It Takes Teamwork: How Endosymbiosis Changed Life on Earth
by the Understanding Evolution team
In 1966, microbiologist Kwang Jeon was studying single-celled organisms called
amoebae, when his amoebae communities were struck by an unexpected plague: a
bacterial infection. Literally thousands of the tiny invaders — named x-bacteria by
Jeon — squeezed inside each amoeba cell, causing the cell to become dangerously
sick. Only a few amoebae survived the epidemic.
The blob-like form of an amoeba
The blob-like form of an amoeba. Photo © 2003 Dr. Peter Weish.
However, several months later, the few surviving amoebae and their descendants
seemed to be unexpectedly healthy. Had the amoebae finally managed to fight off the
x-bacterial infection? Jeon and his colleagues were surprised to find that the
answer was no — the x-bacteria were still thriving inside their amoebae hosts, but
they no longer made the amoebae sick. There were more surprises when Jeon used
antibiotics to kill the bacteria inside an amoeba — the host amoeba also died! The
amoebae could no longer live without their former attackers. Jeon discovered that
this was because the bacteria make a protein that the amoebae need to survive. The
nature of the relationship between the two species had changed entirely: from
attack and defense to cooperation.
In this case study we will explore these key questions:
- What is endosymbiosis?
- What role did endosymbiosis play in the evolution of eukaryotes?
- How does endosymbiosis change our view of the branching pattern on the tree of
life?
When two become one
Jeon’s colonies of amoebae seem perfectly happy living with their permanent guests,
the x-bacteria, inside of them. This kind of relationship — two or more different
species living in close association — is called symbiosis.
Three kinds of symbiosis:
mutualism
a symbiosis in which both organisms benefit
commensalism
a symbiosis in which one organism benefits without helping or harming the other
parasitism
a symbiosis in which one organism benefits at the expense of the other
Bacteria team up.Each amoeba and its x-bacteria work together for mutual benefit —
but they are still separate organisms. Each bacterium or amoeba divides on its own,
gets its own energy, uses its own genes, and makes its own proteins (mostly!).
However, with their close relationship, it seems possible that after many years of
evolving together, these cells could become not just a team, but a single
integrated organism with a common set of genes and proteins. A future scientist
discovering the descendents of Jeon’s amoebae might not guess that this one
“amoebacterium” was once two distinct organisms.
Evidence like this points to the likelihood that the “merging” of two simple
organisms has also happened under natural conditions. Long ago in evolutionary
history, two cells formed a symbiotic team that, over millions of years, evolved
into a single organism. The result of this union was the first eukaryotic cell —
the type of cell that makes up the human body. We humans owe our existence to two
bacteria that teamed up in a symbiotic relationship over a billion years ago!
From prokaryotes to eukaryotes
3 domainsLiving things have evolved into three large clusters of closely related
organisms, called “domains”: Archaea, Bacteria, and Eukaryota. Archaea and Bacteria
are small, relatively simple cells surrounded by a membrane and a cell wall, with a
circular strand of DNA containing their genes. They are called prokaryotes.
Virtually all the life we see each day — including plants and animals — belongs to
the third domain, Eukaryota. Eukaryotic cells are more complex than prokaryotes,
and the DNA is linear and found within a nucleus. Eukaryotic cells boast their own
personal “power plants”, called mitochondria. These tiny organelles in the cell not
only produce chemical energy, but also hold the key to understanding the evolution
of the eukaryotic cell.
typical prokaryote and eukaryote cells
The complex eukaryotic cell ushered in a whole new era for life on Earth, because
these cells evolved into multicellular organisms. But how did the eukaryotic cell
itself evolve? How did a humble bacterium make this evolutionary leap from a simple
prokaryotic cell to a more complex eukaryotic cell? The answer seems to be
symbiosis — in other words, teamwork.
Evidence supports the idea that eukaryotic cells are actually the descendents of
separate prokaryotic cells that joined together in a symbiotic union. In fact, the
mitochondrion itself seems to be the “great-great-great-great-great-great-great-
great-great granddaughter” of a free-living bacterium that was engulfed by another
cell, perhaps as a meal, and ended up staying as a sort of permanent houseguest.
