplasts
secondary endosymbiosis Uptake by an ancestral eukaryotic cell of another single-celled eukaryote, usually an alga, thus providing
chloroplasts at second-hand
 7. Symbiotic Origin of Eukaryotic Cells 5

PRIMARY ENDOSYMBIOSIS FIGURE 26.23

Photosynthetic Primary versus Secondary
Mitochondrion eukaryote (alga) Endosymbiosis
Nucleus Primary endosymbiosis yields
organelles with two membranes.
Ancestral In this example, the original
host cell independent cyanobacterium has
Cytoplasmic a cytoplasmic membrane, which is
membrane Chloroplast retained, and an outer membrane,
with double which is lost during symbiosis.
membrane
Cyanobacterium When the two cells associate, the
host-cell cytoplasmic membrane
SECONDARY ENDOSYMBIOSIS surrounds the cyanobacterium,
which is therefore left surrounded
Mitochondrion Nucleomorph
(degenerate by two membranes. In contrast to
Nucleus nucleus) primary endosymbiosis, secondary
endosymbiosis occurs when
an ancestral host cell engulfs a
Ancestral photosynthetic eukaryotic alga.
host cell The alga already has a chloroplast
with two membranes as well as a
nucleus and other organelles. Since
Chloroplast Chloroplast
the host cell only needs the energy
with four with four
membranes membranes from the chloroplast, the other
captured organelles degenerate
Photosynthetic and eventually disappear. However,
eukaryote (alga) the membranes often remain and
the chloroplast is left with four
membranes, rather than two.

the remains of this nucleus are still to be found lying between the two pairs of mem-
branes (Fig. 26.23). This structure is termed a nucleomorph and can be seen in cryp-
tomonad algae where it represents the remains of the nucleus of a red alga that was
swallowed by an amoeba-like ancestor. The nucleomorph contains three vestigial
linear chromosomes totaling 550 kb of DNA. These carry genes for rRNA that is
incorporated into a few eukaryotic type ribosomes that are also located in the space
between the two pairs of membranes.
Cells resulting from secondary endosymbiosis are composites of four or five
original genomes. These include the primary ancestral eukaryote nucleus and its
mitochondrion, plus the nucleus, mitochondrion, and chloroplast from the second-
ary endosymbiont. Many genes from the subordinate genomes have been lost during
evolution and no trace has ever been found of the secondary mitochondrion. Some
genes from the secondary endosymbiont nucleus have been transferred to the pri-
mary eukaryotic nucleus. The protein products of about 30 of these are made on
ribosomes belonging to the primary nucleus and shipped from the primary eukary-
otic cytoplasm back into the nucleomorph compartment. In turn, the nucleomorph
contains genes for proteins that are made on the 80 S ribosomes in the nucleomorph
compartment and transported across the inner two membranes into the chloroplast.
Finally, there are proteins now encoded by the primary nucleus that must be trans-
located across both sets of double membranes from the primary cytoplasm into the
chloroplast!

nucleomorph Degenerate remains of the nucleus of a symbiotic eukaryote that was incorporated by secondary endosymbiosis into another
eukaryotic cell
6 CHAPTER TWENTY SIX • Molecular Evolution

Box 26.1 Is Malaria Really a Plant?

Malaria is a disease that affects many millions of people world wide and is responsible for
The malarial plastid or “apicoplast” is thought to derive from secondary endosymbio- sis.

rps 2 F

rpo C2 clp C

tuf A
W rps 7
S G rps 12
G
Q rps 11
rpl 36
rpo C1
rps 5
rpl 6
rps 8
rpl 14
rps 17
rpl 16
PEK
Plasmodium falciparum
rpo B DSY
plDNA (35 kb)
rps 3
rps 19
rpl 7
rpl 7
rpl 4

M
L* C
Hrps 4

V T
T R M
L LSU rRNA
AI R'
N

LSU rRNA SSU rRNA

M
R V
R
N SSU rRNA L
A
I

FIGURE 26.24
Plastid Genome of Plasmodium
The circular genome of the plastid of Plasmodium has genes for rRNA, tRNA, and protein synthesis. The tR

Apicomplexa Phylum of parasitic single-celled eukaryotes, including malaria, which contain both mitochondria and degenerate non-photosyn-
thetic chloroplasts
apicoplast Degenerate non-photosynthetic chloroplast found in members of the Apicomplexa
Plasmodium The malaria parasite, a protozoan belonging to the Apicomplexa