The host cell profited from the chemical energy the mitochondrion produced, and the
mitochondrion benefited from the protected, nutrient-rich environment surrounding
it. This kind of “internal” symbiosis — one organism taking up permanent residence
inside another and eventually evolving into a single lineage — is called
endosymbiosis.
Evidence for endosymbiosis
Biologist Lynn Margulis first made the case for endosymbiosis in the 1960s, but for
many years other biologists were skeptical. Although Jeon watched his amoebae
become infected with the x-bacteria and then evolve to depend upon them, no one was
around over a billion years ago to observe the events of endosymbiosis. Why should
we think that a mitochondrion used to be a free-living organism in its own right?
It turns out that many lines of evidence support this idea. Most important are the
many striking similarities between prokaryotes (like bacteria) and mitochondria:
Membranes — Mitochondria have their own cell membranes, just like a prokaryotic
cell does.
DNA — Each mitochondrion has its own circular DNA genome, like a bacteria’s genome,
but much smaller. This DNA is passed from a mitochondrion to its offspring and is
separate from the “host” cell’s genome in the nucleus.bacteria/mitochondria
structural comparison
Reproduction — Mitochondria multiply by pinching in half — the same process used by
bacteria. Every new mitochondrion must be produced from a parent mitochondrion in
this way; if a cell’s mitochondria are removed, it can’t build new ones from
scratch.bacteria/mitochondria reproduction
When you look at it this way, mitochondria really resemble tiny bacteria making
their livings inside eukaryotic cells! Based on decades of accumulated evidence,
the scientific community supports Margulis’s ideas: endosymbiosis is the best
explanation for the evolution of the eukaryotic cell.
What’s more, the evidence for endosymbiosis applies not only to mitochondria, but
to other cellular organelles as well. Chloroplasts are like tiny green factories
within plant cells that help convert energy from sunlight into sugars, and they
have many similarities to mitochondria. The evidence suggests that these
chloroplast organelles were also once free-living bacteria.
The endosymbiotic event that generated mitochondria must have happened early in the
history of eukaryotes, because all eukaryotes have them. Then, later, a similar
event brought chloroplasts into some eukaryotic cells, creating the lineage that
led to plants.
the origins of mitochondria and chloroplasts
Despite their many similarities, mitochondria (and chloroplasts) aren’t free-living
bacteria anymore. The first eukaryotic cell evolved more than a billion years ago.
Since then, these organelles have become completely dependent on their host cells.
For example, many of the key proteins needed by the mitochondrion are imported from
the rest of the cell. Sometime during their long-standing relationship, the genes
that code for these proteins were transferred from the mitochondrion to its host’s
genome. Scientists consider this mixing of genomes to be the irreversible step at
which the two independent organisms become a single individual.
Grabbing take-out: Paramecium bursaria packs a lunch
Paramecium bursaria
P. bursaria, photo © www.micrographia.com.
Paramecium bursaria, a single-celled eukaryote that swims around in pond water, may
not have its own chloroplasts, but it does manage to “borrow” them in a rather
unusual way. P. bursaria swallows photosynthetic green algae, but it stores them
instead of digesting them. In fact, the normally clear paramecium can pack so many
algae into its body that it even looks green! When P. bursaria swims into the
light, the algae photosynthesize sugar, and both cells share lunch on the go. But
P. bursaria doesn’t exploit its algae. Not only does the agile paramecium chauffeur
its algae into well-lit areas, it also shares the food it finds with its algae if
they are forced to live in the dark.
Finding our roots
Many aspects of endosymbiosis still aren’t understood. For example, how did the
first endosymbiont get inside its host? Was the host trying to eat the
endosymbiont? If so, why wasn’t the endosymbiont digested? Or was the endosymbiont
trying to infect and exploit the host, the way the x-bacteria originally invaded
their amoebae hosts?
We might get closer to answering these questions if we knew what species of
bacteria came together to make the original eukaryotic cell. Biologists are in the
process of figuring this out using evidence provided by comparing the gene
sequences of different organisms. In general, the more similar two organisms’ genes
are, the more recently their two lineages split apart from one another. If two
organisms have been evolving separately for a very long time, and have a distant
common ancestor, they are likely to have evolved lots of differences between their
gene sequences. So by comparing the genetic sequences of different organisms,
biologists can piece together their family tree — who is related to whom.
More closely related organisms have more genetic similarity.
Biologists have even studied the DNA in mitochondria to figure out how they are
related to free-living bacteria. These studies have shown, first, that all
mitochondria are descended from one original mitochondrion — that is, endosymbiosis
of a mitochondrion only happened once — and second, that mitochondria are most
closely related to a bacteria called Rickettsia prowazekii. This parasitic
bacterium causes epidemic typhus and is primarily transmitted through bites of
animals such as fleas.
As DNA sequencing projects decode more genomes letter by letter, we are learning
that eukaryotic “host” cells have a lot in common, genomically speaking, with the
strangest, least-understood domain of life: the Archaea. These prokaryotes went
completely undiscovered until 1977 because they live in such extreme environments
as Antarctica, the Dead Sea, deep-sea vents, hot springs, and sewage sludge. Yet
one of them may have been the first endosymbiotic host, and our own ancestor.
Hot in Here: Archaea like it Extreme
Organisms that like extreme conditions are called “extremophiles,” and Archaea are
some of the best. Certain species of Archaea live in solutions that are more acidic
than vinegar. Others survive in environments that are as basic as ammonia. If you
stuck your arm into the homes of some extremophiles, you’d blister and burn your
skin right off! Archaea are most famous for tolerating incredible heat — many
species live at temperatures above the boiling point of water! How do they survive,
when their cells are made of membranes, proteins, and DNA, just like ours? Archaea,
and other extremophile organisms, make specialized salts and proteins. Like
antifreeze in a car radiator, these compounds keep the cell’s metabolism from
freezing up or breaking down, even when conditions outside get hostile. Archaea
demonstrate that even billions of years ago, when volcanic Earth was boiling hot,
life could have existed. In fact, if life is ever found on other planets in our
solar system, it may well resemble extremophiles like the Archaea.
How important is endosymbiosis?
Endosymbiosis explains the origin of mitochondria and chloroplasts, but could it
also explain other features of the eukaryotic cell? Maybe. Endosymbiotic origins
have been suggested for many structures, including flagella (structures like the
tail of a sperm), cilia (hair-like structures that help in locomotion), and even
the nucleus — the cell’s command center! However, scientists are still actively
debating whether or not these structures evolved through endosymbiosis. The jury is
out while more evidence is gathered.
Flagellum, cilia, nucleus
In her theory of endosymbiosis, Lynn Margulis emphasizes that during the history of
life, symbiosis has played a role not just once or twice, but over and over again.
Instead of the traditional tree of life branching out from a few common ancestors
to many descendent species, Margulis proposes that branches have separated, and
then come together again many times as individuals of different species set up
symbiotic relationships and formed new organisms. This process formed an
interconnected tree of life in which organisms have multiple ancestors, even from
different domains. As eukaryotes, our ancestors include both the bacteria that
became mitochondria, and the archaebacterium that was the host cell.
The tree of life branches and converges.
Why have endosymbiosis and symbiosis been so important to evolution? Why cooperate
at all? The answer to these questions points us to one of the basic processes of
evolution: natural selection. As Darwin observed, organisms that are fit enough to
succeed in the game of survival have a good chance of passing on their genes to the
next generation. Any survival or reproductive advantage can help a species out-
compete another species or simply avoid becoming extinct itself. It seems likely
that the first eukaryotic cells gained a slight edge over their neighbors when the
mitochondria, a rich source of energy, moved in with them. Like Kwang Jeon’s x-
bacteria and amoebae, the mitochondria and their hosts relied more and more on each
other in order to survive. Eventually, neither could succeed alone — but as a team
they produced millions of descendants, establishing a whole new domain of life.
7.8: The Endosymbiotic Theory
CCBY
7.7: Flagella and Cilia
7.E: The Eukaryotic Cell (Exercises)
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Gary Kaiser
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Learning Objectives
Briefly describe what is meant by the endosymbiotic theory.
Give some evidence supporting the theory that mitochondria and chloroplasts may
have arisen from prokaryotic organisms.
It is thought that life arose on earth around four billion years ago. The
endosymbiotic theory states that some of the organelles in today's eukaryotic cells
were once prokaryotic microbes. In this theory, the first eukaryotic cell was
probably an amoeba-like cell that got nutrients by phagocytosis and contained a
nucleus that formed when a piece of the cytoplasmic membrane pinched off around the
chromosomes. Some of these amoeba-like organisms ingested prokaryotic cells that
then survived within the organism and developed a symbiotic relationship.
Mitochondria formed when bacteria capable of aerobic respiration were ingested;
chloroplasts formed when photosynthetic bacteria were ingested. They eventually
lost their cell wall and much of their DNA because they were not of benefit within
the host cell. Mitochondria and chloroplasts cannot grow outside their host cell.
Evidence for this is based on the following:
Chloroplasts are the same size as prokaryotic cells, divide by binary fission, and,
like bacteria, have Fts proteins at their division plane. The mitochondria are the
same size as prokaryotic cells, divide by binary fission, and the mitochondria of
some protists have Fts homologs at their division plane.
Mitochondria and chloroplasts have their own DNA that is circular, not linear.
Mitochondria and chloroplasts have their own ribosomes that have 30S and 50S
subunits, not 40S and 60S.
Several more primitive eukaryotic microbes, such as Giardia and Trichomonas have a
nuclear membrane but no mitochondria.
Although evidence is less convincing, it is also possible that flagella and cilia
may have come from spirochetes.
alt
Figure 7.8.1
: One model for the origin of mitochondria and plastids. This model has an
amitochondriate eukaryote engulfing an aerobe and then a cyanobacterium. from
Kelvinsong
Example 7.8.1
Briefly describe what is meant by the endosymbiotic theory.
Give three points of evidence supporting the theory that mitochondria and
chloroplasts may have arisen from prokaryotic organisms.
Solutions
The endosymbiotic theory states that some of the organelles in eukaryotic cells
were once prokaryotic microbes.
Mitochondria and chloroplasts are the same size as prokaryotic cells and divide by
binary fission.
Mitochondria and chloroplasts have their own DNA which is circular, not linear.
Mitochondria and chloroplasts have their own ribosomes which have 30S and 50S
subunits, not 40S and 60S.
The Evolution of the Cell
The Cells That Changed the Earth
Some of the oldest cells on Earth are single-cell organisms called bacteria. Fossil
records indicate that mounds of bacteria once covered young Earth. Some began
making their own food using carbon dioxide in the atmosphere and energy they
harvested from the sun. This process (called photosynthesis) produced enough oxygen
to change Earth's atmosphere. Soon afterward, new oxygen-breathing life forms came
onto the scene. With a population of increasingly diverse bacterial life, the stage
was set for some amazing things to happen.
Bacteria
Bacteria are single-celled organisms with a circular DNA molecule and no
organelles.
The Endosymbiotic Theory
Endosymbiosis
There is compelling evidence that mitochondria and chloroplasts were once primitive
bacterial cells. This evidence is described in the endosymbiotic theory. How did
this theory get its name? Symbiosis occurs when two different species benefit from
living and working together. When one organism actually lives inside the other it's
called endosymbiosis. The endosymbiotic theory describes how a large host cell and
ingested bacteria could easily become dependent on one another for survival,
resulting in a permanent relationship. Over millions of years of evolution,
mitochondria and chloroplasts have become more specialized and today they cannot
live outside the cell.
It's Just a Theory
In everyday speech, people use the word theory to mean an opinion or speculation
not necessarily based on facts. But in the field of science, a theory is a well
established explanation based on extensive experimentation and observation.
Scientific theories are developed and verified by the scientific community and are
generally accepted as fact.
Mitochondria Have DNA
Mitochondria and chloroplasts have striking similarities to bacteria cells. They
have their own DNA, which is separate from the DNA found in the nucleus of the
cell. And both organelles use their DNA to produce many proteins and enzymes
required for their function. A double membrane surrounds both mitochondria and
chloroplasts, further evidence that each was ingested by a primitive host. The two
organelles also reproduce like bacteria, replicating their own DNA and directing
their own division.
Drawing the History of Evolution
Mitochondrial DNA (mtDNA) has a unique pattern of inheritance. It is passed down
directly from mother to child, and it accumulates changes much more slowly than
other types of DNA. Because of its unique characteristics, mtDNA has provided
important clues about evolutionary history. For example, differences in mtDNA are
examined to estimate how closely related one species is to another.
Migration Patterns
Analysis of mitochondrial DNA from people around the world has revealed many clues
about ancient human migration patterns.
Archaea: Models for Astrobiology
Archaea
Archaea survive today in extremely harsh environments, such as evaporative salt
ponds on the edge of Great Salt Lake (above) and the boiling hot springs of
Yellowstone National Park (right).
Conditions on Earth 4 billion years ago were very different than they are today.
The atmosphere lacked oxygen, and an ozone layer did not yet protect Earth from
harmful radiation. Heavy rains, lightning and volcanic activity were common. Yet
the earliest cells originated in this extreme environment. Today, a group of
single-celled organisms called archaeabacteria, or archaea, still thrive in extreme
habitats.
Astrobiologists are now using archaea to study the origins of life on Earth and
other planets. Because archaea inhabit places previously considered incompatible
with life, they may provide clues that will improve our ability to detect
extraterrestrial life. Interestingly, current research suggests archaea may be
capable of space travel by meteorite. Such an event could have seeded life on Earth
or elsewhere.
7.8: The Endosymbiotic Theory
CCBY
7.7: Flagella and Cilia
7.E: The Eukaryotic Cell (Exercises)
picture_as_pdf
Downloads
Buy Print Copy
Submit Adoption Report
Peer Review
Donate
Gary Kaiser
Community College of Baltimore Country (Cantonsville)
Learning Objectives
Briefly describe what is meant by the endosymbiotic theory.
Give some evidence supporting the theory that mitochondria and chloroplasts may
have arisen from prokaryotic organisms.
It is thought that life arose on earth around four billion years ago. The
endosymbiotic theory states that some of the organelles in today's eukaryotic cells
were once prokaryotic microbes. In this theory, the first eukaryotic cell was
probably an amoeba-like cell that got nutrients by phagocytosis and contained a
nucleus that formed when a piece of the cytoplasmic membrane pinched off around the
chromosomes. Some of these amoeba-like organisms ingested prokaryotic cells that
then survived within the organism and developed a symbiotic relationship.
Mitochondria formed when bacteria capable of aerobic respiration were ingested;
chloroplasts formed when photosynthetic bacteria were ingested. They eventually
lost their cell wall and much of their DNA because they were not of benefit within
the host cell. Mitochondria and chloroplasts cannot grow outside their host cell.
Evidence for this is based on the following:
Chloroplasts are the same size as prokaryotic cells, divide by binary fission, and,
like bacteria, have Fts proteins at their division plane. The mitochondria are the
same size as prokaryotic cells, divide by binary fission, and the mitochondria of
some protists have Fts homologs at their division plane.
Mitochondria and chloroplasts have their own DNA that is circular, not linear.
Mitochondria and chloroplasts have their own ribosomes that have 30S and 50S
subunits, not 40S and 60S.
Several more primitive eukaryotic microbes, such as Giardia and Trichomonas have a
nuclear membrane but no mitochondria.
Although evidence is less convincing, it is also possible that flagella and cilia
may have come from spirochetes.
alt
Figure 7.8.1
: One model for the origin of mitochondria and plastids. This model has an
amitochondriate eukaryote engulfing an aerobe and then a cyanobacterium. from
Kelvinsong
Example 7.8.1
Briefly describe what is meant by the endosymbiotic theory.
Give three points of evidence supporting the theory that mitochondria and
chloroplasts may have arisen from prokaryotic organisms.
Solutions
The endosymbiotic theory states that some of the organelles in eukaryotic cells
were once prokaryotic microbes.
Mitochondria and chloroplasts are the same size as prokaryotic cells and divide by
binary fission.
Mitochondria and chloroplasts have their own DNA which is circular, not linear.
Mitochondria and chloroplasts have their own ribosomes which have 30S and 50S
subunits, not 40S and 60S.
Summary
The endosymbiotic theory states that mitochondria and chlopoplasts in today's
eukaryotic cells were once separate prokaryotic microbes.
